https://microbewiki.kenyon.edu/api.php?action=feedcontributions&user=Bdtruong&feedformat=atommicrobewiki - User contributions [en]2024-03-29T13:13:25ZUser contributionsMediaWiki 1.39.6https://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=18524Staphylococcus haemolyticus2007-06-05T18:07:07Z<p>Bdtruong: /* Current Research */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4]).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4])<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunistic pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5], [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]). This is an ability that is unique among staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). The peptidoglycan structure of ''S.haemolyticus'' has been studied ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]) to find out the factors responsible to this special resistance.<br />
<br />
Like those of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,[http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues and their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, when ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]).<br />
<br />
''S.haemolyticus''’ GGI are related in function and characteristics to other relative staphylococci virulent factors, such as delta-lysin in ''S.aureus'' and SLUSH (''Staphylococcus lugdunensis'' synergistic hemolysin) in ''S.lugdunensis'', the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of ''S.haemolyticus'' genome. Recent results have shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract 12]).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find more effective strategies against the development of ''S.haemolyticus'' antibiotic-resistant strains. Studies are aiming at a promising strategy in which different types of antibiotics are used synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases.] Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species.] J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics.] J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus.] Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro.] J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning.] FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus.] J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435.] J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=18520Staphylococcus haemolyticus2007-06-05T18:06:35Z<p>Bdtruong: /* Current Research */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4]).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4])<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunistic pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5], [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]). This is an ability that is unique among staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). The peptidoglycan structure of ''S.haemolyticus'' has been studied ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]) to find out the factors responsible to this special resistance.<br />
<br />
Like those of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,[http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues and their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, when ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]).<br />
<br />
''S.haemolyticus''’ GGI are related in function and characteristics to other relative staphylococci virulent factors, such as delta-lysin in ''S.aureus'' and SLUSH (''Staphylococcus lugdunensis'' synergistic hemolysin) in ''S.lugdunensis'', the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of ''S.haemolyticus'' genome. Recent results have shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract 12]).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find more effective strategies against the development of ''S.haemolyticus'' antibiotic-resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases.] Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species.] J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics.] J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus.] Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro.] J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning.] FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus.] J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435.] J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=18517Staphylococcus haemolyticus2007-06-05T18:05:58Z<p>Bdtruong: /* Current Research */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4]).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4])<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunistic pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5], [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]). This is an ability that is unique among staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). The peptidoglycan structure of ''S.haemolyticus'' has been studied ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]) to find out the factors responsible to this special resistance.<br />
<br />
Like those of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,[http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues and their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, when ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]).<br />
<br />
''S.haemolyticus''’ GGI are related in function and characteristics to other relative staphylococci virulent factors, such as delta-lysin in ''S.aureus'' and SLUSH (''Staphylococcus lugdunensis'' synergistic hemolysin) in ''S.lugdunensis'', the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of ''S.haemolyticus'' genome. Recent results have shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract 12]).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases.] Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species.] J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics.] J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus.] Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro.] J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning.] FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus.] J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435.] J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=18514Staphylococcus haemolyticus2007-06-05T18:05:42Z<p>Bdtruong: /* Current Research */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4]).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4])<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunistic pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5], [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]). This is an ability that is unique among staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). The peptidoglycan structure of ''S.haemolyticus'' has been studied ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]) to find out the factors responsible to this special resistance.<br />
<br />
Like those of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,[http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues and their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, when ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]).<br />
<br />
''S.haemolyticus''’ GGI are related in function and characteristics to other relative staphylococci virulent factors, such as delta-lysin in ''S.aureus'' and SLUSH (''Staphylococcus lugdunensis'' synergistic hemolysin) in ''S.lugdunensis'', the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of ''S.haemolyticus'' genome. Recent results shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract 12]).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases.] Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species.] J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics.] J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus.] Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro.] J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning.] FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus.] J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435.] J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=18512Staphylococcus haemolyticus2007-06-05T18:04:55Z<p>Bdtruong: /* Current Research */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4]).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4])<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunistic pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5], [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]). This is an ability that is unique among staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). The peptidoglycan structure of ''S.haemolyticus'' has been studied ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]) to find out the factors responsible to this special resistance.<br />
<br />
Like those of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,[http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues and their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, when ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]).<br />
<br />
''S.haemolyticus''’ GGI are related in function and characteristics to other relative staphylococci virulent factors, such as delta-lysin in ''S.aureus'' and SLUSH (''Staphylococcus lugdunensis'' synergistic hemolysin) in ''S.lugdunensis'', the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of S.haemolyticus genome. Recent results shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract 12]).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases.] Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species.] J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics.] J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus.] Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro.] J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning.] FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus.] J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435.] J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=18506Staphylococcus haemolyticus2007-06-05T18:03:40Z<p>Bdtruong: /* Pathology */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4]).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4])<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunistic pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5], [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]). This is an ability that is unique among staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). The peptidoglycan structure of ''S.haemolyticus'' has been studied ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]) to find out the factors responsible to this special resistance.<br />
<br />
Like those of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,[http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues and their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, when ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]).<br />
<br />
''S.haemolyticus''’ GGI are related in function and characteristics to other relative staphylococci virulent factors, such as delta-lysin in ''S.aureus'' and SLUSH (''Staphylococcus lugdunensis'' synergistic hemolysin) in ''S.lugdunensis'', the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of S.haemolyticus genome. Recent results shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract 12]).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases.] Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species.] J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics.] J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus.] Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro.] J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning.] FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus.] J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435.] J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=18497Staphylococcus haemolyticus2007-06-05T18:01:27Z<p>Bdtruong: /* Pathology */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4]).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4])<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunistic pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5], [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]). This is an ability that is unique among staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). The peptidoglycan structure of ''S.haemolyticus'' has been studied ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]) to find out the factors responsible to this special resistance.<br />
<br />
Like those of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,[http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues and their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, when ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]).<br />
<br />
S.haemolyticus’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (Staphylococcus lugdunensis synergistic hemolysin) in S.lugdunensis, the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of S.haemolyticus genome. Recent results shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract 12]).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases.] Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species.] J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics.] J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus.] Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro.] J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning.] FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus.] J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435.] J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=18492Staphylococcus haemolyticus2007-06-05T18:00:22Z<p>Bdtruong: /* Pathology */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4]).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4])<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunistic pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5], [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]). This is an ability that is unique among staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). The peptidoglycan structure of ''S.haemolyticus'' has been studied ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]) to find out the factors responsible to this special resistance.<br />
<br />
Like those of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,[http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues and their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]).<br />
<br />
S.haemolyticus’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (Staphylococcus lugdunensis synergistic hemolysin) in S.lugdunensis, the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of S.haemolyticus genome. Recent results shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract 12]).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases.] Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species.] J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics.] J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus.] Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro.] J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning.] FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus.] J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435.] J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=18478Staphylococcus haemolyticus2007-06-05T17:58:28Z<p>Bdtruong: /* Cell Wall */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4]).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4])<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunistic pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5], [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]). This is an ability that is unique among staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). The peptidoglycan structure of ''S.haemolyticus'' has been studied ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]) to find out the factors responsible to this special resistance.<br />
<br />
Like those of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,[http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]).<br />
<br />
S.haemolyticus’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (Staphylococcus lugdunensis synergistic hemolysin) in S.lugdunensis, the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of S.haemolyticus genome. Recent results shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract 12]).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases.] Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species.] J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics.] J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus.] Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro.] J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning.] FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus.] J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435.] J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=18472Staphylococcus haemolyticus2007-06-05T17:57:31Z<p>Bdtruong: /* Genome structure */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4]).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4])<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunistic pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5], [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]). This is an ability that is unique among staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). The peptidoglycan structure of ''S.haemolyticus'' has been studied ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]) to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,[http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]).<br />
<br />
S.haemolyticus’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (Staphylococcus lugdunensis synergistic hemolysin) in S.lugdunensis, the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of S.haemolyticus genome. Recent results shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract 12]).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases.] Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species.] J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics.] J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus.] Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro.] J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning.] FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus.] J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435.] J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=18454Staphylococcus haemolyticus2007-06-05T17:54:14Z<p>Bdtruong: /* Current Research */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4]).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4])<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5], [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]). This is an ability that is unique among staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). The peptidoglycan structure of ''S.haemolyticus'' has been studied ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]) to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,[http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]).<br />
<br />
S.haemolyticus’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (Staphylococcus lugdunensis synergistic hemolysin) in S.lugdunensis, the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of S.haemolyticus genome. Recent results shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract 12]).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases.] Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species.] J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics.] J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus.] Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro.] J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning.] FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus.] J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435.] J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=18450Staphylococcus haemolyticus2007-06-05T17:53:11Z<p>Bdtruong: /* Application to Biotechnology */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4]).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4])<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5], [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]). This is an ability that is unique among staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). The peptidoglycan structure of ''S.haemolyticus'' has been studied ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]) to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,[http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]).<br />
<br />
S.haemolyticus’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (Staphylococcus lugdunensis synergistic hemolysin) in S.lugdunensis, the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract 10]).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract 11]).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu (3), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” (3).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above (3), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of S.haemolyticus genome. Recent results shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 (12).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics (13).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases.] Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species.] J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics.] J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus.] Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro.] J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning.] FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus.] J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435.] J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=18446Staphylococcus haemolyticus2007-06-05T17:52:28Z<p>Bdtruong: /* Pathology */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4]).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4])<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5], [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]). This is an ability that is unique among staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). The peptidoglycan structure of ''S.haemolyticus'' has been studied ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]) to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,[http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract 9]).<br />
<br />
S.haemolyticus’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (Staphylococcus lugdunensis synergistic hemolysin) in S.lugdunensis, the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons (10). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides (10).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell (11). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria (11).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu (3), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” (3).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above (3), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of S.haemolyticus genome. Recent results shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 (12).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics (13).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases.] Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species.] J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics.] J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus.] Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro.] J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning.] FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus.] J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435.] J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=18437Staphylococcus haemolyticus2007-06-05T17:51:01Z<p>Bdtruong: /* Ecology */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4]).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4])<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5], [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]). This is an ability that is unique among staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). The peptidoglycan structure of ''S.haemolyticus'' has been studied ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]) to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,[http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract 6]).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death (2). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) (2).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro (9). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species (9).<br />
<br />
S.haemolyticus’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (Staphylococcus lugdunensis synergistic hemolysin) in S.lugdunensis, the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons (10). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides (10).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell (11). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria (11).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu (3), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” (3).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above (3), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of S.haemolyticus genome. Recent results shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 (12).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics (13).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases.] Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species.] J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics.] J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus.] Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro.] J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning.] FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus.] J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435.] J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=18432Staphylococcus haemolyticus2007-06-05T17:50:24Z<p>Bdtruong: /* Metabolism */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4]).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4])<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5], [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]). This is an ability that is unique among staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). The peptidoglycan structure of ''S.haemolyticus'' has been studied ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]) to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures (3). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,6). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human (6).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death (2). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) (2).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro (9). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species (9).<br />
<br />
S.haemolyticus’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (Staphylococcus lugdunensis synergistic hemolysin) in S.lugdunensis, the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons (10). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides (10).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell (11). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria (11).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu (3), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” (3).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above (3), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of S.haemolyticus genome. Recent results shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 (12).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics (13).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases.] Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species.] J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics.] J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus.] Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro.] J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning.] FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus.] J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435.] J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=18426Staphylococcus haemolyticus2007-06-05T17:49:35Z<p>Bdtruong: /* Cell Wall */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4]).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4])<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5], [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract 13]). This is an ability that is unique among staphylococci ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). The peptidoglycan structure of ''S.haemolyticus'' has been studied ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]) to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract 5]).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids (3). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline (3).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci (3). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species (3). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures (3). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,6). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human (6).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death (2). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) (2).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro (9). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species (9).<br />
<br />
S.haemolyticus’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (Staphylococcus lugdunensis synergistic hemolysin) in S.lugdunensis, the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons (10). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides (10).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell (11). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria (11).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu (3), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” (3).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above (3), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of S.haemolyticus genome. Recent results shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 (12).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics (13).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases.] Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species.] J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics.] J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus.] Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro.] J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning.] FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus.] J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435.] J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=18416Staphylococcus haemolyticus2007-06-05T17:48:04Z<p>Bdtruong: /* Genome structure */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4]).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4])<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin (5, 13). This is an ability that is unique among staphylococci (5). The peptidoglycan structure of ''S.haemolyticus'' has been studied (5) to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) (5). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' (5). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' (5).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids (3). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline (3).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci (3). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species (3). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures (3). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,6). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human (6).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death (2). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) (2).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro (9). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species (9).<br />
<br />
S.haemolyticus’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (Staphylococcus lugdunensis synergistic hemolysin) in S.lugdunensis, the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons (10). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides (10).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell (11). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria (11).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu (3), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” (3).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above (3), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of S.haemolyticus genome. Recent results shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 (12).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics (13).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases.] Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species.] J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics.] J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus.] Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro.] J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning.] FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus.] J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435.] J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=18406Staphylococcus haemolyticus2007-06-05T17:46:47Z<p>Bdtruong: /* Description and significance */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4]).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract 3]). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract 4])<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp (3).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome (3). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability (3). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host (3). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) (3). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance (3). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors (3). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin (5, 13). This is an ability that is unique among staphylococci (5). The peptidoglycan structure of ''S.haemolyticus'' has been studied (5) to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) (5). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' (5). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' (5).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids (3). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline (3).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci (3). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species (3). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures (3). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,6). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human (6).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death (2). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) (2).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro (9). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species (9).<br />
<br />
S.haemolyticus’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (Staphylococcus lugdunensis synergistic hemolysin) in S.lugdunensis, the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons (10). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides (10).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell (11). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria (11).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu (3), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” (3).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above (3), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of S.haemolyticus genome. Recent results shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 (12).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics (13).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases.] Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species.] J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics.] J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus.] Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro.] J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning.] FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus.] J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435.] J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=18399Staphylococcus haemolyticus2007-06-05T17:44:41Z<p>Bdtruong: /* Description and significance */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection ([http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract 2]). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (3). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (2). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time (2). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics(2). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (4).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (3). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism (4)<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp (3).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome (3). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability (3). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host (3). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) (3). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance (3). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors (3). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin (5, 13). This is an ability that is unique among staphylococci (5). The peptidoglycan structure of ''S.haemolyticus'' has been studied (5) to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) (5). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' (5). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' (5).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids (3). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline (3).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci (3). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species (3). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures (3). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,6). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human (6).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death (2). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) (2).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro (9). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species (9).<br />
<br />
S.haemolyticus’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (Staphylococcus lugdunensis synergistic hemolysin) in S.lugdunensis, the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons (10). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides (10).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell (11). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria (11).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu (3), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” (3).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above (3), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of S.haemolyticus genome. Recent results shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 (12).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics (13).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases.] Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species.] J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics.] J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus.] Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro.] J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning.] FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus.] J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435.] J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=18349Staphylococcus haemolyticus2007-06-05T17:33:18Z<p>Bdtruong: /* References */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection (2). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (3). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (2). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time (2). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics(2). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (4).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (3). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism (4)<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp (3).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome (3). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability (3). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host (3). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) (3). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance (3). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors (3). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin (5, 13). This is an ability that is unique among staphylococci (5). The peptidoglycan structure of ''S.haemolyticus'' has been studied (5) to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) (5). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' (5). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' (5).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids (3). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline (3).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci (3). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species (3). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures (3). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,6). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human (6).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death (2). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) (2).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro (9). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species (9).<br />
<br />
S.haemolyticus’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (Staphylococcus lugdunensis synergistic hemolysin) in S.lugdunensis, the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons (10). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides (10).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell (11). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria (11).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu (3), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” (3).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above (3), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of S.haemolyticus genome. Recent results shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 (12).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics (13).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases.] Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species.] J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics.] J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=3138972&dopt=abstract The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus.] Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=7962630&dopt=abstract Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro.] J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=10518741&dopt=abstract Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning.] FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=1624418&dopt=abstract Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus.] J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17237177&dopt=abstract Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435.] J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=16801450&dopt=abstract Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus.] Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=18325Staphylococcus haemolyticus2007-06-05T17:27:12Z<p>Bdtruong: /* References */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection (2). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (3). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (2). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time (2). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics(2). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (4).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (3). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism (4)<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp (3).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome (3). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability (3). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host (3). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) (3). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance (3). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors (3). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin (5, 13). This is an ability that is unique among staphylococci (5). The peptidoglycan structure of ''S.haemolyticus'' has been studied (5) to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) (5). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' (5). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' (5).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids (3). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline (3).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci (3). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species (3). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures (3). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,6). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human (6).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death (2). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) (2).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro (9). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species (9).<br />
<br />
S.haemolyticus’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (Staphylococcus lugdunensis synergistic hemolysin) in S.lugdunensis, the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons (10). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides (10).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell (11). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria (11).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu (3), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” (3).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above (3), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of S.haemolyticus genome. Recent results shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 (12).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics (13).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases.] Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species. J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics. J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus. Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro. J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning. FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus. J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435. J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Methanococcus_jannaschii&diff=15167Methanococcus jannaschii2007-06-05T01:51:32Z<p>Bdtruong: New page: Hello</p>
<hr />
<div>Hello</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Methanococcus&diff=15166Methanococcus2007-06-05T01:51:21Z<p>Bdtruong: /* Species: */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
[[Methanogens]] overview<br />
[[Image:methanococcus_6.gif|thumb|300px|right|''Methanocaldococcus jannaschii'' courtesy of [http://biology.berkeley.edu/EML/sem.html B. Boonyaratanakornkit & D.S. Clark, Chemical Engineering, G. Vrdoljak Electron Microscope Lab, University of California Berkeley.]]]<br />
<br />
==Classification==<br />
<br />
<br />
===Higher order taxa:===<br />
<br />
Archaea; Euryarchaeota; Methanococci; Methanococcales; Methanocaldococcaceae<br />
<br />
===Species:===<br />
<br />
[[Methanococcus jannaschii|''Methanococcus jannaschii'']], ''M. maripaludis, M. voltae'' PS<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?id=2190 Taxonomy] [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=genome&cmd=Retrieve&dopt=Overview&list_uids=10928 Genome]'''<br />
|}<br />
<br />
==Description and Significance==<br />
<br />
''M. jannaschii'' was the first Archaea to have it's genome sequenced, which opened the doors for comparison between the genomes of the three domains. It was originally located from a sediment sample collected from the sea floor at the base of a "white smoker" chimney on the East Pacific Rise.<br />
<br />
''Methanocaldococcus jannaschii'' was formally in the genus ''Methanococcus'', but due to it's ability to grow in high temperatures it was re-classified. There were a few thermophilic species in ''Methanococcus'' that were reorganized, and this reorganization was supported by a low 16S rRNA sequence similarity between the thermophilics and mesophilics. The species left in ''Methanococcus'' are mesophilic and are related on the genus level by close DNA reassociation levels.<br />
<br />
No difference is noted in the G + C content of thermophilics and mesophilics of ''Methanococcus''. However, there is a difference in the proteins. The thermophilic proteins have higher residue volume, higher residue hydrophobicity, more charged amino acids, and fewer uncharged polar residues than the mesophilic proteins.<br />
<br />
==Genome Structure==<br />
<br />
''M. jannaschii'' contains a main circular chromosome consisting of 1,664,970 bp, a large extra-chromosomal element with 58,407 bp, and a small extra-chromosomal element of 16,550 bp. There were similarities found in the genome with other domains, but there were also unique sequences.<br />
<br />
The organism ''M. voltae'' was found to contain a system of gene transfer similar to general transduction except that the bacteriophage component (in terms of virus replication) is defective or absent. VTA (''voltae'' transfer agent) is responsible for the transfer and it's 4.4kb fragments of DNA are resistant to DNase.<br />
<br />
[[Image:methanococcus_1.JPG]]<br />
[[Image:methanococcus_2.JPG]]<br />
Partially purified, concentrated filtrates from ''M''. ''voltae'' PS in two different preparations. Courtesy of Eiserling et al. (1999).<br />
<br />
The majority of genes in ''M. jannaschii'' cannot be identified with Bacterial genomes or eukaryotic sequenced data. Of the similarities, ''Methanocaldococcus'' shared the anabolic genes with Bacteria (especially those involved in energy production and nitrogen fixation) and shared more of the cellular information processing and secretion systems with Eukaryotes.<br />
<br />
==Cell Structure and Metabolism==<br />
<br />
This autotrophic organism is strictly anaerobic and gets it's energy by the reduction of CO<font size="-1"><sub>2</sub></font> with H<font size="-1"><sub>2</sub></font> to generate methane. In addition to it's anabolic pathway, it also contains several scavenging molecules that most likely play a role in importing small organic compounds such as amino acids. Although ''Methanococcus'' spp. have a nifH-like gene they cannot fix N<font size="-1"><sub>2</sub></font>, with the exception of ''M. maripaludis'' that can.<br />
<br />
Structurally, the species consist of having two bundles of flagella at the same cellular pole along with no cell membrane, but a thin S-layer covering the plasma membrane. They are cocci in shape, about 1.0 microns in diameter, and are Gram stain negative.<br />
<br />
All Archaea have lipids with links between the head group and side chains, making the lipids more resistant to heat and acidity than bacterial and eukaryotic ester-linked lipids.The glycerol headgroup with two ether-linked side-chains (instead of esters) is known as ''archaeol''.:<br />
<br />
[[Image:eee.jpg|thumb|300px|center|Different archaea have different derivatives of archaeol, and ''M. jannaschii'' has been found to contain almost exclusively polar archaeal derivatives including maceocyclic archaeol (see image below), an archaeal core lipid. One can see that it has an additional link at the end of its sidechains.]]<br />
<br />
[[Image:f2f.jpg|thumb|300px|center|Images from excellent article by [http://www.chemistry.montana.edu/chem524/pdf/Julian%20lipids%20RCM%20%202004.pdf Sturt ''et al''.]]]<br />
<br />
For more information on Intact Polar Lipids (IPLs) in Archaea, including tetraethers, check the description [http://biology.kenyon.edu/Microbial_Biorealm/archaea/sulfolobus/sulfolobus.html here].<br />
<br />
==Ecology==<br />
<br />
''M. jannaschii'' can grow in habitats with pressure up to more than 200 atm and a temperature range between 48 and 94<font size="-1"><sup>o</sup></font>C, with an optimum growth temperature being 85<font size="-1"><sup>o</sup></font>C.<br />
<br />
==References==<br />
<br />
Bult et al. 1996. Complete Genome Sequence of the Methanogenic Archaeon, ''Methanococcus jannaschii''. Science 273: 1058-1072.<br />
<br />
Eiserling et al. 1999. [http://vir.sgmjournals.org/cgi/reprint/80/12/3305.pdf Bacteriophage-like particles associated with the gene transfer agent of ''Methanococcus voltae'' PS]. Journal of General Virology 80: 3305-3308.<br />
<br />
Haney et al. 1999. [http://www.pnas.org/cgi/reprint/96/7/3578.pdf Thermal adaptation analyzed by comparison of protein sequences from mesophilic and extremely thermophilic ''Methanococcus'' species.] PNAS 96: 3578-3583.<br />
<br />
[http://www.chemistry.montana.edu/chem524/pdf/Julian%20lipids%20RCM%20%202004.pdf Sturt, Helen F.; Roger E. Summons, Kristin Smith, Marcus Elvert, and Kai-Uwe Hinrichs. "Intact polar membrane lipids in prokaryotes and sediments deciphered by high-performance liquid chromatography/electrospray ionization multistage mass spectrometry—new biomarkers for biogeochemistry and microbial ecology." Rapid Commun. Mass Spectrom. 2004; 18: 617–628.]</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Methanococcus&diff=15157Methanococcus2007-06-05T01:50:16Z<p>Bdtruong: /* Species: */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
[[Methanogens]] overview<br />
[[Image:methanococcus_6.gif|thumb|300px|right|''Methanocaldococcus jannaschii'' courtesy of [http://biology.berkeley.edu/EML/sem.html B. Boonyaratanakornkit & D.S. Clark, Chemical Engineering, G. Vrdoljak Electron Microscope Lab, University of California Berkeley.]]]<br />
<br />
==Classification==<br />
<br />
<br />
===Higher order taxa:===<br />
<br />
Archaea; Euryarchaeota; Methanococci; Methanococcales; Methanocaldococcaceae<br />
<br />
===Species:===<br />
<br />
[[Methanocaldococcus jannaschii|''Methanococcus jannaschii'']], ''M. maripaludis, M. voltae'' PS<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?id=2190 Taxonomy] [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=genome&cmd=Retrieve&dopt=Overview&list_uids=10928 Genome]'''<br />
|}<br />
<br />
==Description and Significance==<br />
<br />
''M. jannaschii'' was the first Archaea to have it's genome sequenced, which opened the doors for comparison between the genomes of the three domains. It was originally located from a sediment sample collected from the sea floor at the base of a "white smoker" chimney on the East Pacific Rise.<br />
<br />
''Methanocaldococcus jannaschii'' was formally in the genus ''Methanococcus'', but due to it's ability to grow in high temperatures it was re-classified. There were a few thermophilic species in ''Methanococcus'' that were reorganized, and this reorganization was supported by a low 16S rRNA sequence similarity between the thermophilics and mesophilics. The species left in ''Methanococcus'' are mesophilic and are related on the genus level by close DNA reassociation levels.<br />
<br />
No difference is noted in the G + C content of thermophilics and mesophilics of ''Methanococcus''. However, there is a difference in the proteins. The thermophilic proteins have higher residue volume, higher residue hydrophobicity, more charged amino acids, and fewer uncharged polar residues than the mesophilic proteins.<br />
<br />
==Genome Structure==<br />
<br />
''M. jannaschii'' contains a main circular chromosome consisting of 1,664,970 bp, a large extra-chromosomal element with 58,407 bp, and a small extra-chromosomal element of 16,550 bp. There were similarities found in the genome with other domains, but there were also unique sequences.<br />
<br />
The organism ''M. voltae'' was found to contain a system of gene transfer similar to general transduction except that the bacteriophage component (in terms of virus replication) is defective or absent. VTA (''voltae'' transfer agent) is responsible for the transfer and it's 4.4kb fragments of DNA are resistant to DNase.<br />
<br />
[[Image:methanococcus_1.JPG]]<br />
[[Image:methanococcus_2.JPG]]<br />
Partially purified, concentrated filtrates from ''M''. ''voltae'' PS in two different preparations. Courtesy of Eiserling et al. (1999).<br />
<br />
The majority of genes in ''M. jannaschii'' cannot be identified with Bacterial genomes or eukaryotic sequenced data. Of the similarities, ''Methanocaldococcus'' shared the anabolic genes with Bacteria (especially those involved in energy production and nitrogen fixation) and shared more of the cellular information processing and secretion systems with Eukaryotes.<br />
<br />
==Cell Structure and Metabolism==<br />
<br />
This autotrophic organism is strictly anaerobic and gets it's energy by the reduction of CO<font size="-1"><sub>2</sub></font> with H<font size="-1"><sub>2</sub></font> to generate methane. In addition to it's anabolic pathway, it also contains several scavenging molecules that most likely play a role in importing small organic compounds such as amino acids. Although ''Methanococcus'' spp. have a nifH-like gene they cannot fix N<font size="-1"><sub>2</sub></font>, with the exception of ''M. maripaludis'' that can.<br />
<br />
Structurally, the species consist of having two bundles of flagella at the same cellular pole along with no cell membrane, but a thin S-layer covering the plasma membrane. They are cocci in shape, about 1.0 microns in diameter, and are Gram stain negative.<br />
<br />
All Archaea have lipids with links between the head group and side chains, making the lipids more resistant to heat and acidity than bacterial and eukaryotic ester-linked lipids.The glycerol headgroup with two ether-linked side-chains (instead of esters) is known as ''archaeol''.:<br />
<br />
[[Image:eee.jpg|thumb|300px|center|Different archaea have different derivatives of archaeol, and ''M. jannaschii'' has been found to contain almost exclusively polar archaeal derivatives including maceocyclic archaeol (see image below), an archaeal core lipid. One can see that it has an additional link at the end of its sidechains.]]<br />
<br />
[[Image:f2f.jpg|thumb|300px|center|Images from excellent article by [http://www.chemistry.montana.edu/chem524/pdf/Julian%20lipids%20RCM%20%202004.pdf Sturt ''et al''.]]]<br />
<br />
For more information on Intact Polar Lipids (IPLs) in Archaea, including tetraethers, check the description [http://biology.kenyon.edu/Microbial_Biorealm/archaea/sulfolobus/sulfolobus.html here].<br />
<br />
==Ecology==<br />
<br />
''M. jannaschii'' can grow in habitats with pressure up to more than 200 atm and a temperature range between 48 and 94<font size="-1"><sup>o</sup></font>C, with an optimum growth temperature being 85<font size="-1"><sup>o</sup></font>C.<br />
<br />
==References==<br />
<br />
Bult et al. 1996. Complete Genome Sequence of the Methanogenic Archaeon, ''Methanococcus jannaschii''. Science 273: 1058-1072.<br />
<br />
Eiserling et al. 1999. [http://vir.sgmjournals.org/cgi/reprint/80/12/3305.pdf Bacteriophage-like particles associated with the gene transfer agent of ''Methanococcus voltae'' PS]. Journal of General Virology 80: 3305-3308.<br />
<br />
Haney et al. 1999. [http://www.pnas.org/cgi/reprint/96/7/3578.pdf Thermal adaptation analyzed by comparison of protein sequences from mesophilic and extremely thermophilic ''Methanococcus'' species.] PNAS 96: 3578-3583.<br />
<br />
[http://www.chemistry.montana.edu/chem524/pdf/Julian%20lipids%20RCM%20%202004.pdf Sturt, Helen F.; Roger E. Summons, Kristin Smith, Marcus Elvert, and Kai-Uwe Hinrichs. "Intact polar membrane lipids in prokaryotes and sediments deciphered by high-performance liquid chromatography/electrospray ionization multistage mass spectrometry—new biomarkers for biogeochemistry and microbial ecology." Rapid Commun. Mass Spectrom. 2004; 18: 617–628.]</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Methanococcus&diff=15153Methanococcus2007-06-05T01:49:12Z<p>Bdtruong: /* Species: */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
[[Methanogens]] overview<br />
[[Image:methanococcus_6.gif|thumb|300px|right|''Methanocaldococcus jannaschii'' courtesy of [http://biology.berkeley.edu/EML/sem.html B. Boonyaratanakornkit & D.S. Clark, Chemical Engineering, G. Vrdoljak Electron Microscope Lab, University of California Berkeley.]]]<br />
<br />
==Classification==<br />
<br />
<br />
===Higher order taxa:===<br />
<br />
Archaea; Euryarchaeota; Methanococci; Methanococcales; Methanocaldococcaceae<br />
<br />
===Species:===<br />
<br />
[[Methanocaldococcus jannaschii]] (''Methanococcus jannaschii''), ''M. maripaludis, M. voltae'' PS<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?id=2190 Taxonomy] [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=genome&cmd=Retrieve&dopt=Overview&list_uids=10928 Genome]'''<br />
|}<br />
<br />
==Description and Significance==<br />
<br />
''M. jannaschii'' was the first Archaea to have it's genome sequenced, which opened the doors for comparison between the genomes of the three domains. It was originally located from a sediment sample collected from the sea floor at the base of a "white smoker" chimney on the East Pacific Rise.<br />
<br />
''Methanocaldococcus jannaschii'' was formally in the genus ''Methanococcus'', but due to it's ability to grow in high temperatures it was re-classified. There were a few thermophilic species in ''Methanococcus'' that were reorganized, and this reorganization was supported by a low 16S rRNA sequence similarity between the thermophilics and mesophilics. The species left in ''Methanococcus'' are mesophilic and are related on the genus level by close DNA reassociation levels.<br />
<br />
No difference is noted in the G + C content of thermophilics and mesophilics of ''Methanococcus''. However, there is a difference in the proteins. The thermophilic proteins have higher residue volume, higher residue hydrophobicity, more charged amino acids, and fewer uncharged polar residues than the mesophilic proteins.<br />
<br />
==Genome Structure==<br />
<br />
''M. jannaschii'' contains a main circular chromosome consisting of 1,664,970 bp, a large extra-chromosomal element with 58,407 bp, and a small extra-chromosomal element of 16,550 bp. There were similarities found in the genome with other domains, but there were also unique sequences.<br />
<br />
The organism ''M. voltae'' was found to contain a system of gene transfer similar to general transduction except that the bacteriophage component (in terms of virus replication) is defective or absent. VTA (''voltae'' transfer agent) is responsible for the transfer and it's 4.4kb fragments of DNA are resistant to DNase.<br />
<br />
[[Image:methanococcus_1.JPG]]<br />
[[Image:methanococcus_2.JPG]]<br />
Partially purified, concentrated filtrates from ''M''. ''voltae'' PS in two different preparations. Courtesy of Eiserling et al. (1999).<br />
<br />
The majority of genes in ''M. jannaschii'' cannot be identified with Bacterial genomes or eukaryotic sequenced data. Of the similarities, ''Methanocaldococcus'' shared the anabolic genes with Bacteria (especially those involved in energy production and nitrogen fixation) and shared more of the cellular information processing and secretion systems with Eukaryotes.<br />
<br />
==Cell Structure and Metabolism==<br />
<br />
This autotrophic organism is strictly anaerobic and gets it's energy by the reduction of CO<font size="-1"><sub>2</sub></font> with H<font size="-1"><sub>2</sub></font> to generate methane. In addition to it's anabolic pathway, it also contains several scavenging molecules that most likely play a role in importing small organic compounds such as amino acids. Although ''Methanococcus'' spp. have a nifH-like gene they cannot fix N<font size="-1"><sub>2</sub></font>, with the exception of ''M. maripaludis'' that can.<br />
<br />
Structurally, the species consist of having two bundles of flagella at the same cellular pole along with no cell membrane, but a thin S-layer covering the plasma membrane. They are cocci in shape, about 1.0 microns in diameter, and are Gram stain negative.<br />
<br />
All Archaea have lipids with links between the head group and side chains, making the lipids more resistant to heat and acidity than bacterial and eukaryotic ester-linked lipids.The glycerol headgroup with two ether-linked side-chains (instead of esters) is known as ''archaeol''.:<br />
<br />
[[Image:eee.jpg|thumb|300px|center|Different archaea have different derivatives of archaeol, and ''M. jannaschii'' has been found to contain almost exclusively polar archaeal derivatives including maceocyclic archaeol (see image below), an archaeal core lipid. One can see that it has an additional link at the end of its sidechains.]]<br />
<br />
[[Image:f2f.jpg|thumb|300px|center|Images from excellent article by [http://www.chemistry.montana.edu/chem524/pdf/Julian%20lipids%20RCM%20%202004.pdf Sturt ''et al''.]]]<br />
<br />
For more information on Intact Polar Lipids (IPLs) in Archaea, including tetraethers, check the description [http://biology.kenyon.edu/Microbial_Biorealm/archaea/sulfolobus/sulfolobus.html here].<br />
<br />
==Ecology==<br />
<br />
''M. jannaschii'' can grow in habitats with pressure up to more than 200 atm and a temperature range between 48 and 94<font size="-1"><sup>o</sup></font>C, with an optimum growth temperature being 85<font size="-1"><sup>o</sup></font>C.<br />
<br />
==References==<br />
<br />
Bult et al. 1996. Complete Genome Sequence of the Methanogenic Archaeon, ''Methanococcus jannaschii''. Science 273: 1058-1072.<br />
<br />
Eiserling et al. 1999. [http://vir.sgmjournals.org/cgi/reprint/80/12/3305.pdf Bacteriophage-like particles associated with the gene transfer agent of ''Methanococcus voltae'' PS]. Journal of General Virology 80: 3305-3308.<br />
<br />
Haney et al. 1999. [http://www.pnas.org/cgi/reprint/96/7/3578.pdf Thermal adaptation analyzed by comparison of protein sequences from mesophilic and extremely thermophilic ''Methanococcus'' species.] PNAS 96: 3578-3583.<br />
<br />
[http://www.chemistry.montana.edu/chem524/pdf/Julian%20lipids%20RCM%20%202004.pdf Sturt, Helen F.; Roger E. Summons, Kristin Smith, Marcus Elvert, and Kai-Uwe Hinrichs. "Intact polar membrane lipids in prokaryotes and sediments deciphered by high-performance liquid chromatography/electrospray ionization multistage mass spectrometry—new biomarkers for biogeochemistry and microbial ecology." Rapid Commun. Mass Spectrom. 2004; 18: 617–628.]</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Methanococcus&diff=15147Methanococcus2007-06-05T01:48:26Z<p>Bdtruong: /* Species: */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
[[Methanogens]] overview<br />
[[Image:methanococcus_6.gif|thumb|300px|right|''Methanocaldococcus jannaschii'' courtesy of [http://biology.berkeley.edu/EML/sem.html B. Boonyaratanakornkit & D.S. Clark, Chemical Engineering, G. Vrdoljak Electron Microscope Lab, University of California Berkeley.]]]<br />
<br />
==Classification==<br />
<br />
<br />
===Higher order taxa:===<br />
<br />
Archaea; Euryarchaeota; Methanococci; Methanococcales; Methanocaldococcaceae<br />
<br />
===Species:===<br />
<br />
[Methanocaldococcus jannaschii] (''Methanococcus jannaschii''), ''M. maripaludis, M. voltae'' PS<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?id=2190 Taxonomy] [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=genome&cmd=Retrieve&dopt=Overview&list_uids=10928 Genome]'''<br />
|}<br />
<br />
==Description and Significance==<br />
<br />
''M. jannaschii'' was the first Archaea to have it's genome sequenced, which opened the doors for comparison between the genomes of the three domains. It was originally located from a sediment sample collected from the sea floor at the base of a "white smoker" chimney on the East Pacific Rise.<br />
<br />
''Methanocaldococcus jannaschii'' was formally in the genus ''Methanococcus'', but due to it's ability to grow in high temperatures it was re-classified. There were a few thermophilic species in ''Methanococcus'' that were reorganized, and this reorganization was supported by a low 16S rRNA sequence similarity between the thermophilics and mesophilics. The species left in ''Methanococcus'' are mesophilic and are related on the genus level by close DNA reassociation levels.<br />
<br />
No difference is noted in the G + C content of thermophilics and mesophilics of ''Methanococcus''. However, there is a difference in the proteins. The thermophilic proteins have higher residue volume, higher residue hydrophobicity, more charged amino acids, and fewer uncharged polar residues than the mesophilic proteins.<br />
<br />
==Genome Structure==<br />
<br />
''M. jannaschii'' contains a main circular chromosome consisting of 1,664,970 bp, a large extra-chromosomal element with 58,407 bp, and a small extra-chromosomal element of 16,550 bp. There were similarities found in the genome with other domains, but there were also unique sequences.<br />
<br />
The organism ''M. voltae'' was found to contain a system of gene transfer similar to general transduction except that the bacteriophage component (in terms of virus replication) is defective or absent. VTA (''voltae'' transfer agent) is responsible for the transfer and it's 4.4kb fragments of DNA are resistant to DNase.<br />
<br />
[[Image:methanococcus_1.JPG]]<br />
[[Image:methanococcus_2.JPG]]<br />
Partially purified, concentrated filtrates from ''M''. ''voltae'' PS in two different preparations. Courtesy of Eiserling et al. (1999).<br />
<br />
The majority of genes in ''M. jannaschii'' cannot be identified with Bacterial genomes or eukaryotic sequenced data. Of the similarities, ''Methanocaldococcus'' shared the anabolic genes with Bacteria (especially those involved in energy production and nitrogen fixation) and shared more of the cellular information processing and secretion systems with Eukaryotes.<br />
<br />
==Cell Structure and Metabolism==<br />
<br />
This autotrophic organism is strictly anaerobic and gets it's energy by the reduction of CO<font size="-1"><sub>2</sub></font> with H<font size="-1"><sub>2</sub></font> to generate methane. In addition to it's anabolic pathway, it also contains several scavenging molecules that most likely play a role in importing small organic compounds such as amino acids. Although ''Methanococcus'' spp. have a nifH-like gene they cannot fix N<font size="-1"><sub>2</sub></font>, with the exception of ''M. maripaludis'' that can.<br />
<br />
Structurally, the species consist of having two bundles of flagella at the same cellular pole along with no cell membrane, but a thin S-layer covering the plasma membrane. They are cocci in shape, about 1.0 microns in diameter, and are Gram stain negative.<br />
<br />
All Archaea have lipids with links between the head group and side chains, making the lipids more resistant to heat and acidity than bacterial and eukaryotic ester-linked lipids.The glycerol headgroup with two ether-linked side-chains (instead of esters) is known as ''archaeol''.:<br />
<br />
[[Image:eee.jpg|thumb|300px|center|Different archaea have different derivatives of archaeol, and ''M. jannaschii'' has been found to contain almost exclusively polar archaeal derivatives including maceocyclic archaeol (see image below), an archaeal core lipid. One can see that it has an additional link at the end of its sidechains.]]<br />
<br />
[[Image:f2f.jpg|thumb|300px|center|Images from excellent article by [http://www.chemistry.montana.edu/chem524/pdf/Julian%20lipids%20RCM%20%202004.pdf Sturt ''et al''.]]]<br />
<br />
For more information on Intact Polar Lipids (IPLs) in Archaea, including tetraethers, check the description [http://biology.kenyon.edu/Microbial_Biorealm/archaea/sulfolobus/sulfolobus.html here].<br />
<br />
==Ecology==<br />
<br />
''M. jannaschii'' can grow in habitats with pressure up to more than 200 atm and a temperature range between 48 and 94<font size="-1"><sup>o</sup></font>C, with an optimum growth temperature being 85<font size="-1"><sup>o</sup></font>C.<br />
<br />
==References==<br />
<br />
Bult et al. 1996. Complete Genome Sequence of the Methanogenic Archaeon, ''Methanococcus jannaschii''. Science 273: 1058-1072.<br />
<br />
Eiserling et al. 1999. [http://vir.sgmjournals.org/cgi/reprint/80/12/3305.pdf Bacteriophage-like particles associated with the gene transfer agent of ''Methanococcus voltae'' PS]. Journal of General Virology 80: 3305-3308.<br />
<br />
Haney et al. 1999. [http://www.pnas.org/cgi/reprint/96/7/3578.pdf Thermal adaptation analyzed by comparison of protein sequences from mesophilic and extremely thermophilic ''Methanococcus'' species.] PNAS 96: 3578-3583.<br />
<br />
[http://www.chemistry.montana.edu/chem524/pdf/Julian%20lipids%20RCM%20%202004.pdf Sturt, Helen F.; Roger E. Summons, Kristin Smith, Marcus Elvert, and Kai-Uwe Hinrichs. "Intact polar membrane lipids in prokaryotes and sediments deciphered by high-performance liquid chromatography/electrospray ionization multistage mass spectrometry—new biomarkers for biogeochemistry and microbial ecology." Rapid Commun. Mass Spectrom. 2004; 18: 617–628.]</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=15033Staphylococcus haemolyticus2007-06-05T01:26:49Z<p>Bdtruong: /* References */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection (2). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (3). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (2). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time (2). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics(2). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (4).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (3). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism (4)<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp (3).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome (3). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability (3). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host (3). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) (3). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance (3). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors (3). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin (5, 13). This is an ability that is unique among staphylococci (5). The peptidoglycan structure of ''S.haemolyticus'' has been studied (5) to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) (5). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' (5). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' (5).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids (3). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline (3).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci (3). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species (3). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures (3). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,6). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human (6).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death (2). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) (2).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro (9). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species (9).<br />
<br />
S.haemolyticus’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (Staphylococcus lugdunensis synergistic hemolysin) in S.lugdunensis, the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons (10). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides (10).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell (11). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria (11).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu (3), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” (3).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above (3), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of S.haemolyticus genome. Recent results shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 (12).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics (13).<br />
<br />
==References==<br />
1. Tristan, A., Lina, G., Etienne, J. & Vandenesch, F. in (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 572-586 (ASM Press, Washington, D.C, 2006).<br />
<br />
2. Falcone, M. et al. Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases. Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species. J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics. J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
6. Watson, D. C., Yaguchi, M., Bisaillon, J. G., Beaudet, R. & Morosoli, R. The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus. Biochem. J. 252, 87-93 (1988).<br />
<br />
7. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
8. Novick, R. in Gram-positive Pathogens (eds Fischetti, V., Novick, R., Ferretti, J., Portnoy, D. & Rood, J.) 496-510 (ASM Press, Washington, D.C, 2006).<br />
<br />
9. Molnar, C., Hevessy, Z., Rozgonyi, F. & Gemmell, C. G. Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro. J. Clin. Pathol. 47, 743-748 (1994).<br />
<br />
10. Oh, B., Kim, H., Lee, J., Kang, S. & Oh, T. Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning. FEMS Microbiol. Lett. 179, 385-392 (1999).<br />
<br />
11. Hansson, M. et al. Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus. J. Bacteriol. 174, 4239-4245 (1992).<br />
<br />
12. Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F. & Hiramatsu, K. Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435. J. Bacteriol. 189, 2921-2925 (2007).<br />
<br />
13. Vignaroli, C., Biavasco, F. & Varaldo, P. E. Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 50, 2577-2582 (2006).</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=15028Staphylococcus haemolyticus2007-06-05T01:25:33Z<p>Bdtruong: /* Current Research */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection (2). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (3). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (2). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time (2). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics(2). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (4).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (3). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism (4)<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp (3).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome (3). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability (3). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host (3). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) (3). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance (3). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors (3). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin (5, 13). This is an ability that is unique among staphylococci (5). The peptidoglycan structure of ''S.haemolyticus'' has been studied (5) to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) (5). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' (5). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' (5).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids (3). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline (3).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci (3). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species (3). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures (3). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,6). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human (6).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death (2). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) (2).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro (9). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species (9).<br />
<br />
S.haemolyticus’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (Staphylococcus lugdunensis synergistic hemolysin) in S.lugdunensis, the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons (10). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides (10).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell (11). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria (11).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Keiichi Hiramatsu (3), is a very important and significant step in tackling this problem. The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” (3).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above (3), Dr. Hiramatsu’s group continues to examine the other types of genetic rearrangement that are also responsible for the frequent structural alteration of S.haemolyticus genome. Recent results shed light on a new genetic shuffling mechanism of ''S.haemolyticus'', in which “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in the clinical strain JCSC1435 (12).<br />
<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, or the combination of vancomycin and beta-lactams antitbiotics (13).<br />
<br />
==References==<br />
Billot-Klein, D., Gutmann, L., Bryant, D., Bell, D., Van Heijenoort, J., Grewal, J., and Shlaes, D.M. (1996). Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics. J. Bacteriol. 15, 4696-4703.<br />
<br />
Brooks, G., Butel, J., and Morse, S. (2001). The Staphylococci. In Medical Microbiology, (New York: McGraw-Hill) pp. 197-202.<br />
<br />
Falcone, M., Campanile, F., Giannella, M., Borbone, S., Stefani, S., and Venditti, M. (2007). Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases. Diagn. Microbiol. Infect. Dis. 3, 325-331.<br />
<br />
Froggatt, J.W., Johnston, J.L., Galetto, D.W., and Archer, G.L. (1989). Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 4, 460-466.<br />
Hansson, M., Stahl, S., Nguyen, T.N., Bachi, T., Robert, A., Binz, H., Sjolander, A., and Uhlen, M. (1992). Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus. J. Bacteriol. 13, 4239-4245.<br />
<br />
Molnar, C., Hevessy, Z., Rozgonyi, F., and Gemmell, C.G. (1994). Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro. J. Clin. Pathol. 8, 743-748.<br />
<br />
Novick, R. (2006). Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation. In Gram-positive Pathogens, V. Fischetti, R. Novick, J. Ferretti, D. Portnoy and J. Rood eds., (Washington, D.C: ASM Press) pp. 496-510.<br />
<br />
Oh, B., Kim, H., Lee, J., Kang, S., and Oh, T. (1999). Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning. FEMS Microbiol. Lett. 2, 385-392.<br />
Takeuchi, F., Watanabe, S., Baba, T., Yuzawa, H., Ito, T., Morimoto, Y., Kuroda, M., Cui, L., Takahashi, M., Ankai, A. et al. (2005). Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species. J. Bacteriol. 21, 7292-7308.<br />
<br />
Tristan, A., Lina, G., Etienne, J., and Vandenesch, F. (2006). Biology and Pathogenicity of Staphylococci other than Staphylococcus aureus and Staphylococcus epidermis. V. Fischetti, R. Novick, J. Ferretti, D. Portnoy and J. Rood eds., (Washington, D.C: ASM Press) pp. 572-586.<br />
<br />
Vignaroli, C., Biavasco, F., and Varaldo, P.E. (2006). Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 7, 2577-2582.<br />
<br />
Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F., and Hiramatsu, K. (2007). Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435. J. Bacteriol. 7, 2921-2925.<br />
<br />
Watson, D.C., Yaguchi, M., Bisaillon, J.G., Beaudet, R., and Morosoli, R. (1988). The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus. Biochem. J. 1, 87-93.</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=15015Staphylococcus haemolyticus2007-06-05T01:19:36Z<p>Bdtruong: /* Application to Biotechnology */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection (2). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (3). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (2). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time (2). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics(2). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (4).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (3). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism (4)<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp (3).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome (3). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability (3). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host (3). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) (3). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance (3). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors (3). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin (5, 13). This is an ability that is unique among staphylococci (5). The peptidoglycan structure of ''S.haemolyticus'' has been studied (5) to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) (5). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' (5). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' (5).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids (3). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline (3).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci (3). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species (3). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures (3). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,6). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human (6).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death (2). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) (2).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro (9). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species (9).<br />
<br />
S.haemolyticus’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (Staphylococcus lugdunensis synergistic hemolysin) in S.lugdunensis, the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons (10). Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides (10).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell (11). It was the first time such system could be constructed in a Gram-positive species. This technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria (11).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Dr. Keiichi Hiramatsu, is a very important and significant step in tackling this problem(Takeuchi,Watanabe, et al, 2005). The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” (Takeuchi,Watanabe, et al, 2005).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above (Takeuchi,Watanabe, et al, 2005), Dr. Hiramatsu’s group is examining the other types of genetic rearrangement that are also responsible for the frequent structural alteration of ''S.haemolyticus'' genome. Recent results indicated that “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in clinical strain JCSC1435 of ''S.haemolyticus'' (Watanabe,Ito, et al, 2007).<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, and the combination of vancomycin and beta-lactams antitbiotics (Vignaroli,Biavasco and Varaldo, 2006).<br />
<br />
==References==<br />
Billot-Klein, D., Gutmann, L., Bryant, D., Bell, D., Van Heijenoort, J., Grewal, J., and Shlaes, D.M. (1996). Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics. J. Bacteriol. 15, 4696-4703.<br />
<br />
Brooks, G., Butel, J., and Morse, S. (2001). The Staphylococci. In Medical Microbiology, (New York: McGraw-Hill) pp. 197-202.<br />
<br />
Falcone, M., Campanile, F., Giannella, M., Borbone, S., Stefani, S., and Venditti, M. (2007). Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases. Diagn. Microbiol. Infect. Dis. 3, 325-331.<br />
<br />
Froggatt, J.W., Johnston, J.L., Galetto, D.W., and Archer, G.L. (1989). Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 4, 460-466.<br />
Hansson, M., Stahl, S., Nguyen, T.N., Bachi, T., Robert, A., Binz, H., Sjolander, A., and Uhlen, M. (1992). Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus. J. Bacteriol. 13, 4239-4245.<br />
<br />
Molnar, C., Hevessy, Z., Rozgonyi, F., and Gemmell, C.G. (1994). Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro. J. Clin. Pathol. 8, 743-748.<br />
<br />
Novick, R. (2006). Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation. In Gram-positive Pathogens, V. Fischetti, R. Novick, J. Ferretti, D. Portnoy and J. Rood eds., (Washington, D.C: ASM Press) pp. 496-510.<br />
<br />
Oh, B., Kim, H., Lee, J., Kang, S., and Oh, T. (1999). Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning. FEMS Microbiol. Lett. 2, 385-392.<br />
Takeuchi, F., Watanabe, S., Baba, T., Yuzawa, H., Ito, T., Morimoto, Y., Kuroda, M., Cui, L., Takahashi, M., Ankai, A. et al. (2005). Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species. J. Bacteriol. 21, 7292-7308.<br />
<br />
Tristan, A., Lina, G., Etienne, J., and Vandenesch, F. (2006). Biology and Pathogenicity of Staphylococci other than Staphylococcus aureus and Staphylococcus epidermis. V. Fischetti, R. Novick, J. Ferretti, D. Portnoy and J. Rood eds., (Washington, D.C: ASM Press) pp. 572-586.<br />
<br />
Vignaroli, C., Biavasco, F., and Varaldo, P.E. (2006). Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 7, 2577-2582.<br />
<br />
Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F., and Hiramatsu, K. (2007). Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435. J. Bacteriol. 7, 2921-2925.<br />
<br />
Watson, D.C., Yaguchi, M., Bisaillon, J.G., Beaudet, R., and Morosoli, R. (1988). The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus. Biochem. J. 1, 87-93.</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=15010Staphylococcus haemolyticus2007-06-05T01:18:02Z<p>Bdtruong: /* Pathology */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection (2). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (3). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (2). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time (2). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics(2). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (4).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (3). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism (4)<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp (3).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome (3). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability (3). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host (3). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) (3). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance (3). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors (3). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin (5, 13). This is an ability that is unique among staphylococci (5). The peptidoglycan structure of ''S.haemolyticus'' has been studied (5) to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) (5). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' (5). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' (5).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids (3). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline (3).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci (3). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species (3). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures (3). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,6). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human (6).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (7). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (7). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci (7). ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints (1). In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death (2). Common clinical symptoms of a ''S.haemolyticus'' infection are fever and an increase in white blood cell population (leukocytosis) (2).<br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (8). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but have been recently discovered in the other non-aureus coagulase-negative staphylococci as well (1). In one studies published in 1994, for example, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro (9). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species (9).<br />
<br />
S.haemolyticus’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (Staphylococcus lugdunensis synergistic hemolysin) in S.lugdunensis, the latter of which shows significant similarities in structure with GGI (1). These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons. Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides (Oh,Kim, et al, 1999).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell (Hansson,Stahl, et al, 1992). It was the first time such system could be constructed in a Gram-positive species. The technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria (Hansson,Stahl, et al, 1992).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Dr. Keiichi Hiramatsu, is a very important and significant step in tackling this problem(Takeuchi,Watanabe, et al, 2005). The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” (Takeuchi,Watanabe, et al, 2005).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above (Takeuchi,Watanabe, et al, 2005), Dr. Hiramatsu’s group is examining the other types of genetic rearrangement that are also responsible for the frequent structural alteration of ''S.haemolyticus'' genome. Recent results indicated that “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in clinical strain JCSC1435 of ''S.haemolyticus'' (Watanabe,Ito, et al, 2007).<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, and the combination of vancomycin and beta-lactams antitbiotics (Vignaroli,Biavasco and Varaldo, 2006).<br />
<br />
==References==<br />
Billot-Klein, D., Gutmann, L., Bryant, D., Bell, D., Van Heijenoort, J., Grewal, J., and Shlaes, D.M. (1996). Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics. J. Bacteriol. 15, 4696-4703.<br />
<br />
Brooks, G., Butel, J., and Morse, S. (2001). The Staphylococci. In Medical Microbiology, (New York: McGraw-Hill) pp. 197-202.<br />
<br />
Falcone, M., Campanile, F., Giannella, M., Borbone, S., Stefani, S., and Venditti, M. (2007). Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases. Diagn. Microbiol. Infect. Dis. 3, 325-331.<br />
<br />
Froggatt, J.W., Johnston, J.L., Galetto, D.W., and Archer, G.L. (1989). Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 4, 460-466.<br />
Hansson, M., Stahl, S., Nguyen, T.N., Bachi, T., Robert, A., Binz, H., Sjolander, A., and Uhlen, M. (1992). Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus. J. Bacteriol. 13, 4239-4245.<br />
<br />
Molnar, C., Hevessy, Z., Rozgonyi, F., and Gemmell, C.G. (1994). Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro. J. Clin. Pathol. 8, 743-748.<br />
<br />
Novick, R. (2006). Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation. In Gram-positive Pathogens, V. Fischetti, R. Novick, J. Ferretti, D. Portnoy and J. Rood eds., (Washington, D.C: ASM Press) pp. 496-510.<br />
<br />
Oh, B., Kim, H., Lee, J., Kang, S., and Oh, T. (1999). Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning. FEMS Microbiol. Lett. 2, 385-392.<br />
Takeuchi, F., Watanabe, S., Baba, T., Yuzawa, H., Ito, T., Morimoto, Y., Kuroda, M., Cui, L., Takahashi, M., Ankai, A. et al. (2005). Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species. J. Bacteriol. 21, 7292-7308.<br />
<br />
Tristan, A., Lina, G., Etienne, J., and Vandenesch, F. (2006). Biology and Pathogenicity of Staphylococci other than Staphylococcus aureus and Staphylococcus epidermis. V. Fischetti, R. Novick, J. Ferretti, D. Portnoy and J. Rood eds., (Washington, D.C: ASM Press) pp. 572-586.<br />
<br />
Vignaroli, C., Biavasco, F., and Varaldo, P.E. (2006). Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 7, 2577-2582.<br />
<br />
Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F., and Hiramatsu, K. (2007). Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435. J. Bacteriol. 7, 2921-2925.<br />
<br />
Watson, D.C., Yaguchi, M., Bisaillon, J.G., Beaudet, R., and Morosoli, R. (1988). The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus. Biochem. J. 1, 87-93.</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=14992Staphylococcus haemolyticus2007-06-05T01:12:43Z<p>Bdtruong: /* Ecology */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection (2). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (3). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (2). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time (2). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics(2). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (4).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (3). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism (4)<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp (3).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome (3). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability (3). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host (3). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) (3). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance (3). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors (3). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin (5, 13). This is an ability that is unique among staphylococci (5). The peptidoglycan structure of ''S.haemolyticus'' has been studied (5) to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) (5). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' (5). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' (5).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids (3). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline (3).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci (3). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species (3). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (1). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (1). ''S.haemolyticus'' survive successfully on the drier regions of the body (1), while it can also be found frequently in human blood cultures (3). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI (1). The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (1,6). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human (6).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (Brooks,Butel and Morse, 2001). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (Brooks,Butel and Morse, 2001). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci. (Brooks,Butel and Morse, 2001) ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints. (Tristan,Lina, et al, 2006) In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death. (Falcone,Campanile, et al, 2007) Common clinical symptoms of ''S.haemolyticus'' are fever and an increase in white blood cell population (leukocytosis) (Falcone,Campanile, et al, 2007). <br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (Novick, 2006). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but are now discovered in the other non-aureus, coagulast-negative staphylococci as well (Tristan,Lina, et al, 2006). In one studies published in 1994, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro (Molnar,Hevessy, et al, 1994). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species.<br />
<br />
''S.haemolyticus''’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (''Staphylococcus lugdunensis synergistic hemolysin'') in ''S.lugdunensis'', the latter of which shows significant similarities in structure with GGI. (Tristan,Lina, et al, 2006) These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons. Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides (Oh,Kim, et al, 1999).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell (Hansson,Stahl, et al, 1992). It was the first time such system could be constructed in a Gram-positive species. The technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria (Hansson,Stahl, et al, 1992).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Dr. Keiichi Hiramatsu, is a very important and significant step in tackling this problem(Takeuchi,Watanabe, et al, 2005). The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” (Takeuchi,Watanabe, et al, 2005).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above (Takeuchi,Watanabe, et al, 2005), Dr. Hiramatsu’s group is examining the other types of genetic rearrangement that are also responsible for the frequent structural alteration of ''S.haemolyticus'' genome. Recent results indicated that “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in clinical strain JCSC1435 of ''S.haemolyticus'' (Watanabe,Ito, et al, 2007).<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, and the combination of vancomycin and beta-lactams antitbiotics (Vignaroli,Biavasco and Varaldo, 2006).<br />
<br />
==References==<br />
Billot-Klein, D., Gutmann, L., Bryant, D., Bell, D., Van Heijenoort, J., Grewal, J., and Shlaes, D.M. (1996). Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics. J. Bacteriol. 15, 4696-4703.<br />
<br />
Brooks, G., Butel, J., and Morse, S. (2001). The Staphylococci. In Medical Microbiology, (New York: McGraw-Hill) pp. 197-202.<br />
<br />
Falcone, M., Campanile, F., Giannella, M., Borbone, S., Stefani, S., and Venditti, M. (2007). Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases. Diagn. Microbiol. Infect. Dis. 3, 325-331.<br />
<br />
Froggatt, J.W., Johnston, J.L., Galetto, D.W., and Archer, G.L. (1989). Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 4, 460-466.<br />
Hansson, M., Stahl, S., Nguyen, T.N., Bachi, T., Robert, A., Binz, H., Sjolander, A., and Uhlen, M. (1992). Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus. J. Bacteriol. 13, 4239-4245.<br />
<br />
Molnar, C., Hevessy, Z., Rozgonyi, F., and Gemmell, C.G. (1994). Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro. J. Clin. Pathol. 8, 743-748.<br />
<br />
Novick, R. (2006). Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation. In Gram-positive Pathogens, V. Fischetti, R. Novick, J. Ferretti, D. Portnoy and J. Rood eds., (Washington, D.C: ASM Press) pp. 496-510.<br />
<br />
Oh, B., Kim, H., Lee, J., Kang, S., and Oh, T. (1999). Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning. FEMS Microbiol. Lett. 2, 385-392.<br />
Takeuchi, F., Watanabe, S., Baba, T., Yuzawa, H., Ito, T., Morimoto, Y., Kuroda, M., Cui, L., Takahashi, M., Ankai, A. et al. (2005). Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species. J. Bacteriol. 21, 7292-7308.<br />
<br />
Tristan, A., Lina, G., Etienne, J., and Vandenesch, F. (2006). Biology and Pathogenicity of Staphylococci other than Staphylococcus aureus and Staphylococcus epidermis. V. Fischetti, R. Novick, J. Ferretti, D. Portnoy and J. Rood eds., (Washington, D.C: ASM Press) pp. 572-586.<br />
<br />
Vignaroli, C., Biavasco, F., and Varaldo, P.E. (2006). Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 7, 2577-2582.<br />
<br />
Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F., and Hiramatsu, K. (2007). Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435. J. Bacteriol. 7, 2921-2925.<br />
<br />
Watson, D.C., Yaguchi, M., Bisaillon, J.G., Beaudet, R., and Morosoli, R. (1988). The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus. Biochem. J. 1, 87-93.</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=14986Staphylococcus haemolyticus2007-06-05T01:11:03Z<p>Bdtruong: /* Metabolism */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection (2). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (3). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (2). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time (2). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics(2). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (4).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (3). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism (4)<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp (3).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome (3). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability (3). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host (3). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) (3). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance (3). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors (3). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin (5, 13). This is an ability that is unique among staphylococci (5). The peptidoglycan structure of ''S.haemolyticus'' has been studied (5) to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) (5). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' (5). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' (5).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids (3). Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline (3).<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci (3). However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species (3). This is another example demonstrating the flexibility of S.haemolyticus genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (Tristan,Lina, et al, 2006). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (Tristan,Lina, et al, 2006). ''S.haemolyticus'' survive successfully on the drier regions of the body (Tristan,Lina, et al, 2006), while it can also be found frequently in human blood cultures (Takeuchi,Watanabe, et al, 2005). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI. The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (Tristan,Lina, et al, 2006). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human (Watson,Yaguchi, et al, 1988).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (Brooks,Butel and Morse, 2001). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (Brooks,Butel and Morse, 2001). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci. (Brooks,Butel and Morse, 2001) ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints. (Tristan,Lina, et al, 2006) In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death. (Falcone,Campanile, et al, 2007) Common clinical symptoms of ''S.haemolyticus'' are fever and an increase in white blood cell population (leukocytosis) (Falcone,Campanile, et al, 2007). <br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (Novick, 2006). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but are now discovered in the other non-aureus, coagulast-negative staphylococci as well (Tristan,Lina, et al, 2006). In one studies published in 1994, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro (Molnar,Hevessy, et al, 1994). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species.<br />
<br />
''S.haemolyticus''’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (''Staphylococcus lugdunensis synergistic hemolysin'') in ''S.lugdunensis'', the latter of which shows significant similarities in structure with GGI. (Tristan,Lina, et al, 2006) These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons. Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides (Oh,Kim, et al, 1999).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell (Hansson,Stahl, et al, 1992). It was the first time such system could be constructed in a Gram-positive species. The technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria (Hansson,Stahl, et al, 1992).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Dr. Keiichi Hiramatsu, is a very important and significant step in tackling this problem(Takeuchi,Watanabe, et al, 2005). The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” (Takeuchi,Watanabe, et al, 2005).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above (Takeuchi,Watanabe, et al, 2005), Dr. Hiramatsu’s group is examining the other types of genetic rearrangement that are also responsible for the frequent structural alteration of ''S.haemolyticus'' genome. Recent results indicated that “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in clinical strain JCSC1435 of ''S.haemolyticus'' (Watanabe,Ito, et al, 2007).<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, and the combination of vancomycin and beta-lactams antitbiotics (Vignaroli,Biavasco and Varaldo, 2006).<br />
<br />
==References==<br />
Billot-Klein, D., Gutmann, L., Bryant, D., Bell, D., Van Heijenoort, J., Grewal, J., and Shlaes, D.M. (1996). Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics. J. Bacteriol. 15, 4696-4703.<br />
<br />
Brooks, G., Butel, J., and Morse, S. (2001). The Staphylococci. In Medical Microbiology, (New York: McGraw-Hill) pp. 197-202.<br />
<br />
Falcone, M., Campanile, F., Giannella, M., Borbone, S., Stefani, S., and Venditti, M. (2007). Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases. Diagn. Microbiol. Infect. Dis. 3, 325-331.<br />
<br />
Froggatt, J.W., Johnston, J.L., Galetto, D.W., and Archer, G.L. (1989). Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 4, 460-466.<br />
Hansson, M., Stahl, S., Nguyen, T.N., Bachi, T., Robert, A., Binz, H., Sjolander, A., and Uhlen, M. (1992). Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus. J. Bacteriol. 13, 4239-4245.<br />
<br />
Molnar, C., Hevessy, Z., Rozgonyi, F., and Gemmell, C.G. (1994). Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro. J. Clin. Pathol. 8, 743-748.<br />
<br />
Novick, R. (2006). Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation. In Gram-positive Pathogens, V. Fischetti, R. Novick, J. Ferretti, D. Portnoy and J. Rood eds., (Washington, D.C: ASM Press) pp. 496-510.<br />
<br />
Oh, B., Kim, H., Lee, J., Kang, S., and Oh, T. (1999). Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning. FEMS Microbiol. Lett. 2, 385-392.<br />
Takeuchi, F., Watanabe, S., Baba, T., Yuzawa, H., Ito, T., Morimoto, Y., Kuroda, M., Cui, L., Takahashi, M., Ankai, A. et al. (2005). Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species. J. Bacteriol. 21, 7292-7308.<br />
<br />
Tristan, A., Lina, G., Etienne, J., and Vandenesch, F. (2006). Biology and Pathogenicity of Staphylococci other than Staphylococcus aureus and Staphylococcus epidermis. V. Fischetti, R. Novick, J. Ferretti, D. Portnoy and J. Rood eds., (Washington, D.C: ASM Press) pp. 572-586.<br />
<br />
Vignaroli, C., Biavasco, F., and Varaldo, P.E. (2006). Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 7, 2577-2582.<br />
<br />
Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F., and Hiramatsu, K. (2007). Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435. J. Bacteriol. 7, 2921-2925.<br />
<br />
Watson, D.C., Yaguchi, M., Bisaillon, J.G., Beaudet, R., and Morosoli, R. (1988). The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus. Biochem. J. 1, 87-93.</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=14812Staphylococcus haemolyticus2007-06-05T00:17:24Z<p>Bdtruong: /* Cell Wall */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection (2). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (3). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (2). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time (2). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics(2). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (4).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (3). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism (4)<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp (3).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome (3). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability (3). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host (3). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) (3). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance (3). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors (3). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin (5, 13). This is an ability that is unique among staphylococci (5). The peptidoglycan structure of ''S.haemolyticus'' has been studied (5) to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of S.haemolyticus is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the cross bridge structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2) (5). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus'' (5). This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus'' (5).<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids. Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline. (Takeuchi,Watanabe, et al, 2005)<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci (Takeuchi,Watanabe, et al, 2005) However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species (Takeuchi,Watanabe, et al, 2005). Again, this can also be considered another example demonstrating the flexibility of ''S.haemolyticus'' genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (Tristan,Lina, et al, 2006). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (Tristan,Lina, et al, 2006). ''S.haemolyticus'' survive successfully on the drier regions of the body (Tristan,Lina, et al, 2006), while it can also be found frequently in human blood cultures (Takeuchi,Watanabe, et al, 2005). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI. The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (Tristan,Lina, et al, 2006). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human (Watson,Yaguchi, et al, 1988).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (Brooks,Butel and Morse, 2001). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (Brooks,Butel and Morse, 2001). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci. (Brooks,Butel and Morse, 2001) ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints. (Tristan,Lina, et al, 2006) In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death. (Falcone,Campanile, et al, 2007) Common clinical symptoms of ''S.haemolyticus'' are fever and an increase in white blood cell population (leukocytosis) (Falcone,Campanile, et al, 2007). <br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (Novick, 2006). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but are now discovered in the other non-aureus, coagulast-negative staphylococci as well (Tristan,Lina, et al, 2006). In one studies published in 1994, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro (Molnar,Hevessy, et al, 1994). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species.<br />
<br />
''S.haemolyticus''’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (''Staphylococcus lugdunensis synergistic hemolysin'') in ''S.lugdunensis'', the latter of which shows significant similarities in structure with GGI. (Tristan,Lina, et al, 2006) These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons. Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides (Oh,Kim, et al, 1999).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell (Hansson,Stahl, et al, 1992). It was the first time such system could be constructed in a Gram-positive species. The technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria (Hansson,Stahl, et al, 1992).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Dr. Keiichi Hiramatsu, is a very important and significant step in tackling this problem(Takeuchi,Watanabe, et al, 2005). The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” (Takeuchi,Watanabe, et al, 2005).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above (Takeuchi,Watanabe, et al, 2005), Dr. Hiramatsu’s group is examining the other types of genetic rearrangement that are also responsible for the frequent structural alteration of ''S.haemolyticus'' genome. Recent results indicated that “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in clinical strain JCSC1435 of ''S.haemolyticus'' (Watanabe,Ito, et al, 2007).<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, and the combination of vancomycin and beta-lactams antitbiotics (Vignaroli,Biavasco and Varaldo, 2006).<br />
<br />
==References==<br />
Billot-Klein, D., Gutmann, L., Bryant, D., Bell, D., Van Heijenoort, J., Grewal, J., and Shlaes, D.M. (1996). Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics. J. Bacteriol. 15, 4696-4703.<br />
<br />
Brooks, G., Butel, J., and Morse, S. (2001). The Staphylococci. In Medical Microbiology, (New York: McGraw-Hill) pp. 197-202.<br />
<br />
Falcone, M., Campanile, F., Giannella, M., Borbone, S., Stefani, S., and Venditti, M. (2007). Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases. Diagn. Microbiol. Infect. Dis. 3, 325-331.<br />
<br />
Froggatt, J.W., Johnston, J.L., Galetto, D.W., and Archer, G.L. (1989). Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 4, 460-466.<br />
Hansson, M., Stahl, S., Nguyen, T.N., Bachi, T., Robert, A., Binz, H., Sjolander, A., and Uhlen, M. (1992). Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus. J. Bacteriol. 13, 4239-4245.<br />
<br />
Molnar, C., Hevessy, Z., Rozgonyi, F., and Gemmell, C.G. (1994). Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro. J. Clin. Pathol. 8, 743-748.<br />
<br />
Novick, R. (2006). Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation. In Gram-positive Pathogens, V. Fischetti, R. Novick, J. Ferretti, D. Portnoy and J. Rood eds., (Washington, D.C: ASM Press) pp. 496-510.<br />
<br />
Oh, B., Kim, H., Lee, J., Kang, S., and Oh, T. (1999). Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning. FEMS Microbiol. Lett. 2, 385-392.<br />
Takeuchi, F., Watanabe, S., Baba, T., Yuzawa, H., Ito, T., Morimoto, Y., Kuroda, M., Cui, L., Takahashi, M., Ankai, A. et al. (2005). Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species. J. Bacteriol. 21, 7292-7308.<br />
<br />
Tristan, A., Lina, G., Etienne, J., and Vandenesch, F. (2006). Biology and Pathogenicity of Staphylococci other than Staphylococcus aureus and Staphylococcus epidermis. V. Fischetti, R. Novick, J. Ferretti, D. Portnoy and J. Rood eds., (Washington, D.C: ASM Press) pp. 572-586.<br />
<br />
Vignaroli, C., Biavasco, F., and Varaldo, P.E. (2006). Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 7, 2577-2582.<br />
<br />
Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F., and Hiramatsu, K. (2007). Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435. J. Bacteriol. 7, 2921-2925.<br />
<br />
Watson, D.C., Yaguchi, M., Bisaillon, J.G., Beaudet, R., and Morosoli, R. (1988). The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus. Biochem. J. 1, 87-93.</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=14805Staphylococcus haemolyticus2007-06-05T00:14:08Z<p>Bdtruong: /* Genome structure */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection (2). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (3). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (2). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time (2). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics(2). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (4).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (3). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism (4)<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp (3).<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome (3). However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is therefore called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability (3). On the other hand, the region is most likely responsible for the diversification of staphylococci species and enables the bacteria to successfully colonize and infect the human host (3). <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs) (3). These ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance (3). <br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors (3). However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
(Billot-Klein,Gutmann, et al, 1996)<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains of ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin. This is an ability that is unique among staphylococci. The peptidoglycan structure of ''S.haemolyticus'' has been studied to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of ''S.haemolyticus'' is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the probable cross bridges structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus''. This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus''.<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids. Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline. (Takeuchi,Watanabe, et al, 2005)<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci (Takeuchi,Watanabe, et al, 2005) However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species (Takeuchi,Watanabe, et al, 2005). Again, this can also be considered another example demonstrating the flexibility of ''S.haemolyticus'' genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (Tristan,Lina, et al, 2006). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (Tristan,Lina, et al, 2006). ''S.haemolyticus'' survive successfully on the drier regions of the body (Tristan,Lina, et al, 2006), while it can also be found frequently in human blood cultures (Takeuchi,Watanabe, et al, 2005). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI. The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (Tristan,Lina, et al, 2006). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human (Watson,Yaguchi, et al, 1988).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (Brooks,Butel and Morse, 2001). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (Brooks,Butel and Morse, 2001). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci. (Brooks,Butel and Morse, 2001) ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints. (Tristan,Lina, et al, 2006) In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death. (Falcone,Campanile, et al, 2007) Common clinical symptoms of ''S.haemolyticus'' are fever and an increase in white blood cell population (leukocytosis) (Falcone,Campanile, et al, 2007). <br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (Novick, 2006). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but are now discovered in the other non-aureus, coagulast-negative staphylococci as well (Tristan,Lina, et al, 2006). In one studies published in 1994, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro (Molnar,Hevessy, et al, 1994). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species.<br />
<br />
''S.haemolyticus''’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (''Staphylococcus lugdunensis synergistic hemolysin'') in ''S.lugdunensis'', the latter of which shows significant similarities in structure with GGI. (Tristan,Lina, et al, 2006) These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons. Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides (Oh,Kim, et al, 1999).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell (Hansson,Stahl, et al, 1992). It was the first time such system could be constructed in a Gram-positive species. The technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria (Hansson,Stahl, et al, 1992).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Dr. Keiichi Hiramatsu, is a very important and significant step in tackling this problem(Takeuchi,Watanabe, et al, 2005). The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” (Takeuchi,Watanabe, et al, 2005).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above (Takeuchi,Watanabe, et al, 2005), Dr. Hiramatsu’s group is examining the other types of genetic rearrangement that are also responsible for the frequent structural alteration of ''S.haemolyticus'' genome. Recent results indicated that “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in clinical strain JCSC1435 of ''S.haemolyticus'' (Watanabe,Ito, et al, 2007).<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, and the combination of vancomycin and beta-lactams antitbiotics (Vignaroli,Biavasco and Varaldo, 2006).<br />
<br />
==References==<br />
Billot-Klein, D., Gutmann, L., Bryant, D., Bell, D., Van Heijenoort, J., Grewal, J., and Shlaes, D.M. (1996). Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics. J. Bacteriol. 15, 4696-4703.<br />
<br />
Brooks, G., Butel, J., and Morse, S. (2001). The Staphylococci. In Medical Microbiology, (New York: McGraw-Hill) pp. 197-202.<br />
<br />
Falcone, M., Campanile, F., Giannella, M., Borbone, S., Stefani, S., and Venditti, M. (2007). Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases. Diagn. Microbiol. Infect. Dis. 3, 325-331.<br />
<br />
Froggatt, J.W., Johnston, J.L., Galetto, D.W., and Archer, G.L. (1989). Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 4, 460-466.<br />
Hansson, M., Stahl, S., Nguyen, T.N., Bachi, T., Robert, A., Binz, H., Sjolander, A., and Uhlen, M. (1992). Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus. J. Bacteriol. 13, 4239-4245.<br />
<br />
Molnar, C., Hevessy, Z., Rozgonyi, F., and Gemmell, C.G. (1994). Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro. J. Clin. Pathol. 8, 743-748.<br />
<br />
Novick, R. (2006). Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation. In Gram-positive Pathogens, V. Fischetti, R. Novick, J. Ferretti, D. Portnoy and J. Rood eds., (Washington, D.C: ASM Press) pp. 496-510.<br />
<br />
Oh, B., Kim, H., Lee, J., Kang, S., and Oh, T. (1999). Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning. FEMS Microbiol. Lett. 2, 385-392.<br />
Takeuchi, F., Watanabe, S., Baba, T., Yuzawa, H., Ito, T., Morimoto, Y., Kuroda, M., Cui, L., Takahashi, M., Ankai, A. et al. (2005). Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species. J. Bacteriol. 21, 7292-7308.<br />
<br />
Tristan, A., Lina, G., Etienne, J., and Vandenesch, F. (2006). Biology and Pathogenicity of Staphylococci other than Staphylococcus aureus and Staphylococcus epidermis. V. Fischetti, R. Novick, J. Ferretti, D. Portnoy and J. Rood eds., (Washington, D.C: ASM Press) pp. 572-586.<br />
<br />
Vignaroli, C., Biavasco, F., and Varaldo, P.E. (2006). Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 7, 2577-2582.<br />
<br />
Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F., and Hiramatsu, K. (2007). Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435. J. Bacteriol. 7, 2921-2925.<br />
<br />
Watson, D.C., Yaguchi, M., Bisaillon, J.G., Beaudet, R., and Morosoli, R. (1988). The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus. Biochem. J. 1, 87-93.</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=14791Staphylococcus haemolyticus2007-06-05T00:08:44Z<p>Bdtruong: /* Description and significance */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood (1). <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of opportunistic infection (2). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (3). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (2). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time (2). Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics(2). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (4).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (3). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs), which might be responsible for the frequent genomic arrangement observed in this organism (4)<br />
<br />
==Genome structure==<br />
<br />
(Takeuchi,Watanabe, et al, 2005)<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp.<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome. However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability. On the other hand, the region is most likely responsible for the diversification of staphylococci species which enables them to successfully colonize and infect the human host. <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs). The ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance. <br />
While 6% of the orfs found in the more virulent S.aureus are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors. However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
(Billot-Klein,Gutmann, et al, 1996)<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains of ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin. This is an ability that is unique among staphylococci. The peptidoglycan structure of ''S.haemolyticus'' has been studied to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of ''S.haemolyticus'' is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the probable cross bridges structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus''. This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus''.<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids. Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline. (Takeuchi,Watanabe, et al, 2005)<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci (Takeuchi,Watanabe, et al, 2005) However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species (Takeuchi,Watanabe, et al, 2005). Again, this can also be considered another example demonstrating the flexibility of ''S.haemolyticus'' genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (Tristan,Lina, et al, 2006). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (Tristan,Lina, et al, 2006). ''S.haemolyticus'' survive successfully on the drier regions of the body (Tristan,Lina, et al, 2006), while it can also be found frequently in human blood cultures (Takeuchi,Watanabe, et al, 2005). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI. The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (Tristan,Lina, et al, 2006). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human (Watson,Yaguchi, et al, 1988).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (Brooks,Butel and Morse, 2001). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (Brooks,Butel and Morse, 2001). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci. (Brooks,Butel and Morse, 2001) ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints. (Tristan,Lina, et al, 2006) In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death. (Falcone,Campanile, et al, 2007) Common clinical symptoms of ''S.haemolyticus'' are fever and an increase in white blood cell population (leukocytosis) (Falcone,Campanile, et al, 2007). <br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (Novick, 2006). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but are now discovered in the other non-aureus, coagulast-negative staphylococci as well (Tristan,Lina, et al, 2006). In one studies published in 1994, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro (Molnar,Hevessy, et al, 1994). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species.<br />
<br />
''S.haemolyticus''’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (''Staphylococcus lugdunensis synergistic hemolysin'') in ''S.lugdunensis'', the latter of which shows significant similarities in structure with GGI. (Tristan,Lina, et al, 2006) These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons. Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides (Oh,Kim, et al, 1999).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell (Hansson,Stahl, et al, 1992). It was the first time such system could be constructed in a Gram-positive species. The technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria (Hansson,Stahl, et al, 1992).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Dr. Keiichi Hiramatsu, is a very important and significant step in tackling this problem(Takeuchi,Watanabe, et al, 2005). The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” (Takeuchi,Watanabe, et al, 2005).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above (Takeuchi,Watanabe, et al, 2005), Dr. Hiramatsu’s group is examining the other types of genetic rearrangement that are also responsible for the frequent structural alteration of ''S.haemolyticus'' genome. Recent results indicated that “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in clinical strain JCSC1435 of ''S.haemolyticus'' (Watanabe,Ito, et al, 2007).<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, and the combination of vancomycin and beta-lactams antitbiotics (Vignaroli,Biavasco and Varaldo, 2006).<br />
<br />
==References==<br />
Billot-Klein, D., Gutmann, L., Bryant, D., Bell, D., Van Heijenoort, J., Grewal, J., and Shlaes, D.M. (1996). Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics. J. Bacteriol. 15, 4696-4703.<br />
<br />
Brooks, G., Butel, J., and Morse, S. (2001). The Staphylococci. In Medical Microbiology, (New York: McGraw-Hill) pp. 197-202.<br />
<br />
Falcone, M., Campanile, F., Giannella, M., Borbone, S., Stefani, S., and Venditti, M. (2007). Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases. Diagn. Microbiol. Infect. Dis. 3, 325-331.<br />
<br />
Froggatt, J.W., Johnston, J.L., Galetto, D.W., and Archer, G.L. (1989). Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 4, 460-466.<br />
Hansson, M., Stahl, S., Nguyen, T.N., Bachi, T., Robert, A., Binz, H., Sjolander, A., and Uhlen, M. (1992). Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus. J. Bacteriol. 13, 4239-4245.<br />
<br />
Molnar, C., Hevessy, Z., Rozgonyi, F., and Gemmell, C.G. (1994). Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro. J. Clin. Pathol. 8, 743-748.<br />
<br />
Novick, R. (2006). Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation. In Gram-positive Pathogens, V. Fischetti, R. Novick, J. Ferretti, D. Portnoy and J. Rood eds., (Washington, D.C: ASM Press) pp. 496-510.<br />
<br />
Oh, B., Kim, H., Lee, J., Kang, S., and Oh, T. (1999). Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning. FEMS Microbiol. Lett. 2, 385-392.<br />
Takeuchi, F., Watanabe, S., Baba, T., Yuzawa, H., Ito, T., Morimoto, Y., Kuroda, M., Cui, L., Takahashi, M., Ankai, A. et al. (2005). Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species. J. Bacteriol. 21, 7292-7308.<br />
<br />
Tristan, A., Lina, G., Etienne, J., and Vandenesch, F. (2006). Biology and Pathogenicity of Staphylococci other than Staphylococcus aureus and Staphylococcus epidermis. V. Fischetti, R. Novick, J. Ferretti, D. Portnoy and J. Rood eds., (Washington, D.C: ASM Press) pp. 572-586.<br />
<br />
Vignaroli, C., Biavasco, F., and Varaldo, P.E. (2006). Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 7, 2577-2582.<br />
<br />
Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F., and Hiramatsu, K. (2007). Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435. J. Bacteriol. 7, 2921-2925.<br />
<br />
Watson, D.C., Yaguchi, M., Bisaillon, J.G., Beaudet, R., and Morosoli, R. (1988). The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus. Biochem. J. 1, 87-93.</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=13454Staphylococcus haemolyticus2007-06-04T02:23:42Z<p>Bdtruong: /* References */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood. (Tristan,Lina, et al, 2006) <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of infection (Falcone,Campanile, et al, 2007). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (Takeuchi,Watanabe, et al, 2005). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (Falcone,Campanile, et al, 2007). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time. (Falcone,Campanile, et al, 2007) Common antibiotics that are subject to resistance in S haemolyticus include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics (Falcone,Campanile, et al, 2007). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (Froggatt,Johnston, et al, 1989).<br />
<br />
In order to study the multi-drug resistant ability of Staphylococcus haemolyticus and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (Takeuchi,Watanabe, et al, 2005). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs). These sequences might be responsible for the frequent genomic arrangement observed in this organism.(Froggatt,Johnston, et al, 1989)<br />
<br />
==Genome structure==<br />
<br />
(Takeuchi,Watanabe, et al, 2005)<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp.<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome. However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability. On the other hand, the region is most likely responsible for the diversification of staphylococci species which enables them to successfully colonize and infect the human host. <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs). The ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance. <br />
While 6% of the orfs found in the more virulent S.aureus are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors. However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
(Billot-Klein,Gutmann, et al, 1996)<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains of ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin. This is an ability that is unique among staphylococci. The peptidoglycan structure of ''S.haemolyticus'' has been studied to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of ''S.haemolyticus'' is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the probable cross bridges structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus''. This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus''.<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids. Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline. (Takeuchi,Watanabe, et al, 2005)<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci (Takeuchi,Watanabe, et al, 2005) However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species (Takeuchi,Watanabe, et al, 2005). Again, this can also be considered another example demonstrating the flexibility of ''S.haemolyticus'' genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (Tristan,Lina, et al, 2006). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (Tristan,Lina, et al, 2006). ''S.haemolyticus'' survive successfully on the drier regions of the body (Tristan,Lina, et al, 2006), while it can also be found frequently in human blood cultures (Takeuchi,Watanabe, et al, 2005). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI. The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (Tristan,Lina, et al, 2006). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human (Watson,Yaguchi, et al, 1988).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (Brooks,Butel and Morse, 2001). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (Brooks,Butel and Morse, 2001). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci. (Brooks,Butel and Morse, 2001) ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints. (Tristan,Lina, et al, 2006) In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death. (Falcone,Campanile, et al, 2007) Common clinical symptoms of ''S.haemolyticus'' are fever and an increase in white blood cell population (leukocytosis) (Falcone,Campanile, et al, 2007). <br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (Novick, 2006). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but are now discovered in the other non-aureus, coagulast-negative staphylococci as well (Tristan,Lina, et al, 2006). In one studies published in 1994, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro (Molnar,Hevessy, et al, 1994). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species.<br />
<br />
''S.haemolyticus''’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (''Staphylococcus lugdunensis synergistic hemolysin'') in ''S.lugdunensis'', the latter of which shows significant similarities in structure with GGI. (Tristan,Lina, et al, 2006) These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons. Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides (Oh,Kim, et al, 1999).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell (Hansson,Stahl, et al, 1992). It was the first time such system could be constructed in a Gram-positive species. The technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria (Hansson,Stahl, et al, 1992).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Dr. Keiichi Hiramatsu, is a very important and significant step in tackling this problem(Takeuchi,Watanabe, et al, 2005). The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” (Takeuchi,Watanabe, et al, 2005).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above (Takeuchi,Watanabe, et al, 2005), Dr. Hiramatsu’s group is examining the other types of genetic rearrangement that are also responsible for the frequent structural alteration of ''S.haemolyticus'' genome. Recent results indicated that “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in clinical strain JCSC1435 of ''S.haemolyticus'' (Watanabe,Ito, et al, 2007).<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, and the combination of vancomycin and beta-lactams antitbiotics (Vignaroli,Biavasco and Varaldo, 2006).<br />
<br />
==References==<br />
Billot-Klein, D., Gutmann, L., Bryant, D., Bell, D., Van Heijenoort, J., Grewal, J., and Shlaes, D.M. (1996). Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics. J. Bacteriol. 15, 4696-4703.<br />
<br />
Brooks, G., Butel, J., and Morse, S. (2001). The Staphylococci. In Medical Microbiology, (New York: McGraw-Hill) pp. 197-202.<br />
<br />
Falcone, M., Campanile, F., Giannella, M., Borbone, S., Stefani, S., and Venditti, M. (2007). Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases. Diagn. Microbiol. Infect. Dis. 3, 325-331.<br />
<br />
Froggatt, J.W., Johnston, J.L., Galetto, D.W., and Archer, G.L. (1989). Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 4, 460-466.<br />
Hansson, M., Stahl, S., Nguyen, T.N., Bachi, T., Robert, A., Binz, H., Sjolander, A., and Uhlen, M. (1992). Expression of recombinant proteins on the surface of the coagulase-negative bacterium Staphylococcus xylosus. J. Bacteriol. 13, 4239-4245.<br />
<br />
Molnar, C., Hevessy, Z., Rozgonyi, F., and Gemmell, C.G. (1994). Pathogenicity and virulence of coagulase negative staphylococci in relation to adherence, hydrophobicity, and toxin production in vitro. J. Clin. Pathol. 8, 743-748.<br />
<br />
Novick, R. (2006). Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation. In Gram-positive Pathogens, V. Fischetti, R. Novick, J. Ferretti, D. Portnoy and J. Rood eds., (Washington, D.C: ASM Press) pp. 496-510.<br />
<br />
Oh, B., Kim, H., Lee, J., Kang, S., and Oh, T. (1999). Staphylococcus haemolyticus lipase: biochemical properties, substrate specificity and gene cloning. FEMS Microbiol. Lett. 2, 385-392.<br />
Takeuchi, F., Watanabe, S., Baba, T., Yuzawa, H., Ito, T., Morimoto, Y., Kuroda, M., Cui, L., Takahashi, M., Ankai, A. et al. (2005). Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species. J. Bacteriol. 21, 7292-7308.<br />
<br />
Tristan, A., Lina, G., Etienne, J., and Vandenesch, F. (2006). Biology and Pathogenicity of Staphylococci other than Staphylococcus aureus and Staphylococcus epidermis. V. Fischetti, R. Novick, J. Ferretti, D. Portnoy and J. Rood eds., (Washington, D.C: ASM Press) pp. 572-586.<br />
<br />
Vignaroli, C., Biavasco, F., and Varaldo, P.E. (2006). Interactions between glycopeptides and beta-lactams against isogenic pairs of teicoplanin-susceptible and -resistant strains of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 7, 2577-2582.<br />
<br />
Watanabe, S., Ito, T., Morimoto, Y., Takeuchi, F., and Hiramatsu, K. (2007). Precise excision and self-integration of a composite transposon as a model for spontaneous large-scale chromosome inversion/deletion of the Staphylococcus haemolyticus clinical strain JCSC1435. J. Bacteriol. 7, 2921-2925.<br />
<br />
Watson, D.C., Yaguchi, M., Bisaillon, J.G., Beaudet, R., and Morosoli, R. (1988). The amino acid sequence of a gonococcal growth inhibitor from Staphylococcus haemolyticus. Biochem. J. 1, 87-93.</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=13453Staphylococcus haemolyticus2007-06-04T02:22:45Z<p>Bdtruong: /* Current Research */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood. (Tristan,Lina, et al, 2006) <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of infection (Falcone,Campanile, et al, 2007). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (Takeuchi,Watanabe, et al, 2005). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (Falcone,Campanile, et al, 2007). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time. (Falcone,Campanile, et al, 2007) Common antibiotics that are subject to resistance in S haemolyticus include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics (Falcone,Campanile, et al, 2007). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (Froggatt,Johnston, et al, 1989).<br />
<br />
In order to study the multi-drug resistant ability of Staphylococcus haemolyticus and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (Takeuchi,Watanabe, et al, 2005). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs). These sequences might be responsible for the frequent genomic arrangement observed in this organism.(Froggatt,Johnston, et al, 1989)<br />
<br />
==Genome structure==<br />
<br />
(Takeuchi,Watanabe, et al, 2005)<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp.<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome. However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability. On the other hand, the region is most likely responsible for the diversification of staphylococci species which enables them to successfully colonize and infect the human host. <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs). The ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance. <br />
While 6% of the orfs found in the more virulent S.aureus are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors. However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
(Billot-Klein,Gutmann, et al, 1996)<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains of ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin. This is an ability that is unique among staphylococci. The peptidoglycan structure of ''S.haemolyticus'' has been studied to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of ''S.haemolyticus'' is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the probable cross bridges structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus''. This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus''.<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids. Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline. (Takeuchi,Watanabe, et al, 2005)<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci (Takeuchi,Watanabe, et al, 2005) However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species (Takeuchi,Watanabe, et al, 2005). Again, this can also be considered another example demonstrating the flexibility of ''S.haemolyticus'' genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (Tristan,Lina, et al, 2006). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (Tristan,Lina, et al, 2006). ''S.haemolyticus'' survive successfully on the drier regions of the body (Tristan,Lina, et al, 2006), while it can also be found frequently in human blood cultures (Takeuchi,Watanabe, et al, 2005). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI. The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (Tristan,Lina, et al, 2006). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human (Watson,Yaguchi, et al, 1988).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (Brooks,Butel and Morse, 2001). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (Brooks,Butel and Morse, 2001). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci. (Brooks,Butel and Morse, 2001) ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints. (Tristan,Lina, et al, 2006) In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death. (Falcone,Campanile, et al, 2007) Common clinical symptoms of ''S.haemolyticus'' are fever and an increase in white blood cell population (leukocytosis) (Falcone,Campanile, et al, 2007). <br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (Novick, 2006). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but are now discovered in the other non-aureus, coagulast-negative staphylococci as well (Tristan,Lina, et al, 2006). In one studies published in 1994, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro (Molnar,Hevessy, et al, 1994). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species.<br />
<br />
''S.haemolyticus''’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (''Staphylococcus lugdunensis synergistic hemolysin'') in ''S.lugdunensis'', the latter of which shows significant similarities in structure with GGI. (Tristan,Lina, et al, 2006) These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons. Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides (Oh,Kim, et al, 1999).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell (Hansson,Stahl, et al, 1992). It was the first time such system could be constructed in a Gram-positive species. The technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria (Hansson,Stahl, et al, 1992).<br />
<br />
==Current Research==<br />
Although ''Staphylococcus haemolyticus'' is relatively less virulent than some other staphylococci such as ''S.aureus'', the ability of the species to acquire multi-antibiotic resistance has made it a serious threat to worldwide healthcare facilities. Whole-genome sequencing of ''S.haemolyticus'', carried out by a research group at at Jutendo University in Tokyo, Japan and led by Dr. Dr. Keiichi Hiramatsu, is a very important and significant step in tackling this problem(Takeuchi,Watanabe, et al, 2005). The information provided by the genome sequence will not only allow further examinations of the species’ characteristic bacterial lifestyle, but also facilitates the “development of novel immunotherapeutic and chemotherapeutic approaches to control them” (Takeuchi,Watanabe, et al, 2005).<br />
<br />
After discovering the presence of abundant IS copies in the chromosome as mentioned above (Takeuchi,Watanabe, et al, 2005), Dr. Hiramatsu’s group is examining the other types of genetic rearrangement that are also responsible for the frequent structural alteration of ''S.haemolyticus'' genome. Recent results indicated that “precise excision and self-integration of a composite transposon” (ISSha1) lead to a large-scale chromosome inversion/deletion found in clinical strain JCSC1435 of ''S.haemolyticus'' (Watanabe,Ito, et al, 2007).<br />
Beside genomic and genetic approaches, clinical investigations combined with molecular approaches are being carried out to find effective strategy against the development of ''S.haemolyticus'' antibiotic resistant strains. Studies are aiming at a promising strategy of using different types of antibiotics synergistically to fight against specific antibiotic-resistant strains. Some examples are the combination of glycopeptide and beta-lactams antibiotics against methicillin- and teicoplanin-resistant staphylococci strains, and the combination of vancomycin and beta-lactams antitbiotics (Vignaroli,Biavasco and Varaldo, 2006).<br />
<br />
==References==<br />
1. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases]. Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species]. J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus]. Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics]. J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
<br />
Edited by Bao D. Truong, student of [mailto:ralarsen@ucsd.edu Rachel Larsen]</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=13451Staphylococcus haemolyticus2007-06-04T02:21:18Z<p>Bdtruong: /* Application to Biotechnology */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood. (Tristan,Lina, et al, 2006) <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of infection (Falcone,Campanile, et al, 2007). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (Takeuchi,Watanabe, et al, 2005). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (Falcone,Campanile, et al, 2007). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time. (Falcone,Campanile, et al, 2007) Common antibiotics that are subject to resistance in S haemolyticus include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics (Falcone,Campanile, et al, 2007). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (Froggatt,Johnston, et al, 1989).<br />
<br />
In order to study the multi-drug resistant ability of Staphylococcus haemolyticus and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (Takeuchi,Watanabe, et al, 2005). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs). These sequences might be responsible for the frequent genomic arrangement observed in this organism.(Froggatt,Johnston, et al, 1989)<br />
<br />
==Genome structure==<br />
<br />
(Takeuchi,Watanabe, et al, 2005)<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp.<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome. However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability. On the other hand, the region is most likely responsible for the diversification of staphylococci species which enables them to successfully colonize and infect the human host. <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs). The ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance. <br />
While 6% of the orfs found in the more virulent S.aureus are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors. However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
(Billot-Klein,Gutmann, et al, 1996)<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains of ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin. This is an ability that is unique among staphylococci. The peptidoglycan structure of ''S.haemolyticus'' has been studied to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of ''S.haemolyticus'' is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the probable cross bridges structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus''. This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus''.<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids. Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline. (Takeuchi,Watanabe, et al, 2005)<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci (Takeuchi,Watanabe, et al, 2005) However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species (Takeuchi,Watanabe, et al, 2005). Again, this can also be considered another example demonstrating the flexibility of ''S.haemolyticus'' genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (Tristan,Lina, et al, 2006). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (Tristan,Lina, et al, 2006). ''S.haemolyticus'' survive successfully on the drier regions of the body (Tristan,Lina, et al, 2006), while it can also be found frequently in human blood cultures (Takeuchi,Watanabe, et al, 2005). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI. The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (Tristan,Lina, et al, 2006). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human (Watson,Yaguchi, et al, 1988).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (Brooks,Butel and Morse, 2001). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (Brooks,Butel and Morse, 2001). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci. (Brooks,Butel and Morse, 2001) ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints. (Tristan,Lina, et al, 2006) In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death. (Falcone,Campanile, et al, 2007) Common clinical symptoms of ''S.haemolyticus'' are fever and an increase in white blood cell population (leukocytosis) (Falcone,Campanile, et al, 2007). <br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (Novick, 2006). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but are now discovered in the other non-aureus, coagulast-negative staphylococci as well (Tristan,Lina, et al, 2006). In one studies published in 1994, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro (Molnar,Hevessy, et al, 1994). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species.<br />
<br />
''S.haemolyticus''’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (''Staphylococcus lugdunensis synergistic hemolysin'') in ''S.lugdunensis'', the latter of which shows significant similarities in structure with GGI. (Tristan,Lina, et al, 2006) These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
''Staphylococcus haemolyticus'', together with its related staphylococci like ''S.aureus'' and ''S.epidermis'', possesses a class of lipase enzymes, which involve in the hydrolysis process of long chain triacylglycerons. Thanks to the enzymes’ uniquely useful properties such as chain length selectivity and chiral selectivity, they are widely used in the industrial production and synthesis of fatty acids, fats, oils, esters and peptides (Oh,Kim, et al, 1999).<br />
<br />
A close relative of ''S.haemolyticus'', the coagulase-negative ''Staphylococcus xylosus'', has been used to construct a host-vector system that can express recombinant proteins on the surface of the bacterial cell (Hansson,Stahl, et al, 1992). It was the first time such system could be constructed in a Gram-positive species. The technique greatly facilitates the study of receptors, substrate-binding proteins and other antigenic determinants expressed on the surface of bacteria (Hansson,Stahl, et al, 1992).<br />
<br />
==Current Research==<br />
<br />
Enter summaries of the most recent research here--at least three required<br />
<br />
==References==<br />
1. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases]. Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species]. J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus]. Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics]. J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
<br />
Edited by Bao D. Truong, student of [mailto:ralarsen@ucsd.edu Rachel Larsen]</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=13450Staphylococcus haemolyticus2007-06-04T02:20:17Z<p>Bdtruong: /* Pathology */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood. (Tristan,Lina, et al, 2006) <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of infection (Falcone,Campanile, et al, 2007). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (Takeuchi,Watanabe, et al, 2005). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (Falcone,Campanile, et al, 2007). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time. (Falcone,Campanile, et al, 2007) Common antibiotics that are subject to resistance in S haemolyticus include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics (Falcone,Campanile, et al, 2007). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (Froggatt,Johnston, et al, 1989).<br />
<br />
In order to study the multi-drug resistant ability of Staphylococcus haemolyticus and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (Takeuchi,Watanabe, et al, 2005). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs). These sequences might be responsible for the frequent genomic arrangement observed in this organism.(Froggatt,Johnston, et al, 1989)<br />
<br />
==Genome structure==<br />
<br />
(Takeuchi,Watanabe, et al, 2005)<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp.<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome. However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability. On the other hand, the region is most likely responsible for the diversification of staphylococci species which enables them to successfully colonize and infect the human host. <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs). The ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance. <br />
While 6% of the orfs found in the more virulent S.aureus are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors. However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
(Billot-Klein,Gutmann, et al, 1996)<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains of ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin. This is an ability that is unique among staphylococci. The peptidoglycan structure of ''S.haemolyticus'' has been studied to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of ''S.haemolyticus'' is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the probable cross bridges structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus''. This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus''.<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids. Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline. (Takeuchi,Watanabe, et al, 2005)<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci (Takeuchi,Watanabe, et al, 2005) However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species (Takeuchi,Watanabe, et al, 2005). Again, this can also be considered another example demonstrating the flexibility of ''S.haemolyticus'' genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (Tristan,Lina, et al, 2006). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (Tristan,Lina, et al, 2006). ''S.haemolyticus'' survive successfully on the drier regions of the body (Tristan,Lina, et al, 2006), while it can also be found frequently in human blood cultures (Takeuchi,Watanabe, et al, 2005). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI. The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (Tristan,Lina, et al, 2006). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human (Watson,Yaguchi, et al, 1988).<br />
<br />
==Pathology==<br />
Staphylococci in general cause disease through their ability to spread widely in tissues ad their production of extracellular substances (Brooks,Butel and Morse, 2001). One example of such substances is coagulase, an enzyme-like protein produced by ''S.aureus'' that may deposit fibrin on the surface of the bacteria, altering their ingestion and destruction by phagocytic cells (Brooks,Butel and Morse, 2001). <br />
<br />
Traditionally, production of coagulase is considered to represent the invasive pathogenic potential among staphylococci. (Brooks,Butel and Morse, 2001) ''S.haemolyticus'', however, is a coagulase-negative species. Therefore, like other non-aureus staphylococci, its pathogenic characters were not well-studied until recently, as ''S.haemolyticus'' is emerging to be a major cause of nosocomial infections (infections acquired during treatment at a hospital for another disease). Reported cases of infections caused by ''S.haemolyticus'' include septicemia (dysfunction of organ systems resulting from immune response to a severe infection), peritonitis (inflammation of the serous membrane lining abdominal cavity), and infections of urinary tract, wound, bone and joints. (Tristan,Lina, et al, 2006) In rare cases, ''S.haemolyticus'' has also been reported to cause infective endocarditis, inflammation of the inner of the heart (the endocardium), which might lead to severe complications such as heart failure or death. (Falcone,Campanile, et al, 2007) Common clinical symptoms of ''S.haemolyticus'' are fever and an increase in white blood cell population (leukocytosis) (Falcone,Campanile, et al, 2007). <br />
<br />
Being the most common pathogen among staphylococci, virulent factors of ''S.aureus'' have been well-known. Important among them are different classes of enterotoxin (toxins released in lower-intestine, causing food poisoning), toxic shock syndrome toxin, and hemolysin (substances allow the bacteria to break down red-blood cells) (Novick, 2006). Some of these substances used to be considered to belong exclusively to ''S.aureus'', but are now discovered in the other non-aureus, coagulast-negative staphylococci as well (Tristan,Lina, et al, 2006). In one studies published in 1994, all strains of ''S.haemolyticus'' under investigation produced hemolysins in vitro (Molnar,Hevessy, et al, 1994). Investigators therefore suggested that hemolysins might be the important factor responsible for the high virulence of this staphylococcus species.<br />
<br />
''S.haemolyticus''’ GGI are related in function and some properties to other relative staphylococci virulent factor, such as delta-lysin in S.aureus and SLUSH (''Staphylococcus lugdunensis synergistic hemolysin'') in ''S.lugdunensis'', the latter of which shows significant similarities in structure with GGI. (Tristan,Lina, et al, 2006) These findings suggest a connection between pathogenesis pathways and virulent factors of common staphylococcal pathogens.<br />
<br />
==Application to Biotechnology==<br />
Does this organism produce any useful compounds or enzymes? What are they and how are they used?<br />
<br />
==Current Research==<br />
<br />
Enter summaries of the most recent research here--at least three required<br />
<br />
==References==<br />
1. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases]. Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species]. J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus]. Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics]. J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
<br />
Edited by Bao D. Truong, student of [mailto:ralarsen@ucsd.edu Rachel Larsen]</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=13446Staphylococcus haemolyticus2007-06-04T02:17:40Z<p>Bdtruong: /* Ecology */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood. (Tristan,Lina, et al, 2006) <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of infection (Falcone,Campanile, et al, 2007). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (Takeuchi,Watanabe, et al, 2005). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (Falcone,Campanile, et al, 2007). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time. (Falcone,Campanile, et al, 2007) Common antibiotics that are subject to resistance in S haemolyticus include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics (Falcone,Campanile, et al, 2007). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (Froggatt,Johnston, et al, 1989).<br />
<br />
In order to study the multi-drug resistant ability of Staphylococcus haemolyticus and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (Takeuchi,Watanabe, et al, 2005). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs). These sequences might be responsible for the frequent genomic arrangement observed in this organism.(Froggatt,Johnston, et al, 1989)<br />
<br />
==Genome structure==<br />
<br />
(Takeuchi,Watanabe, et al, 2005)<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp.<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome. However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability. On the other hand, the region is most likely responsible for the diversification of staphylococci species which enables them to successfully colonize and infect the human host. <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs). The ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance. <br />
While 6% of the orfs found in the more virulent S.aureus are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors. However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
(Billot-Klein,Gutmann, et al, 1996)<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains of ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin. This is an ability that is unique among staphylococci. The peptidoglycan structure of ''S.haemolyticus'' has been studied to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of ''S.haemolyticus'' is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the probable cross bridges structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus''. This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus''.<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids. Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline. (Takeuchi,Watanabe, et al, 2005)<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci (Takeuchi,Watanabe, et al, 2005) However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species (Takeuchi,Watanabe, et al, 2005). Again, this can also be considered another example demonstrating the flexibility of ''S.haemolyticus'' genome.<br />
<br />
==Ecology==<br />
<br />
''Staphylococcus haemolyticus'' can be found on the skins and in the bodies of a wide range of mammals, including prosimians, monkeys, domestic animals, and human (Tristan,Lina, et al, 2006). The most common natural habitats of the bacteria on human are in the axillae (underarm area), in the perineum (pubic area), and in the inguinal area (Tristan,Lina, et al, 2006). ''S.haemolyticus'' survive successfully on the drier regions of the body (Tristan,Lina, et al, 2006), while it can also be found frequently in human blood cultures (Takeuchi,Watanabe, et al, 2005). <br />
<br />
It has been known that ''S.haemolyticus'' produces gonococcal growth inhibitor, GGI. The substance was first discovered to cause cytoplasmic leakage in gonococcal cells and eventually lead to cell death (Tristan,Lina, et al, 2006). Remarkably, this substance can also lyse erythrocytes, especially those of horse and human (Watson,Yaguchi, et al, 1988).<br />
<br />
==Pathology==<br />
How does this organism cause disease? Human, animal, plant hosts? Virulence factors, as well as patient symptoms.<br />
<br />
==Application to Biotechnology==<br />
Does this organism produce any useful compounds or enzymes? What are they and how are they used?<br />
<br />
==Current Research==<br />
<br />
Enter summaries of the most recent research here--at least three required<br />
<br />
==References==<br />
1. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases]. Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species]. J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus]. Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics]. J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
<br />
Edited by Bao D. Truong, student of [mailto:ralarsen@ucsd.edu Rachel Larsen]</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=13445Staphylococcus haemolyticus2007-06-04T02:16:04Z<p>Bdtruong: /* Metabolism */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood. (Tristan,Lina, et al, 2006) <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of infection (Falcone,Campanile, et al, 2007). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (Takeuchi,Watanabe, et al, 2005). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (Falcone,Campanile, et al, 2007). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time. (Falcone,Campanile, et al, 2007) Common antibiotics that are subject to resistance in S haemolyticus include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics (Falcone,Campanile, et al, 2007). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (Froggatt,Johnston, et al, 1989).<br />
<br />
In order to study the multi-drug resistant ability of Staphylococcus haemolyticus and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (Takeuchi,Watanabe, et al, 2005). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs). These sequences might be responsible for the frequent genomic arrangement observed in this organism.(Froggatt,Johnston, et al, 1989)<br />
<br />
==Genome structure==<br />
<br />
(Takeuchi,Watanabe, et al, 2005)<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp.<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome. However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability. On the other hand, the region is most likely responsible for the diversification of staphylococci species which enables them to successfully colonize and infect the human host. <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs). The ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance. <br />
While 6% of the orfs found in the more virulent S.aureus are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors. However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
(Billot-Klein,Gutmann, et al, 1996)<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains of ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin. This is an ability that is unique among staphylococci. The peptidoglycan structure of ''S.haemolyticus'' has been studied to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of ''S.haemolyticus'' is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the probable cross bridges structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus''. This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus''.<br />
<br />
===Metabolism===<br />
Whole-genome sequencing of ''S.haemolyticus'' (strain JCSC1435) revealed some orfs encoding metabolic genes unique to the species, such as those involved in transport of ribose and ribitol or biosynthesis of essential components of nucleic acids and cell wall techoic acids. Thanks to these unique orfs, ''S.haemolyticus'' has a relative great biosynthetic capacity. Strain JCSC1435 only requires arginine for growth, while the ''S.aureus'' strain N315 requires the availability of many different amino acids: alanine, glycine, isoleucine, arginine, valine and proline. (Takeuchi,Watanabe, et al, 2005)<br />
<br />
''S.haemolyticus'' (strain JCSC1435) also possesses the ability to ferment mannitol, a metabolic character also found in some other “non-aureus” staphylococci (Takeuchi,Watanabe, et al, 2005) However, genetic analysis suggested that certain strains of ''S.haemolyticus'' might have gained this ability through horizontal gene transfer of the mannitol PTS locus from other bacterial species (Takeuchi,Watanabe, et al, 2005). Again, this can also be considered another example demonstrating the flexibility of ''S.haemolyticus'' genome.<br />
<br />
==Ecology==<br />
Describe any interactions with other organisms (included eukaryotes), contributions to the environment, effect on environment, etc.<br />
<br />
==Pathology==<br />
How does this organism cause disease? Human, animal, plant hosts? Virulence factors, as well as patient symptoms.<br />
<br />
==Application to Biotechnology==<br />
Does this organism produce any useful compounds or enzymes? What are they and how are they used?<br />
<br />
==Current Research==<br />
<br />
Enter summaries of the most recent research here--at least three required<br />
<br />
==References==<br />
1. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases]. Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species]. J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus]. Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics]. J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
<br />
Edited by Bao D. Truong, student of [mailto:ralarsen@ucsd.edu Rachel Larsen]</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=13443Staphylococcus haemolyticus2007-06-04T02:14:33Z<p>Bdtruong: /* Cell Wall */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood. (Tristan,Lina, et al, 2006) <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of infection (Falcone,Campanile, et al, 2007). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (Takeuchi,Watanabe, et al, 2005). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (Falcone,Campanile, et al, 2007). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time. (Falcone,Campanile, et al, 2007) Common antibiotics that are subject to resistance in S haemolyticus include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics (Falcone,Campanile, et al, 2007). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (Froggatt,Johnston, et al, 1989).<br />
<br />
In order to study the multi-drug resistant ability of Staphylococcus haemolyticus and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (Takeuchi,Watanabe, et al, 2005). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs). These sequences might be responsible for the frequent genomic arrangement observed in this organism.(Froggatt,Johnston, et al, 1989)<br />
<br />
==Genome structure==<br />
<br />
(Takeuchi,Watanabe, et al, 2005)<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp.<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome. However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability. On the other hand, the region is most likely responsible for the diversification of staphylococci species which enables them to successfully colonize and infect the human host. <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs). The ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance. <br />
While 6% of the orfs found in the more virulent S.aureus are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors. However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
<br />
(Billot-Klein,Gutmann, et al, 1996)<br />
<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains of ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin. This is an ability that is unique among staphylococci. The peptidoglycan structure of ''S.haemolyticus'' has been studied to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of ''S.haemolyticus'' is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the probable cross bridges structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus''. This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus''.<br />
<br />
===Metabolism===<br />
<br />
==Ecology==<br />
Describe any interactions with other organisms (included eukaryotes), contributions to the environment, effect on environment, etc.<br />
<br />
==Pathology==<br />
How does this organism cause disease? Human, animal, plant hosts? Virulence factors, as well as patient symptoms.<br />
<br />
==Application to Biotechnology==<br />
Does this organism produce any useful compounds or enzymes? What are they and how are they used?<br />
<br />
==Current Research==<br />
<br />
Enter summaries of the most recent research here--at least three required<br />
<br />
==References==<br />
1. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases]. Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species]. J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus]. Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics]. J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
<br />
Edited by Bao D. Truong, student of [mailto:ralarsen@ucsd.edu Rachel Larsen]</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=13442Staphylococcus haemolyticus2007-06-04T02:13:49Z<p>Bdtruong: /* Cell Wall */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood. (Tristan,Lina, et al, 2006) <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of infection (Falcone,Campanile, et al, 2007). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (Takeuchi,Watanabe, et al, 2005). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (Falcone,Campanile, et al, 2007). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time. (Falcone,Campanile, et al, 2007) Common antibiotics that are subject to resistance in S haemolyticus include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics (Falcone,Campanile, et al, 2007). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (Froggatt,Johnston, et al, 1989).<br />
<br />
In order to study the multi-drug resistant ability of Staphylococcus haemolyticus and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (Takeuchi,Watanabe, et al, 2005). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs). These sequences might be responsible for the frequent genomic arrangement observed in this organism.(Froggatt,Johnston, et al, 1989)<br />
<br />
==Genome structure==<br />
<br />
(Takeuchi,Watanabe, et al, 2005)<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp.<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome. However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability. On the other hand, the region is most likely responsible for the diversification of staphylococci species which enables them to successfully colonize and infect the human host. <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs). The ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance. <br />
While 6% of the orfs found in the more virulent S.aureus are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors. However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics that interfere with the peptidoglycan biosynthesis process. However, some strains of ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin. This is an ability that is unique among staphylococci. The peptidoglycan structure of ''S.haemolyticus'' has been studied to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of ''S.haemolyticus'' is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. In the resistant strains, studies have found cross bridges that contain an additional serine in place of glycine (so the probable cross bridges structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2). Furthermore, the presense of a novel cystoplasmic peptidoglycan precursor, UDP-muramyl-tetrapeptide-D-lactate, has been detected in strains of ''S.haemolyticus''. This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in ''S.haemolyticus''.<br />
<br />
===Metabolism===<br />
<br />
==Ecology==<br />
Describe any interactions with other organisms (included eukaryotes), contributions to the environment, effect on environment, etc.<br />
<br />
==Pathology==<br />
How does this organism cause disease? Human, animal, plant hosts? Virulence factors, as well as patient symptoms.<br />
<br />
==Application to Biotechnology==<br />
Does this organism produce any useful compounds or enzymes? What are they and how are they used?<br />
<br />
==Current Research==<br />
<br />
Enter summaries of the most recent research here--at least three required<br />
<br />
==References==<br />
1. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases]. Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species]. J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus]. Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics]. J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
<br />
Edited by Bao D. Truong, student of [mailto:ralarsen@ucsd.edu Rachel Larsen]</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=13440Staphylococcus haemolyticus2007-06-04T02:11:42Z<p>Bdtruong: /* Genome structure */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood. (Tristan,Lina, et al, 2006) <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of infection (Falcone,Campanile, et al, 2007). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (Takeuchi,Watanabe, et al, 2005). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (Falcone,Campanile, et al, 2007). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time. (Falcone,Campanile, et al, 2007) Common antibiotics that are subject to resistance in S haemolyticus include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics (Falcone,Campanile, et al, 2007). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (Froggatt,Johnston, et al, 1989).<br />
<br />
In order to study the multi-drug resistant ability of Staphylococcus haemolyticus and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (Takeuchi,Watanabe, et al, 2005). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs). These sequences might be responsible for the frequent genomic arrangement observed in this organism.(Froggatt,Johnston, et al, 1989)<br />
<br />
==Genome structure==<br />
<br />
(Takeuchi,Watanabe, et al, 2005)<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp.<br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome. However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (oriC), is called “oriC environ”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability. On the other hand, the region is most likely responsible for the diversification of staphylococci species which enables them to successfully colonize and infect the human host. <br />
<br />
Besides having the oriC environ region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs). The ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance. <br />
While 6% of the orfs found in the more virulent S.aureus are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors. However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics interfering with the peptidoglycan biosynthesis process. However, some strains of ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin. This is an ability that is unique among staphylococci. The structure of ''S.haemolyticus'' cell wall has been studied to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of ''S.haemolyticus'' is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. Studies have found in resistant strains cross bridges that contain an additional serine in place of glycine (so the probable cross bridges structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2). Furthermore, the presence of a novel cystoplasmic peptidoglycan precursor, ''UDP-muramyl-tetrapeptide-D-lactate'', has been detected in strains of ''S.haemolyticus''. This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in S.haemolyticus.(5)<br />
<br />
===Metabolism===<br />
<br />
==Ecology==<br />
Describe any interactions with other organisms (included eukaryotes), contributions to the environment, effect on environment, etc.<br />
<br />
==Pathology==<br />
How does this organism cause disease? Human, animal, plant hosts? Virulence factors, as well as patient symptoms.<br />
<br />
==Application to Biotechnology==<br />
Does this organism produce any useful compounds or enzymes? What are they and how are they used?<br />
<br />
==Current Research==<br />
<br />
Enter summaries of the most recent research here--at least three required<br />
<br />
==References==<br />
1. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases]. Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species]. J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus]. Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics]. J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
<br />
Edited by Bao D. Truong, student of [mailto:ralarsen@ucsd.edu Rachel Larsen]</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=13434Staphylococcus haemolyticus2007-06-04T02:04:46Z<p>Bdtruong: /* Description and significance */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus Staphylococcus. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood. (Tristan,Lina, et al, 2006) <br />
<br />
Lacking coagulase, an enzyme-like protein that was traditionally associated with virulent potential of staphylococci, coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive ''Staphylococcus aureus''. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of infection (Falcone,Campanile, et al, 2007). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (Takeuchi,Watanabe, et al, 2005). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (Falcone,Campanile, et al, 2007). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time. (Falcone,Campanile, et al, 2007) Common antibiotics that are subject to resistance in S haemolyticus include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics (Falcone,Campanile, et al, 2007). The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin) (Froggatt,Johnston, et al, 1989).<br />
<br />
In order to study the multi-drug resistant ability of Staphylococcus haemolyticus and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (Takeuchi,Watanabe, et al, 2005). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs). These sequences might be responsible for the frequent genomic arrangement observed in this organism.(Froggatt,Johnston, et al, 1989)<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp. <br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome. However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (''oriC''), is called “''oriC environ''”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability. On the other hand, the region is most likely responsible for the diversification of staphylococci species which enables them to successfully colonize and infect the human host.<br />
<br />
Besides having the ''oriC environ'' region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs). The ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance.<br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors. However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.(3)<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics interfering with the peptidoglycan biosynthesis process. However, some strains of ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin. This is an ability that is unique among staphylococci. The structure of ''S.haemolyticus'' cell wall has been studied to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of ''S.haemolyticus'' is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. Studies have found in resistant strains cross bridges that contain an additional serine in place of glycine (so the probable cross bridges structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2). Furthermore, the presence of a novel cystoplasmic peptidoglycan precursor, ''UDP-muramyl-tetrapeptide-D-lactate'', has been detected in strains of ''S.haemolyticus''. This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in S.haemolyticus.(5)<br />
<br />
===Metabolism===<br />
<br />
==Ecology==<br />
Describe any interactions with other organisms (included eukaryotes), contributions to the environment, effect on environment, etc.<br />
<br />
==Pathology==<br />
How does this organism cause disease? Human, animal, plant hosts? Virulence factors, as well as patient symptoms.<br />
<br />
==Application to Biotechnology==<br />
Does this organism produce any useful compounds or enzymes? What are they and how are they used?<br />
<br />
==Current Research==<br />
<br />
Enter summaries of the most recent research here--at least three required<br />
<br />
==References==<br />
1. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases]. Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species]. J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus]. Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics]. J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
<br />
Edited by Bao D. Truong, student of [mailto:ralarsen@ucsd.edu Rachel Larsen]</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=10614Staphylococcus haemolyticus2007-05-03T19:43:25Z<p>Bdtruong: /* References */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus ''Staphylococcus''. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood.(1) <br />
<br />
Coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive Staphylococcus aureus. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of infection (2). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (3). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (2). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time.(2,4) Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics.(2) The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin)(4).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (3). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs). These sequences might be responsible for the frequent genomic arrangement observed in this organism.(4)<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp. <br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome. However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (''oriC''), is called “''oriC environ''”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability. On the other hand, the region is most likely responsible for the diversification of staphylococci species which enables them to successfully colonize and infect the human host.<br />
<br />
Besides having the ''oriC environ'' region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs). The ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance.<br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors. However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.(3)<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics interfering with the peptidoglycan biosynthesis process. However, some strains of ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin. This is an ability that is unique among staphylococci. The structure of ''S.haemolyticus'' cell wall has been studied to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of ''S.haemolyticus'' is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. Studies have found in resistant strains cross bridges that contain an additional serine in place of glycine (so the probable cross bridges structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2). Furthermore, the presence of a novel cystoplasmic peptidoglycan precursor, ''UDP-muramyl-tetrapeptide-D-lactate'', has been detected in strains of ''S.haemolyticus''. This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in S.haemolyticus.(5)<br />
<br />
===Metabolism===<br />
<br />
==Ecology==<br />
Describe any interactions with other organisms (included eukaryotes), contributions to the environment, effect on environment, etc.<br />
<br />
==Pathology==<br />
How does this organism cause disease? Human, animal, plant hosts? Virulence factors, as well as patient symptoms.<br />
<br />
==Application to Biotechnology==<br />
Does this organism produce any useful compounds or enzymes? What are they and how are they used?<br />
<br />
==Current Research==<br />
<br />
Enter summaries of the most recent research here--at least three required<br />
<br />
==References==<br />
1. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
2. Falcone, M. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=17141458&dopt=abstract Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases]. Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species]. J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=2729941&dopt=abstract Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus]. Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics]. J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
<br />
Edited by Bao D. Truong, student of [mailto:ralarsen@ucsd.edu Rachel Larsen]</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=10612Staphylococcus haemolyticus2007-05-03T19:41:44Z<p>Bdtruong: /* References */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus ''Staphylococcus''. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood.(1) <br />
<br />
Coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive Staphylococcus aureus. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of infection (2). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (3). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (2). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time.(2,4) Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics.(2) The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin)(4).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (3). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs). These sequences might be responsible for the frequent genomic arrangement observed in this organism.(4)<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp. <br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome. However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (''oriC''), is called “''oriC environ''”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability. On the other hand, the region is most likely responsible for the diversification of staphylococci species which enables them to successfully colonize and infect the human host.<br />
<br />
Besides having the ''oriC environ'' region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs). The ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance.<br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors. However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.(3)<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics interfering with the peptidoglycan biosynthesis process. However, some strains of ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin. This is an ability that is unique among staphylococci. The structure of ''S.haemolyticus'' cell wall has been studied to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of ''S.haemolyticus'' is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. Studies have found in resistant strains cross bridges that contain an additional serine in place of glycine (so the probable cross bridges structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2). Furthermore, the presence of a novel cystoplasmic peptidoglycan precursor, ''UDP-muramyl-tetrapeptide-D-lactate'', has been detected in strains of ''S.haemolyticus''. This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in S.haemolyticus.(5)<br />
<br />
===Metabolism===<br />
<br />
==Ecology==<br />
Describe any interactions with other organisms (included eukaryotes), contributions to the environment, effect on environment, etc.<br />
<br />
==Pathology==<br />
How does this organism cause disease? Human, animal, plant hosts? Virulence factors, as well as patient symptoms.<br />
<br />
==Application to Biotechnology==<br />
Does this organism produce any useful compounds or enzymes? What are they and how are they used?<br />
<br />
==Current Research==<br />
<br />
Enter summaries of the most recent research here--at least three required<br />
<br />
==References==<br />
1. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
2. Falcone, M. et al. Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases. Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=16237012&dopt=abstract Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species]. J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics]. J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
<br />
Edited by Bao D. Truong, student of [mailto:ralarsen@ucsd.edu Rachel Larsen]</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=10611Staphylococcus haemolyticus2007-05-03T19:40:51Z<p>Bdtruong: /* Description and significance */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus ''Staphylococcus''. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood.(1) <br />
<br />
Coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive Staphylococcus aureus. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of infection (2). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (3). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (2). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time.(2,4) Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics.(2) The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin)(4).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435 (3). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs). These sequences might be responsible for the frequent genomic arrangement observed in this organism.(4)<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp. <br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome. However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (''oriC''), is called “''oriC environ''”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability. On the other hand, the region is most likely responsible for the diversification of staphylococci species which enables them to successfully colonize and infect the human host.<br />
<br />
Besides having the ''oriC environ'' region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs). The ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance.<br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors. However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.(3)<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics interfering with the peptidoglycan biosynthesis process. However, some strains of ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin. This is an ability that is unique among staphylococci. The structure of ''S.haemolyticus'' cell wall has been studied to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of ''S.haemolyticus'' is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. Studies have found in resistant strains cross bridges that contain an additional serine in place of glycine (so the probable cross bridges structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2). Furthermore, the presence of a novel cystoplasmic peptidoglycan precursor, ''UDP-muramyl-tetrapeptide-D-lactate'', has been detected in strains of ''S.haemolyticus''. This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in S.haemolyticus.(5)<br />
<br />
===Metabolism===<br />
<br />
==Ecology==<br />
Describe any interactions with other organisms (included eukaryotes), contributions to the environment, effect on environment, etc.<br />
<br />
==Pathology==<br />
How does this organism cause disease? Human, animal, plant hosts? Virulence factors, as well as patient symptoms.<br />
<br />
==Application to Biotechnology==<br />
Does this organism produce any useful compounds or enzymes? What are they and how are they used?<br />
<br />
==Current Research==<br />
<br />
Enter summaries of the most recent research here--at least three required<br />
<br />
==References==<br />
1. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
2. Falcone, M. et al. Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases. Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species. J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics]. J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
<br />
Edited by Bao D. Truong, student of [mailto:ralarsen@ucsd.edu Rachel Larsen]</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=10609Staphylococcus haemolyticus2007-05-03T19:36:43Z<p>Bdtruong: /* Genome structure */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus ''Staphylococcus''. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood.(1) <br />
<br />
Coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive Staphylococcus aureus. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of infection (2). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (3). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (2). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time.(2) Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics.(2) The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin)(4).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435(4). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs). These sequences might be responsible for the frequent genomic arrangement observed in this organism.(4)<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp. <br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome. However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (''oriC''), is called “''oriC environ''”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability. On the other hand, the region is most likely responsible for the diversification of staphylococci species which enables them to successfully colonize and infect the human host.<br />
<br />
Besides having the ''oriC environ'' region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs). The ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance.<br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors. However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.(3)<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics interfering with the peptidoglycan biosynthesis process. However, some strains of ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin. This is an ability that is unique among staphylococci. The structure of ''S.haemolyticus'' cell wall has been studied to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of ''S.haemolyticus'' is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. Studies have found in resistant strains cross bridges that contain an additional serine in place of glycine (so the probable cross bridges structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2). Furthermore, the presence of a novel cystoplasmic peptidoglycan precursor, ''UDP-muramyl-tetrapeptide-D-lactate'', has been detected in strains of ''S.haemolyticus''. This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in S.haemolyticus.(5)<br />
<br />
===Metabolism===<br />
<br />
==Ecology==<br />
Describe any interactions with other organisms (included eukaryotes), contributions to the environment, effect on environment, etc.<br />
<br />
==Pathology==<br />
How does this organism cause disease? Human, animal, plant hosts? Virulence factors, as well as patient symptoms.<br />
<br />
==Application to Biotechnology==<br />
Does this organism produce any useful compounds or enzymes? What are they and how are they used?<br />
<br />
==Current Research==<br />
<br />
Enter summaries of the most recent research here--at least three required<br />
<br />
==References==<br />
1. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
2. Falcone, M. et al. Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases. Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species. J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics]. J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
<br />
Edited by Bao D. Truong, student of [mailto:ralarsen@ucsd.edu Rachel Larsen]</div>Bdtruonghttps://microbewiki.kenyon.edu/index.php?title=Staphylococcus_haemolyticus&diff=10608Staphylococcus haemolyticus2007-05-03T19:36:06Z<p>Bdtruong: /* References */</p>
<hr />
<div>{{Biorealm Genus}}<br />
<br />
==Classification==<br />
<br />
===Higher order taxa===<br />
<br />
Bacteria; Firmicutes; Bacilli; Bacillales; Staphylococcaceae; [[Staphylococcus]]<br />
<br />
===Species===<br />
<br />
{|<br />
| height="10" bgcolor="#FFDF95" |<br />
'''NCBI: [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=2&lvl=3&lin=f&keep=1&srchmode=1&unlock Taxonomy]'''<br />
|}<br />
<br />
''Staphylococcus haemolyticus''<br />
<br />
==Description and significance==<br />
<br />
''Staphylococus haemolyticus'' is a coagulase-negative member of the genus ''Staphylococcus''. The bacteria can be found on normal human skin flora and can be isolated from axillae, perineum, and ingunial areas of humans. ''S.haemolyticus'' is also the second most common coagulase-negative staphylocci presenting in human blood.(1) <br />
<br />
Coagulase-negative staphylococci are usually considered low-virulent pathogens comparing to the well-known pathogenic coagulase-positive Staphylococcus aureus. However, recent studies indicate that coagulase-negative staphylococci have emerged as a major cause of infection (2). ''Staphylococcus haemolyticus'' itself is also a remarkable opportunistic baterial pathogen that is well-known for its highly antibiotic-resistant phenotype (3). The bacteria can cause meningitis, skin or soft tissue infections, prosthetic join infections, or bacteremia (2). The ability of the bacteria to simultaneously resist against multiple types of antibiotic has been observed and studied for a long time.(2) Common antibiotics that are subject to resistance in ''S haemolyticus'' include methicillin, gentamycin, erythormycin, and uniquely among staphylococci, glycopeptide antibiotics.(2) The resistance genes for each type of anitbiotic can be located on the chromosome (methicillin), on the plasmids (erythromycin) or on both chromosome and plasmids (gentamycin)(4).<br />
<br />
In order to study the multi-drug resistant ability of ''Staphylococcus haemolyticus'' and its pathogenic characters, researchers sequenced the whole genome of one strain, JCSC1435(4). Beside the bacteria’s antibiotic resistance genes, the study of the sequence also revealed a surprising number of homologous insertion sequences (ISs). These sequences might be responsible for the frequent genomic arrangement observed in this organism.(4)<br />
<br />
==Genome structure==<br />
<br />
The genome of ''Staphylococcus haemolyticus'' (strain JCSC1435) includes a circular chromosome of 2,685,015 bp and 3 plasmids of 2,300 bp, 2,366 bp and 8,180 bp. <br />
<br />
Comparative genomic analysis has revealed significant similarities between the genomes of ''S.haemolyticus'' and those of the other two well-known staphylococci, ''S.aureus'' and ''S.epidermis''. Beside the comparable genome sizes, a large proportion of open reading frames (orfs) are conserved both in the sequences and in their order on the chromosome. However, the study also found a region on the chromosome that is unique for each of the 3 organisms. This region, which locate near the chromosome origin of replication (''oriC''), is called “''oriC environ''”. As most of the region could be deleted without affecting growth, it can be concluded that the oriC environ region does not contain genes essential for bacterial viability. On the other hand, the region is most likely responsible for the diversification of staphylococci species which enables them to successfully colonize and infect the human host.<br />
<br />
Besides having the ''oriC environ'' region where rearrangements of the genome can take place frequently, ''S.haemolyticus'' also possess a surprisingly large number of insertion sequences (ISs). The ISs can either inactivate a gene by direct integration into the open reading frame or activate a gene by providing the gene with a potent promoter. By changing the content of the genome, the IS elements might contribute to the innate ability of the bacteria to acquire drug resistance.<br />
<br />
While 6% of the orfs found in the more virulent ''S.aureus'' are pathogenic factors, only 2% of those found in ''S.haemolyticus'' are pathogenic factors. However, it is the ability of ''S.haemolyticus'' to alter its genome content and to acquire resistance to antibotics that makes the species a remarkable and hard-to-control opportunity pathogen.(4)<br />
<br />
==Cell structure and metabolism==<br />
===Cell Wall===<br />
As a gram-positive species like other staphylococci, ''S.haemolyticus'' has a thick peptidoglycan wall outside of its membrane and therefore can be targeted by antibiotics interfering with the peptidoglycan biosynthesis process. However, some strains of ''S.haemolyticus'' have developed resistance to glycopeptide antibiotics such as teicoplanin and vancomycin. This is an ability that is unique among staphylococci. The structure of ''S.haemolyticus'' cell wall has been studied to find out the factors responsible to this special resistance.<br />
<br />
Like that of other staphylococci, the peptidoglycan of ''S.haemolyticus'' is highly cross-linked. The predominant cross bridges are COOH-Gly-Gly-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Gly-Gly-NH2. Studies have found in resistant strains cross bridges that contain an additional serine in place of glycine (so the probable cross bridges structures are COOH-Gly-Ser-Ser-Gly-Gly-NH2 and COOH-Ala-Gly-Ser-Ser-Gly-NH2). Furthermore, the presence of a novel cystoplasmic peptidoglycan precursor, ''UDP-muramyl-tetrapeptide-D-lactate'', has been detected in strains of ''S.haemolyticus''. This precursor and the alterations of cross bridges are believed to interfere with the cooperative binding of glycopeptide antibiotics like vancomycin and teicoplanin to their targets in S.haemolyticus.(5)<br />
<br />
===Metabolism===<br />
<br />
==Ecology==<br />
Describe any interactions with other organisms (included eukaryotes), contributions to the environment, effect on environment, etc.<br />
<br />
==Pathology==<br />
How does this organism cause disease? Human, animal, plant hosts? Virulence factors, as well as patient symptoms.<br />
<br />
==Application to Biotechnology==<br />
Does this organism produce any useful compounds or enzymes? What are they and how are they used?<br />
<br />
==Current Research==<br />
<br />
Enter summaries of the most recent research here--at least three required<br />
<br />
==References==<br />
1. Brooks, G., Butel, J. & Morse, S. in Medical Microbiology 197-202 (McGraw-Hill, New York, 2001).<br />
<br />
2. Falcone, M. et al. Staphylococcus haemolyticus endocarditis: clinical and microbiologic analysis of 4 cases. Diagn. Microbiol. Infect. Dis. 57, 325-331 (2007).<br />
<br />
3. Takeuchi, F. et al. Whole-genome sequencing of staphylococcus haemolyticus uncovers the extreme plasticity of its genome and the evolution of human-colonizing staphylococcal species. J. Bacteriol. 187, 7292-7308 (2005).<br />
<br />
4. Froggatt, J. W., Johnston, J. L., Galetto, D. W. & Archer, G. L. Antimicrobial resistance in nosocomial isolates of Staphylococcus haemolyticus. Antimicrob. Agents Chemother. 33, 460-466 (1989).<br />
<br />
5. Billot-Klein, D. et al. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=retrieve&db=pubmed&list_uids=8755902&dopt=abstract Peptidoglycan synthesis and structure in Staphylococcus haemolyticus expressing increasing levels of resistance to glycopeptide antibiotics]. J. Bacteriol. 178, 4696-4703 (1996).<br />
<br />
<br />
Edited by Bao D. Truong, student of [mailto:ralarsen@ucsd.edu Rachel Larsen]</div>Bdtruong