ANL/MCS-TM-342

Connecting Sequence Data to Virulence Factors in Streptococcus Genomes

Mathematics and Computer Science Division

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ANL/MCS-TM-342

prepared by Jeyashree Alagarsamy Madurai Kamaraj University, Madurai, Tamil Nadu, India Ramya Seetharaman Chandramouli Sivanandham Anna University, Chennai, Tamil Nadu, India Karthickeyan Chella Krishnan Department of Molecular Genetics, University of Cincinnati Ross Overbeek Mathematics and Computer Science Division, Argonne National Laboratory March 31, 2014 *Date

Connecting Sequence Data to Virulence Factors in Streptococcus Genomes

!!

Connecting)Sequence)Data)to)Virulence)Factors)in)Streptococcus*Genomes)

))The!SEED!Project!(http://www.theseed.org/index.php/Main_Page)!was!started!over!a!decade!ago!to!focus!on!creating!more!accurate!annotations!of!prokaryotic!genomes,!along!with!the!tools!needed!to!support!such!an!effort.!!A!number!of!research!teams!have!participated,!and!numerous!projects!were!based!on!the!evolving!technology!that!was!cooperatively!built.!!Some!of!these!teams!sought!funding!to!support!comparative!analysis!of!pathogens,!!and!a!number!did!successfully!acquire!funding!from!NSF,!NIH,!and!DOE.!!While!it!is!often!asserted!that!we!cannot!afford!to!support!manual!annotation!efforts,!we!counter!with!the!following!simple!argument:!!

1. Accurate!automated!annotations!will!directly!impact!the!value!of!the!hundreds!of!thousands!of!genomes!that!will!be!sequenced!during!the!next!few!years;!and!!

2. The!quality!of!automated!annotations!is!directly!related!to!the!availability!of!accurately!annotated!reference!genomes.!

!While!it!is!true!that!most!manual!annotation!efforts!could!have!achieved!far!higher!efficiency,!it!is!also!true!that!extraction!of!value!from!newly!sequenced!genomes!will!depend!heavily!on!a!carefully!curated!set!of!subsystems!built!upon!the!annotations!of!a!set!of!reference!genomes.!!Ultimately,!accurate!annotations!need!to!be!connected!to!the!supporting!literature.!!Since!a!number!of!groups!utilize!the!SEED!technology!for!research!projects!and!since!we!were!initiating!a!pilot!project!relating!to!Streptococcus-pyogenes,!we!decided!to!gather!literature!relating!to!virulence!factors!in!Streptococcus!species,!to!relate!papers!to!specific!genes!in!the!pubSEED!database,!and!to!make!the!connections!we!derived!freely!available.!!We!have!not!connected!all!the!articles!to!specific!genes!because,!in!a!few!cases,!the!actual!strains!that!were!experimentally!characterized!have!not!even!been!sequenced!(in!other!cases,!they!probably!have!been!sequenced,!but!we!could!not!reliably!make!the!needed!connection).!!The!largest!impact!of!this!effort!will!probably!be!via!the!connections!from!genes!to!literature!as!supported!in!pubSEED.!Our!intent,!however,!is!to!provide!the!data!in!a!form!that!makes!it!conveniently!accessible!to!any!annotation!effort.!!We!include!here!a!summary!of!the!Streptococcus-species!for!which!data!was!gathered.!!

Summary'of'the'Streptococcus'Species'Analyzed'

Bacteria(belonging(to(the(genus(Streptococcus(are(all(Gram3positive(and(spherical.((

'

Streptococcus*pneumoniae(is(one(of(the(major(human(pathogenic(bacteria(from(the(Genus(

Streptococcus.(S.+pneumoniae(is(a(major(cause(of(bacterial(Meningitis.(It(also(causes(various(

pneumococcal( diseases( such( as( acute( sinusitis,( otitis( media,( conjunctivitis,( bacteremia,(

sepsis,( osteomyelitis,( septic( arthritis,( endocarditis,( peritonitis,( pericarditis,( cellulitis,( and(

brain(abscess.(S.+pneumoniae(resides(on(mucosal(surfaces(in(nasopharynx(but(can(spread(to(

other(location(causing(infections(in(immune3compromised(people(and(children.((

*

Streptococcus* iniae( is( a( major( fish( pathogen( and( occasionally( infects( humans.( S.+ iniae(

causes(meningoencephalitis,(septicemia,(and(central(nervous(system(damage(in(fishes,(and(

causes( meningitis,( endocarditis,( osteomyelitis,( and( septic( arthritis( in( immune3

compromised(people.(The(site(of(S.+iniae(infection(varies(from(species(to(species(in(case(of(

fishes,(while(source(of(infection(is(not(determined(in(humans.((

'

Streptococcus*mitis(is(closely(related(to(S.+pneumoniae.(S.+mitis(are(not(usually(pathogenic,(

but( commonly( cause( bacterial( endocarditis.( It( resides( in( hard( surfaces( in( the( oral( cavity(

such(as(dental(hard(tissues(as(well(as(mucous(membranes(and(is(part(of(the(oral(flora.(

'

Streptococcus* pyogenes' a.k.a* Group' A' Streptococcus' (GAS)( is( associated( with( wide(

variety(of(disease(conditions(in(humans.(The(majority(of(GAS(diseases(are(associated(with(

skin( and( pharyngeal( mucosa;( however,( occasionally( GAS( also( causes( life3threatening(

conditions( such(as(necrotizing( fasciitis( and( toxic( shock( syndrome.(The(annual(death( rate(

due(to(GAS(infection(is(estimated(to(about(150,000(worldwide.(*

'

Streptococcus* parauberis' is( a( known( pathogenic( bacterium( to( cause( two( economically(

important( diseases:( mastitis( in( dairy( cattle( and( streptococcosis( in( fish.( Only( limited(

research(has(been(done(on(S.+parauberis.((

Streptococcus* uberis' is' responsible( for( a( significant( proportion( of( bovine( mastitis( in(

commercial(dairy(herds.( It(colonizes(body(sites(of( the(cow(including(the(gut,(genital( tract(

and(mammary(gland.('

*

Streptococcus*agalactiae*a.k.a.*Group'B'Streptococcus*(GBS)(is(a(well3known(causative(

agent(for(neonatal(sepsis.(From(a(veterinary(point(of(view,(this(bacterium(is(considered(one(

of(the(major(causes(of(bovine(intramammary(infections,(particularly(in(North(America,(and(

is(a(source(of(economic(loss(for(the(industry.*

*

Streptococcus* suis( is( an( important( pathogen( in( pigs.( S.+ suis+ is( the( name( assigned( to(

streptococci( that( were( formally( called( Lancefield( groups( R,( S,( and( T.( S.+ suis( can( also( be(

communicated( from( pigs( to( humans.( Human( infections( include( meningitis,( septicemia,(

endocarditis,(and(deafness.((

'

Streptococcus*mutans(is(found(in(the(human(oral(cavity.(S.+mutans(is(considered(to(be(the(

most(cariogenic(of(all(the(streptococci.( It(can(thrive(in(high(temperature(such(as(18340OC,(

which(helps(it(to(metabolize(different(kinds(of(carbohydrates,(creating(acidic(environment(

in( the(mouth(causing( tooth( decay.( S.+ mutans+is( implicated( in( the( pathogenesis( of( certain(

cardiovascular(diseases(and(is(the(most(prevalent(bacterial(species(detected(in(extirpated(

heart(valve(tissues,(as(well(as(in(atheromatous(plaques.(

'

Streptococcus* dysgalactiae' subsp.'equisimilis'belongs( to( Lancefield( groups( C( and(G.( It(

causes( infections( in(cows,(and(humans.( It( is( found( in( the(oral(cavity(of(humans.( It( causes(

invasive(streptococcal(infections(such(as(streptococcal(toxic(shock((syndrome(and(mastitis.('

*

Streptococcus* gallolyticus' subsp.' pasteurianus( is( a( Lancefield( group( D( streptococcus,(

formerly(known(as(Streptococcus+bovis'biotype( II.(S.+pasteurianus( are( isolated( from(blood(

cultures(of(patients(with(colonic(cancer.(It(is(also(reported(to(cause(adult(meningitis.(

Streptococcus* gallolyticus' subsp.' Gallolyticus,+ formerly( referred( to( as(Streptococcus+

bovis(biotype(I,( is(a(member(of(group(D(streptococci.( It( is( found(in(the(alimentary(tract(of(

cows,(sheep,(and(humans.(It+is+responsible(for(endocarditis(in(a(patient(suffering(from(colon(

cancer,( but( knowledge( about( the( virulence( factors( is( limited,( and( its( pathogenesis( is( not(

fully(understood.(

'

Results'

The( virulence( factors( for( all( the( species( were( identified( from( the( experimental( proofs(

available( in( the( research( papers( for( different( strains( used( in( this( analysis.(We( chose( the(

virulence(factors(based(on(the(experimental(data(where(wild(type(strain(and(mutant(strain(

(virulence( factor( deleted( from( the( strain)( is( compared( and( data( analyzed( to( identify( the(

function(of(the(virulence(factor.((

(

However,(we(encountered(few(issues(while(collecting(the(data.(

(

1. We(were(unable(to(find(the(Gene(ID/Genome(ID/PubSEED(ID/NCBI(ID(((their(roles(given(in(

PubSEED/NCBI),(even(though(we(were(able(to(find(the(experimental(proof.(The(reason(may(

be( that( the( strain(has(not( yet( been(deposited( into(NCBI/PubSEED(or( it( has(not( yet( been(

sequenced.(

2. In(some(cases,(virulence( factors( for( the(given(strains(were(not(experimentally( tested(but,(

instead,( were( characterized( based( on( sequence( comparison( and( alignment.( In( brief,( the(

query( strain( is( compared( with( a( different( strain( or( a( species( harboring( well3known(

virulence( factors(and( is( in(some(way(related(to( the(query(strain;(based(on(this(alignment(

and(sequence(comparison,(virulence(factors(are(proposed.(These(are(theoretical(data,(but(

we(cannot(ignore(them(because(these(might(be(virulence(factors.(Further(experiments(are(

needed(in(order(to(determine(whether(these(are(actually(virulent(or(not.(

3. The( last( issue(was( a( slight( variant( of( the( second.( Certain( strains( are( proposed( to( harbor(

putative( virulence( factors( but( do( not( have( any( experimental( verification( associated(with(

them.( Instead( they( were( compared( with( other( well3known( virulence( factors( based( on(

sequence(alignments.(These(virulence(factors(are(mentioned(as(hypothetical(genes(in(both(

NCBI(and(PubSEED(and(therefore(warrant(further(experimentation(in(order(to(identify(the(

function(of(these(putative(virulence(factors.((

S.N

o , 2

'0

" '2

'3

'4 '5

'6 ;7

Virulence Factor

Role

ZmpB

zinc metalloprotease

N.:.r'I.A s13'diJ,!W!I AI n.,uram

inid.Jse en host. ufls

pavA adherence and Fibronectiru,.ibrinogen-binding protein

c:bpD

.bp

"",'R

sitB

silO

CopA 1m

b

Sop8

enolase binding piotein 0

Choline binding prolein A

Mn dependent Transcriptional regulatcf

Periplasmc binding plotein

AT? binding protein

Hyaluronate lyase

SodA

Inner mem

brane permease protein

cold sheek protein lam

inin-binding surface praein

Stteptoeoccal GSa peptidase

lyase precursor

Manganese superoxide disrrutlse

laminm

-binding protein C

AM

Ptador

Genom

e 10 365659.3 365659.3

36S6S9.3 36S6S93

36S6S9.3

205S21.3

20592'.3

205921.3

205921.3

2111'0.3

211110.3

211"0.3 208435.3 208435.3

PubSeed role Zincm

etililoproteue zmp8 precursor

Si.llidue

Fibronectin/fibrlnocen-bindine protein [nolue

Choline blndlnC prateln 0

Choline bt\dinc protein A

Mn«pendent

reculilto( MntR

M.an&anese ABC transporter.

PfotdnSitA

MilnCM

lese A8Ctr .. AlP.bindin, protein SitS

MIo",anesc ABC triinsporter,lnner m

embrilne

permeue prDtefn SUD

Cold shod: protefn (spA

Uiminln-bindincsurfilcc proteIn

Hv;aluronilte IVase precur"",

superoJlide dismut;ase

urninin-bindinc surf.ce protcln CAM

Pfilctor

FIgID flgI36S6S9.3.peg.1492

IIgI36S6S9.3.peg, 57'

IIgI36S6S9.3.peg.969 IIgl3G

5659.3 peg,11 67 IIgl36S6S9·3.peg,31

IIgI36S6S9.3.peg,32

1Ig120S921.3.peg.I832

IIgl20592U,peg.I831

IIgl20592' .3, peg.' 830

IIgl205921 ,3,peg, 1829

ngl20592'.3.peg,11B6 IIg1205921 ,3.peg,' 600

fig12111 'O,3,peg.' 33B

figl2' 1t10,3,peg. '300

flgI211110.3.peg.830

figI208435.3.peg.'243 IIgl20B435,3,peg,2047

Organism

Strepb:x:ocx:uI mitis

S1replDcocxu$ mitis

SlreptDc;oo:;:usrN

is Streptoeoocus m

itis StreptDc:oceus m

itis

StreptDcoc;:cus mitis

Strept)COOeu$ aga1acti3e

Stteptoeoc:eus agal3dlae

SlreptDcooc:us agalactiae

StreplDcoccus agalactiao

Streptococeus agalactiae

Strepbxoc:c:us asatactiae

Streptoeoceus agalactiae

Streptoeoceus agalacti3e

StrcllnlD

86 B6

B6 B6 B6

B6

Aro9

Aro9

Aro9

Aro9 Aro9 Aro9

NEM

3'6

NEM

3'6

NEM

3'6 2603VIR 2603VIR

'5

1mb

"'" Ibs fibrinogen.binding protein

1203610.3 IIgl'203670,3.peg,'O

'9 protein

Streplotoc;:cU5 agalactiae GD201 008-001

'9

1mb !am

nin-binding $Urfac:e Pfotein

20 C

5a peptidase C

Sapeptida&e

21 Slrepm

domase 0

Streptodomase 0

22 ThioJ..adivated cytolysin

cytolysin 23

Streptokinase StreplolUna.s.e

24 speS

Streptocoocal cysleine pl'otease 25

SpeH

Streptococcal pyrogenic exotoxin H

27 scpA

cSa peptidase

26 fbp

Fibronectin binding protein 29

mf1

Streptodornase 8 30

sic Streptolysin 0

3'

SpeG

Ex_

in type G

32

SpeC

Streptococcal pyrogenic exotoxin C 33

SpeS cysteine protease (Streptopain)

34 SpeH

Streptocoecal pyrogenic exotoxin H

3S $pel

Streptoooccal pyrogenic exotoxin I 3Ei

SpeJ Streptococ:cal

exotoxin J 37

SmeZ

Streptoc:oc;cal mik)ge,lic exotoxin Z

36 SI<a

Streptokinase 39

Slo StteptotvSn 0

410 .,5lidine kinase

loystem histidine kinase

41 toNo-com

ponent respoA&e regulator response regulator

42 nptoeooeal of

(SIC) Stroptococ::cal inhibitor of eorr¢em

ert (SIC) 43

Streptodomase B; M

itogenic factor 1 Stleptodotnase B; M

itogenic facto( 1 44

SpeH

Stteplooxeal pylogenic exotoxin H

4S streptococ:cal phage D

NAse

Streptoeocc:al phage DN

Ase 46

SpeA Streptococcal pyrogenic exotoll:in A

47 SpeC

Streptoc:oecal pyrogenic exotoxin C

48 Ska

Streptokinase 49

.&10' Streptolysin 0

50 sda

Streptodoroase 0 5

' SagA

Streptotys;n S 52

Sag C Streptolysin S biosynthesis protein C

53 SagH

Streptolysin S export tr.iiIIn5lT'oem

brane permease

54 SpeS

Stteptoeoeeal pyrogenic exotoxin B 55

"spat< Stteptocoooal pyrogenic exotoxin K

56 SSA

Streptoeoeeal superantigen A 57

SiM protein Idherence to fish epithelial cells. and m

acrophage rHistanee

58 FibronectinlFi6rinogen binding protein

Fb binding protein 59

hasA

hyaluronic acid $Vnthesi:s 60

hasB

hyaluronic acid synthesis 61

Enol3Se Enolase

62 m

etal ABC tJ&nsportE!f$

metal ABC

transportef5 63

hasC

UTP-gllJCO

Se-1-phosphate uridytyltl3nslerase 64

19t Proipoprotein

transferase 65

IspA Lipoprotein signal peptidase

66 LPXTG

containing surface proteins M

ediators of host palhogen Interactions 87

LPXTG containing surface proteins

01 host pathogen interactions 68

Adhesion Invasion of hut1'l!ln brain

cens 69

Adhesin A&SOCiated w

ilh IE 70

Adhesin. Glucan producing and binding

Protecting bacterium against host defenses

71 Adtoe.lfll. O

ll,ie;ln proriiXing ar,d IW

IdlflSl Protecting bacterium

against host defenses 72

Adhesln. Glucan producing and binding

Protecting bacterium against host defenses

73 Adhesln. G

lucan producing and binding Protecting bacterium

against host defenses 74

Streptolysin 0 Thiol-activated cytolysin

75 Streplokinase

Streptokinase A precur5()( 76

scpB Streptococcal GSa peptidase

n streptokinase

Streptokinase 78

M protein

antiphagocytc M protein

79 exotoxin G

variant 4 exotoxin G

variant 4 60

sagA bacteriocin-like prepropeptide

81 53gB

streptotysin S biosynthesis protein B 62

sagC

Streptolysin 5 biosynthesis protein C 83

sagO

Streptolysin S biosyrthes.is protein 0 64

sagE Streptolysin S setl'·im

.mwlity prote;n

85 sagF

Streptolysin S biosynthesis protein F 86

sagG

Streptolysin S export ATP-binding protein 87

sagH

Streptolysin S export transmem

brane protein 88

sagl Streptolysin S export transm

embrane prolein

89 fbpS

Fibroned.inltibrinogen-binding protein 90

hemolysin·like protein

RN

A binding methyltransferase FtsJ like

91 SU

AM proteins

Surface exclusion protein 92

hasA H

yaluronan synthase 93

hasB U

DP-glucose dehydrogenase in hyaluronic acid synthesis

94 lP

XTG

containin9 surface proteins M

ediators of Host-pathogen interaction

95

96 97 98 99

Adherence to human endothelial cells and causing infective

Bacterial adhesions endocarditis

pilBconlribule to innate invnune resistance and btofilm form

ation G

tfA C

ell adhesion. dental caries.biofilm form

ation G

tfB Cell adhesion. dental ca.ries.biofilm

formation

AtiA Autolysin

1203670.3 293653,3

293653.3 Streptodornase D

Streptolysin 0 293653.3

Streptokinil5C 471876.6

SCrepotoc:ocnl cvrtelne protuse (StreptoJHIinllEC 471876.6

Streptococx.al dlctO

lfln H (SpeH); Towlmoron

471876.6 Streptokinue

471876.6 pepticS;ase

471876.6 Flbronectln-tHndln, protein

471676.6 Streptodom

ue a; MitoCenlc '.etor 1

471876.6 Thlol..adM

ted cytoiyl1n 471676.6

Streptococal PVf"lI!nlc Elotoxin G ($peG

) 160490.1

SCtt:ptOC'OCCoilI pyrocenlc el(otoxin C CSPeC): TOIIlmoJOn 1 &0490.1

SCfCpotOCDCCill 'VSteine pfO\e;ase (Strcptop.aln) ('EC 1 fi04SO

.1 StreptocO

l:lUI pvroccnic CIIotemn H (5peH): TOIIim

Ofon 160490.1

SUflIlM

acc.llt P\J'tatenle 1U0000MIn J (Spcl) 160490.1

prtal!l!-tIic eaOlOllin J 160490.1

Streptococcal mitocenlc ekotO

'lln Z CSmcZ) 160490.1

Strcptokm.w:

160490.1 Thlol..ainted

160490.1 Two-com

P'1"c"t hbOcl,"c tinua

160490.1 Two-aln\PO

nent sVS(cm respoft5e recuJ:ttor 1$0490.1

Streptococollnhtbttor 01 eompicm

eflt (SIC) 160490.1

Sttt:ptodomascBiM

itoecnlc.fmor1

160491.17 5UtptococalpyrofU

lCnocorh K

160491.17 StrtptocOCCllI nudeue 4; M

ltoeenlc rldO(

1"'0

3.3

Streptocoa;.al pyrorenlcootOlCin A (SpeAl; _TOlfim

Dron 186103.3

Sttt:ptococCilI pyrocenlcexotoJllin C (Spec); TOklmorun

186103.3 StreptotlnHC:

1861 03.3 cytolysm

M Streptolysin 0

186103.3 Streptodornase D

186103.3 streptolysin S precursor (S.cAl

186103.3 Strepto!ys.in S bloSfllthesl$ protein C (SIIcC)

186103.3 StreptotV5/n S el(port trusmem

br.ane permeil5e

198466.3 Sttt!:poIoc:occ;aI't¥ottlne pnrt,e;ll!(:

fEe 198466.3

Streptococcal pyroll!nlc er:otolin K (SpeIC) 198466.3

streptococcal supcr;antlf;en A (SSAI 936154.3

s/M protein

936154.3 Fibronedin/flbrinocen-bindinc IN"aCeln

936154.3 Hy;aluron;an svnth.asc

936154.3 UOP-Cluoo5e dehydroeen;ase in hYilluronlt.cid

936154.3 EncNsc

936154.3 m

etill ABC trilnsporter SCoIb5triltC!-blndinC lipoprotein 936154.3

UTP-c/UCOSC·l-phO

lphiite 936154.3

Prolipoprotein diacvldvcerv/ Irilnsferne 936154.3

UpoorOI.dn ,",en.1 pep'IdIlJC

981540.3 -

Sort;ase A.lPXTG specific 981540.3

Sortasc A. LPXTG 981540.3

Late compell!flce protein Com

GA. access of DNA to 981540.3

N-.ac:etylmuram

ov!-L-;alanine amidase

511691.3 hvpgth'cUcal protein

S11691.3 hypothetical protein

511691.3 hypothetical protein

511691.3 dextnnase precursor

663954.3 ThloJ..aCliv;ated cytol",In

663954.3 streptokinilse

4186410.3 cSa peptid<llse

486410.3 Streptokinue

486410.3 M

proc .• in 486410.3

5trcpCocou.il (5pcG

) 486410.3

StreptotysinS

prc<uraar'tsllAl

greptolpin S biosynthesis protein 8 (Sq8)

486410.3 SCrcptotysinSbiosvnthesh pro\e(n C (SlICe)

486410.3 Strcptotyslns

protc!n D (S,alD) 486410:3

Streptolysin S self·imm

unitv protein 486410.3

S biosvnthes/s protein IS.CF) 466410.3

hport ABC trilnsporter AlP-blndinc protein; 4815410.3 Stteptolysin S export trollnsm

embrilne perm

ease 486410.3

Streptolvsin S export transmem

brane permeuc lsacl)

218495.3 Fibronectin/fibrlnoeen-bindinl protein

21 RNA bindinc m

ethvltr;anster3se ft$.llike 21&495.3

Surface CJlclusion proteIn 218495.3

Hy;aluronolln synthase 218495.3

UOP-elum

se dehvdroeenase in hVilluronk: acid 9815J9.3

en;a protein a-type dornilin-conl;ainlnt: protein

981539.3 981539.3 981539.3 95'539.3 9815J9.3

M;a",anese ABC transporter. periplasm

ic-bindin, proteinSitA

late competence protein ComGA,. ;access or DNA to

Glucosyltr;ansfer;ase GUG GlucosvttnnsleriiSC G

tfG

G0201006-001

fIQI293653.3.peg.3G

2 Streptoa)ocus p)':lgenes

MG

AS

5005

Qgf2936$J.l.pog,l721

S1Jeplo<xx:ou> pyogenas M

GAS 5005 Streptooooeus pyogenes

1 1g12936S3.34'011.""

Streptooooeus .yogen .. ng/4'1U

J76.G.peg.1745

Stre_

... P\'09 ..... IIg/471576."-I>OI1,826

St<ept0000w5 .y

og

"''' !lg/471B76,6.POil.'G

SS S1Jeptoc:oow

s .yogenes IIg/47181'-6 .... 9,'722

S1

J_

py

og

en

es

ng/471S7U

.peg.'721 StreploooQ

;u$ PY"9"nes -

OgI411e7G

.t.peg.1147 S1J.plooooxus .yogen ..

1Ig/47117'-6.peg.I40 Slreplo<:o<xul pyogenes

llg!"7'B'76.ftPCg.,S2 Streptoeoc:cus pyogenes

flg , 604:10.' _,620

Streplo<:o<xul pyogenes 1101160490.1

_,1562

Str

_.y

og

.n ..

IIgl' 604:10.' """,.7&3 S1r.plD

c:cc:ws pyogenes

11911604:1O.I.peg.171O S1Jeptac;ocx:us pyogen ..

l!gllG04:1O

.I.peg,312 S

Ire_

us pyogenes

S1Jeptac;ocx:us .yog .... 1Ig1'604:1O

.I,peg.'521 S

1J_

pyog .... ngl,G

04:1O.1.peg,lZ1

Slreptoc:oows pyogenes

Og)l604!lQ

.\,_'554 SIr • ..,..,.....pyog.nes

IIgl.6049O.1._

15

>5

S1Jepto<;o<;ouspyoganas

Pgl.604!lQ.',peg.1501.

St<ep1o<;<><:ou5 pyogenes 11g11S0490.I.peg.l56C

S1J..,....,..... pyoge ...

IlUII6049',17.peg.I051 S

1J._

.... pyogenes flgI1604".17.peg,71U

S

1J_

pyogenes flul'8G

103.3.peg.333 S

Ir._u

s; pyogenes

Ilgll1161 ou

.peg.m

Str._

usp

yo

gen

es

ngl166103.3.pog.1825 Streptoeoocus pyogenes

11011 1161 O303.P''II.''' Stroptooocxus pyogones

'562 StreploooQ

;u$ pyog ..... •• GS9

Stteptac;ocx:uo pyog_ Ilgll86I03.:J.peg.701

S •• _OOI: ... pyog_ flull8610U

.peg.706 Sl1eptac;ocx:uo pyog .....

IIg/15ao166,3_1851 S

II._

"" pyog_

flgl,0046U,pog.'301

__

us pyDgenos

'i91'88466.3.pog.100S pyogenes

Sheptoc::QC:C:ll5par-auberis IIgI93S'54.3.pe1lo"13

.... beris flgl!l36'5oI.3.peg.1l72

SlI._ poliWbc,;'

OgI$36,54.3.peg.2171

su._ po .... beris r19I93615<C.3

_ .. 2

s..o_p._ O

g/93&I54 .• _

.... ""_

.. ""'''''''' 11jj19361154.3.peg.11,61

S.'pIOCO

OQ

I' po ........ "" flgl936154.3.peg."

3

S •• pIOCOOQI ...... Ubotls

figl.36'54,3,peg.647 Stt.ptoo>oous po"

,"_

fl13l981

pa

teU

fllnuJ liB

15Sl54o.a"",g.993 $

ln>

p""","""_

"L.rio",,, flgl9&

I540.3.pog.B6 _

• ..,..,..... ".. .. una ....

r'llI!l1l1540.',peg.1292 SltO

plOO

OC

<UJ ...... '"

....

IlgJ51 '6SI.e.peg41373

nglS1 16S1.3.peg .1026 Stm

ptoooooul5o mUllIns

flgI611GS1.3.pog.00<

m .....

IIOI511691.3.peg.17.. S ... p"""""""m

u .... fI9lGG3554.3,peg,2043 _

pt.,.".. ... d ...... t..;t;'e"'lB

p. <quW"",,,

rlglG6395'.3.Sleg.tt31

figJ4&64 to.3,peg.9Cn Streptococcus d .... .. toI.Ibsp, ttquisim

lli5 Streptoeoc:cus

(igJ48G4\O.3.peg.190 Streptococc:us

stibsp. equla.imjrJSi flgI46G

410.3.peg.1953 S1reptoeoocus d)'$9abctiae equW

mHiI flg1486410.3.peg.67S

Streptococcus dY'9l1bc1t.ae $Ubq:i. oquW

m'lo

fig148&4tO.3, pov.676 Stteptococ:cus $UbJ9. equislm

llca fiV148G4 1 O.)"peo..£17 Streptoooccu!' dY

'9Wot:\1,c $U

bqJ. equWm

ili. IIgl4864'O

.3,pog.678 Strepto<:oa:us dl'll_tiae

ngl486410.3.peg.679 Slrept.ococeus dysgalactiae subsp. equisimilis

Og14864 t O.3.peg.&80 Streptoeoccus dysgal.Jctiae subsp. equiSoim

ilis IIgI486410.3.peg.6:e1

Streptococeus dysgalactiae subsp. equisirmis

rlgl486410.3.peg.682 Streptoeoocus dysga/aetiae subsp tigl4B64 1 0.3.peg.683 Streptococcus dvsgalaetiae subsp equisim

ilis figl21 849S.-3.peg.1 01 2

Streptoooceus ubcrjs figl21 849S.3.peg. 1167

Streptoc:oecu$ uberis figl21 B495.3.peg.1498

Stleptoooceu$ uberis figl21 B495.3.peg.1557

Streptococcus uberis

MG

AS5O

O5

MG

AS5O

O5

NZ13'

NZ13,

NZ131 N

Z'3'

NZ

'3' N

Z'3'

NZ131 N

Z'3'

SF370 SF370 SF370 SF370 SF370 SF370 SF370 SF370 SF370 SF370 SF370 SF370

Manfredo

Manfredo

MG

AS8232

MG

AS8232

MG

AS8232

MG

AS8232

MG

AS8232

MG

AS8232

MG

AS8232

MG

AS8232

MG

AS315

MG

AS315

MG

AS315

KCTC 11537 KCTC 1,537 KCTC 11537 K

CTC

I1537 KCTC 11537 K

CTC11537 K

CTC11537 KCTC 11537 K

CTC

1,537 A

TCC

43,44 ATC

C43144

ATCC 43144 A

TCC

43'44 N

N2025

NN

2025 N

N2025

NN

2025 A

TCCI23904 ATCC ,2394

GG

S_124 G

GS_'24

GG

S_'24 G

GS_'24

GG

S_'24 G

GS_'24

GG

S_'24 G

GS_'24

GG

S_'24 G

GS_

124 G

GS_'24

GG

S_'24 G

GS_'24 0140J 0'4O

J 0'40J 0'4O

J tlgI21849S.3.peg.1S56

Streptococ:cus uberis 0140J

figI981S39.3.peg.1S71 Streptococcus gallolyticus 5Ubsp gallolyticuli

ATeC 43143

IIgI981539.3.peg 2039 IIgl·81539,3.peg.B7

ligl561539.3.peg. '045 IIgI9B'

047 IIgl56' 539.3.peg.' 363

Streptocoecus gallolytiaJs subsp_ gallolyticus Streptococcus gallolyticus sub5p gallolyticus SlreptQC:9Ccus gaUolytic:us 5Ubsp gallolyticus Streptococcus gallolyticus 5Ubsp gallolytic::us Slreptoc::oeeus gallolyticus

gallolyticus

ATCC

43143 ATC

C43143

ATCC 43'43 ATCC 43,43 A

TCC

43'43

PEG 10 '4Q

2

51'

969 1167

3' 32

'1132 '83' 'S30 '829 1186 ,600

'338 ,300 830

'243 2047

'0'9

1769 362

1727 233 33'

1745 826

16S5 1722 ,72' 1749

140 ,82 520

1562 763

'719 312

1536 152, 123

'554 ,555 1549 '566 ,061 7'B

333 679

'1!2S 141

1562 699 701 706

1881 1301 1005 '957 1123 2172 2

'7'

542 334

2'67 "3

647

'S56 993

86 '292 1313 '026 936

'789 2043 113' 909 '93 '90

1963 '675 676 617 67B 679 6SO 68' 682 683

'012 1161 1498 '551 '556 '571

2039 87

1045 '047 '363

NC

BIID

smL'482

smL())Q

1

smU

S67 sm

i_1159

smL0037

smUX138 SA

IU557

SAK

_'556

SAK....'555

SAK

_'554

SAK_'3'9

gbs,308

gbs'270

gbs080S SA

GI234

SAG

:Ill43

A964_'028

A964_1791

M5O

O5_SoY

_17'5

M5O

O5_SoY

_'4'5

M5tXl5_SpV_ 014'

M5O

O5_SpV

_'684 S.V49_ '6900 SP}'49_0792 SpV

49_'630 SI')'49_16690 S.V49_,66&

; Spy49_1692e SPV

4lLO'46

SI')'MLO

'87 SPv_0711 SPv_2039 SPv_1008

NA

SPv_0435 SPv_I998

SPv_O'67 SPv_2026 SoY

_2027

SPv-.2043 SpyM

5'021 SpyM

50691 SpyM

'B_0393

SpyM'B

_0778 S.yM

'5_2042 SpyM

15_0'65 SpyM

'8_1746 SpyM

IB_0799 SpyM

'8_080' SpyM

18_0S06 SpyM

3_1742 SpyM

3_1205 SpyM

3_0920 STP_I567 STP_0963 STP_1716 STP_t715 STP_0468 'STP 0295 STP::t711 STP_036' STP

'0559

SGPB_09B6

SGPB

_0086 SG

PB_1289

SJT&INN2025_1372 SITlJNN2025_1036 SrnJN

N2025_0945

SmJN

N2025 17811

SDE

'2394j'0255 SoE

'2394_05710 SD

EG_0933

SDEG

_0233 SD

EG_0230

SDE

G_'99'

SDEG

_0705 SD

EG_0706

SDEG

_0707 SO

EG_07O

B SD

EG_0700

SOEG

_07'0 '5D

EG_0711

SDEG

_ 07'2 SoEG

07'3 SU

a1111 SU

B'273

SUB

'635 " ..

SUB

' 696 SG

GB_1565

SGG

B_2030 SG

GB

_0067 SG

GB

_l044 SG

GB_'046

SGG

B_I362

Rofentnce (')

(')

(') (') (1)

(') (2)

[2J

[2J

(2)

(3) (3)

(4)

(4)

(4)

[5] [5]

[6]

[6]

[7J

[8) [B) (9J

['D)

['0) ['OJ ('D

) ['D

) ('0) ['D

) ('D

) [11) [11) [1') [1'1 [1') [11) [111 [11) (11) (111 [11) [11) ['2) ['2) ['3) ['3) ['3) ('3) ['3) ['3) ('3) ('3) [14) ['4) 1'4) ['5] ['5] ['51 [15] [15] ('5] ['5) ['5) ['5) ['6] ['6] ['6] ['6] ['7) ['7) ['1J ['1J ['8) ['8) ['9) ['9) ['91 [19) ['9) [191 (191 ['91 1'91 ('9) ('91 ('91 ['91 (20) [20) (20) [201 (20) (2')

12') [2') (2') (21) (2'1

References

1. O

illl. d .... The G

ettome of Slrepococw

s mlis B

6. 'tIIotwt 15. Com

meM

ar? PlbS ON

E 5(2t. February 2010 .2

B.I Sl.,tdftlrt. and D. HlIrrlngton. E

xp.uion dSrA

lflfQC

O(ll:w

1r1'IacfJuA909

... ndiron.A

nlall ...... lMJwenhotiC. 2OOiI.

95(1t.p.10'·'09. 3.

GI.n1W UndW

II_ Slaham

m .. -Ca1em

am. Ind Thanas ArftchQIJCI. SurtiKtII Prot .... 11 Qf SI,-.,pIoeo«llS ag,IIr.ct.iote..-.d R

Nted Proteinsin OU.ef o",derial PliIhog«ls. diNcal

mladiolog1 M

f_. jll\ 2005. P 101-127.

4. G

enome M

qUeflQ

II dStntplOCDCl:UII;alacti.le ... pllttlDgel"\

necrWal llw

a_. Gauer. Cm

slopM Rusnio.:.

Microbiglgg)' (20::12145(81. 1499-1511

5. Cem

pW. venem

a ... .w:.

compantiw genomic

of an trnergngl'lumilft

I«dype V stmplD

«ccw

"" ..... Tetlelin. Veoa M.a..i",M

Ii. PNAS Septem

b.-17. 2002_ '1'01. "_I'\Q. 'SI _ 12391-1Zl86.. 6.

G«1omt

of ShFIocoa:u.s 9'-=

'iN G0201I):)'&

·0:)1. hd-tod WI CNM tnJrn T

op. 'cfIih MeN\go.nCJ:lPNLlis. G

uat'I" lkl. Wd ZNna. and Ctlengpin" lu.

.... ruric:en SodeCy far fItocab'alogy. 25 SeptemtUIf 20 12.

7. 8eflr-tg lei, SiltY

IrotIdcie. s.-omar lUI:aa-. Ja'T

I. M. frol.neer.ldm

titcaiOl"l and

d GtoupA Sfrep/oc:ocrus eulUe

Pm

riM. inlediatl and im

munly.

tA.no.12

a. Sunby It •

• E\IOUiCl'llol"J Origin M

d Em.wgenc:. of a HIghly &

.icc""" Dine

Mt G

roup A. Horirorlal G

tne TlVilftr The

JouTi., of Infedbn. Di ..... ' 200S: 1i2:771-.a2. 9.

SatrI D. Reid. M

. Ofltfl'l. M

lJU0Q.I5 .,aIytI. at ",Id. by IlJUIJPASl,."tOCllXiCUS"

;ene(ie, tMMTI.-. respon'O

.ai'ld 0eN: ItM

saiptlon. PNot.S .MI. 10. 2001 \d

. De "0. 13

10. W

. M

cSNn, G

.. OInIl Sequence of. N

qltn:cgtM.1nd IiIot\tY

..... 0 strAI 01 Steploc:O

CWIi

JaJmal Of Bacteriolo;y. Dec_ 2008. p. 1773-7765

11. JoN

phJ.FMfeIII.W

J&amM

PN

AS.A

pril10.2001.wI 94."0 a

12. M

mM

w 1: G

. Halden at ., .• C

omplllls G

.-une Gf "cuts Rheum.'e: FIrY

IlI"-AnoeY

led M

5 {IyO

glJrIIJ$ 51 ..... Milnfredo. Jw

mal O

f e.ctGiolGgy. Feb. 2001. P 1473'-1477. 13.

.am" C. Sm

oot. K..-. D. BartIIan. d" G

eJ'ltm. sequ.neoe and com

pat1llt-.. alytl_ ofserotypo M

t8 orwp A. Strep1cm>ccL1S stnIin, niOd.lted v.\th _CIA.

loYIII'" oUtI ....... w

i. 89_nC1 1.4BM

-4m. PN

AS.A

prl2.2002. 14.

SCephen a. a_H

. Gal L.S)fW

at at • Genom

. nqu .. ;o 0( t W

fot}'pe Ml *V

I of FOU9 A Stl1lp(oC

ocau PMo.enc:oded laK

lM.lns hi,gh ....-ulen;. ph.,OCtpe. and don. em

orgrnce. w

i. no. 15. H

107a-1CO

il PNAS.JUy 23 2002. 1$,

Nho. S •• lilt III ..

g.tO

tne

aI'''' beef"w

JCJJmIl of bttSeOoIogy. 2011_tU

(111: p.3l55-33eO

. 16

Wi. Uu T-T. T

q Y-T, W

I H-l. I..kJ til. (2011)

.nd CO

lTqnnCiw

of Two

Sll"I!P'oc:o«w

So.IblpedH: Genom

e Plntidy. A

daptdoft and PlaS O

NE 8($1: e20S19.

17 Fumito. M

....... (2000) "Com

,,""'" genomic an .. )'M

$ of SlteptD

coc:ws m

t.hn. prIMO. ft6lghls lroIo c:htcrnOJomal"5hutling and tpeQ

e5-tpeQlic c:onltnC.-8M

C Genom

ia to_

18. o.W

ts. M. "a

I" VlNI«ICG ptoIlino of Steploc:occus ctyegil1actiae 8'qI.IIsm

lis IlOLIIed tram infected hum."

flIVe,IlS 2 dislntS genetic: .... UII" tN

t do nol: their Clhenut)lP" O

f prop8'\tly ICI ;au .. elM

..... Cliric:.lll irlfediats diMa$M

: .-! Dlichll' oI'lhe Intcliou. 0i1U

SH Society fJ(

2007 414(11): p 14012-14501.

18. ",-"I SN

mcn.\R

.. .,. C«nP'ete a!nG

mG

MqU

lSldng ISld anaI)'Ii. of.

gfOlCl G

dysgatK:IiH

3Ubsp.

IlrXI cavsin; ... ,."tococc" krIric shock

lI)'"ctom.

8MC G

enc:m1cl20". 12:t7

:zo boW

1e"atM;en

BMC Genom

ies 2009. p: 10:54 21.

Un I-ti. Uu T-T. T.no Y-T."

" H-l. Uu YoM. lilt

(20"1 Sc.,ftdlg lind

Anal,.. olTwo PMtIoglinic S

Mlpoax.:us

Genom

. PtaUicity. AcbPM

an •• P

loSO

NE

0{5l: e20S19

.... Gcononwo to

'ogI' Pnetn101y.ntP\11

ilnd h 10 IdlV

llt mm

phmerC

.,d 'COO'IlnS of the prole.-. rH

OO

I'".ble ae1 ........ ,

l1J15l27 1

_

""108 p.HU

• ..o, I(C 14 I. I hg111J15327 pI;.656

31

JUl2

1

p .. A CoIlltIU

__

• elft IildKe ,fN

.",

17t101 1

110111.11 1)ff .... "

" .... C'..,.o_'r.w

'Of ...

figl171101.'PofgllSO

1l

\3

p .. e

ebp.

c",

' .... '

ill''''''' 0&

. bmcil'llll.aflt1l

rceeplO( binds lhe

S .....

•• lC'1Irar ...

daphOipI'lCl)1..:e CpcO

f<il'CoIpw

" Fom!,J,1oQtI

re4usc dhlghtynl\Mlm

l'lCl'} cell W

3D dognld.1.lu:n

mpC

l1.l'IIfO(bltlllm

formilbon

mom

t.&IU for bltI'lm

,_-.on. CdOft,ullcn InnasophilrynlC

a:w

'\O'1bw

.,lo..",,,dilmilQ

otlnl\oghdctstpneutnDtW

.

eydgph'IfHypt PPliI'u ;lind eonInbulcs Itl edo.iulllln oflht upper Hw.111c

Clt ..... g H

umil'" im

munoglobulin'"

" " PcaA

,""dhcW

I. CIQb>q:.on.. ilrtilgo'UZe 1M

Meu

dTH

P·1 'dis

kw"1I ...... r..hon dlKho #lfldO

Ol\

17

" ,. '" " 27

" 29

30

" 12 " 35 ,. " >II '" .0 " " " .. .. ., .. •• .. " .., ., .. 0>

.. 51

58

50

.. 112 03 .. . , .. 41

.. ..

SodA

.... """ ."" .... .... ...... ... , SIC

.,.0 G

bp.

l4iopro(tinreeep(Ofa'ltigtn(L

nlI

p-.p

"'" ......

phO$phoducxm

ut.l ..

... A

.... "'C &

.0' " .. &.oF

.. " ",H .. 1M

.... .,... ...... IlIA

p,,, FlO

F ... T .... .... urn;

""'" 70

EC

Mprd..,

71 FIoJ'O

Mdjnbinding prdtin

n G

k.ocos,.tr.l1fIftM

7loU w

illi ItlChot.cI pttJI.nl

74 P

I .... 15

CoIiIgCr'i tirwr"" a41wst1 18

CoIlgtO

binding prd.tn T7

C(lIilgenbiodin"praein

H.O

laCltjil photphm

P'.y I'd

" IrI metlballtm

olG

d""'" ;,,,,,1.

Segregm

on",d C:CJ1dtns.3l,lDn p

m", B

Hph.iI·O

fIat. tyKe preeunor

MilnllollntseM

olperoldckd'smvt •

•

Betaheftlal)'tla'lopD"'O

nC)'!E'Pf1'I,.n

Strep4op.ntstfeSJlOQXC:;aJ pm

I .. M

QI ..

StJ'cp.oox:e-' p)1ogeric u:ctolOn J

Slteplok"'M

ArtIphigoeytcc:M

pftltllin

ScrtiiSC A. LPX

TG tpltO

fc

1Idh.M1oefn ........ Q

tllpilheMleeim

onal.)'tI' •• hil!lmdy6c:.drvC

)' ldtM

rencem'4lsiot!

oeI mond',...I. hlernd)4ie .diY

il),

dlerenoe .... 'o\ ...

dillP'ftIuR

w .. bn

...th ... 'I[Iwfn

d .....

... Ic:trvtW

Idhl5'el'loe/" ..... .

" df eptheIm

I eel mond.prs. tulem

olytlc:.ildMII)'

&.pta.con, R

e#

_.

TG

dcifwtohderillub

C5;a.,d Irnpahthe IIbiIty of tho 6nfected h

as 10 19h1.1hII int'edlU'l. ,wlb

nc:.

in«c ... resist_a. and dIM

I.1 dis$el'T1nnon

Trilget'TNFalpha production

Pfoa..dibn

... mdD

ltllUallc.11

"'_""" ••

eot1iding IM.iIte ,,"m

Wlo deffl'lH

$ hlendion with host eanpltfnenl f""tbID

ry fadOf' hfH

Ifw

IT .... lllllort •

CIU

IlWU

till" 0Idd .......... MM

ogIf'He: IIcqyililion

prdeof)tic ot eelS. m

odl.ltd.

Adh.-en.oe. "'""do pordno brain n.lO

'GI4fO

Jbir endothelial eeC.

""ilKtiw

endocardti. tM

"doQrdill$

Endoc:lrd •• C

okif'ftQI _er. EnO

oc3rdItrl C

oIcndIf c:;;Jrw;e(. Endoc:.d' ...

CoIa"1dI1 C

lf'ICtf. E

ndoc:nllil BkJ6m

In eob'edill CM

lcef pilfltrlll B

lofihiftllTlllionineOlofectillt:illloerpiililrils

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