Paenibacillus Assamensis
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''Paenibacillus'' is a genus of
facultative anaerobic A facultative anaerobic organism is an organism that makes ATP by aerobic respiration if oxygen is present, but is capable of switching to fermentation if oxygen is absent. Some examples of facultatively anaerobic bacteria are '' Staphylococc ...
,
endospore An endospore is a dormant, tough, and non-reproductive structure produced by some bacteria in the phylum Bacillota. The name "endospore" is suggestive of a spore or seed-like form (''endo'' means 'within'), but it is not a true spore (i.e., no ...
-forming
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
, originally included within the genus ''
Bacillus ''Bacillus'' (Latin "stick") is a genus of Gram-positive, rod-shaped bacteria, a member of the phylum ''Bacillota'', with 266 named species. The term is also used to describe the shape (rod) of other so-shaped bacteria; and the plural ''Bacilli ...
'' and then reclassified as a separate genus in 1993.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. Bacteria belonging to this genus have been detected in a variety of environments, such as: soil, water,
rhizosphere The rhizosphere is the narrow region of soil or substrate that is directly influenced by root secretions and associated soil microorganisms known as the root microbiome. Soil pores in the rhizosphere can contain many bacteria and other microor ...
, vegetable matter, forage and insect larvae, as well as clinical samples.McSpadden Gardener BB: Ecology of Bacillus and Paenibacillus spp. in Agricultural Systems. Phytopathology 2004, 94:1252-1258.Montes MJ, Mercade E, Bozal N, Guinea J: Paenibacillus antarcticus sp. nov., a novel psychrotolerant organism from the Antarctic environment. Int J Syst Evol Microbiol 2004, 54:1521-1526.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. The name reflects: Latin ''paene'' means almost, so the paenibacilli are literally "almost bacilli". The genus includes '' P. larvae'', which causes
American foulbrood American foulbrood (AFB, ''Histolysis infectiosa perniciosa larvae apium'', ''Pestis americana larvae apium''), caused by the spore-forming bacterium ''Paenibacillus larvae'' (reclassified as one species without subspecies differentiation in 20 ...
in
honeybees A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the current cosmo ...
, '' P. polymyxa'', which is capable of
fixing nitrogen Nitrogen fixation is a chemical process by which molecular nitrogen (), with a strong triple covalent bond, in the air is converted into ammonia () or related nitrogenous compounds, typically in soil or aquatic systems but also in industry. Atmos ...
, so is used in agriculture and horticulture, the ''Paenibacillus'' sp. JDR-2 which is a rich source of chemical agents for biotechnology applications, and pattern-forming strains such as '' P. vortex'' and ''P. dendritiformis'' discovered in the early 90s,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: 394-416Ben-Jacob E, Cohen I, Gutnick DL: Cooperative organization of bacterial colonies: from genotype to morphotype. Annu Rev Microbiol 1998, 52:779-806.Ben-Jacob E, Schochet O, Tenenbaum A, Cohen I, Czirok A, Vicsek T: Generic modelling of cooperative growth patterns in bacterial colonies. Nature 1994, 368:46-49.Ben-Jacob E, Shmueli H, Shochet O, Tenenbaum A: Adaptive self-organization during growth of bacterial colonies. Physica A 1992, 187:378-424.Ben-Jacob E, Shochet O, Tenenbaum A, Avidan O: Evolution of complexity during growth of bacterial colonies. In NATO Advanced Research Workshop; Santa Fe, USA. Edited by Cladis PE, Palffy-Muhorey P. Addison-Wesley Publishing Company; 1995: 619-633. which develop complex colonies with intricate architecturesBen-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.Ben-Jacob E, Cohen I, Golding I, Gutnick DL, Tcherpakov M, Helbing D, Ron IG: Bacterial cooperative organization under antibiotic stress. Physica A 2000, 282:247-282.Ben-Jacob E, Cohen I, Levine H: Cooperative self-organization of microorganisms. Adv Phys 2000, 49:395-554.Ben-Jacob E, Levine H: Self-engineering capabilities of bacteria. J R Soc Interface 2005, 3:197-214.Ingham CJ, Ben-Jacob E: Swarming and complex pattern formation in Paenibacillus vortex studied by imaging and tracking cells. BMC Microbiol 2008, 8:36. as shown in the pictures: File:Paenibacillus dendritiformis colony (Chiral morphotype).jpg, A colony generated by the chiral morphotype bacteria of ''P. dendritiformis'': The colony diameter is 5 cm and the colors indicate the bacterial density (bright yellow for high density). The branches are curly with well-defined handedness. File:Paenibacillus vortex colony.jpg, A colony generated by ''P. vortex'' sp. bacteria: The colony diameter is 5 cm and the colors indicate the bacterial density (bright yellow for high density). The bright dots are the vortices described in the text. File:Paenibacillus dendritiformis colony.jpg, A colony generated by the branching (tip splitting) morphotype bacteria of ''P. dendritiformis'': The colony diameter is 6 cm and the colors indicate the bacterial density (darker shade for higher density).


Importance

Interest in ''Paenibacillus'' spp. has been rapidly growing since many were shown to be importantChoi KK, Park CW, Kim SY, Lyoo WS, Lee SH, Lee JW: Polyvinyl alcohol degradation by Microbacterium barkeri KCCM 10507 and Paeniblacillus amylolyticus KCCM 10508 in dyeing wastewater. J Microbiol Biotechnol 2004, 14:1009-1013.Konishi J, Maruhashi K: 2-(2'-Hydroxyphenyl)benzene sulfinate desulfinase from the thermophilic desulfurizing bacterium Paenibacillus sp. strain A11-2: purification and characterization. Appl Microbiol Biotechnol 2003, 62:356-361.Nielsen P, Sorensen J: Multi-target and medium-independent fungal antagonism by hydrolytic enzymes in Paenibacillus polymyxa and Bacillus pumilus strains from barley rhizosphere. Fems Microbiol Ecol 1997, 22:183-192. for agriculture and horticulture (e.g. ''P. polymyxa''), industrial (e.g. ''P. amylolyticus''), and medical applications (e.g. ''P. peoriate''). These bacteria produce various extracellular enzymes such as polysaccharide-degrading enzymes and proteases, which can catalyze a wide variety of synthetic reactions in fields ranging from cosmetics to
biofuel Biofuel is a fuel that is produced over a short time span from biomass, rather than by the very slow natural processes involved in the formation of fossil fuels, such as oil. According to the United States Energy Information Administration (E ...
production. Various ''Paenibacillus'' spp. also produce antimicrobial substances that affect a wide spectrum of micro-organismsGirardin 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.Piuri M, Sanchez-Rivas C, Ruzal SM: A novel antimicrobial activity of a Paenibacillus polymyxa strain isolated from regional fermented sausages. Lett Appl Microbiol 1998, 27:9-13.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. such as fungi, soil bacteria, plant pathogenic bacteria, and even important anaerobic pathogens such as ''
Clostridium botulinum ''Clostridium botulinum'' is a Gram-positive, rod-shaped, anaerobic, spore-forming, motile bacterium with the ability to produce the neurotoxin botulinum. The botulinum toxin can cause botulism, a severe flaccid paralytic disease in humans an ...
''. More specifically, several ''Paenibacillus ''species serve as efficient plant growth-promoting rhizobacteria (PGPR), which competitively colonize plant roots and can simultaneously act as
biofertilizer A biofertilizer is a substance which contains living micro-organisms which, when applied to seeds, plant surfaces, or soil, colonize the rhizosphere or the interior of the plant and promotes growth by increasing the supply or availability of prim ...
s and as antagonists (
biopesticide A Biopesticide is a biological substance or organism that damages, kills, or repels organisms seens as pests. Biological pest management intervention involves predatory, parasitic, or chemical relationships. They are obtained from organisms inclu ...
s) of recognized root pathogens, such as bacteria, fungi, and nematodes.Bloemberg GV, Lugtenberg BJ: Molecular basis of plant growth promotion and biocontrol by rhizobacteria. Curr Opin Plant Biol 2001, 4:343-350. They enhance plant growth by several direct and indirect mechanisms. Direct mechanisms include phosphate solubilization, nitrogen fixation, degradation of environmental pollutants, and hormone production. Indirect mechanisms include controlling phytopathogens by competing for resources such as iron, amino acids and sugars, as well as by producing antibiotics or lytic enzymes.Kloepper JW, Leong J, Teintze M, Schroth MN: Enhanced plant growth by siderophores produced by plant growth-promoting rhizobacteria. Nature 1980, 286:885-886.Ryu CM, Farag MA, Hu CH, Reddy MS, Wei HX, Pare PW, Kloepper JW: Bacterial volatiles promote growth in Arabidopsis. Proc Natl Acad Sci U S A 2003, 100:4927-4932. Competition for iron also serves as a strong selective force determining the microbial population in the rhizosphere. Several studies show that PGPR exert their plant growth-promoting activity by depriving native microflora of iron. Although iron is abundant in nature, the extremely low solubility of Fe at pH 7 means that most organisms face the problem of obtaining enough iron from their environments. To fulfill their requirements for iron, bacteria have developed several strategies, including the reduction of ferric to ferrous ions, the secretion of high-affinity iron-chelating compounds, called
siderophores Siderophores (Greek: "iron carrier") are small, high-affinity iron-chelating compounds that are secreted by microorganisms such as bacteria and fungi. They help the organism accumulate iron. Although a widening range of siderophore functions is no ...
, and the uptake of heterologous siderophores. ''P. vortex's'' genome, for example, harbors many genes which are employed in these strategies, in particular it has the potential to produce siderophores under iron-limiting conditions. Despite the increasing interest in ''Paenibacillus'' spp., genomic information of these bacteria is lacking. More extensive genome sequencing could provide fundamental insights into pathways involved in complex social behavior of bacteria, and can discover a source of genes with biotechnological potential. ''
Candidatus In prokaryote nomenclature, ''Candidatus'' (Latin for candidate of Roman office) is used to name prokaryotic phyla that are well characterized but yet-uncultured. Contemporary sequencing approaches, such as 16S sequencing or metagenomics, provide m ...
'' Paenibacillus glabratella causes white nodules and high mortality of ''
Biomphalaria glabrata ''Biomphalaria glabrata'' is a species of air-breathing freshwater snail, an aquatic pulmonate gastropod mollusk in the family Planorbidae, the ram's horn snails. ''Biomphalaria glabrata'' is an intermediate snail host for the trematode ''Sch ...
'' freshwater snails. This is potentially important because ''Biomphalaria glabrata'' is an intermediate host of
schistosomiasis Schistosomiasis, also known as snail fever, bilharzia, and Katayama fever, is a disease caused by parasitic flatworms called schistosomes. The urinary tract or the intestines may be infected. Symptoms include abdominal pain, diarrhea, bloody s ...
.


Pattern formation, self-organization, and social behaviors

Several ''Paenibacillus'' species can form complex patterns on semisolid surfaces. Development of such complex colonies require
self-organization Self-organization, also called spontaneous order in the social sciences, is a process where some form of overall order arises from local interactions between parts of an initially disordered system. The process can be spontaneous when suffi ...
and cooperative behavior of individual cells while employing sophisticated chemical communication called
quorum sensing In biology, quorum sensing or quorum signalling (QS) is the ability to detect and respond to cell population density by gene regulation. As one example, QS enables bacteria to restrict the expression of specific genes to the high cell densities at ...
.Bassler BL, Losick R: Bacterially speaking. Cell 2006, 125:237-246.Ben-Jacob E, Becker I, Shapira Y, Levine H: Bacterial linguistic communication and social intelligence. Trends Microbiol 2004, 12:366-372.Dunny GM, Brickman TJ, Dworkin M: Multicellular behavior in bacteria: communication, cooperation, competition and cheating. Bioessays 2008, 30:296-298. Pattern formation and self-organization in microbial systems is an intriguing phenomenon and reflects social behaviors of bacteriaGalperin MY, Gomelsky M: Bacterial Signal Transduction Modules: from Genomics to Biology. ASM News 2005, 71:326-333. that might provide insights into the evolutionary development of the collective action of cells in higher organisms.Aguilar C, Vlamakis H, Losick R, Kolter R: Thinking about Bacillus subtilis as a multicellular organism. Curr Opin Microbiol 2007, 10:638-643.Dwyer DJ, Kohanski MA, Collins JJ: Networking opportunities for bacteria. Cell 2008, 135:1153-1156.Kolter R, Greenberg EP: Microbial sciences: the superficial life of microbes. Nature 2006, 441:300-302. Shapiro JA: Thinking about bacterial populations as multicellular organisms. Annu Rev Microbiol 1998, 52:81-104. Shapiro JA, Dworkin M: Bacteria as multicellular organisms. 1st edn: Oxford University Press, USA; 1997.


Pattern forming in ''P. vortex''

One of the most fascinating pattern forming ''Paenibacillus'' species is ''P. vortex'', self-lubricating,
flagella A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have f ...
-driven bacteria. ''P. vortex'' organizes its colonies by generating modules, each consisting of many bacteria, which are used as building blocks for the colony as a whole. The modules are groups of bacteria that move around a common center at about 10 µm/s.


Pattern forming in ''P. dendritiformis''

An additional intriguing pattern forming ''Paenibacillus'' species is ''P. dendritiformis'', which generates two different morphotypes – the branching (or tip-splitting) morphotype and the chiral morphotype that is marked by curly branches with well-defined handedness (see pictures). These two pattern-forming ''Paenibacillus'' strains exhibit many distinct physiological and genetic traits, including
β-galactosidase β-Galactosidase (EC 3.2.1.23, lactase, beta-gal or β-gal; systematic name β-D-galactoside galactohydrolase), is a glycoside hydrolase enzyme that catalyst, catalyzes hydrolysis of terminal non-reducing β-D-galactose residues in β-D-galactosi ...
-like activity causing colonies to turn blue on
X-gal X-gal (also abbreviated BCIG for 5-bromo-4-chloro-3-indolyl-β--galactopyranoside) is an organic compound consisting of galactose linked to a substituted indole. The compound was synthesized by Jerome Horwitz and collaborators in 1964. The formal c ...
plates and
multiple drug resistance Multiple drug resistance (MDR), multidrug resistance or multiresistance is antimicrobial resistance shown by a species of microorganism to at least one antimicrobial drug in three or more antimicrobial categories. Antimicrobial categories are c ...
(MDR) (including septrin,
penicillin Penicillins (P, PCN or PEN) are a group of β-lactam antibiotics originally obtained from ''Penicillium'' moulds, principally '' P. chrysogenum'' and '' P. rubens''. Most penicillins in clinical use are synthesised by P. chrysogenum using ...
,
kanamycin Kanamycin A, often referred to simply as kanamycin, is an antibiotic used to treat severe bacterial infections and tuberculosis. It is not a first line treatment. It is used by mouth, injection into a vein, or injection into a muscle. Kanamycin ...
,
chloramphenicol Chloramphenicol is an antibiotic useful for the treatment of a number of bacterial infections. This includes use as an eye ointment to treat conjunctivitis. By mouth or by injection into a vein, it is used to treat meningitis, plague, cholera, a ...
,
ampicillin Ampicillin is an antibiotic used to prevent and treat a number of bacterial infections, such as respiratory tract infections, urinary tract infections, meningitis, salmonellosis, and endocarditis. It may also be used to prevent group B strepto ...
,
tetracycline Tetracycline, sold under various brand names, is an oral antibiotic in the tetracyclines family of medications, used to treat a number of infections, including Acne vulgaris, acne, cholera, brucellosis, plague (disease), plague, malaria, and sy ...
,
spectinomycin Spectinomycin, sold under the tradename Trobicin among others, is an antibiotic useful for the treatment of gonorrhea infections. It is given by injection into a muscle. Common side effects include pain at the area of injection, rash, nausea, ...
,
streptomycin Streptomycin is an antibiotic medication used to treat a number of bacterial infections, including tuberculosis, ''Mycobacterium avium'' complex, endocarditis, brucellosis, ''Burkholderia'' infection, plague, tularemia, and rat bite fever. Fo ...
, and
mitomycin C Mitomycin C is a mitomycin that is used as a chemotherapeutic agent by virtue of its antitumour activity. Medical uses It is given intravenously to treat upper gastro-intestinal cancers (e.g. esophageal carcinoma), anal cancers, and breast c ...
). Colonies that are grown on surfaces in Petri dishes exhibit several-fold higher drug resistance in comparison to growth in liquid media. This particular resistance is believed to be due to a
surfactant Surfactants are chemical compounds that decrease the surface tension between two liquids, between a gas and a liquid, or interfacial tension between a liquid and a solid. Surfactants may act as detergents, wetting agents, emulsifiers, foaming ...
-like liquid front that actually forms a particular pattern on the Petri plate.


References


External links


''Paenibacillus'' TaxonomyGenome sequence of the pattern forming ''Paenibacillus vortex'' bacterium reveals potential for thriving in complex environments - manuscriptProf. Eshel Ben-Jacob home pageSpecific PCR for Paenibacillus genus based on rpoB geneUse of rpoB gene analysis for identification of nitrogen-fixing Paenibacillus species as an alternative to the 16S rRNA gene''Paenibacillus'' at Bac''Dive'' - the Bacterial Diversity Metadatabase
{{Taxonbar, from=Q1952884 Paenibacillaceae Bacteria genera