ライフサイエンスコーパス: pilin

1 ilus systems and possess only a single minor pilin.

2 f PilE with other minor pilins and the major pilin.

3 ls were not restored by expression of either pilin.

4 flexible and structurally divergent type IV pilin.

5 the C-terminal donor strands of the incoming pilin.

6 on pilus assembly and function of paralogous pilins.

7 nal regions of other non-conductive type IVa pilins.

8 as the N terminus of archaeal PilA adhesion pilins.

9 pilus (TFP) retraction motors and the minor pilins.

10 y immunoblotting with antibodies against Spa pilins.

11 , including 27 of 29 experimentally verified pilins.

12 lar bacteriocins, cyanobactins, and circular pilins.

13 heir N-terminal portion to bacterial Type IV pilins.

14 , with the only exception being the circular pilins.

15 reviously described in several Gram-positive pilins.

16 t are structurally homologous to the type IV pilins.

17 defining a monophyletic group among type IV pilins.

18 ic or hostile interactions (i.e. hemolysins, pilins, adhesins), and exoenzymes with a potential mixot

19 f the filament, allowing water to access the pilin alpha1 domains as reported for VC-T4P in deuterium

20 SMD simulations revealed that the buried pilin alpha1 domains maintain hydrophobic contacts with

21 We sequenced the major pilin and minor adhesin subunit genes of a geographicall

22 lE structure identifies an authentic Type IV pilin and provides a framework for understanding the rol

23 ion, suggesting that interaction between the pilin and sensor kinase is necessary but not sufficient

24 sequenced archaeal genomes encode predicted pilins and conserved pilus biosynthesis components, rece

25 The roles of the pilins and of the pilus in transformation are discussed.

26 guinis In particular, we focused on the five pilins and pilin-like proteins involved in Tfp biology i

27 accordingly, while elevated levels of shaft pilins and SrtC2 produce long pili and block coaggregati

28 showing interaction of PilE with other minor pilins and the major pilin.

29 inal domain of CdiA-CT(536) interacts with F pilin, and that pilus retraction is critical for toxin i

30 wo O-OTases, one devoted uniquely to type IV pilin, and the other one responsible for glycosylation o

31 For pilin antigenic variation (Av) to occur, a guanine quart

32 m of the pilE gene and both are required for pilin antigenic variation (Av).

33 ects on the genome in the well-characterized pilin antigenic variation systems of Neisseria gonorrhoe

34 Combined results define a unified pilin architecture, specialized subdomain roles in pilus

35 erence to ECM proteins; however, all the Emp pilins are important for E. faecium to cause infection i

36 ools for automated identification of type IV pilins are not currently available.

37 estingly, P. aeruginosa strains with group I pilins are O-glycosylated through the TfpO glycosyltrans

38 Low levels of the minor pilins are optimal for pilus assembly, and CofB can be d

39 eria occurs by a two-step mechanism, whereby pilins are polymerized and then covalently anchored to t

40 Strains lacking both pilins are resistant to a broad panel of SSVs as well as

41 e helical parameters, emphasizing that major pilins are the principal determinants of the fibre struc

42 t nonpiliated mutants, which retain the CtpA pilin, are proficient in attachment and adherence.

43 Accessory pilins at the tip and base have unique structural featur

44 and 3 causes a decrease in the frequency of pilin Av comparable to that obtained with a functional k

45 Since pilin Av frequencies were not altered when modulating R-

46 w that the reduced sRNA transcription lowers pilin Av frequencies.

47 These data suggest that the decrease in pilin Av observed in the RecQ HRDC domain 2 and 3 deleti

48 air enzymes required for efficient levels of pilin Av, and RecQ family members have been shown to bin

49 s three HRDC domains, which are critical for pilin Av.

50 t stable R-loops are not required to promote pilin Av.

51 cessfully refined to more accurately predict pilins based on the in vivo verification of PilA[1-6].

52 cytochromes responsible for EET, rather than pilin-based structures as previously thought.

53 filaments formed from monomers of the major pilin, BcpA, capped by the minor pilin, BcpB.

54 f the major pilin, BcpA, capped by the minor pilin, BcpB.

55 conserved, consistent with them encompassing pilin-binding domains required for initial contact with

56 accessible portion of the single copy of the pilin-binding maturation protein present in virions.

57 We propose that CdiA-CT(536) mimics the pilin-binding maturation proteins of leviviruses, allowi

58 in the pilMNOPQ operon, encoding the type IV pilin biogenesis apparatus.

59 ation/membrane anchor segment with the major pilins but are much less abundant.

60 re the diverse C-terminal region of the PilA pilins but specifically involves the conserved H-domain.

61 motility regulation requires the presence of pilins, but not assembled pili.

62 PCR and for the expression of the respective pilins by immunoblotting with antibodies against Spa pil

63 ar to be low, as PilFind predicted only four pilin candidates in eleven bacterial species (>13,000 pr

64 ns in TBG+ operons, as well as 53 additional pilin candidates in operons lacking biosynthetic genes i

65 d is similar in structure to the CFA/I major pilin CfaB.

66 nction of the protein, designated Spc (short pilin chaperone), encoded by the gene immediately downst

67 show that the enterotoxigenic E. coli minor pilins CofB and LngB are required for assembly of their

68 inor pilins directly interact, and the minor pilin ComGG interacts with most of the known proteins re

69 ccal type IV pili bind DNA through the minor pilin ComP via an electropositive stripe that is predict

70 data support a model whereby the PilY1/minor pilin complex functions as a priming complex in T4aPM fo

71 Comparisons with other chaperone-pilin complexes indicated that the interactive surfaces

72 transcript and protein levels of the primary pilin component pilE via mechanisms that were both conta

73 4F), which are helical assemblies of type IV pilins, constitute a superfamily of filamentous nanomach

74 mechanisms and extend the emerging model of pilin construction in S. pnuenomae.

75 A pilin-deficient mutant and a genetically complemented st

76 In contrast, the U(IV) in the pilin-deficient mutant cells also required an additional

77 efects; however, H. volcanii lacking all six pilins (DeltapilA[1-6]) does not adhere to glass or plas

78 Mutagenesis studies revealed that the pilin-dependent motility regulatory mechanism does not r

79 We demonstrate that the minor pilins directly interact, and the minor pilin ComGG inte

80 pneumoniae (ComGC(SP)), revealing that this pilin displays extensive structural conservation.

81 bacterial adhesin FimH, where the C-terminal pilin domain exerts negative allosteric control over bin

82 So far, it has been assumed that the pilin domain is essential for the allosteric propagation

83 ve the structure of PapD in complex with the pilin domain of isoform II of PapG (PapGIIp).

84 domain at the tip of the pilus followed by a pilin domain that comprises the initial polymerizing sub

85 eta strand of PapD temporarily completes the pilin domain's fold, preventing premature, nonproductive

86 l-length protein and, even in absence of the pilin domain, conducted the cross-talk between allosteri

87 formation upon separation from the anchoring pilin domain.

88 librium between the FimH lectin and the FimH pilin domain.

89 When the lectin and pilin domains are separated under shear stress, the FimH

90 Prior to assembly, periplasmic pilin domains bind to a chaperone, PapD.

91 e shear-induced separation of the lectin and pilin domains results in a shift from a low affinity to

92 As in pilin domains, the bonds are placed at a strategic posit

93 sulted in three distinct classes of type IVa pilins, each found in multiple species.

94 prepilin amassed in the periplasm for rapid pilin elaboration and subsequent BRP production to promo

95 comprehensive list of putative Gram-positive pilins encoded by operons containing highly conserved pi

96 Cells lacking adhesion pilins exhibit a severe motility defect, however, expres

97 Based on pilin features, T4P are classified into type IVa pili (T

98 A group of pilin-like proteins, the minor pilins FimU and PilVWXE, prime pilus assembly and are in

99 Deletion of the associated pilin (flp) decreased NT biofilm and aggregate formation

100 unique structural features, including a dual-pilin fold and the incorporation of a structural zinc io

101 GC(SS) and ComGC(SP) exhibit a novel type IV pilin fold that is purely helical.

102 ure of PilEDelta1-28 shows a typical type IV pilin fold, demonstrating how it may be incorporated int

103 f PilE1, solved by NMR, revealed a classical pilin-fold with a highly unusual flexible C terminus.

104 re-ComGG must be associated with other minor pilins for processing to take place.

105 Many genes involved in adhesion, pilin formation, and others that are important for exoel

106 ructure into the fiber model of the type IVa pilin from Neisseria gonorrhoeae.

107 We investigated a set of 20 Bacteroidia pilins from the human microbiome whose structures and me

108 rsor polypeptides; however, how the unfolded pilins gain their native conformation is presently unkno

109 ich forms the pilus shaft, and two secondary pilins, GBS104 and GBS52, which are incorporated into th

110 o identified a pilin-like motif in the minor pilin GBS52, which might aid its incorporation at the pi

111 The GBS PI-1 pilus is made of one major pilin, GBS80, which forms the pilus shaft, and two secon

112 us allelic differences occurred in the CFA/I pilin gene cfaB (two changes) and adhesin gene cfaE (thr

113 inactivated, mutation of each minor or major pilin gene greatly reduced the bacterium's ability to ad

114 In-frame deletion of the ctpA pilin gene in the ctpCD, ctpE, ctpF, ctpH, and ctpI muta

115 f C. diphtheriae clinical isolates for their pilin gene pool by PCR and for the expression of the res

116 omonas aeruginosa Transcription of the major pilin gene-pilA-is controlled by the PilS-PilR two-compo

117 nsiderably affected in transcription of lngA pilin gene.

118 letion mutants lacking up to five of the six pilin genes display no significant adhesion defects; how

119 as obtained by replacing the pulE-K putative pilin genes of the Klebsiella oxytoca type II secretion

120 All the pilin genes were inactivated, mutation of each minor or

121 (SSV9) conferred by chromosomal deletions of pilin genes, pilA1 and pilA2 that are individually able

122 At the filament surface, gaps between pilin globular head domains in both the native and pulle

123 eover, by deleting genes responsible for the pilin glycan structure, we show that the number of sacch

124 sorting between bacteria carrying different pilin glycoforms.

125 ets produce morphologically distinct type IV pilin glycoproteins.

126 By interfering with different steps of the pilin glycosylation and phosphoform modification pathway

127 The bacterial pilins have a single N-terminal hydrophobic alpha-helix,

128 Mutations in ctpA (a pilin homologue), ctpB, and ctpG decreased early attachm

129 nslational modifications frequently found in pilins, i.e. a methylated N terminus; (iii) are found in

130 ity in Gram-positive bacteria and facilitate pilin identification, we compiled a comprehensive list o

131 cation of otherwise poorly conserved type IV pilins in any species, regardless of their association w

132 are unusual because they are composed of two pilins in comparable amounts, rather than one as normall

133 Based on sequence homology to pilins in Gram-negative bacteria, C. perfringens appears

134 Export of cell surface pilins in Gram-positive bacteria likely occurs by the tr

135 n pilins, PilA1 and PilA2, the most abundant pilins in pili of wild-type and DeltaaglB strains, are m

136 fied 55 of 58 manually curated Gram-negative pilins in TBG+ operons, as well as 53 additional pilin c

137 wever, expression of any one of the adhesion pilins in trans can rescue the motility and adhesion.

138 results, the expression of any one of these pilins in trans is sufficient to produce functional pili

139 7,000 proteins), we identified 286 candidate pilins, including 214 in operons containing TFP biosynth

140 Minor pilins influence pilus assembly and retraction, but thei

141 ssembly mechanism of the central, structural pilins involves proteinase-assisted removal of their N-t

142 o show that the overall fold of the archaeal pilin is remarkably similar to that of archaeal flagelli

143 residue at position 5 (E5) of mature (pseudo)pilins is essential for assembly.

144 BapA1 possesses nine putative pilin isopeptide linker domains which are crucial for pi

145 ll-length fiber-forming and membrane protein pilins leave unanswered questions regarding pilin struct

146 We have also identified a pilin-like motif in the minor pilin GBS52, which might a

147 Here, we show that a Lys residue within the pilin-like motif of the EbpC subunit was necessary for E

148 etween the sortase recognition motif and the pilin-like motif of the pilus subunits.

149 hile incorporation of EbpB only required its pilin-like motif.

150 sively dedicated to the glycosylation of the pilin-like protein ComP.

151 of N-terminally truncated CofB, revealing a pilin-like protein with an extended C-terminal region co

152 articular, we focused on the five pilins and pilin-like proteins involved in Tfp biology in S. sangui

153 A group of pilin-like proteins, the minor pilins FimU and PilVWXE,

154 se that specifically processes precursors of pilin-like proteins.

155 ene conversion reactions between many silent pilin loci and the expressed pilin locus (pilE) allow fo

156 een many silent pilin loci and the expressed pilin locus (pilE) allow for numerous pilus variants per

157 ge via the prepilin peptidase, essential for pilin membrane extraction and assembly, is followed by N

158 he three N-terminal aromatic residues of the pilin monomer are arranged in a potentially electrically

159 ut coupled steps, an orderly crosslinking of pilin monomers and subsequent anchoring of the polymer t

160 th a previously described approach, in which pilin monomers are assembled using randomized structural

161 rder to determine if it was feasible for the pilin monomers of G. sulfurreducens to assemble into a c

162 exposure but instead allowed the release of pilin monomers or short oligomers to the milieu.

163 Mutation of the major pilin MshA and MshB, a minor pilin, resulted in their lo

164 s of ecological interest include two type IV pilins, multiple extracytoplasmic function-sigma factors

165 The DeltactpA DeltapilA double pilin mutant displayed a diminished biovolume and lower

166 lowed by N-methylation of the mature (pseudo)pilin N terminus.

167 cterium Geobacter sulfurreducens and also in pilin nanofilaments (known as microbial nanowires) extra

168 endages composed of protein subunits, called pilins, noncovalently assembled into helical fibers.

169 e, PibD, show the reduction in the number of pilins observed in TEM and reduced surface adherence but

170 logical studies to demonstrate that the FimA pilin of Actinomyces oris contains two disulfide bonds.

171 Our studies also indicate that unlike the pilin of the pathogenic Neisseria species, which exhibit

172 tein interactions among the competence minor pilins of Bacillus subtilis through in vitro binding stu

173 tness mediated by O-glycosylation of group 1 pilins on Tfp in the P. aeruginosa clinical isolate 1244

174 amounts of T4P by a mutant lacking the minor pilin operon was traced to expression of minor pseudopil

175 s and 1 synonymous point mutations among all pilin or adhesin gene copies, implying that each fimbria

176 s, the pairwise nucleotide diversity for the pilin or adhesin genes ranged from 35-43%.

177 rises stacked beta-sandwich adhesin (AD) and pilin (PD) domains, with the putative receptor-binding p

178 Here, we identify a pilin peptidase iamP, and LysR-type regulator (LRTR) iam

179 stabilization and for processing by the pre-pilin peptidase.

180 ling sequence CIP for "cell-cycle initiating pilin" peptide.

181 y couple the priming subcomplex to the major pilin PilA, allowing for efficient pilus assembly.

182 ulate wild-type cellular levels of the major pilin PilA, but not extracellular PilA, indicating dysre

183 ace structures on the bacterium, the type IV pilins PilA and MshA, in adherence to diatom-derived chi

184 tional proteins processed by PilD, the minor pilins PilA, PilB, and PilC.

185 ave now characterized six Haloferax volcanii pilins, PilA[1-6], each containing an identical 30-amino

186 Data presented here demonstrate that these pilins (PilA1-A6) also play an important role in regulat

187 erax volcanii has a specific set of adhesion pilins (PilA1-A6) that, although diverse, contain an abs

188 show that of six Haloferax volcanii adhesion pilins, PilA1 and PilA2, the most abundant pilins in pil

189 Wide pili are composed of the major pilin PilA4, while narrow pili are composed of a so-far

190 ystal structure of the N. meningitidis major pilin PilE and a approximately 6 A cryo-electron microsc

191 We showed previously that minor pilin PilE depends on the putative priming subcomplex Pi

192 hat were genetically deficient in the native pilin, PilE, but supplemented with inducible expression

193 in S. sanguinis We found that the two major pilins, PilE1 and PilE2, (i) follow widely conserved pri

194 Although trained on Gram-positive pilins, PilFind identified 55 of 58 manually curated Gra

195 nsional structure of a Gram-positive Type IV pilin, PilJ, demonstrate its incorporation into Type IV

196 Here, we report a role for the minor pilins, PilW and PilX, components of the type IV pilus a

197 ally similar to Neisseria meningitidis minor pilins PilXNm and PilVNm, recently suggested via charact

198 and is proposed to couple the putative minor pilin-PilY1 complex to the major subunit.

199 tion pathways, we investigated the effect of pilin post-translational modification on the shape and d

200 We conclude that different pilin post-translational modifications moderately affect

201 cient assembly of T4aP composed of the major pilin PpdD, as shown by shearing assays and immunofluore

202 Finally, the pilin prediction program, FlaFind, which was trained pri

203 re we show that Pseudomonas aeruginosa minor pilins prime pilus assembly and traffic the pilus-associ

204 s of several virulence phenotypes, including pilin production, biofilm formation, and NAD glycohydrol

205 y connects a priming subcomplex to the major pilin, promoting efficient assembly of T4aP.

206 "competence pili" are composed of the major pilin protein ComGC and exclusively assembled during bac

207 try that PilA is indeed the majority type IV pilin protein expressed by NTHI.

208 t are assembled by polymerization of a major pilin protein in the periplasm of a wide range of bacter

209 lts provide the first structure of a type IV pilin protein involved in the formation of competence-in

210 ulobacter crescentus We identify the type IV pilin protein PilA as the primary signaling input that c

211 PilMN, PilMNO, and PilMNO bound to the major pilin protein PilA4, to chart the assembly of the inner

212 tative priming subcomplex PilVWX and the non-pilin protein PilY1 for incorporation into pili, and tha

213 synthesis operon (cpa, which encodes a minor pilin protein).

214 gnificantly greater expression of pilA, PilA/pilin protein, and likely, fully assembled Tfp than NTHI

215 4P are multifunctional polymers of the major pilin protein, which share a conserved hydrophobic N ter

216 al sequence of the previously isolated major pilin protein.

217 ression, and relative abundances of PilA and pilin protein.

218 marily composed of helically arranged single pilin-protein subunits, with a unique biomechanical abil

219 In each system, pilin proteins (major and minor) are required to make th

220 proteins belonging to the type IVa family of pilin proteins and are mainly comprised of one subunit p

221 ation; furthermore, both the major and minor pilin proteins are essential for this process.

222 Pilin proteins assemble into Type IV pili (T4P), surface

223 They are helical fibres of pilin proteins assembled by a multi-component macromolec

224 parisons of genomic and structural data with pilin proteins from other species of soil gammaproteobac

225 and mshA genes, all of which encode type IV pilin proteins that aid in attachment to surfaces, were

226 acteria, consisting of a helical assembly of pilin proteins.

227 emarkable structural diversity among type IV pilin proteins.

228 n of the base subunit would only require the pilin recognition motif.

229 ne the structural and thermodynamic basis of pilin recognition.

230 ess, although the precise roles of the minor pilins remain unclear.

231 retraction of pili filaments, the monomeric pilin reservoir in the inner membrane is sensed by the 1

232 on of the major pilin MshA and MshB, a minor pilin, resulted in their loss.

233 Alterations to pilin sequence can also block phage infection, but glyco

234 nformatics without previous knowledge of the pilin sequence-an approach that should prove useful for

235 f the N-terminal domain of bacterial type IV pilin, showing once again how proteins can be repurposed

236 Both series lacked 126 genes encoding pilins, siderophores, and virulence factors whose inacti

237 Among these is a set of minor pilins, so named because they share their hydrophobic N-

238 Here we show that the shaft pilin SpaA harbors a disulfide bond in vivo and alanine

239 the SpaA-type pilus, consisting of the shaft pilin SpaA, tip pilin SpaC and minor pilin SpaB.

240 e shaft pilin SpaA, tip pilin SpaC and minor pilin SpaB.

241 lus, consisting of the shaft pilin SpaA, tip pilin SpaC and minor pilin SpaB.

242 Particular emphasis is placed on 'pilin' specific class C sortases that construct structur

243 l in pilus assembly and that they govern the pilin-specific and housekeeping sortase specificity.

244 With the exception of ADP1, the pilin-specific OTases in Acinetobacter resemble the TfpO

245 In Corynebacterium diphtheriae, the pilin-specific sortase SrtA catalyses polymerization of

246 , but not microcolony formation, and neither pilin stimulates surface adhesion in DeltapilA[1-6]Delta

247 Here we report pilin structures of full-length DnFimA from the sheep pa

248 pilins leave unanswered questions regarding pilin structures, assembly, functions, and vaccine poten

249 ments in SrtA that partake in recognition of pilin substrates and regulate pilus length by inducing t

250 Oral administration of anti-CFA/I minor pilin subunit (CfaE) antibodies conferred significant pr

251 hyperimmune bIgG raised against CFA/I minor pilin subunit (CfaE) tip adhesin or colonization factor

252 NTHi adhesive proteins, protein E (PE) and a pilin subunit (PilA), was developed.

253 ing a Lom-like protein (pVAX-31), a putative pilin subunit (pVAX-12), and a fragment of the type III

254 on, in which the N terminus of each incoming pilin subunit fits into a hydrophobic groove in the term

255 a cysteine substitution is made in the major pilin subunit for subsequent labeling with thiol-reactiv

256 accessory genes (usher, chaperone, and minor pilin subunit genes) with AAF/III, as well as the signal

257 sence of the chaperone CfaA, CfaB, the major pilin subunit of CFA/I fimbriae, is able to spontaneousl

258 and structural analyses of ComGC, the major pilin subunit of Com pili.

259 ibit both phase and antigenic variation, the pilin subunit of M. catarrhalis appears to be more highl

260 the recipient pilE gene, encoding the major pilin subunit of the pilus.

261 The major pilin subunit of the T4P bears multiple post-translation

262 In particular, expression of the pilin subunit pilA requires the PilSR two-component sign

263 ensor PilJ directly interacts with the major pilin subunit PilA.

264 ermined a crystal structure of the CS1 major pilin subunit, CooA, to a 1.6-A resolution.

265 the high resolution structures of the major pilin subunit, PilA, from three Acinetobacter strains, d

266 pili are extracellular polymers of the major pilin subunit.

267 mino-terminal extension on the next incoming pilin subunit.

268 product of the pilA gene would be the major pilin subunit.

269 es named sortases: One for polymerization of pilin subunits and another for anchoring pili to peptido

270 ature, nonproductive interactions with other pilin subunits and facilitating subunit folding.

271 Pilin subunits are recognized by SrtC proteins through a

272 erminal alpha-helices, which also anchor the pilin subunits in the inner membrane prior to pilus asse

273 the study by Xu et al., the structures of 20 pilin subunits of human oral and gut Bacteroidales are e

274 determine the orientation of the assembled F-pilin subunits relative to the cell envelope, providing

275 he CNA(1) isopeptide bond assembled deformed pilin subunits that failed to associate as bundles.

276 f the prepilin peptidase PilD and (iii) that pilin subunits with a C-terminally appended hexa-histidi

277 bacteria, C. perfringens appears to have two pilin subunits, PilA1 and PilA2.

278 T4aP are composed primarily of major pilin subunits, which are repeatedly assembled and disas

279 n parallel by the G1 PapD strand, typical of pilin subunits.

280 a 3D model elaborated from two other type 4 pilin subunits.

281 polymerized and depolymerized from a pool of pilin subunits.

282 initially membrane-embedded pseudopilin and pilin subunits.

283 lightly slower dynamics than the rest of the pilin, suggesting that the first helix is involved in fo

284 acteroidales are elucidated, revealing a new pilin superfamily, assembled into pili by a distinct fif

285 ginosa express one of five different type IV pilins (T4P) (5) , two of which are glycosylated with O-

286 reover, we show that TcpF interacts with the pilin TcpA, suggesting that these proteins are secreted

287 The TCP filament is a polymer of major pilins, TcpA, and one or more minor pilin, TcpB.

288 of major pilins, TcpA, and one or more minor pilin, TcpB.

289 Pili are proteinaceous polymers of linked pilins that protrude from the cell surface of many bacte

290 of all 10 T4PM core components and the minor pilins, thereby providing insights into pilus assembly,

291 d complex that is primed for addition of the pilin to the nascent pilus fiber.

292 shorter Tff than bona fide Tfp-forming major pilins, underlining the evolutionary relatedness among d

293 more highly conserved as there are no major pilin variants produced by a single strain and only two

294 d on the similarity of PilA to other type IV pilins, we further predicted that the product of the pil

295 Genes coding for heat shock proteins and pilins were also induced in Deltasll1130 We observed tha

296 The N-terminal sequences of predicted pilins were exploited to develop PilFind, a rule-based a

297 Unlike T4 pilins, where E5 residue substitutions also abolish N-me

298 ili are composed of a so-far uncharacterized pilin which we name PilA5.

299 s contrast with that of pre-ComGC, the major pilin, which is accessible to externally added protease

300 strate that the PilY1 protein and four minor pilins, which are widely conserved in T4aP systems, are