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system for exchange of DNA) and Ted (Thermoproteales system for exchange of DNA), appears to be responsible for the transfer of cellular DNA between members of the same species. It has been suggested that in these archaea the conjugation machinery has been fully domesticated for promoting DNA repair through homologous recombination rather than spread of mobile genetic elements.
38:
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376:
674:. This is because the presence of pili greatly enhances bacteria's ability to bind to body tissues, which then increases replication rates and ability to interact with the host organism. If a species of bacteria has multiple strains but only some are pathogenic, it is likely that the pathogenic strains will have pili while the nonpathogenic strains do not.
145:. They are also fragile and constantly replaced, sometimes with pili of different composition, resulting in altered antigenicity. Specific host responses to old pili structures are not effective on the new structure. Recombination between genes of some (but not all) pili code for variable (V) and constant (C) regions of the pili (similar to
243:. The connection established by the F-pilus is extremely mechanically and thermochemically resistant thanks to the robust properties of the F-pilus, which ensures successful gene transfer in a variety of environments. Not all bacteria can make conjugative pili, but conjugation can occur between bacteria of different species.
578:
This family was originally identified as "type IV fimbriae" by their appearance under the microscope. This classification survived as it happens to correspond to a clade. It has been shown that some archaeal type IV pilins can exist in 4 different conformations, yielding two pili with dramatically
311:
This appendage ranges from 3–10 nanometers in diameter and can be as much as several micrometers long. Fimbriae are used by bacteria to adhere to one another and to adhere to animal cells and some inanimate objects. A bacterium can have as many as 1,000 fimbriae. Fimbriae are only visible with the
259:
encode pili structurally similar to the bacterial conjugative pili. However, unlike in bacteria, where conjugation apparatus typically mediates the transfer of mobile genetic elements, such as plasmids or transposons, the conjugative machinery of hyperthermophilic archaea, called Ced (Crenarchaeal
608:
adhesions found on the bacteria's fimbriae. This process of bacteria adhering to a host cell can result in the colonization of that host cell as more and more bacteria collect around it, and is integral to the continued survival of the bacteria, enabling them to infect tissues and entire organs.
607:
This figure depicts fimbriae adhesion. In this process the fimbriae of a bacterial cell (right) adhere to specific proteins, called receptors, found on the outer membrane of a host cell (left). They do this by a specific interaction between the receptors of the host cell and the perfectly matched
511:
forces. The external ends of the pili adhere to a solid substrate, either the surface to which the bacterium is attached or to other bacteria. Then, when the pili contract, they pull the bacterium forward like a grappling hook. Movement produced by type IV pili is typically jerky, so it is called
250:
Proposed conjugation mechanisms between donor and recipient cells in archaea (left) and bacteria (right). The schematic shows how ssDNA substrates are generated by the HerA-NurA machinery in the donor archaeal cells and by the plasmid-encoded relaxosome in bacteria. The figure is reproduced from
542:
systems. Besides archaella, many archaea produce adhesive type 4 pili, which enable archaeal cells to adhere to different substrates. The N-terminal alpha-helical portions of the archaeal type 4 pilins and archaellins are homologous to the corresponding regions of bacterial T4P; however, the
161:, there has been much interest in the study of pili as organelle of adhesion and as vaccine components. The first detailed study of pili was done by Brinton and co-workers who demonstrated the existence of two distinct phases within one bacterial strain: pileated (p+) and non-pileated)
1070:
Beltran, Leticia C.; Cvirkaite-Krupovic, Virginija; Miller, Jessalyn; Wang, Fengbin; Kreutzberger, Mark A. B.; Patkowski, Jonasz B.; Costa, Tiago R. D.; Schouten, Stefan; Levental, Ilya; Conticello, Vincent P.; Egelman, Edward H.; Krupovic, Mart (2023-02-07).
571:. Menningococcal type IV pili bind DNA through the minor pilin ComP via an electropositive stripe that is predicted to be exposed on the filament's surface. ComP displays an exquisite binding preference for selective DUSs. The distribution of DUSs within the
231:, which establishes direct contact and the formation of a controlled pore that allows transfer of DNA from the donor to the recipient. Typically, the DNA transferred consists of the genes required to make and transfer pili (often encoded on a
579:
different structures. Remarkably, the two pili were produced by the same secretion machinery. However, which of the two pili is formed appears to depend on the growth conditions, suggesting that the two pili are functionally distinct.
481:
The secretin protein, PilQ, found on the outer membrane of the cell is necessary for the development/extension of the pilus. PilC is the first proteins to form the pilus and are responsible for overall attachment of the pilus.
486:
Once the Type IV Pilus attaches or interacts with what it needs to, it begins to retract. This occurs with the PilT beginning to degrade the last parts of the PilA in the pilus. The mechanism of PilT is very similar to PilF.
896:
Patkowski, Jonasz B.; Dahlberg, Tobias; Amin, Himani; Gahlot, Dharmender K.; Vijayrajratnam, Sukhithasri; Vogel, Joseph P.; Francis, Matthew S.; Baker, Joseph L.; Andersson, Magnus; Costa, Tiago R. D. (5 April 2023).
1005:
Gschwind, Rémi; Petitjean, Marie; Fournier, Claudine; Lao, Julie; Clermont, Olivier; Nordmann, Patrice; Mellmann, Alexander; Denamur, Erick; Poirel, Laurent; Ruppé, Etienne (2024-04-03). Uhlemann, Anne-Catrin (ed.).
1974:
Craig, Lisa; Taylor, Ronald (2014). "Chapter 1: The Vibrio cholerae Toxin
Coregulated Pilus: Structure, Assembly, and Function with Implications for Vaccine Design". In Barocchi, Michèle; Telford, John (eds.).
2000:
Rinaudo, Daniela; Moschioni, Monica (2014). "Chapter 13: Pilus-based
Vaccine Development in Streptococci: Variability, Diversity, and Immunological Resposes". In Barocchi, Michèle; Telford, John (eds.).
352:, consists of many aerobic bacteria that adhere to the surface by their fimbriae. Thus, fimbriae allow the aerobic bacteria to remain both on the broth, from which they take nutrients, and near the air.
767:
149:
diversity). As the primary antigenic determinants, virulence factors and impunity factors on the cell surface of a number of species of Gram negative and some Gram positive bacteria, including
437:(T4SS). They can be classified into the F-like type (after the F-pilus) and the P-like type. Like their secretion counterparts, the pilus injects material, DNA in this case, into another cell.
719:. The gene for this toxin, once incorporated into the bacterium's genome, is expressed when the gene coding for the pilus is expressed (hence the name "toxin mediated pilus").
359:, as they attach bacteria to host surfaces for colonization during infection. Fimbriae are either located at the poles of a cell or are evenly spread over its entire surface.
1443:
Wang, F; Cvirkaite-Krupovic, V; Kreutzberger, MAB; Su, Z; de
Oliveira, GAP; Osinski, T; Sherman, N; DiMaio, F; Wall, JS; Prangishvili, D; Krupovic, M; Egelman, EH (2019).
227:
in diameter. During conjugation, a pilus emerging from the donor bacterium ensnares the recipient bacterium, draws it in close, and eventually triggers the formation of a
219:. Bacteria produce long extracellular appendages called sex pili, which connect two neighbouring cells and serve as a physical conduit for transfer of DNA. Adapted from
1901:
169:
A few names are given to different types of pili by their function. The classification does not always overlap with the structural or evolutionary-based types, as
1749:
Cookson, AL; Cooley, WA; Woodward, MJ (2002), "The role of type 1 and curli fimbriae of Shiga toxin-producing
Escherichia coli in adherence to abiotic surfaces",
300:. The term "fimbria" can refer to many different (structural) types of pilus. Indeed, many different types of pili have been used for adhesion, a case of
239:; however, other pieces of DNA are often co-transferred and this can result in dissemination of genetic traits throughout a bacterial population, such as
2057:
Georgiadou, Michaella; Pelicic, Vladimir (2014). "Chapter 5: Type IV Pili: Functions & Biogenesis". In
Barocchi, Michèle; Telford, John (eds.).
677:
The development of attachment pili may then result in the development of further virulence traits. Fimbriae are one of the primary mechanisms of
829:"Proteinaceous determinants of surface colonization in bacteria: bacterial adhesion and biofilm formation from a protein secretion perspective"
2066:
2010:
1984:
1427:
772:
1920:
Epstein, EA; Reizian, MA; Chapman, MR (2009), "Spatial clustering of the curlin secretion lipoprotein requires curli fiber assembly.",
2456:
2196:
785:
Brinton, Charles (1954). "Electrophoresis and phage susceptibility studies on a filament-producing variant of the E. coli bacterium".
296:) is a term used for a short pilus that is used to attach the bacterium to a surface, sometimes also called an "attachment pilus" or
2794:
1359:
193:, because they allow for the exchange of genes via the formation of "mating pairs". Perhaps the most well-studied is the F-pilus of
701:
bacteria. Their presence greatly enhances the bacteria's ability to attach to the host and cause disease. Nonpathogenic strains of
549:
is the process by which a recipient bacterial cell takes up DNA from a neighboring cell and integrates this DNA into its genome by
1492:
Cehovin A, Simpson PJ, McDowell MA, Brown DR, Noschese R, Pallett M, Brady J, Baldwin GS, Lea SM, Matthews SJ, Pelicic V (2013).
392:
899:"The F-pilus biomechanical adaptability accelerates conjugative dissemination of antimicrobial resistance and biofilm formation"
2446:
2373:
2570:
2301:
2689:
362:
This term was also used in a lax sense to refer to all pili, by those who use "pilus" to specifically refer to sex pili.
57:
Both cells recircularize their plasmids, synthesize second strands, and reproduce pili; both cells are now viable donors.
878:
469:, removes a leader sequence, thus making the Pre-PilA shorter and into PilA, the main building-block protein of Pili.
1697:
Liu, J; Eastep, GN; Cvirkaite-Krupovic, V; Rich-New, ST; Kreutzberger, MAB; Egelman, EH; Krupovic, M; Wang, F (2024).
200:
2411:
575:
genome favors certain genes, suggesting that there is a bias for genes involved in genomic maintenance and repair.
308:
system does not treat fimbriae as a distinct type of appendage, using the generic pilus (GO:0009289) type instead.
948:"Phylum barrier and Escherichia coli intra-species phylogeny drive the acquisition of antibiotic-resistance genes"
495:
546:
2565:
2488:
2451:
2247:
2168:
2158:
2148:
620:." Curli are a type of fimbriae. Curli are composed of proteins called curlins. Some of the genes involved are
550:
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131:
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91:
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104:
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1505:
1153:
1084:
689:
301:
170:
1837:"Pyelonephritic Escherichia coli expressing P fimbriae decrease immune response of the mouse kidney"
1354:. Foster, John Watkins (Fourth ed.). New York: W. W. Norton & Company. pp. 1000–1002.
603:
591:
is responsible for moving many types of fimbriae out of the cell, including type 1 fimbriae and the
2784:
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2531:
2526:
2419:
2389:
2242:
2227:
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560:
313:
190:
53:
The mobile plasmid is nicked and a single strand of DNA is then transferred to the recipient cell.
1209:"Pili in Gram-negative and Gram-positive bacteria — structure, assembly and their role in disease"
2676:
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2384:
2296:
2289:
1248:
522:
513:
946:
Petitjean, Marie; Condamine, Bénédicte; Burdet, Charles; Denamur, Erick; Ruppé, Etienne (2021).
879:"Gut bacteria use super-polymers to dodge antibiotics | Imperial News | Imperial College London"
1786:"Everything You Always Wanted to Know About Salmonella Type 1 Fimbriae, but Were Afraid to Ask"
2789:
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320:
31:
2164:
543:
C-terminal beta-strand-rich domains appear to be unrelated in bacterial and archaeal pilins.
491:
Degradation of the pilus into the components to be utilized and synthesized into PilA again.
2468:
2344:
2279:
2113:
2103:
1937:
1929:
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The Tra (transfer) family includes all known sex pili (as of 2010). They are related to the
341:
334:
273:
211:
195:
2154:
1396:
122:
Dozens of these structures can exist on the bacterial and archaeal surface. Some bacteria,
2694:
2593:
2478:
2354:
2284:
2237:
1877:
1835:
Rice JC, Peng T, Spence JS, Wang HQ, Goldblum RM, Corthésy B, Nowicki BJ (December 2005).
1699:"Two distinct archaeal type IV pili structures formed by proteins with identical sequence"
702:
664:
112:
1008:"Inter-phylum circulation of a beta-lactamase-encoding gene: a rare but observable event"
656:
Pili are responsible for virulence in the pathogenic strains of many bacteria, including
2099:
1723:
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1509:
1279:
1235:
1208:
1157:
1088:
1040:
1007:
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401:
Please expand the section to include this information. Further details may exist on the
2349:
2313:
1942:
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1671:
1647:"Evolution of the chaperone/usher assembly pathway: fimbrial classification goes Greek"
1646:
1622:
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1444:
1327:
1302:
1184:
1141:
1113:
1072:
982:
947:
855:
828:
587:
Another type are called type 1 fimbriae. They contain FimH adhesins at the "tips". The
146:
2084:"Type 1 fimbrial expression enhances Escherichia coli virulence for the urinary tract"
1577:
1552:
206:
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2703:
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305:
228:
127:
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135:
1613:
1252:
349:
246:
2621:
2082:
Connell I, Agace W, Klemm P, Schembri M, Mărild S, Svanborg C (September 1996).
2028:"Textbook of Bacteriology: Bacterial Structure in Relationship to Pathogenicity"
1073:"Archaeal DNA-import apparatus is homologous to bacterial conjugation machinery"
748:
708:
559:(also called meningococcus), DNA transformation requires the presence of short
324:
323:
which attach them to some sort of substratum so that the bacteria can withstand
236:
1714:
1146:
Proceedings of the
National Academy of Sciences of the United States of America
1096:
914:
530:. Bacterial type IV pili are similar in structure to the component proteins of
268:
2728:
2713:
2631:
2364:
2027:
1553:"Biased distribution of DNA uptake sequences towards genome maintenance genes"
1460:
1225:
743:
592:
531:
402:
1802:
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1031:
973:
845:
17:
2723:
2554:
2512:
2424:
2318:
2108:
1762:
1518:
1266:
Ottow, JC (1975). "Ecology, physiology, and genetics of fimbriae and pili".
1166:
733:
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224:
78:
1951:
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1821:
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1537:
1478:
1404:
1336:
1244:
1193:
1122:
1049:
991:
932:
864:
806:
516:, as opposed to other forms of bacterial motility such as that produced by
375:
2127:
1662:
1287:
964:
99:) can be used interchangeably, although some researchers reserve the term
2232:
2209:
2174:
1568:
1551:
Davidsen T, Rødland EA, Lagesen K, Seeberg E, Rognes T, Tønjum T (2004).
1023:
567:
of the donor DNA. Specific recognition of DUSs is mediated by a type IV
508:
466:
116:
1933:
1445:"An extensively glycosylated archaeal pilus survives extreme conditions"
1318:
37:
2662:
2626:
2262:
738:
613:
356:
329:
256:
232:
142:
82:
1602:"Meningococcal carriage and disease--population biology and evolution"
2708:
2463:
1784:
Kolenda, Rafal; Ugorski, Maciej; Grzymajlo, Krzysztof (14 May 2019).
457:
Pre-PilA is made in the cytoplasm and moves into the inner membrane.
707:
first evolved pili, allowing them to bind to human tissues and form
1303:"A variety of bacterial pili involved in horizontal gene transfer"
827:
Chagnot, C; Zorgani, MA; Astruc, T; Desvaux, M (14 October 2013).
617:
602:
568:
494:
477:-Binding protein that provides energy for Type IV Pili Assembly.
396:
285:
267:
245:
205:
123:
108:
66:
36:
49:
Pilus attaches to recipient cell, brings the two cells together.
2178:
1136:
van
Wolferen, Marleen; Wagner, Alexander; van der Does, Chris;
369:
182:
27:
A proteinaceous hair-like appendage on the surface of bacteria
1418:
Jarrell; et al. (2009). "Archaeal
Flagella and Pili".
1383:
Mattick JS (2002). "Type IV pili and twitching motility".
2059:
Bacterial Pili: Structure, Synthesis, and Role in
Disease
2003:
Bacterial Pili: Structure, Synthesis, and Role in
Disease
1977:
Bacterial Pili: Structure, Synthesis, and Role in Disease
77:) is a hair-like appendage found on the surface of many
1494:"Specific DNA recognition mediated by a type IV pilin"
189:. They are sometimes called "sex pili", in analogy to
1420:
Pili and Flagella: Current Research and Future Trends
2675:
2655:
2607:
2496:
2487:
2439:
2332:
2270:
2256:
2216:
1692:
1690:
1878:"Colonization and Invasion by Bacterial Pathogens"
711:. These pili then served as binding sites for the
107:. All conjugative pili are primarily composed of
534:(archaeal flagella), and both are related to the
1915:
1913:
1911:
1744:
1742:
461:Pre-PilA is inserted into the inner membrane.
822:
820:
818:
816:
399:), LPXTG including type 3 pilus (T3P; spaHIG).
2190:
1841:Journal of the American Society of Nephrology
8:
1900:: CS1 maint: multiple names: authors list (
41:Schematic drawing of bacterial conjugation.
563:(DUSs) which are 9-10 monomers residing in
393:extracellular nucleation-precipitation pili
355:Fimbriae are required for the formation of
344:form a very thin layer at the surface of a
181:Conjugative pili allow for the transfer of
2493:
2267:
2197:
2183:
2175:
2005:. C.A.B. International. pp. 182–202.
1651:Microbiology and Molecular Biology Reviews
499:Type IVa pilus machine architectural model
2167:at the U.S. National Library of Medicine
2157:at the U.S. National Library of Medicine
2147:at the U.S. National Library of Medicine
2117:
2107:
1941:
1852:
1811:
1801:
1722:
1670:
1621:
1576:
1527:
1517:
1468:
1326:
1234:
1224:
1207:Proft, T.; Baker, E. N. (February 2009).
1183:
1165:
1112:
1039:
981:
963:
922:
854:
844:
538:(T2SS); they are unified by the group of
2061:. C.A.B. International. pp. 71–84.
444:
1979:. C.A.B. International. pp. 1–16.
1352:Microbiology : an evolving science
760:
366:Types by assembling system or structure
1893:
1397:10.1146/annurev.micro.56.012302.160938
520:. However, some bacteria, for example
387:about mention of other types: various
2052:
2050:
2048:
1969:
1967:
1965:
1963:
1961:
1142:"The archaeal Ced system imports DNA"
1012:Antimicrobial Agents and Chemotherapy
7:
2759:
1213:Cellular and Molecular Life Sciences
1065:
1063:
1061:
1059:
327:and obtain nutrients. For example,
316:. They may be straight or flexible.
185:between bacteria, in the process of
1280:10.1146/annurev.mi.29.100175.000455
289:
70:
25:
715:that carries the disease-causing
612:"Gram-negative bacteria assemble
2758:
2749:
2748:
451:Type IV Pilus Twitching Motility
374:
223:A sex pilus is typically 6 to 7
2447:Bacterial cellular morphologies
1645:Nuccio SP, et al. (2007).
103:for the appendage required for
1882:www.textbookofbacteriology.net
1600:Caugant DA, Maiden MC (2009).
134:on pili at the start of their
1:
1614:10.1016/j.vaccine.2009.04.061
1268:Annual Review of Microbiology
787:Biochimica et Biophysica Acta
95:(Latin for 'fringe'; plural:
30:For the sea snail genus, see
2088:Proc. Natl. Acad. Sci. U.S.A
1876:WI, Kenneth Todar, Madison.
1498:Proc. Natl. Acad. Sci. U.S.A
799:10.1016/0006-3002(54)90011-6
773:Dorland's Medical Dictionary
45:Donor cell produces pilus.
2811:
2690:Bacteria (classifications)
2412:Primary nutritional groups
1715:10.1038/s41467-024-45062-z
1608:. 27 Suppl 2 (4): B64–70.
1422:. Caister Academic Press.
1350:Joan, Slonczewski (2017).
1097:10.1038/s41467-023-36349-8
915:10.1038/s41467-023-37600-y
426:
29:
2744:
1790:Frontiers in Microbiology
1461:10.1038/s41564-019-0458-x
1226:10.1007/s00018-008-8477-4
833:Frontiers in Microbiology
395:built by T8SS (including
2795:Prokaryotic cell anatomy
2566:Bacterial outer membrane
2169:Medical Subject Headings
2159:Medical Subject Headings
2149:Medical Subject Headings
2032:Textbook of Bacteriology
1803:10.3389/fmicb.2019.01017
1301:Filloux, A (July 2010).
846:10.3389/fmicb.2013.00303
551:homologous recombination
536:Type II secretion system
435:type IV secretion system
389:chaperone-Usher fimbriae
2109:10.1073/pnas.93.18.9827
1763:10.1078/1438-4221-00203
1519:10.1073/pnas.1218832110
1307:Journal of Bacteriology
1167:10.1073/pnas.1513740113
713:lysogenic bacteriophage
589:chaperone-usher pathway
348:. This layer, called a
333:uses them to attach to
235:), and so is a kind of
2561:Gram-negative bacteria
2540:Gram-positive bacteria
1854:10.1681/ASN.2005030243
668:, and many strains of
616:surface fibers called
609:
556:Neisseria meningitidis
547:Genetic transformation
500:
492:
385:is missing information
278:
252:
220:
58:
2416:Substrate preference
1703:Nature Communications
1663:10.1128/MMBR.00014-07
1077:Nature Communications
965:10.1099/mgen.0.000489
903:Nature Communications
606:
498:
448:
271:
249:
241:antibiotic resistance
209:
187:bacterial conjugation
105:bacterial conjugation
40:
2397:Microbial metabolism
1385:Annu. Rev. Microbiol
1138:Albers, Sonja-Verena
1024:10.1128/aac.01459-23
690:Bordetella pertussis
561:DNA uptake sequences
302:convergent evolution
171:convergent evolution
2647:Non-motile bacteria
2243:Pathogenic bacteria
2100:1996PNAS...93.9827C
1934:10.1128/JB.01244-08
1751:Int J Med Microbiol
1510:2013PNAS..110.3065C
1449:Nature Microbiology
1319:10.1128/JB.00424-10
1158:2016PNAS..113.2496V
1089:2023NatCo..14..666B
729:Bacterial nanowires
314:electron microscope
191:sexual reproduction
2576:Lipopolysaccharide
1569:10.1093/nar/gkh255
952:Microbial Genomics
614:functional amyloid
610:
523:Myxococcus xanthus
514:twitching motility
503:Some pili, called
501:
493:
279:
255:Hyperthermophilic
253:
221:
151:Enterobacteriaceae
59:
2772:
2771:
2671:
2670:
2617:Bacterial capsule
2583:Periplasmic space
2550:Lipoteichoic acid
2435:
2434:
2407:Microbial ecology
2402:Nitrogen fixation
2165:Fimbriae+Proteins
2068:978-1-78064-255-0
2012:978-1-78064-255-0
1986:978-1-78064-255-0
1557:Nucleic Acids Res
1429:978-1-904455-48-6
420:
419:
335:mannose receptors
319:Fimbriae possess
199:, encoded by the
165:Types by function
32:Pilus (gastropod)
16:(Redirected from
2802:
2762:
2761:
2752:
2751:
2700:Former groupings
2494:
2345:Human microbiome
2268:
2199:
2192:
2185:
2176:
2132:
2131:
2121:
2111:
2079:
2073:
2072:
2054:
2043:
2042:
2040:
2038:
2026:Todar, Kenneth.
2023:
2017:
2016:
1997:
1991:
1990:
1971:
1956:
1955:
1945:
1917:
1906:
1905:
1899:
1891:
1889:
1888:
1873:
1867:
1866:
1856:
1832:
1826:
1825:
1815:
1805:
1781:
1775:
1774:
1757:(3–4): 195–205,
1746:
1737:
1736:
1726:
1694:
1685:
1684:
1674:
1642:
1636:
1635:
1625:
1597:
1591:
1590:
1580:
1548:
1542:
1541:
1531:
1521:
1489:
1483:
1482:
1472:
1455:(8): 1401–1410.
1440:
1434:
1433:
1415:
1409:
1408:
1380:
1374:
1373:
1347:
1341:
1340:
1330:
1298:
1292:
1291:
1263:
1257:
1256:
1238:
1228:
1204:
1198:
1197:
1187:
1169:
1152:(9): 2496–2501.
1133:
1127:
1126:
1116:
1067:
1054:
1053:
1043:
1002:
996:
995:
985:
967:
943:
937:
936:
926:
893:
887:
886:
875:
869:
868:
858:
848:
824:
811:
810:
782:
776:
765:
540:Type IV filament
528:gliding motility
507:(T4P), generate
415:
412:
406:
378:
370:
342:aerobic bacteria
291:
274:Escherichia coli
212:Escherichia coli
196:Escherichia coli
177:Conjugative pili
155:Pseudomonadaceae
113:fibrous proteins
72:
21:
2810:
2809:
2805:
2804:
2803:
2801:
2800:
2799:
2775:
2774:
2773:
2768:
2740:
2695:Bacterial phyla
2679:
2667:
2651:
2609:
2603:
2594:Arabinogalactan
2499:
2483:
2431:
2328:
2272:
2260:
2252:
2238:Lysogenic cycle
2219:
2212:
2203:
2155:Bacterial+Pilus
2141:
2136:
2135:
2094:(18): 9827–32.
2081:
2080:
2076:
2069:
2056:
2055:
2046:
2036:
2034:
2025:
2024:
2020:
2013:
1999:
1998:
1994:
1987:
1973:
1972:
1959:
1919:
1918:
1909:
1892:
1886:
1884:
1875:
1874:
1870:
1847:(12): 3583–91.
1834:
1833:
1829:
1783:
1782:
1778:
1748:
1747:
1740:
1696:
1695:
1688:
1644:
1643:
1639:
1599:
1598:
1594:
1550:
1549:
1545:
1491:
1490:
1486:
1442:
1441:
1437:
1430:
1417:
1416:
1412:
1382:
1381:
1377:
1362:
1349:
1348:
1344:
1300:
1299:
1295:
1265:
1264:
1260:
1206:
1205:
1201:
1135:
1134:
1130:
1069:
1068:
1057:
1018:(4): e0145923.
1004:
1003:
999:
945:
944:
940:
895:
894:
890:
877:
876:
872:
826:
825:
814:
784:
783:
779:
766:
762:
757:
725:
665:Vibrio cholerae
654:
601:
585:
583:Type 1 fimbriae
573:N. meningitides
443:
431:
425:
416:
410:
407:
400:
391:built by T7SS,
379:
368:
266:
179:
167:
35:
28:
23:
22:
15:
12:
11:
5:
2808:
2806:
2798:
2797:
2792:
2787:
2777:
2776:
2770:
2769:
2767:
2766:
2756:
2745:
2742:
2741:
2739:
2738:
2737:
2736:
2731:
2726:
2721:
2711:
2706:
2697:
2692:
2686:
2684:
2673:
2672:
2669:
2668:
2666:
2665:
2659:
2657:
2653:
2652:
2650:
2649:
2644:
2639:
2634:
2629:
2624:
2619:
2613:
2611:
2605:
2604:
2602:
2601:
2596:
2585:
2580:
2579:
2578:
2573:
2557:
2552:
2547:
2536:
2535:
2534:
2529:
2524:
2510:
2504:
2502:
2491:
2485:
2484:
2482:
2481:
2476:
2471:
2466:
2461:
2460:
2459:
2454:
2452:cell structure
2443:
2441:
2437:
2436:
2433:
2432:
2430:
2429:
2428:
2427:
2425:Saccharophilic
2422:
2414:
2409:
2404:
2399:
2394:
2393:
2392:
2387:
2382:
2377:
2367:
2362:
2357:
2352:
2342:
2336:
2334:
2330:
2329:
2327:
2326:
2321:
2316:
2314:Microaerophile
2311:
2310:
2309:
2304:
2294:
2293:
2292:
2287:
2276:
2274:
2265:
2254:
2253:
2251:
2250:
2245:
2240:
2235:
2230:
2224:
2222:
2214:
2213:
2204:
2202:
2201:
2194:
2187:
2179:
2173:
2172:
2162:
2152:
2140:
2139:External links
2137:
2134:
2133:
2074:
2067:
2044:
2018:
2011:
1992:
1985:
1957:
1928:(2): 608–615,
1907:
1868:
1827:
1776:
1738:
1686:
1657:(4): 551–575.
1637:
1592:
1543:
1504:(8): 3065–70.
1484:
1435:
1428:
1410:
1391:(1): 289–314.
1375:
1360:
1342:
1313:(13): 3243–5.
1293:
1258:
1219:(4): 613–635.
1199:
1140:(2016-03-01).
1128:
1055:
997:
938:
888:
870:
812:
793:(4): 533–542.
777:
759:
758:
756:
753:
752:
751:
746:
741:
736:
731:
724:
721:
695:Staphylococcus
653:
650:
600:
597:
584:
581:
565:coding regions
442:
439:
427:Main article:
424:
421:
418:
417:
382:
380:
373:
367:
364:
298:adhesive pilus
288:for 'fringe',
265:
262:
178:
175:
166:
163:
147:immunoglobulin
128:bacteriophages
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
2807:
2796:
2793:
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2715:
2712:
2710:
2707:
2705:
2704:Schizomycetes
2701:
2698:
2696:
2693:
2691:
2688:
2687:
2685:
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2569:
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2558:
2556:
2553:
2551:
2548:
2546:
2545:Teichoic acid
2543:
2541:
2537:
2533:
2530:
2528:
2525:
2523:
2520:
2519:
2518:
2517:Peptidoglycan
2514:
2511:
2509:
2508:Cell membrane
2506:
2505:
2503:
2501:
2495:
2492:
2490:
2486:
2480:
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2275:
2269:
2266:
2264:
2259:
2255:
2249:
2246:
2244:
2241:
2239:
2236:
2234:
2231:
2229:
2226:
2225:
2223:
2221:
2215:
2211:
2207:
2200:
2195:
2193:
2188:
2186:
2181:
2180:
2177:
2170:
2166:
2163:
2160:
2156:
2153:
2150:
2146:
2143:
2142:
2138:
2129:
2125:
2120:
2115:
2110:
2105:
2101:
2097:
2093:
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2060:
2053:
2051:
2049:
2045:
2033:
2029:
2022:
2019:
2014:
2008:
2004:
1996:
1993:
1988:
1982:
1978:
1970:
1968:
1966:
1964:
1962:
1958:
1953:
1949:
1944:
1939:
1935:
1931:
1927:
1923:
1916:
1914:
1912:
1908:
1903:
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1615:
1611:
1607:
1603:
1596:
1593:
1588:
1584:
1579:
1574:
1570:
1566:
1563:(3): 1050–8.
1562:
1558:
1554:
1547:
1544:
1539:
1535:
1530:
1525:
1520:
1515:
1511:
1507:
1503:
1499:
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1471:
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1001:
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984:
979:
975:
971:
966:
961:
958:(8): 000489.
957:
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942:
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934:
930:
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920:
916:
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908:
904:
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892:
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883:Imperial News
880:
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871:
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852:
847:
842:
838:
834:
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821:
819:
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764:
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747:
745:
742:
740:
737:
735:
732:
730:
727:
726:
722:
720:
718:
714:
710:
709:microcolonies
706:
705:
700:
699:Streptococcus
696:
692:
691:
686:
685:
680:
675:
673:
672:
671:Streptococcus
667:
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619:
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447:
440:
438:
436:
430:
429:Transfer gene
422:
414:
411:December 2020
404:
398:
394:
390:
386:
383:This section
381:
377:
372:
371:
365:
363:
360:
358:
353:
351:
347:
346:broth culture
343:
338:
336:
332:
331:
326:
322:
317:
315:
309:
307:
306:Gene Ontology
303:
299:
295:
287:
283:
276:
275:
270:
263:
261:
258:
248:
244:
242:
238:
234:
230:
229:mating bridge
226:
218:
214:
213:
208:
204:
202:
198:
197:
192:
188:
184:
176:
174:
172:
164:
162:
160:
159:Neisseriaceae
156:
152:
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144:
139:
137:
133:
129:
125:
120:
118:
114:
110:
106:
102:
98:
94:
93:
88:
85:. The terms
84:
80:
76:
68:
64:
56:
52:
48:
44:
39:
33:
19:
18:Type IV pilus
2734:Mendosicutes
2719:Gracilicutes
2699:
2636:
2599:Mycolic acid
2589:Mycobacteria
2587:
2559:
2538:
2474:Coccobacilli
2374:in pregnancy
2340:Extremophile
2324:Aerotolerant
2258:Biochemistry
2220:microbiology
2206:Microbiology
2091:
2087:
2077:
2058:
2035:. Retrieved
2031:
2021:
2002:
1995:
1976:
1925:
1921:
1885:. Retrieved
1881:
1871:
1844:
1840:
1830:
1793:
1789:
1779:
1754:
1750:
1706:
1702:
1654:
1650:
1640:
1605:
1595:
1560:
1556:
1546:
1501:
1497:
1487:
1452:
1448:
1438:
1419:
1413:
1388:
1384:
1378:
1351:
1345:
1310:
1306:
1296:
1271:
1267:
1261:
1216:
1212:
1202:
1149:
1145:
1131:
1080:
1076:
1015:
1011:
1000:
955:
951:
941:
906:
902:
891:
882:
873:
836:
832:
790:
786:
780:
771:
763:
703:
698:
694:
688:
682:
676:
669:
663:
658:
655:
645:
641:
637:
633:
629:
625:
621:
611:
586:
577:
572:
554:
545:
521:
505:type IV pili
504:
502:
488:
483:
478:
470:
462:
458:
454:
450:
449:
441:Type IV pili
432:
408:
384:
361:
354:
339:
328:
325:shear forces
318:
310:
297:
293:
281:
280:
272:
254:
222:
210:
201:F sex factor
194:
180:
168:
158:
154:
150:
140:
136:reproductive
121:
115:, which are
100:
96:
90:
86:
74:
69:for 'hair';
62:
60:
54:
50:
46:
42:
2622:Slime layer
2302:Facultative
2290:Facultative
2037:24 November
1922:J Bacteriol
1709:(1): 5049.
909:(1): 1879.
749:PilZ domain
704:V. cholerae
237:selfish DNA
217:conjugation
215:undergoing
2785:Organelles
2779:Categories
2729:Mollicutes
2724:Firmicutes
2714:Prokaryota
2632:Glycocalyx
2457:plasticity
2420:Lipophilic
2273:preference
2248:Resistance
1887:2016-12-03
1274:: 79–108.
1083:(1): 666.
755:References
744:P fimbriae
593:P fimbriae
526:, exhibit
312:use of an
130:attach to
117:oligomeric
2682:evolution
2656:Composite
2555:Endospore
2513:Cell wall
2489:Structure
2380:Placental
2319:Nanaerobe
2297:Anaerobic
2228:Infection
2145:Sex+Pilus
1370:951925510
1176:1091-6490
1105:2041-1723
1032:0066-4804
974:2057-5858
734:Flagellum
679:virulence
652:Virulence
532:archaella
467:peptidase
403:talk page
173:occurs.
143:antigenic
141:Pili are
132:receptors
2790:Bacteria
2754:Category
2677:Taxonomy
2610:envelope
2500:envelope
2390:Salivary
2307:Obligate
2285:Obligate
2233:Exotoxin
2210:Bacteria
1952:19011034
1896:cite web
1863:16236807
1822:31139165
1796:: 1017.
1771:12398210
1733:38877064
1724:11178852
1681:18063717
1632:19464092
1587:14960717
1538:23386723
1479:31110358
1405:12142488
1337:20418394
1245:18953686
1236:11131518
1194:26884154
1123:36750723
1050:38441061
1041:10989005
992:34435947
933:37019921
924:10076315
865:24133488
807:13230101
723:See also
518:flagella
473:PilF, a
465:PilD, a
423:Transfer
350:pellicle
321:adhesins
294:fimbriae
264:Fimbriae
97:fimbriae
79:bacteria
2764:Commons
2663:Biofilm
2642:Fimbria
2627:S-layer
2608:Outside
2469:Bacilli
2385:Uterine
2370:Vaginal
2280:Aerobic
2263:ecology
2218:Medical
2128:8790416
2096:Bibcode
1943:2620823
1813:6527747
1672:2168650
1623:2719693
1606:Vaccine
1529:3581936
1506:Bibcode
1470:6656605
1328:2897649
1288:1180526
1185:4780597
1154:Bibcode
1114:9905601
1085:Bibcode
983:8549366
856:3796261
839:: 303.
768:"pilus"
739:Sortase
684:E. coli
659:E. coli
357:biofilm
330:E. coli
282:Fimbria
257:archaea
233:plasmid
138:cycle.
124:viruses
92:fimbria
83:archaea
2709:Monera
2479:Spiral
2271:Oxygen
2171:(MeSH)
2161:(MeSH)
2151:(MeSH)
2126:
2116:
2065:
2009:
1983:
1950:
1940:
1861:
1820:
1810:
1769:
1731:
1721:
1679:
1669:
1630:
1620:
1585:
1578:373393
1575:
1536:
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1467:
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1368:
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1335:
1325:
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1253:860681
1251:
1243:
1233:
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1121:
1111:
1103:
1048:
1038:
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990:
980:
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853:
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644:, and
553:. In
509:motile
304:. The
157:, and
2637:Pilus
2591:only:
2571:Porin
2563:only:
2542:only:
2464:Cocci
2440:Shape
2360:Mouth
2333:Other
2119:38514
1249:S2CID
717:toxin
618:curli
599:Curli
569:pilin
397:curli
340:Some
286:Latin
109:pilin
101:pilus
87:pilus
67:Latin
63:pilus
2680:and
2498:Cell
2365:Skin
2355:Lung
2261:and
2124:PMID
2063:ISBN
2039:2017
2007:ISBN
1981:ISBN
1948:PMID
1902:link
1859:PMID
1818:PMID
1767:PMID
1729:PMID
1677:PMID
1628:PMID
1583:PMID
1534:PMID
1475:PMID
1424:ISBN
1401:PMID
1366:OCLC
1356:ISBN
1333:PMID
1284:PMID
1241:PMID
1190:PMID
1172:ISSN
1119:PMID
1101:ISSN
1046:PMID
1028:ISSN
988:PMID
970:ISSN
929:PMID
861:PMID
803:PMID
697:and
681:for
646:CsgG
642:CsgF
638:CsgE
634:CsgD
630:CsgC
626:CsgB
622:CsgA
89:and
81:and
75:pili
2532:DAP
2527:NAG
2522:NAM
2350:Gut
2114:PMC
2104:doi
1938:PMC
1930:doi
1926:191
1849:doi
1808:PMC
1798:doi
1759:doi
1755:292
1719:PMC
1711:doi
1667:PMC
1659:doi
1618:PMC
1610:doi
1573:PMC
1565:doi
1524:PMC
1514:doi
1502:110
1465:PMC
1457:doi
1393:doi
1323:PMC
1315:doi
1311:192
1276:doi
1231:PMC
1221:doi
1180:PMC
1162:doi
1150:113
1109:PMC
1093:doi
1036:PMC
1020:doi
978:PMC
960:doi
919:PMC
911:doi
851:PMC
841:doi
795:doi
770:at
475:NTP
337:.
290:pl.
183:DNA
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