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Paenibacillus vortex

Paenibacillus vortex is a species of pattern-forming bacteria, first discovered in the early 1990s by Eshel Ben-Jacob's group at Tel Aviv University.[1] It is a social microorganism that forms colonies with complex and dynamic architectures. P. vortex is mainly found in heterogeneous and complex environments, such as the rhizosphere, the soil region directly influenced by plant roots.

Paenibacillus vortex
Figure 1: Colony organization of the P. vortex bacteria when grown on 15g/L peptone and 2.25% (w/v) agar for four days. The bright yellow dots are the vortices. The colonies were grown in a Petri dish size 8.8cm and stained with Coomassie dyes (Brilliant Blue). The colors were inverted to emphasize higher densities using the brighter shades of yellow.
Scientific classification
Domain:
Phylum:
Class:
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Genus:
Binomial name
Paenibacillus vortex
Synonyms
Bacillus vortex
Ash et al. 1994

The genus Paenibacillus comprises facultative anaerobic, endospore-forming bacteria originally included within the genus Bacillus and then reclassified as a separate genus in 1993.[2] Bacteria in the genus have been detected in a variety of environments such as: soil, water, vegetable matter, forage and insect larvae, as well as clinical samples.[3][4][5][6] Paenibacillus spp., including P. vortex, produce extracellular enzymes that catalyze a variety of synthetic reactions in industrial, agricultural and medical applications.[7][8][9] Various Paenibacillus spp. also produce antimicrobial substances that can affect micro-organisms such as fungi, in addition to soil and plant pathogenic bacteria.[10][11][12]

Social Motility

Paenibacillus vortex possesses advanced social motility employing cell-cell attractive and repulsive chemotactic signalling and physical links. When grown on soft surfaces (e.g. agar), the collective motility is reflected by the formation of foraging swarms[13] that act as arms sent out in search of food. These swarms have an aversion to crossing each other’s trail and collectively change direction when food is sensed. The “swarming intelligence” P. vortex, is further marked by the fact that of the swarms can even split and reunite when detecting scattered patches of nutrients.[13]

Pattern Formation and Social Behaviors

 
Figure 2: Scanning electron microscope (SEM) observation of P. vortex illustrating a typical bacteria arrangement in the center of a vortex. Notable, that each individual bacterium is curved. Scale bar in is 5µm.

P. vortex is a social microorganism: when grown on under growth conditions that mimic natural environments such as hard surfaces it forms colonies of 109-1012 cells with remarkably complex and dynamic architectures (Figure 1).[1][14][15] Being part of a large cooperative, the bacteria can better compete for food resources and be protected against antibacterial assaults.[13][14] Under laboratory growth conditions, similar to other social bacteria, P. vortex colonies behave much like a multi-cellular organism, with cell differentiation and task distribution.[16][17][18][19] P. vortex is marked by its ability to generate special aggregates of dense bacteria that are pushed forward by repulsive chemotactic signals sent from the cells at the back.[15][20][21][22][23][24] These rotating aggregates termed vortices (Figure 2), pave the way for the colony to expand. The vortices serve as building blocks of colonies with special modular organization (Figure 1). Accomplishing such intricate cooperative ventures requires sophisticated cell-cell communication,[14][19][25][26][27] including semantic and pragmatic aspects of linguistics.[19] Communicating with each other using a variety of chemical signals, bacteria exchange information regarding population size, a myriad of individual environmental measurements at different locations, their internal states and their phenotypic and epigenetic adjustments. The bacteria collectively sense the environment and execute distributed information processing to glean and assess relevant information.[14][19][28] The information is then used by the bacteria for reshaping the colony while redistributing tasks and cell epigenetic differentiations, for collective decision-making and for turning on and off defense and offense mechanisms needed to thrive in competitive environments, faculties that can be perceived as social intelligence of bacteria.[19]

 
A typical colonial pattern generated by P. vortex when grown on 2.25% (w/v) agar with 2% (w/v) peptone in a 9 cm Petri dish.

Genome Sequence of the Paenibacillus vortex

The genome sequence of the P. vortex [29] is now available [GenBank: ADHJ00000000). The genome was sequenced by a hybrid approach using 454 Life Sciences and Illumina, achieving a total of 289X coverage, with 99.8% sequence identity between the two methods. The sequencing results were validated using a custom designed Agilent microarray expression chip submitted to EMBL-EBI [ArrayExpress: E-MEXP-3019 which represented the coding and the non-coding regions. Analysis of the P. vortex genome revealed 6,437 open reading frames (ORFs) and 73 non-coding RNA genes. The analysis also unveiled the P. vortex potential to produce a wealth of enzymes and proteases as well as a great variety of antimicrobial substances that affect a wide range of microorganisms. The possession of these advanced defense and offense strategies render Paenibacillus vortex as a rich source of useful genes for agricultural, medical, industrial and biofuel applications.

Comparative Genomics and Social-IQ Score

Comparative genomic analysis revealed that bacteria successful in heterogeneous and competitive environments often contain extensive signal transduction and regulatory networks.[30][31][32] Detailed comparative genomic analysis with a dataset of 500 complete bacterial genomes revealed that P. vortex has the third highest number of signal transduction genes, slightly below two other Paenibacillus species, Paenibacillus sp. JDR-2 and Paenibacillus sp. Y412MC10.[29] The comparative genomic analysis further revealed that these three Paenibacillus species also have the highest “Social-IQ” score among all 500 sequenced bacteria, over 3 standard deviations higher than average. The score is based on the number of genes which afford bacteria abilities to communicate and process environmental information (two-component and transcription-factor genes), to make decisions and to synthesize offensive (toxic) and defensive (neutralizing) agents as needed during chemical warfare with other microorganisms.[29] Defined this way, the Social-IQ score provides a measure of the genome capacity for social intelligence, hence it helps realizing social intelligence of bacteria.

References

  1. ^ a b Ben-Jacob E, Shochet O, Tenenbaum A, Avidan O (1995). Cladis PE, Palffy-Muhorey P. (eds.). "Evolution of complexity during growth of bacterial colonies". NATO Advanced Research Workshop. Santa Fe, USA: Addison-Wesley Publishing Company: 619–633.
  2. ^ Ash C, Priest FG, Collins MD: Molecular identification of rRNA group 3 bacilli (Ash, Farrow, Wallbanks and Collins) using a PCR probe test. Proposal for the creation of a new genus Paenibacillus. Antonie van Leeuwenhoek 1993, 64:253-260.
  3. ^ Lal, Sadhana; Tabacchioni, Silvia (2009). "Ecology and biotechnological potential of Paenibacillus polymyxa: a minireview". Indian Journal of Microbiology. Springer Science and Business Media LLC. 49 (1): 2–10. doi:10.1007/s12088-009-0008-y. ISSN 0046-8991. PMC 3450047. PMID 23100748.
  4. ^ McSpadden Gardener, Brian B. (2004). "Ecology of Bacillus and Paenibacillus spp. in Agricultural Systems". Phytopathology. Scientific Societies. 94 (11): 1252–1258. doi:10.1094/phyto.2004.94.11.1252. ISSN 0031-949X. PMID 18944463.
  5. ^ Montes, M. J. (2004-09-01). "Paenibacillus antarcticus sp. nov., a novel psychrotolerant organism from the Antarctic environment". International Journal of Systematic and Evolutionary Microbiology. Microbiology Society. 54 (5): 1521–1526. doi:10.1099/ijs.0.63078-0. ISSN 1466-5026. PMID 15388704.
  6. ^ Ouyang J, Pei Z, Lutwick L, Dalal S, Yang L, Cassai N, Sandhu K, Hanna B, Wieczorek RL, Bluth M, Pincus MR: Case report: Paenibacillus thiaminolyticus: a new cause of human infection, inducing bacteremia in a patient on hemodialysis. Ann Clin Lab Sci 2008, 38:393-400.
  7. ^ Konishi, J.; Maruhashi, K. (2003-09-01). "2-(2'-Hydroxyphenyl)benzene sulfinate desulfinase from the thermophilic desulfurizing bacterium Paenibacillus sp. strain A11-2: purification and characterization". Applied Microbiology and Biotechnology. Springer Science and Business Media LLC. 62 (4): 356–361. doi:10.1007/s00253-003-1331-6. ISSN 0175-7598. PMID 12743754. S2CID 7956236.
  8. ^ Raza W, Yang W, Shen QR: Paenibacillus polymyxa: Antibiotics, Hydrolytic Enzymes and Hazard Assessment. J Plant Pathol 2008, 90:419-430.
  9. ^ Watanapokasin RY, Boonyakamol A, Sukseree S, Krajarng A, Sophonnithiprasert T, Kanso S, Imai T: Hydrogen production and anaerobic decolorization of wastewater containing Reactive Blue 4 by a bacterial consortium of Salmonella subterranea and Paenibacillus polymyxa. Biodegradation 2009, 20:411-418.
  10. ^ Dijksterhuis J, Sanders M, Gorris LG, Smid EJ: Antibiosis plays a role in the context of direct interaction during antagonism of Paenibacillus polymyxa towards Fusarium oxysporum. J Appl Microbiol 1999, 86:13-21.
  11. ^ Girardin H, Albagnac C, Dargaignaratz C, Nguyen-The C, Carlin F: Antimicrobial activity of foodborne Paenibacillus and Bacillus spp. against Clostridium botulinum. J Food Prot 2002, 65:806-813.
  12. ^ von der Weid I, Alviano DS, Santos AL, Soares RM, Alviano CS, Seldin L: Antimicrobial activity of Paenibacillus peoriae strain NRRL BD-62 against a broad spectrum of phytopathogenic bacteria and fungi. J Appl Microbiol 2003, 95:1143-1151.
  13. ^ a b c Ingham CJ, Ben-Jacob E: Swarming and complex pattern formation in Paenibacillus vortex studied by imaging and tracking cells. BMC Microbiol 2008, 8:36.
  14. ^ a b c d Ben-Jacob E: Bacterial self-organization: co-enhancement of complexification and adaptability in a dynamic environment. Phil Trans R Soc Lond A 2003, 361:1283-1312.
  15. ^ a b Ben-Jacob E, Cohen I, Gutnick DL: Cooperative organization of bacterial colonies: from genotype to morphotype. Annu Rev Microbiol 1998, 52:779-806.
  16. ^ Aguilar C, Vlamakis H, Losick R, Kolter R: Thinking about Bacillus subtilis as a multicellular organism. Curr Opin Microbiol 2007, 10:638-643.
  17. ^ Dunny GM, Brickman TJ, Dworkin M: Multicellular behavior in bacteria: communication, cooperation, competition and cheating. Bioessays 2008, 30:296-298.
  18. ^ Shapiro JA, Dworkin M: Bacteria as multicellular organisms. 1st edn: Oxford University Press, USA; 1997.
  19. ^ a b c d e Ben-Jacob E, Becker I, Shapira Y, Levine H: Bacterial linguistic communication and social intelligence. Trends Microbiol 2004, 12:366-372.
  20. ^ Ben-Jacob E: From snowflake formation to growth of bacterial colonies II: Cooperative formation of complex colonial patterns. Contem Phys 1997, 38:205 - 241.
  21. ^ Ben-Jacob E, Cohen I: Cooperative formation of bacterial patterns. In Bacteria as Multicellular Organisms Edited by Shapiro JA, Dworkin M. New York: Oxford University Press; 1997: 394-416
  22. ^ Ben-Jacob E, Cohen I, Czirók A, Vicsek T, Gutnick DL: Chemomodulation of cellular movement, collective formation of vortices by swarming bacteria, and colonial development. Physica A 1997, 238:181-197.
  23. ^ Cohen I, Czirok A, Ben-Jacob E: Chemotactic-based adaptive self-organization during colonial development. Physica A 1996, 233:678-698.
  24. ^ Czirók, András; Ben-Jacob, Eshel; Cohen, Inon; Vicsek, Tamás (1996-08-01). "Formation of complex bacterial colonies via self-generated vortices". Physical Review E. American Physical Society (APS). 54 (2): 1791–1801. Bibcode:1996PhRvE..54.1791C. doi:10.1103/physreve.54.1791. ISSN 1063-651X. PMID 9965259.
  25. ^ Bassler BL, Losick R: Bacterially speaking. Cell 2006, 125:237-246.
  26. ^ Bischofs IB, Hug JA, Liu AW, Wolf DM, Arkin AP: Complexity in bacterial cell-cell communication: quorum signal integration and subpopulation signaling in the Bacillus subtilis phosphorelay. Proc Natl Acad Sci U S A 2009, 106:6459-6464.
  27. ^ Kolter R, Greenberg EP: Microbial sciences: the superficial life of microbes. Nature 2006, 441:300-302.
  28. ^ Dwyer DJ, Kohanski MA, Collins JJ: Networking opportunities for bacteria. Cell 2008, 135:1153-1156.
  29. ^ a b c Sirota-Madi A, Olender T, Helman Y, Ingham C, Brainis I, Roth D, Hagi E, Brodsky L, Leshkowitz D, Galatenko V, et al: Genome sequence of the pattern forming Paenibacillus vortex bacterium reveals potential for thriving in complex environments. BMC Genomics, 11:710.
  30. ^ Alon U: An Introduction to Systems Biology: Design Principles of Biological circuits. London, UK: CRC Press; 2006.
  31. ^ Galperin MY, Gomelsky M: Bacterial Signal Transduction Modules: from Genomics to Biology. ASM News 2005, 71:326-333.
  32. ^ Whitworth DE, Cock PJ: Two-component systems of the myxobacteria: structure, diversity and evolutionary relationships. Microbiology 2008, 154:360-372.

External links

  • Genome sequence of the pattern forming Paenibacillus vortex bacterium reveals potential for thriving in complex environments.
  • Realizing Social Intelligence of Bacteria
  • Bacterial Art

paenibacillus, vortex, species, pattern, forming, bacteria, first, discovered, early, 1990s, eshel, jacob, group, aviv, university, social, microorganism, that, forms, colonies, with, complex, dynamic, architectures, vortex, mainly, found, heterogeneous, compl. Paenibacillus vortex is a species of pattern forming bacteria first discovered in the early 1990s by Eshel Ben Jacob s group at Tel Aviv University 1 It is a social microorganism that forms colonies with complex and dynamic architectures P vortex is mainly found in heterogeneous and complex environments such as the rhizosphere the soil region directly influenced by plant roots Paenibacillus vortexFigure 1 Colony organization of the P vortex bacteria when grown on 15g L peptone and 2 25 w v agar for four days The bright yellow dots are the vortices The colonies were grown in a Petri dish size 8 8cm and stained with Coomassie dyes Brilliant Blue The colors were inverted to emphasize higher densities using the brighter shades of yellow Scientific classificationDomain BacteriaPhylum BacillotaClass BacilliOrder BacillalesFamily PaenibacillaceaeGenus PaenibacillusBinomial namePaenibacillus vortexSynonymsBacillus vortex Ash et al 1994The genus Paenibacillus comprises facultative anaerobic endospore forming bacteria originally included within the genus Bacillus and then reclassified as a separate genus in 1993 2 Bacteria in the genus have been detected in a variety of environments such as soil water vegetable matter forage and insect larvae as well as clinical samples 3 4 5 6 Paenibacillus spp including P vortex produce extracellular enzymes that catalyze a variety of synthetic reactions in industrial agricultural and medical applications 7 8 9 Various Paenibacillus spp also produce antimicrobial substances that can affect micro organisms such as fungi in addition to soil and plant pathogenic bacteria 10 11 12 Contents 1 Social Motility 2 Pattern Formation and Social Behaviors 3 Genome Sequence of the Paenibacillus vortex 4 Comparative Genomics and Social IQ Score 5 References 6 External linksSocial Motility EditPaenibacillus vortex possesses advanced social motility employing cell cell attractive and repulsive chemotactic signalling and physical links When grown on soft surfaces e g agar the collective motility is reflected by the formation of foraging swarms 13 that act as arms sent out in search of food These swarms have an aversion to crossing each other s trail and collectively change direction when food is sensed The swarming intelligence P vortex is further marked by the fact that of the swarms can even split and reunite when detecting scattered patches of nutrients 13 Pattern Formation and Social Behaviors Edit Figure 2 Scanning electron microscope SEM observation of P vortex illustrating a typical bacteria arrangement in the center of a vortex Notable that each individual bacterium is curved Scale bar in is 5µm P vortex is a social microorganism when grown on under growth conditions that mimic natural environments such as hard surfaces it forms colonies of 109 1012 cells with remarkably complex and dynamic architectures Figure 1 1 14 15 Being part of a large cooperative the bacteria can better compete for food resources and be protected against antibacterial assaults 13 14 Under laboratory growth conditions similar to other social bacteria P vortex colonies behave much like a multi cellular organism with cell differentiation and task distribution 16 17 18 19 P vortex is marked by its ability to generate special aggregates of dense bacteria that are pushed forward by repulsive chemotactic signals sent from the cells at the back 15 20 21 22 23 24 These rotating aggregates termed vortices Figure 2 pave the way for the colony to expand The vortices serve as building blocks of colonies with special modular organization Figure 1 Accomplishing such intricate cooperative ventures requires sophisticated cell cell communication 14 19 25 26 27 including semantic and pragmatic aspects of linguistics 19 Communicating with each other using a variety of chemical signals bacteria exchange information regarding population size a myriad of individual environmental measurements at different locations their internal states and their phenotypic and epigenetic adjustments The bacteria collectively sense the environment and execute distributed information processing to glean and assess relevant information 14 19 28 The information is then used by the bacteria for reshaping the colony while redistributing tasks and cell epigenetic differentiations for collective decision making and for turning on and off defense and offense mechanisms needed to thrive in competitive environments faculties that can be perceived as social intelligence of bacteria 19 A typical colonial pattern generated by P vortex when grown on 2 25 w v agar with 2 w v peptone in a 9 cm Petri dish Genome Sequence of the Paenibacillus vortex EditThe genome sequence of the P vortex 29 is now available GenBank ADHJ00000000 The genome was sequenced by a hybrid approach using 454 Life Sciences and Illumina achieving a total of 289X coverage with 99 8 sequence identity between the two methods The sequencing results were validated using a custom designed Agilent microarray expression chip submitted to EMBL EBI ArrayExpress E MEXP 3019 which represented the coding and the non coding regions Analysis of the P vortex genome revealed 6 437 open reading frames ORFs and 73 non coding RNA genes The analysis also unveiled the P vortex potential to produce a wealth of enzymes and proteases as well as a great variety of antimicrobial substances that affect a wide range of microorganisms The possession of these advanced defense and offense strategies render Paenibacillus vortex as a rich source of useful genes for agricultural medical industrial and biofuel applications Comparative Genomics and Social IQ Score EditComparative genomic analysis revealed that bacteria successful in heterogeneous and competitive environments often contain extensive signal transduction and regulatory networks 30 31 32 Detailed comparative genomic analysis with a dataset of 500 complete bacterial genomes revealed that P vortex has the third highest number of signal transduction genes slightly below two other Paenibacillus species Paenibacillus sp JDR 2 and Paenibacillus sp Y412MC10 29 The comparative genomic analysis further revealed that these three Paenibacillus species also have the highest Social IQ score among all 500 sequenced bacteria over 3 standard deviations higher than average The score is based on the number of genes which afford bacteria abilities to communicate and process environmental information two component and transcription factor genes to make decisions and to synthesize offensive toxic and defensive neutralizing agents as needed during chemical warfare with other microorganisms 29 Defined this way the Social IQ score provides a measure of the genome capacity for social intelligence hence it helps realizing social intelligence of bacteria References Edit a b Ben Jacob E Shochet O Tenenbaum A Avidan O 1995 Cladis PE Palffy Muhorey P eds Evolution of complexity during growth of bacterial colonies NATO Advanced Research Workshop Santa Fe USA Addison Wesley Publishing Company 619 633 Ash C Priest FG Collins MD Molecular identification of rRNA group 3 bacilli Ash Farrow Wallbanks and Collins using a PCR probe test Proposal for the creation of a new genus Paenibacillus Antonie van Leeuwenhoek 1993 64 253 260 Lal Sadhana Tabacchioni Silvia 2009 Ecology and biotechnological potential of Paenibacillus polymyxa a minireview Indian Journal of Microbiology Springer Science and Business Media LLC 49 1 2 10 doi 10 1007 s12088 009 0008 y ISSN 0046 8991 PMC 3450047 PMID 23100748 McSpadden Gardener Brian B 2004 Ecology of Bacillus and Paenibacillus spp in Agricultural Systems Phytopathology Scientific Societies 94 11 1252 1258 doi 10 1094 phyto 2004 94 11 1252 ISSN 0031 949X PMID 18944463 Montes M J 2004 09 01 Paenibacillus antarcticus sp nov a novel psychrotolerant organism from the Antarctic environment International Journal of Systematic and Evolutionary Microbiology Microbiology Society 54 5 1521 1526 doi 10 1099 ijs 0 63078 0 ISSN 1466 5026 PMID 15388704 Ouyang J Pei Z Lutwick L Dalal S Yang L Cassai N Sandhu K Hanna B Wieczorek RL Bluth M Pincus MR Case report Paenibacillus thiaminolyticus a new cause of human infection inducing bacteremia in a patient on hemodialysis Ann Clin Lab Sci 2008 38 393 400 Konishi J Maruhashi K 2003 09 01 2 2 Hydroxyphenyl benzene sulfinate desulfinase from the thermophilic desulfurizing bacterium Paenibacillus sp strain A11 2 purification and characterization Applied Microbiology and Biotechnology Springer Science and Business Media LLC 62 4 356 361 doi 10 1007 s00253 003 1331 6 ISSN 0175 7598 PMID 12743754 S2CID 7956236 Raza W Yang W Shen QR Paenibacillus polymyxa Antibiotics Hydrolytic Enzymes and Hazard Assessment J Plant Pathol 2008 90 419 430 Watanapokasin RY Boonyakamol A Sukseree S Krajarng A Sophonnithiprasert T Kanso S Imai T Hydrogen production and anaerobic decolorization of wastewater containing Reactive Blue 4 by a bacterial consortium of Salmonella subterranea and Paenibacillus polymyxa Biodegradation 2009 20 411 418 Dijksterhuis J Sanders M Gorris LG Smid EJ Antibiosis plays a role in the context of direct interaction during antagonism of Paenibacillus polymyxa towards Fusarium oxysporum J Appl Microbiol 1999 86 13 21 Girardin H Albagnac C Dargaignaratz C Nguyen The C Carlin F Antimicrobial activity of foodborne Paenibacillus and Bacillus spp against Clostridium botulinum J Food Prot 2002 65 806 813 von der Weid I Alviano DS Santos AL Soares RM Alviano CS Seldin L Antimicrobial activity of Paenibacillus peoriae strain NRRL BD 62 against a broad spectrum of phytopathogenic bacteria and fungi J Appl Microbiol 2003 95 1143 1151 a b c Ingham CJ Ben Jacob E Swarming and complex pattern formation in Paenibacillus vortex studied by imaging and tracking cells BMC Microbiol 2008 8 36 a b c d Ben Jacob E Bacterial self organization co enhancement of complexification and adaptability in a dynamic environment Phil Trans R Soc Lond A 2003 361 1283 1312 a b Ben Jacob E Cohen I Gutnick DL Cooperative organization of bacterial colonies from genotype to morphotype Annu Rev Microbiol 1998 52 779 806 Aguilar C Vlamakis H Losick R Kolter R Thinking about Bacillus subtilis as a multicellular organism Curr Opin Microbiol 2007 10 638 643 Dunny GM Brickman TJ Dworkin M Multicellular behavior in bacteria communication cooperation competition and cheating Bioessays 2008 30 296 298 Shapiro JA Dworkin M Bacteria as multicellular organisms 1st edn Oxford University Press USA 1997 a b c d e Ben Jacob E Becker I Shapira Y Levine H Bacterial linguistic communication and social intelligence Trends Microbiol 2004 12 366 372 Ben Jacob E From snowflake formation to growth of bacterial colonies II Cooperative formation of complex colonial patterns Contem Phys 1997 38 205 241 Ben Jacob E Cohen I Cooperative formation of bacterial patterns In Bacteria as Multicellular Organisms Edited by Shapiro JA Dworkin M New York Oxford University Press 1997 394 416 Ben Jacob E Cohen I Czirok A Vicsek T Gutnick DL Chemomodulation of cellular movement collective formation of vortices by swarming bacteria and colonial development Physica A 1997 238 181 197 Cohen I Czirok A Ben Jacob E Chemotactic based adaptive self organization during colonial development Physica A 1996 233 678 698 Czirok Andras Ben Jacob Eshel Cohen Inon Vicsek Tamas 1996 08 01 Formation of complex bacterial colonies via self generated vortices Physical Review E American Physical Society APS 54 2 1791 1801 Bibcode 1996PhRvE 54 1791C doi 10 1103 physreve 54 1791 ISSN 1063 651X PMID 9965259 Bassler BL Losick R Bacterially speaking Cell 2006 125 237 246 Bischofs IB Hug JA Liu AW Wolf DM Arkin AP Complexity in bacterial cell cell communication quorum signal integration and subpopulation signaling in the Bacillus subtilis phosphorelay Proc Natl Acad Sci U S A 2009 106 6459 6464 Kolter R Greenberg EP Microbial sciences the superficial life of microbes Nature 2006 441 300 302 Dwyer DJ Kohanski MA Collins JJ Networking opportunities for bacteria Cell 2008 135 1153 1156 a b c Sirota Madi A Olender T Helman Y Ingham C Brainis I Roth D Hagi E Brodsky L Leshkowitz D Galatenko V et al Genome sequence of the pattern forming Paenibacillus vortex bacterium reveals potential for thriving in complex environments BMC Genomics 11 710 Alon U An Introduction to Systems Biology Design Principles of Biological circuits London UK CRC Press 2006 Galperin MY Gomelsky M Bacterial Signal Transduction Modules from Genomics to Biology ASM News 2005 71 326 333 Whitworth DE Cock PJ Two component systems of the myxobacteria structure diversity and evolutionary relationships Microbiology 2008 154 360 372 External links EditGenome sequence of the pattern forming Paenibacillus vortex bacterium reveals potential for thriving in complex environments Prof Eshel Ben Jacob s Home Page Realizing Social Intelligence of Bacteria Bacterial Art Retrieved from https en wikipedia org w index php title Paenibacillus vortex amp oldid 1111462267, wikipedia, wiki, book, books, library,

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