Bdellovibrio bacteriovorus: Difference between revisions
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''Bdellovibrios'' were discovered by Stolp and Petzhold in 1962, in an attempt to isolate bacteriophage from soil samples. Stolp and Petzhold observed unique plaques in their samples that took several days to develop and continued to grow for over a week, instead of plaques caused by bacteriophages that would appear within hours. A closer inspection of the plaques under a light microscope revealed cells that were small, highly motile, and vibrio-shaped. These cells were ''Bdellovibrios''. | ''Bdellovibrios'' were discovered by Stolp and Petzhold in 1962, in an attempt to isolate bacteriophage from soil samples. Stolp and Petzhold observed unique plaques in their samples that took several days to develop and continued to grow for over a week, instead of plaques caused by bacteriophages that would appear within hours. A closer inspection of the plaques under a light microscope revealed cells that were small, highly motile, and vibrio-shaped. These cells were ''Bdellovibrios''. | ||
After the discovery of ''Bdellovibrios'' further observations revealed many interesting and unique properties. One property that makes ''Bdellovibrios'' interesting is that it is a parasite to other Gram-negative bacterias. ''Bdellovibrios'' have biphasic life-cycles that include an attack phase, and a free living and mobile phase.(2) The attack phase is when it attaches to another Gram-negative bacteria and imbeds itself into its periplasm, it then procedes to grow and replicate itself by degrading the host bacterium from the inside out.(1) In the free living and mobile phase, ''Bdellovibrios'' move about in search of host or prey bacteria so it can intiate attack phase again.(2) Each of these phases are of interest to researchers because | After the discovery of ''Bdellovibrios'' further observations revealed many interesting and unique properties. One property that makes ''Bdellovibrios'' interesting is that it is a parasite to other Gram-negative bacterias. ''Bdellovibrios'' have biphasic life-cycles that include an attack phase, and a free living and mobile phase.(2) The attack phase is when it attaches to another Gram-negative bacteria and imbeds itself into its periplasm, it then procedes to grow and replicate itself by degrading the host bacterium from the inside out.(1) In the free living and mobile phase, ''Bdellovibrios'' move about in search of host or prey bacteria so it can intiate attack phase again.(2) Each of these phases are of interest to researchers because they reveal unique cell-cell interactions and unusual cell metabolism.(1) | ||
''Bdellovibrio bacteriovorus'' is a small, curved, and highly motile Gram-negative bacteria approximately 0.2 to 0.5μm wide and 0.5 to 2.5μm long.(3) It has been found in many environments that include soil, sewage, and other terrestial and aquatic habitats. ''B. bacteriovorus'' has been observed to only attack Gram-negative bacteria which includes many plant, animal and human pathogens making it an execellent candidate as a biocontrol agent.(2) The study of its degradative enzymes and host targeting system has shed some insight in possible designs for new antimicrobial agents.(3) | ''Bdellovibrio bacteriovorus'' is a small, curved, and highly motile Gram-negative bacteria approximately 0.2 to 0.5μm wide and 0.5 to 2.5μm long.(3) It has been found in many environments that include soil, sewage, and other terrestial and aquatic habitats. ''B. bacteriovorus'' has been observed to only attack Gram-negative bacteria which includes many plant, animal and human pathogens making it an execellent candidate as a biocontrol agent.(2) The study of its degradative enzymes and host targeting system has shed some insight in possible designs for new antimicrobial agents.(3) | ||
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==Genome structure== | ==Genome structure== | ||
The sequencing of the ''B. bacteriovorus HD100'' genome was completed on 01/31/2004. The complete genome consists of a single circular chromosome that is 3,782,950 nucleotides long. The entire genome has a 50% GC content and contains 3629 genes which | The sequencing of the ''B. bacteriovorus HD100'' genome was completed on 01/31/2004. The complete genome consists of a single circular chromosome that is 3,782,950 nucleotides long. The entire genome has a 50% GC content and contains 3629 genes which code for 3587 proteins and 42 structural RNAs.(4) | ||
Of the large number of proteins that the ''B. bacteriovorus'' genome encodes, many of them are degradative and lytic enzymes(2). These enzymes are necessary for ''B. bacteriovorus'' to breakdown its host or prey cell. Some of these enzymes | Of the large number of proteins that the ''B. bacteriovorus'' genome encodes, many of them are degradative and lytic enzymes(2). These enzymes are necessary for ''B. bacteriovorus'' to breakdown its host or prey cell. Some of these enzymes need to be secreted out of ''B. bacteriovorus'' and into its prey's cell, while degradative products needs to be transported into the ''B. bacteriovorus'' from the prey cell cytoplasm. Therefore, the genome of ''B. bacteriovorus'' also includes a large amount of transport proteins. In fact, ''B. bacteriovorus'' has the potential protein secretory capabilities of at least five types of outer membrane secretion systems and four types of inner membrane secretion systems.(5) | ||
Other than encoding for degradative enzymes and transport systems, the chromosome also contains multiple biosynthetic gene clusters that are necessary for the formation of the flagella and pilus. These gene clusters are essential for the motility ''B. bacteriovorus'', which is important to its life cycle because it directly relates to its ability to find and approach the prey cell. However, the organism sheds its flagellum once it makes irreversible contact with its prey cell suggesting that the flagellum is not required for it to grow.(2) | Other than encoding for degradative enzymes and transport systems, the chromosome also contains multiple biosynthetic gene clusters that are necessary for the formation of the flagella and pilus. These gene clusters are essential for the motility ''B. bacteriovorus'', which is important to its life cycle because it directly relates to its ability to find and approach the prey cell. However, the organism sheds its flagellum once it makes irreversible contact with its prey cell suggesting that the flagellum is not required for it to grow.(2) | ||
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The general cell structure of ''B. bacteriovorus'' is small, curved, and highly motile Gram-negative bacteria approximately 0.2 to 0.5 μm wide and 0.5 to 2.5 μm long.(3) It has a flagellum that is membrane bound, allowing it to swim up to 100 body length per second while it looks for its prey.(9) | The general cell structure of ''B. bacteriovorus'' is small, curved, and highly motile Gram-negative bacteria approximately 0.2 to 0.5 μm wide and 0.5 to 2.5 μm long.(3) It has a flagellum that is membrane bound, allowing it to swim up to 100 body length per second while it looks for its prey.(9) | ||
The unique biphasic life-cycle of ''B. bacteriovorus'' is directly related to its metabolism. In the free-living or mobile phase, ''B. bacteriovorus'' actively seeks other Gram-negative bacteria by chemotaxic means. During the mobile phase, most of the metabolic activity is | The unique biphasic life-cycle of ''B. bacteriovorus'' is directly related to its metabolism. In the free-living or mobile phase, ''B. bacteriovorus'' actively seeks other Gram-negative bacteria by chemotaxic means. During the mobile phase, most of the metabolic activity is involved in motility and prey detection.(7) In the attack phase, ''B. bacteriovorus'' burrows itself into the periplasm of the prey cell and begins to degrade the prey cell from the inside out. A large variety of metabolic activities occur during this phase.(2) | ||
In the attack phase, ''B. bacteriovorus'' begins by attaching itself to the outer membrane of the prey cell via hook-like structures.(8) It then proceeds to secrete enzymes which | In the attack phase, ''B. bacteriovorus'' begins by attaching itself to the outer membrane of the prey cell via hook-like structures.(8) It then proceeds to secrete enzymes which break down a large enough hole in the outer membrane for ''B. bacteriovorus'' to enter. After entry into the prey cell, ''B. bacteriovorus'' sheds its flagellum and seals the hole in the outer membrane of the prey cell.(1) ''B. bacteriovorus'' then secretes degradative enzymes that act on the inner membrane of the prey cell allowing it to access the prey's cytoplasm. The degradation of the inner membrane causes the prey cell to become a bdelloplast, host cell of ''Bdellovibrios'' with a stable spherical structure, and also the destruction of the host cell's ability to generate energy.(1) At this point, ''B. bacteriovorus'' degrades and metabolizes the macromolecules and nutrients from the host's cytoplamic material allowing it to grow and begin DNA replication. ''B. bacteriovorus'' subsequently becomes a long filament and then fragments into individual mobile progenies which will lyse the remaining bdelloplast.(1) | ||
The metabolism of ''B. bacteriovorus'' has many interesting areas. One of these areas is that ''B. bacteriovorus'' can survive while lacking numerous amino acid biosynthetic pathways. The reason for this is that ''B. bacteriovorus'' utilizes the metabolites of its prey for its own protein synthesis.(2) ''B. bacteriovorus'' is very efficient at utilizing the DNA, RNA, and other pre-existing precursors from the host cell. It can break down or rearrange these precursors for the biosynthesis of homologous polymers. The uptake of nucleoside monophosphates and glycerol phoshphates greatly reduces the amount of energy that ''B. bacteriovorus'' has to produce during nucleic acid synthesis and lipid synthesis. There | The metabolism of ''B. bacteriovorus'' has many interesting areas. One of these areas is that ''B. bacteriovorus'' can survive while lacking numerous amino acid biosynthetic pathways. The reason for this is that ''B. bacteriovorus'' utilizes the metabolites of its prey for its own protein synthesis.(2) ''B. bacteriovorus'' is very efficient at utilizing the DNA, RNA, and other pre-existing precursors from the host cell. It can break down or rearrange these precursors for the biosynthesis of homologous polymers. The uptake of nucleoside monophosphates and glycerol phoshphates greatly reduces the amount of energy that ''B. bacteriovorus'' has to produce during nucleic acid synthesis and lipid synthesis. There has also been evidence that ''B. bacteriovorus'' incorporates intact fatty acid molecules from its host. Experiments have shown that the energy efficiency of ''B. bacteriovorus'' can be 0.8 to 1.5 times more efficient than bacteria growing in rich media.(6) | ||
==Ecology== | ==Ecology== | ||
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''B. bacteriovorus'' seems to be pretty ubiquitous in nature and manmade habitats. They have been found in soil samples, rhizosphere of plant roots, rivers, oceans, sewage, intestines and feces of birds and mammals, and even in oyster shells and the gills of crabs.(9) ''B. bacteriovorus'' are able to thrive in almost any habitat, the general requirements are that there needs to be oxygen and some other Gram-negative bacteria present in its environment. Its optimal temperature is between 28-30C, making ''B. bacteriovorus'' a mesophile.(2) | ''B. bacteriovorus'' seems to be pretty ubiquitous in nature and manmade habitats. They have been found in soil samples, rhizosphere of plant roots, rivers, oceans, sewage, intestines and feces of birds and mammals, and even in oyster shells and the gills of crabs.(9) ''B. bacteriovorus'' are able to thrive in almost any habitat, the general requirements are that there needs to be oxygen and some other Gram-negative bacteria present in its environment. Its optimal temperature is between 28-30C, making ''B. bacteriovorus'' a mesophile.(2) | ||
''B. bacteriovorus'' | ''B. bacteriovorus'' is somewhat tolerant to its environment. It can live in a range of salinity and is resistant to pollution of its environment.(2) During its attack phase or as bdelloplasts, ''B. bacteriovorus'' can tolerate dry spells and short periods of anaerobic environment. This makes ''B. bacteriovorus'' more tolerant to environmental changes than other permanantly aerobic bacteria.(9) | ||
''B. bacteriovorus'' has also been associated with biofilms. It seems that the rich environment of biofilms supply ''B. bacteriovorus'' with many possible host cells. Research has shown that ''B. bacteriovorus'' can significantly decrease ''E. Coli'' biofilm that is grown on stainless steel.(9) | ''B. bacteriovorus'' has also been associated with biofilms. It seems that the rich environment of biofilms supply ''B. bacteriovorus'' with many possible host cells. Research has shown that ''B. bacteriovorus'' can significantly decrease ''E. Coli'' biofilm that is grown on stainless steel.(9) | ||
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==Application to Biotechnology== | ==Application to Biotechnology== | ||
The major applications of ''B. bacteriovorus'' to biotechnology | The major applications of ''B. bacteriovorus'' to biotechnology seem to be its potential as a biocontrol agent. ''B. bacteriovorus'' has been used in certain water treatment plants to reduce the number of Gram-negative organisms in the water. Agriculturally, ''B. bacteriovorus'' has been used to limit or prevent the start and spread of plant pathogens with certain crops. The use of ''B. bacteriovorus'' in agriculture requires extensive knowledge of the crop and its ecology; ''B. bacteriovorus'' may attack growth promoting Gram-negative organisms needed for healthy crops such as rhizobacterias.(9) | ||
==Current Research== | ==Current Research== | ||
Currently, the main field of research with ''B. bacteriovorus'' is its high potential as a biocontrol agent. Many experiments are performed to test how it specifically | Currently, the main field of research with ''B. bacteriovorus'' is its high potential as a biocontrol agent. Many experiments are performed to test how it specifically recognizes its prey. There is also research on how ''B. bacteriovorus'' recognizes and prefers one prey over another.(10) Molecular research on the degradative enzymes of ''B. bacteriovorus'' and its targets provide insight to evolutionary successful point of attack to Gram-negative bacteria.(3) The ultimate goal in much of this research is to provide possible elements to create more specific antimicrobial agents. It is also highly possible that a slight change in its genetic code can allow ''B. bacteriovorus'' to truly become a ''living antibiotic''. | ||
===Prey Selection of ''B. bacteriovorus''=== | ===Prey Selection of ''B. bacteriovorus''=== | ||
When introduced to multiple preys, ''B. bacteriovorus'' preferentially lysed | When introduced to multiple preys, ''B. bacteriovorus'' preferentially lysed one prey over the others. This research showed that ''B. bacteriovorus'' may not randomly infect prey cells, but differentially prefer to infect one prey over another.(10) | ||
The identification and characterization of the genetic control of the bdellovibrio developmental cycle and its temporal gene expression | The identification and characterization of the genetic control of the bdellovibrio developmental cycle and its temporal gene expression will help identify which genes are responsible for prey selection and the proteins that these genes will encode.(11) | ||
==References== | ==References== | ||
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(10)Rogosky AM, Moak PL, Emmert EA. "Differential predation by Bdellovibrio bacteriovorus 109J." Current Microbiology, Feb 2006;52(2), p. 81-85 | (10)Rogosky AM, Moak PL, Emmert EA. "Differential predation by Bdellovibrio bacteriovorus 109J." Current Microbiology, Feb 2006;52(2), p. 81-85 | ||
(11)Tudor John, "Control of predation and development in Bdellovibrio bacteriovorus." http://www.sju.edu/biology/Teaching_Postdoctoral_Fellowsh/Bdellovibrio_bacteriovorus_Res/bdellovibrio_bacteriovorus_res.html | |||
Edited by [mailto:hlcheng@ucsd.edu Hiu Cheng], student of [mailto:ralarsen@ucsd.edu Rachel Larsen] | Edited by [mailto:hlcheng@ucsd.edu Hiu Cheng], student of [mailto:ralarsen@ucsd.edu Rachel Larsen] | ||
Edited by KLB | |||
Latest revision as of 20:46, 10 September 2010
A Microbial Biorealm page on the genus Bdellovibrio bacteriovorus
Classification
Higher order taxa
Bacteria; Proteobacteria; Deltaproteobacteria; Bdellovibrionales; Bdellovibrionaceae; Bdellovibrio
Species
Bdellovibro bacteriovous
Strain
Bdellovibrio bacteriovorus: HD100, 109J
Description and significance
Bdellovibrios were discovered by Stolp and Petzhold in 1962, in an attempt to isolate bacteriophage from soil samples. Stolp and Petzhold observed unique plaques in their samples that took several days to develop and continued to grow for over a week, instead of plaques caused by bacteriophages that would appear within hours. A closer inspection of the plaques under a light microscope revealed cells that were small, highly motile, and vibrio-shaped. These cells were Bdellovibrios.
After the discovery of Bdellovibrios further observations revealed many interesting and unique properties. One property that makes Bdellovibrios interesting is that it is a parasite to other Gram-negative bacterias. Bdellovibrios have biphasic life-cycles that include an attack phase, and a free living and mobile phase.(2) The attack phase is when it attaches to another Gram-negative bacteria and imbeds itself into its periplasm, it then procedes to grow and replicate itself by degrading the host bacterium from the inside out.(1) In the free living and mobile phase, Bdellovibrios move about in search of host or prey bacteria so it can intiate attack phase again.(2) Each of these phases are of interest to researchers because they reveal unique cell-cell interactions and unusual cell metabolism.(1)
Bdellovibrio bacteriovorus is a small, curved, and highly motile Gram-negative bacteria approximately 0.2 to 0.5μm wide and 0.5 to 2.5μm long.(3) It has been found in many environments that include soil, sewage, and other terrestial and aquatic habitats. B. bacteriovorus has been observed to only attack Gram-negative bacteria which includes many plant, animal and human pathogens making it an execellent candidate as a biocontrol agent.(2) The study of its degradative enzymes and host targeting system has shed some insight in possible designs for new antimicrobial agents.(3)
Genome structure
The sequencing of the B. bacteriovorus HD100 genome was completed on 01/31/2004. The complete genome consists of a single circular chromosome that is 3,782,950 nucleotides long. The entire genome has a 50% GC content and contains 3629 genes which code for 3587 proteins and 42 structural RNAs.(4)
Of the large number of proteins that the B. bacteriovorus genome encodes, many of them are degradative and lytic enzymes(2). These enzymes are necessary for B. bacteriovorus to breakdown its host or prey cell. Some of these enzymes need to be secreted out of B. bacteriovorus and into its prey's cell, while degradative products needs to be transported into the B. bacteriovorus from the prey cell cytoplasm. Therefore, the genome of B. bacteriovorus also includes a large amount of transport proteins. In fact, B. bacteriovorus has the potential protein secretory capabilities of at least five types of outer membrane secretion systems and four types of inner membrane secretion systems.(5)
Other than encoding for degradative enzymes and transport systems, the chromosome also contains multiple biosynthetic gene clusters that are necessary for the formation of the flagella and pilus. These gene clusters are essential for the motility B. bacteriovorus, which is important to its life cycle because it directly relates to its ability to find and approach the prey cell. However, the organism sheds its flagellum once it makes irreversible contact with its prey cell suggesting that the flagellum is not required for it to grow.(2)
Cell structure and metabolism
The general cell structure of B. bacteriovorus is small, curved, and highly motile Gram-negative bacteria approximately 0.2 to 0.5 μm wide and 0.5 to 2.5 μm long.(3) It has a flagellum that is membrane bound, allowing it to swim up to 100 body length per second while it looks for its prey.(9)
The unique biphasic life-cycle of B. bacteriovorus is directly related to its metabolism. In the free-living or mobile phase, B. bacteriovorus actively seeks other Gram-negative bacteria by chemotaxic means. During the mobile phase, most of the metabolic activity is involved in motility and prey detection.(7) In the attack phase, B. bacteriovorus burrows itself into the periplasm of the prey cell and begins to degrade the prey cell from the inside out. A large variety of metabolic activities occur during this phase.(2)
In the attack phase, B. bacteriovorus begins by attaching itself to the outer membrane of the prey cell via hook-like structures.(8) It then proceeds to secrete enzymes which break down a large enough hole in the outer membrane for B. bacteriovorus to enter. After entry into the prey cell, B. bacteriovorus sheds its flagellum and seals the hole in the outer membrane of the prey cell.(1) B. bacteriovorus then secretes degradative enzymes that act on the inner membrane of the prey cell allowing it to access the prey's cytoplasm. The degradation of the inner membrane causes the prey cell to become a bdelloplast, host cell of Bdellovibrios with a stable spherical structure, and also the destruction of the host cell's ability to generate energy.(1) At this point, B. bacteriovorus degrades and metabolizes the macromolecules and nutrients from the host's cytoplamic material allowing it to grow and begin DNA replication. B. bacteriovorus subsequently becomes a long filament and then fragments into individual mobile progenies which will lyse the remaining bdelloplast.(1)
The metabolism of B. bacteriovorus has many interesting areas. One of these areas is that B. bacteriovorus can survive while lacking numerous amino acid biosynthetic pathways. The reason for this is that B. bacteriovorus utilizes the metabolites of its prey for its own protein synthesis.(2) B. bacteriovorus is very efficient at utilizing the DNA, RNA, and other pre-existing precursors from the host cell. It can break down or rearrange these precursors for the biosynthesis of homologous polymers. The uptake of nucleoside monophosphates and glycerol phoshphates greatly reduces the amount of energy that B. bacteriovorus has to produce during nucleic acid synthesis and lipid synthesis. There has also been evidence that B. bacteriovorus incorporates intact fatty acid molecules from its host. Experiments have shown that the energy efficiency of B. bacteriovorus can be 0.8 to 1.5 times more efficient than bacteria growing in rich media.(6)
Ecology
B. bacteriovorus seems to be pretty ubiquitous in nature and manmade habitats. They have been found in soil samples, rhizosphere of plant roots, rivers, oceans, sewage, intestines and feces of birds and mammals, and even in oyster shells and the gills of crabs.(9) B. bacteriovorus are able to thrive in almost any habitat, the general requirements are that there needs to be oxygen and some other Gram-negative bacteria present in its environment. Its optimal temperature is between 28-30C, making B. bacteriovorus a mesophile.(2)
B. bacteriovorus is somewhat tolerant to its environment. It can live in a range of salinity and is resistant to pollution of its environment.(2) During its attack phase or as bdelloplasts, B. bacteriovorus can tolerate dry spells and short periods of anaerobic environment. This makes B. bacteriovorus more tolerant to environmental changes than other permanantly aerobic bacteria.(9)
B. bacteriovorus has also been associated with biofilms. It seems that the rich environment of biofilms supply B. bacteriovorus with many possible host cells. Research has shown that B. bacteriovorus can significantly decrease E. Coli biofilm that is grown on stainless steel.(9)
Pathology
B. bacteriovorus is not a known pathogen to humans. It is a known pathogen to Gram-negative bacteria, making it a possible biocontrol agent to many human pathogens.(2)
Application to Biotechnology
The major applications of B. bacteriovorus to biotechnology seem to be its potential as a biocontrol agent. B. bacteriovorus has been used in certain water treatment plants to reduce the number of Gram-negative organisms in the water. Agriculturally, B. bacteriovorus has been used to limit or prevent the start and spread of plant pathogens with certain crops. The use of B. bacteriovorus in agriculture requires extensive knowledge of the crop and its ecology; B. bacteriovorus may attack growth promoting Gram-negative organisms needed for healthy crops such as rhizobacterias.(9)
Current Research
Currently, the main field of research with B. bacteriovorus is its high potential as a biocontrol agent. Many experiments are performed to test how it specifically recognizes its prey. There is also research on how B. bacteriovorus recognizes and prefers one prey over another.(10) Molecular research on the degradative enzymes of B. bacteriovorus and its targets provide insight to evolutionary successful point of attack to Gram-negative bacteria.(3) The ultimate goal in much of this research is to provide possible elements to create more specific antimicrobial agents. It is also highly possible that a slight change in its genetic code can allow B. bacteriovorus to truly become a living antibiotic.
Prey Selection of B. bacteriovorus
When introduced to multiple preys, B. bacteriovorus preferentially lysed one prey over the others. This research showed that B. bacteriovorus may not randomly infect prey cells, but differentially prefer to infect one prey over another.(10) The identification and characterization of the genetic control of the bdellovibrio developmental cycle and its temporal gene expression will help identify which genes are responsible for prey selection and the proteins that these genes will encode.(11)
References
(1) Thomashow Michael F., Cotter Todd W., ‘‘Bdellovibrio Host Dependence: the Search for Signal Molecules and Genes That Regulate the Intraperiplasmic Growth Cycle’’, Journal of Bacteriology Sept.1992, p. 5767-5771
(2)NCBI Genome Project Database
(3)European Bioinformatics Institute Database
(5)Barabote RD, Rendulic S, Schuster SC, Saier MH Jr. “Comprehensive analysis of transport proteins encoded within the genome of Bdellovibrio bacteriovorus’’ Genomics, 2007 Aug 14
(6)Rittenberg Sydney C., Hespell Robert B. “Energy Efficiency of Intraperiplasmic Growth of ‘‘Bdellovibrio bacteriovorus’’, Journal of Bacteriology Mar. 1975, p.1158-1165
(7)Lambert, C., Smith, M. C. M., & Sockett, R. E., “ A novel assay to monitor predator-prey interaction for Bdellovibrio bacteriovorus 109 J reveals a role for methylaccepting chemotaxis proteins in predation.” Environmental Biology, 5(2), 127.(2003, February).
(8)Fox, J. F. . Bdellovibrio make fast food of gram-negative bacteria . In Microbe Magazine. (2006) http://www.asm.org/
(9)YAIR Shemesh, YAACOV Davidov, SUSAN Koval, JURKEVITCH Edouard "Small eats big: ecology and diversity of Bdellovibrio and like organisms, and their dynamics in predator-prey interactions", Agronomie 23, (2003), p.433–439, http://www.agronomy-journal.org/index.php?option=article&access=standard&Itemid=129&url=/articles/agro/pdf/2003/05/A3505.pdf
(10)Rogosky AM, Moak PL, Emmert EA. "Differential predation by Bdellovibrio bacteriovorus 109J." Current Microbiology, Feb 2006;52(2), p. 81-85
(11)Tudor John, "Control of predation and development in Bdellovibrio bacteriovorus." http://www.sju.edu/biology/Teaching_Postdoctoral_Fellowsh/Bdellovibrio_bacteriovorus_Res/bdellovibrio_bacteriovorus_res.html
Edited by Hiu Cheng, student of Rachel Larsen
Edited by KLB
