Halloween T4P.cdr

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1: Dept. of Microbiology and Molecular Genetics, University of California, Davis, USA; 2: McMaster University, Ontario, Canada. Abstract is. < se. : uw. 0 III. א of. א.
Diversity and Evolution of Myxobacterial Type IVa Pilus Systems Gaurav Sharma1, Lori L. Burrows2, and Mitchell Singer1* 1: Dept. of Microbiology and Molecular Genetics, University of California, Davis, USA; 2: McMaster University, Ontario, Canada

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737294327 Desulforegula conservatrix 29757022 3 Desulfuri vibrio alkaliphilu s AHT2 85860123 Syntrophus aciditrophicus SB LL peg 07960 736613256 Desulfoverm 40991201 iculus halop 9 Geoba hilus cter sulfu 522165 230 Geo rreducen s KN400 45194 p sy chroba 7873 D ct er esulfo 25840 electrod capsa 6 iphilus sulfex 7490 253 Desu lf igens 7 DSM 1 4044 0970 Geo ohalobium 0523 9378 a re lk tbaen aliba 3742 8 Pe se DS cter fe M 569 rrihy 425 86948 B lobacter 2 d 228 c a ri a c ti r cus terio bino 13 B AsF licus v Ad1 peg 0 dellovib orax m DSM arin 2380 rio b Ad peg 0 694 u s SJ acte C riov As peg 646 orus Vu K peg 0614 HD1 00 As i peg 0651 Ad K pe 0698 g Ad 1 pe 28 Cc C p g 30 67 M pe eg 2 34 Ms fB p g 63 788 M pe eg 93 M h p g 6 687 M f p eg 46 7 M xDZ eg 7 112 0 M xD 2 36 6 AG xD ZF peg 8 K 1 pe 16 pe 311 g 22 g 7 0 08 p 58 28 eg 7 7 57 91

C Y Cc VI m p C c e Sa AP pe g 0 m p g 33 Ho y pe eg 7 373 72 Pp pe g 4 91 8 So 6 P p g 02 So ce5 p p eg 423 7 01 6 p eg 22 6 57 eg 8 53 2 Cc m peg 047 26 CA c pe 057 66 g P 9 LL peg 555 8 CA peg 530 6 P 08 0 M peg 82 So ce5 ir pe 591 3 g 4 6 So0 157 peg 0 3796 peg 476 057 5 97

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1 56 08 836 4 eg 01 19 I p eg 00 427 YV p g 6 C YB pe g 0 649 9 C B pe 07 82 CY YB eg g 02 6 C G p pe 57 8 A B g 6 32 CY pe eg 0 426 Sa p g 4 04 HM pe 00 3 HM peg 000 7 Sa peg 657 5 Sa peg 610 Sa peg 3600 HM peg 070 HM peg 7 64 Sa peg 11 13 6 Sa peg 7 27 HM peg 76 4 HM eg 256 0 Cc p eg 0528 AG p 578 eg 5 Cc p g 00837 e AG p eg 06159 p CYVI g 3316 HM pe 3166 Cc peg 03 Sa peg 35 HM peg 0965 Mh peg 1241 Cc peg 3440 Mh peg 1240 MxDZ2 peg 3475 MxDZF1 peg 7478 MxDK1622 peg 568 1 Sa my pe g 4028 Samy peg 4029 Ho peg 4235 Pp peg LL pe 8225 g Mir p 08824 eg 37 Cc 97 m c So01 peg 555 Soc 57 peg 4 0 e Ccm 56 peg 5796 CA c peg 04764 Pp eg 5555 530 1

70 3 ro 297 ph 3 AC 35 ob 81 5 AB E 0 D a C3 829 .1 H N cte es B P 4 AL AN3 342 9.1 ahe 23 r fu ulfo 1.1 K1 59 Ce lla 20 m ba a 76 3 5 84 1.1 Hah llvib che 3.1 rox cte r .1 S Bu ulf ella rio juen Vib ida cu 11 u j c rk ri h ap s rio ns rv 67 49 NP hold cell eju on is K ch MP atu 98 er a d ens icu CT ol O s 45 4S ia en s 4 C e B y it is K Ue 2 rae N 758 ce WP ntro ph P 41 .1 S pac rific CT da 396 BA 00 o alm ia A an C 2 10 46 N3 34 ba AL s 6 5 6 ct 011 T 6 0 o K2 s 3 7 093 15.1 9.1 er fu .1 Es nell CC kB 96 Su 404 0.1 Clo ma ch a e 254 26 lfu n 493 Bu e r rkh ricel strid oxid rich teri 16 302 404 la d ium an ia ca old P 491 s c eri 678 eloba a c enitr perf MP oli 409 c Pel i 912 oba ter ca epaci fican ringe OB 2 r 97 G aA ss cte bin ns 409 r T k o e 912 c C B li oba 298 cter arbino cus D C 254 26 4044 Geo licu SM su 16 9378 bac 7 Pe ter s lfurred s DSM 2380 loba u u cen lfur 238 cter r s edu 0 K carb NP 2 6444 inoli cens K N400 5324 8862 cus 6.1 P N40 3D D seud YP 20 omon eferriso SM 238 0 7607.1 m 0 as ae Neiss 52216 rugin a camin eria g 3519 G i os onorr eopsy hoeae a PAO1 771609 chrob 441 Des F a A cter ele 1090 ulfonat ctro ronum 52216350 thioauto diphilus 9 Geops trophic ychrobac um ter electr odiphilu APJ13998 s .1 Aeromo nas aquatic ABC32577.1 Ha a hella chejuensi s KCTC 2396 404493798 Pelobact er carbinolicus DSM 2380 CAD85657.1 Nitrosomonas europaea ATCC 19718 YP 208495.1 Neisseria gonorrhoeae FA 1090 YP 208251.1 Neisseria gonorrhoeae FA 1090 erae NP 230504.1 Vibrio chol 2 MxDZF1 peg 603 4701 g MxDZ2 pe 51 Mh peg 59 4 ila LSv5 chroph PAO1 lea psy ginosa lerae esulfota ru D e a 20 nas cho o 512453 o m o ri d Vib ticus Pseu 0505.1 ferrihydri M2 3240.1 NP 23 NP 25 HR s cter m a u b li n ic oalka utotroph etivora s 6 Ge a ac an 7689 ella cetivor 11 rium r te u 7490 c lf a ba su 104 ix sulfo rella 6 De r SM 9 De 7122 Desulfu ima D servat rix 6985 6449 arit on ervat ix 2243 554 c m 7 8 a a 7 s ul pe n atr g o 740 v ip e H la c lfor ser xd3 244 esu oregu a con ns H SB 399 f l 4D a us ni l 8 r 327 u u 4 o g s v e ic mi 108 De for leo 2 ph a 653 072 sul us o itro a c HT 0 d A De 110 m c 30 oc s aci riso ilus 238 8 653 3 c 3 o r lf 29 hu efe liph SM 971 6 u 7 p s 3 o 7 De 7 D lka s D C 1 239 7 ntr 503 3 Sy 8507 rio a licu ATC TC 070 7 6 520 ib C no a 44 18 64 furiv arbi pae is K peg g 06 05 158 5860 7 5 l c 8 su ter euro ens AsK pe g 0 90 8 e u c e C 01 9 2 D oba nas hej Ad d1 p eg 006 368 22 l c o 0 e 7 A I p eg 09 om ella 3P 75 V p g 29 80 tros ah H 93 CY YVI pe Ni 44 .1 1 6 . C AG 40 62 257 6 5 C3 D8 AB CA nt

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771 611 9 4 0 D e s ulfo 750 nat 061 ro 4044 565 9196 Des num th 1 u i P o lf 7034 e a u lo u ro b t 1 m o a 1 c t 5 o ro t 97 G eops er carbin nas ac phicu et m ychr obac olicus D oxidan ADV4 KY s te S 5 J 6 8 8 0 6 .1 Nit 286.1 r electro M 2380 ratifr dip Su a c to r sals lfurovum hilus APJ13 uginis ri 718.1 DSM ftiae Ae 16511 ro m o na NP 23 00 63.1 Vib s aquatica rio cho NP 23 lerae 01 53 ADP96183 .1 Vibrio .1 Marinob cholerae acter adha ABC3 erens HP15 27 93 .1 Ha hella chejuen sis KCTC 2396 APJ16883.1 Aeromo nas aquatica APJ13716.1 Aeromonas aquatica ACE83084.1 Cellvibrio japonicus Ueda107 BAN36482.1 Sulfuricella denitrificans skB26 m thioautotrophicum 771610316 Desulfonatronu autotrophicum ulfonatronum thio M 2380 DS 771610349 Des carbinolicus Pelobacter DSM 2380 404492468 inolicus cter carb g 00192 6 Peloba CYB pe 40449246

4 Sv5 ila L rans o oph chr aticiv idans psy x 4 m alea us aro aceto LSv5 4 lfot 5 d s hila LSv 4 esu hab ona 16 D ophor urom chrop hila Sv5 6 L 453 p y lf 2 tr s 512 hro hila kB c 4 esu Syn ap s c D e p l y 245 23 ota a ps chro cans us N M1 512 616 ulf e y l 0 l fi P s i 655 De 750 ota ea ps nitri oph lum 2396 7 l ulf 07 e l i 453 4 Des lfota lla d s ha eiph TC a10 07 l 512 0 ce su cu KC ed 1 453 3 De lfuri coc etro sis s U eda AO1 5 p 512 89 so Su en cu U P 1 i 5 o ium eju oni cus sa HP in i 24 6.1 itr b 9 N 51 yli a ch jap oni gino ens cam in 3 61 .1 th o ap ru er a am d N3 792 ll j Me ahe ibri 3 BA a m c x e E1 H 7.1 rio s a dh so a s H llv AD 302 Ce lvib na r a rri om an 1.1 M9 3195 79.1 Cel mo acte efe rris vor o b D fe o 3 AB 1 C d . 5 o 8 AB De ole E8 050 seu rin 46 AC E84 1 P Ma 836 470 us . c 6 AC 215 72.1 737 83 oc c 3 7 73 ulfo es D

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Myxococcales Cystobacterineae Nannocystineae Sorangiineae Alphaproteobacteria Betaproteobacteria DeltaProteobacteria Epsilonproteobacteria Gammaproteobacteria Firmicutes Actinobacteria

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Gliding mo lity is an evolu onarily dis nc ve feature between myxobacteria and Deltaproteobacteria. How gliding mo lity evolved in myxobacteria, it has been a longlas ng ques on. Myxobacteria lack flagella but have type IV pili, which help these organisms to glide on the surface. Using various homology search strategies, we iden fied the genes encoding key T4aP proteins and examined their genomic organiza on, generated phylogene c trees of conserved proteins, and explored their synteny in 28 myxobacterial and 66 outgroup genomes, including members of the Proteobacteria, Firmicutes, and Ac nobacteria. Phylogenomic, gene organiza on and syntenic studies of myxobacterial T4P described here reveal the diversity of these systems and propose the evolu onary model of their emergence. Based on this work, we propose that the suborder Cystobacterineae from order Myxococcales and orders Bdellovibrionales and Desulfuromonadales of class Deltaproteobacteria have common ancestors. Similarly, suborders Nannocys neae and Sorangiineae from order Myxococcales and rest of the Deltaproteobacteria orders coevolved separately from the suborder Cystobacterineae. Altogether, current analysis confidently proposes that the T4aP machinery within the three suborders of the order Myxococcales had evolved bilaterally from the order Deltaproteobacteria, thereby producing two dis nct and different genomic organiza ons of T4P within a single diverse o rd e r. .

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