HevIew

lntroduction
Thodiscovoryoponicillinin19?8byAloxandor
Flomingwasarovoluionaryovoninhohisoryo
modicino.Fonicillinandmanyohoranibioicsdis-
covorodinlaoryoarshavosavodcounlosshuman
livosrominociousdisoasosproviouslyhougho
boincurablo.Anibioicsarounarguablyonooho
ma|orscioniicandmodicaladvancosoho?0h
conury.Howovor,hoiniialwidosproadandinado-
quaousooanibioicshaslodohoomorgoncoo
rosisanbacoria,whicharoprosonlyapublichoalh
concorn.Mohicillin-rosisan5/ao|,/cccccasaaas
MF8A and vancomycin-rosisan onorococci
VFEbocamoasoriousproblomnoonlybocauso
ohoirpahogoniciybualsobocausohososrains
canransorhoiranibioic-rosisangonosoohor
bacoria.Thoroisanurgonnoodoranowclasso
anibacorialcompoundshaaroacivoagainsho
mulidrug-rosisansrainsandwouldnodovolop
bacorialrosisanco8ang and Elocha,?008,
Asadu..amanand8onomoo,?009.Eacoriocins
aronovolcompoundswihpoonialanibacorialap-
plicaionsandarodiscussodinhisroviow.
Eacoriocinsaroanibacorialpopidosproducod
bybacoriaocompooagainsbacoriaohosamo
spociosorohorgonoraCooroal.,?005b.
EACTEA8Eisanopon-accossdaabasodosignod
orhocharacori.aionobacoriocins,andhonum-
boroonriosconinuosogrow.Thodaabasocon-
ains?00bacoriocinsoquoncosJuly,?011,mos
owhicharohoproducsoGram-posiivobacoria,
paricularlylacicacidbacoriaLAEHammamio
al.,?010.LAEaroaracivobacoriocinproducors
AntIbacterIaIPeptIdes

BacterIocIns

:AnOvervIewof
TheIrDIverseCharacterIstIcsandAppIIcatIons
MAMlNlSHlE
1
,JUN-lCHlNAGAO
2
,ANDKENJlSONOMOTO
1,3
1
///s/c^c/^oo//J/c/ca/a//cc//c/c,a^J//cass|/s/,
/oa/^/c///csc/^ca^J//c/c|^c/c,/aca//,c/^/ca//aCaJaa/5c|cc/
/,as|a|^/s//,0101/a/c?a////as|//a/a/ac/a1?b1'aoa^
?
5c//c^c//^/c//c^//c/c,/oa/^/c//a^c//c^a///csc/^c/a/ac/a/^/a/c//
?1b11aaa5anaa/a/a/ac/a1401'aoa^
3
/oa/^/c//a^c//c^a///a/c//c/s/^//c^c|//c/a^//,as|a|^/s//,
0101/a/c?a////as|//a/a/ac/a1?b1'aoa^
Focoivod14Augus,?011/Accopod5Ocobor,?011
BacterIocInsarerIbosomaIIysynthesIzedantIbacterIaIpeptIdesproducedbybacterIathat
InhIbItthegrowthofsImIIarorcIoseIyreIatedbacterIaIstraIns.AnumberofbacterIocInsfrom
awIdevarIetyofbacterIahavebeendIscovered,andtheIrdIversestructureshavebeenre-
ported.GrowIngevIdencesuggeststhatbacterIocInshavedIversestructures,modesofac-
tIon,mechanIsmsofbIosynthesIsandseIf-ImmunIty,andgenereguIatIon.BacterIocInsare
consIderedasanattractIvecompoundInfoodandpharmaceutIcaIIndustrIestopreventfood
spoIIageandpathogenIcbacterIaIgrowth.Furthermore,eIucIdatIonoftheIrbIosynthesIshas
IedtotheuseofbacterIocIn-controIIedgene-expressIonsystemsandthebIosynthetIcen-
zymesofIantIbIotIcs,acIassofbacterIocIns,astooIstodesIgnnoveIpeptIdes.lnthIsrevIew,
wesummarIzeanddIscusscurrentIyknownInformatIononbacterIocInsproducedbyGram-
posItIvebacterIaandtheIrappIIcatIons.
/,ncJsAnibacorialpopido/Fopidobiosynhosis/Eacoriocin/Lanibioic.
ioconlrolScience,!01!,Vol.17,No.1,116

csoc^J/^aa/|c1//a-1?04?01/a//
sc^cc/caa/,as|aaac/o
bocausoohoirGFA8gonorallyrocogni.odas
saosausasdosignaodbyhoU.8.Foodand
DrugAdminisraionFDA,indicainghoirsaoand
oasyapplicaionasoodprosorvaivosorpharmacou-
icals.isinA,aroprosonaivoandwoll-sudiodLAE
bacoriocin,wasirsdiscovorodin19?8Fogorsand
Whiior,19?8andhasboonusodasacommorcial
oodprosorvaivoinmorohan50counriosCoor
oal.,?005b.on-LAEbacoriocinsaroalsoworhy
oaonion,orinsanco,somoohobacoriocins
producodbyhogonus5/ao|,/cccccasworoshown
oboacivoagainsmulidrug-rosisan5aaasand
coagulaso-nogaivosaphylococciinvolvodinhuman
inocionsascimonooal.,?006.Colicinsor
microcinsproducodby/sc|/c|/acc//aroablooin-
hibihogrowhoGram-nogaivobacoria,whilo
mosLAE-dorivodbacoriocinscannoDuquosnoo
al.,?007.Howovor,hoormbacoriocinisnowusod
mainlyodosignaoanibacorialpopidosproducod
byGram-posiivobacoria.Thisroviowwillsummari.o
anddiscussbacoriocinsproducodbyGram-posiivo
bacoria,includinghoirclassiicaion,modooacion,
biosynhosis mochanism,gono rogulaion,sol-
immuniymochanism,andpoonialorindusrialap-
plicaion.
CIassification
Alhoughhoirsidoniiodbacoriocinwascolicin
producodby/cc//in19?5Duquosnooal.,?007,
LAEbacoriocinshavoboonhomossudiodhus
ar.LAEarooonoundinoods,havingunknowingly
boonuili.odinhomanuacurooormonodoods
suchaschooso,yogur,kimchi,andradiionalor-
monodoodsinoachvariousaroasohoworld.LAE
bacoriocinsalsoplayanimporanroloininhibiing
hogrowhospoilago-causingbacoria.
Coor o al.?005b havo classiiod LAE
bacoriocinsino?disinccaogoriosTablo1.
Classiscomprisodoso-callod

lanibioics,

which
sands orlanhionino-conaining anibioics,and
class includos non-lanhionino-conaining
bacoriocins.Largo,hoa-labiloprooins,oonmuroin
hydrolasos,worooxcludodromproviouslyclassiiod
bacoriocinsandnamodbacoriolysins.Horo,non-
LAEbacoriocinswillalsobodiscussod.
CIasslbacteriocinsIantibiotics
Classbacoriocins,hoso-callodlanibioics,aro
smallpopidos19-38aminoacidshaconainun-
usualaminoacidsinroducodbypos-ranslaional
on.ymaicmodiicaionsagaooal.,?006,Fioldo
al.,?010,LooandKim,?011Fig.1.Unusual
amino acids such as dohydraod amino acids,
lanhionino,and3-mohyllanhionino,whichormmul-
iploringsrucuros,disinguishlanibioicsromohor
anibacorial popidos and conribuo o ho
lanibioics'srucuralsabiliyagainshoa,awido
rangoopHandprooolysis,andohoirincroasod
olorancoooxidaion8ahloal.,1995,Eiorbaumo
al.,1996.
Lanibioicsworoproviouslyclassiiodasoihor
ypoAorypoEaccordingohoirsrucuroandunc-
ion.Elongaodamphiphilicpopidossuchasnisin
rom /ac/cccccas /ac//s and Fop5 rom
5/ao|,/cccccaso/J/J/s5woroypoA,whoroas
globularpopidossuchasmorsacidinrom/ac///as
sp.andacagardinorom^c//^co/a^ssp.woroypo
E.TypoAlanibioicsworourhordividodinowo
subypos.ypoA,whichhavoanolongaodorm,
suchasnisinandsubilinrom/ac///assa/////sATCC
K.SNMTFTAL.
Classiicaionobacoriocins
Classiicaion Fomarks Examplos
ClassLanibioics
Classlanibioics
Classlanibioics
Lanhionino-conaining
bacoriocins
Unusualaminoacidsinroducod
byLanEandLanC
Unusualaminoacidsinroducod
byLanM
isinA,8ubilin,Epidormin,
Fop5
Lacicin481,ukacin8K-1,
Morsacidin,Lacicin3147
wo-popidobacoriocin
Class
Classa
Classb
Classc
Classd
on-lanhionino-conaining
bacoriocins
Fodiocin-likobacoriocins
8pociicagains//s//a
c^cc,/c^s
Two-popidobacoriocins
Cyclicbacoriocins
8inglo-popidonon-podiocin-liko
linoarbacoriocins
FodiocinFA-1/AcH,Loucocin
A
LacococcinG,LacococcinO
EnorocinA8-48,GassoricinA,
lacocyclicinO
LacococcinA,LacicinO
6633,ypoA,whichhavoanunbridgod-orminal
ailandaglobularC-orminalring,suchaslacicin
481 rom //ac//s and nukacin 8K-1 rom
5/ao|,/cccccasna^/8K-1Jungand8ahl,1991,
Duouroal.,?007.Howovor,ashonumboro
lanibioicsincroasodandhoircharacorisicsworo
olucidaod, his classiicaion bocamo uncloar.
Foconly,onhobasisohoirsrucuroandmodoo
acion,lanibioicsworodividodino11subgroups
isin,Epidormin,8ropin,Fop5,Lacicin 481,
Morsacidin,LnA,Cyolysin,Lacocin8,Cinnamycin,
and8ublancingroupsroprosoninghonamoso
woll-knownlanibioicswihdioronoaurosCoor
oal.,?005a.Thisclassiicaionpoinodouhaa
numboronauralvariansolanibioicsoxis,andi
islikolyhahoroaroalargonumboroohorvarians
hahavonoyoboonisolaod.naddiion,hooxis-
oncoonauralvarianssuggosshahoroisloxi-
biliyrogardingsomoohoaminoacidsacorain
posiions,andimayhusbopossiblooonginoor
popidoswihouroducinghoiraciviy.
Lanibioicsaroalsoclassiiodonhobasisoho
pahwaybywhichmauraionohopopidooccurs
Tablo1.Thisdosignaionmainlyincorporaosdi-
oroncosinholoadorpopidosoquoncoandbiosyn-
hoicoporonsrucuroFagand8ahl,?00?,Willoy
andvandorDonk,?007.Classlanibioicsmainly
consisowhahavoboonclassicallycallodypoA
lanibioics,hopropopidosowhicharomodiiodby
?disincon.ymos,LanEandLanC.LanEdohydraos
ho8orandThrrosiduosopropopidos,andLanC
modiaoscycli.aionodohydraodrosiduoswih
inramolocularCys.Classlanibioicsaroclassically
callodypoAandypoElanibioics.Modiicaion
on.ymo LanM oxhibis boh dohydraion and
cycli.aionaciviios.
CIassllbacteriocins
Classbacoriocinsaronon-lanhionino-conaining
bacoriocins.Unlikolanibioics,hoyaronopos-
ranslaionallymodiiodbydodicaodon.ymos.Tho
hoorogonoousnauroohisclassobacoriocins
makosurhorclassiicaiondiicul,andhussovoral
groupingapproachoshavoboonproposodoso
al.,1996,vanEolkumand8ilos,?000,Cooroal.,
?005b.Horo,woollowhoclassiicaionbyCooro
ACTFkCNSFkMCkAM-STVFACTFkA
.8rucuroobacoriocins.ClassiicaionisbasodonhoproposalbyCooroal.?005b.Classiicaiono
lanibioicsisbasodonhobiosynhosispahwaysopopidosproposodbyWilloyandvandorDonk?007.Thoshadod
rosiduosroprosonunusualaminoacids.Dha,dohydroalanino,Dhb,dohydrobuyrino,A-8-A,lanhionino,Abu-8-A,3-
mohyllanhionino,OEu,?-oxobuyryl,D-A,D-alanino,M,ormylmohionino
al.?005b,whichsubdividosclassbacoriocins
ino4groupsclassosa,b,c,anddTablo1.
Classabacoriocinsaropodiocin-likopopidos
haarohighlyacivoandspociicagainshoood
pahogon //s//a c^cc,/c^s.Fodiocin FA-
1/AcH rom FJ/cccccas sp.or /ac/c/ac///as
o/a^/aa conainsho-orminalconsonsusso-
quoncoYGGVXC,acharacorisicoclassa
bacoriocinsHondorsonoal.,199?,Molaghoal.,
199?,Dridoroal.,?006.
Classbbacoriocinsarowo-popidobacoriocins
consisingo?unmodiiodpopidos,bohowhich
aroroquirodoranibacorialaciviy.Whonosodin-
dividually,hosobacoriocinsdisplayvorylow,iany,
anibacorialaciviy.Eachpopidopairisoncododby
ad|aconoponroadingramosinhosamooporon,
andranscribodandsynhosi.odogohorOppogard
oal.,?007.LacococcinG rom //ac//ssubsp.
/ac//sLMG?081isson-Moyoroal.,199?washo
irsobocharacori.odobolongohisclass,ohor
popidossuchasAEF-118Flynnoal.,?00?,
planaricinC8Maldonadooal.,?003,onorocin
1071Fran.oal.,?00?,andlacococcinOZondo
oal.,?006havoboonidoniiod asclassb
bacoriocins.
Classcbacoriocinsarocyclicpopidoswhoso-
andC-orminiarocovalonlylinkodbyapopidobond
vanEolkumoal.,?011.EnorocinA8-48producod
byavarioyoonorococciwashoirsdiscovorod
classcbacoriocinMarino.-Euonooal.,1994,
8amyn o al., 1994. LAE-dorivod class c
bacoriocinsincludogassoricinArom/ac/c/ac///as
ass/LA39Kawaioal.,1998andhoroconly
idoniiod uborolysin rom 5/o/cccccas a//s
Wirawanoal.,?007,andlacocyclicinO rom
/ac/cccccassp.8awaoal.,?009.on-LAE-
dorivodbacoriocinssuchasbuyrivibriocinAF10
rom/a/,////c////sc/^sAF10Kalmokooal.,
?003havoboonroconlyidoniiod.
Thoromainingbacoriocinshahavonosigniican
soquonco similariy wih ho ohor class
bacoriocinsarocaogori.odasclassdbacoriocins.
Thisgroupincludossinglo-popidonon-podiocin-liko
linoarbacoriocinsCooroal.,?005b,isson-
Moyoroal.,?009.Thosobacoriocinsarogroalydi-
vorso,andhoroaroowproposalsosubdividohoso
popidosaccordingoloadorpopidosoquoncos.Tho
loadorpopido soquoncos odivorgicin A rom
a^c/ac//a J/^sWorobooal.,1995,
lacococcin97?rom//ac//sFLA97?Marino.o
al.,1999,andsomoohorbacoriocinshavoacloav-
agosio,whichacsasasignalpopidohaaccossos
hogonoralsocrooryscpahwayDiopandos,
?00?.Onhoohorhand,somoclassdbacoriocins
suchasonorocinL50rom /^/cccccas/ac/a
L50Cinasoal.,1998,auroocinA70rom 5
aaasA70o.oal.,?001,andlacicinOand
lacicinZrom//ac//sFu|iaoal.,?007,waanio
al.,?007arosynhosi.odwihouan-orminal
loadorsoquonco,andhusarocallod

loadorloss
bacoriocins.

Howovor,classdbacoriocinshado
noiinohosc-dopondonorloadorlosssubgroup
sillromain,howovor,aslongasscionisssoarchor
novolbacoriocins,imayboimpossibloosricly
classiyanddoinohom.
Modeofaction
Eocausohoroisanoxpocaionhabacoriocins
willsorvoasindusrialoodprosorvaivosorasanal-
ornaivoopharmacouicalanibioics,horiskodo-
volopingrosisansrainsshouldboavoidodbyusing
hom inolligonly,basodon

scioniicovidonco.

Thorooro,acivorosoarchhasboondodicaodo
sudyhobacoriocinmodooaciononhomolocular
lovol.Thomochanismobacoriocinaciviyagains
argobacoriashouldprovidoinormaiononhowo
provonhoomorgoncoobacoriocin-rosisanbaco-
ria.Eacoriocinssuporiciallyrosomblooachohorin
hahoyarooonmombranopormoabili.ing,cai-
onic,andamphiphilicorhydrophobicisson-Moyor
oal.,?009.Howovor,rosoarchhasshownhahoro
aroavarioyomodosoacion,asinglobacoriocin
possossosmorohanonomodooacionoaack
hoargobacoriaWiodomannoal.,?001,Haspor
oal.,?006.
Thomodosoacionobacoriocinsarolinkodo
hoprimarysrucurosohopopidos.hasboonro-
porodhasovorallanibioicsargolipid,amom-
brano-bound collwallprocursor,as a docking
moloculoEroukinkanddoKrui|,?006.isinisa
poro-orminglanibioichakillshoargosrainby
pormoabili.inghoplasmamombrano,loadingoho
oluxoinracollularmoloculosFig.?A.hasboon
shownhahohighaciviyonisininhonanomolar
rangoishorosuloahigh-ainiybindingolipid
Eroukinkoal.,1999.ThoMFsrucurooho
nisin-lipidcomploxrovoalodhaho-orminuso
nisinbindsohopyrophosphaoolipidHsuoal.,
?004.Muaioninhohingorogioncausosinaciva-
ioninporoormaion,suggosinghoimporancoo
hisrogioninporoormaion.Thismuan,howovor,
roainsanibacorialaciviyosomooxon,showing
hanisincankillbacoriabyinhibiingpopidoglycan
synhosishroughinoracionwihlipidWiodomann
oal.,?001.wasdomonsraodhanisinromovos
lipidromhocolldivisionsioandhusblockscoll
wallsynhosisHasporoal.,?006.Muacin1140is
abacoricidallanibioichasharosmanysrucural
K.SNMTFTAL.
andsoquoncosimilariioswihho-orminusonisin
lipid-bindingdomain.Alhoughhomuacin1140-
lipidcomploxisassabloasnisin-lipid,his
lanibioiccannoormporos,huswarraning
urhorsudyoismodooacion8mihoal.,
?008.
Lacicin3147isawo-popidolanibioic,wihpop-
idosLnA1andLnA?acingsynorgisicallyopro-
ducoanibioicaciviy.wasproposodhaLnA1
irsinoracsspociicallywihlipid,honholipid-
LnA1comploxisabloorocruiLnA?,andinally,a
3-compononcomploxinhibiscollwallbiosynhosis
andormsporosWiodomannoal.,?006Fig.?E.
Morsacidinisalipid-bindingpopidohainhibis
collwallsynhosisbudoosnoinducoporoorma-
ion.Thobindingsioonlipidwassuggosodobo
hoorminalGlcAcsugar,whichisdioronrom
nisinEro

.oal.,1998.
ClassabacoriocinssuchaspodiocinFA-1/AcH
usocompononsohomannosophosphoransoraso
sysomman-FT8osuscopiblocollsasrocopors
andoxorhoiraciviybydissipaionohoproon
moivoorcoviamombranoporoormaionFamnah
oal.,?004,Diopoal.,?007,isson-Moyoroal.,
ACTFkCNSFkMCkAM-STVFACTFkA
.Modoloargo-collkillingornisinAA,lacicin3147E,andlacococcinAC.
ATho-orminusonisinAbindshopyrophosphaoolipidandpormoabili.osho
plasmamombrano,rosulinginporoormaion,whichloadsohooluxoinracollularmolo-
culos.ELnA1andLnA?inoracsynorgisicallyoproducoanibioicaciviyinhowo-
popidolanibioiclacicin3147.LnA1irsinoracsspociicallywihlipid,holipid-LnA1
comploxishonabloorocruiLnA?,andinally,ahigh-ainiy3-compononcomploxorms
poros.CLacococcinAomployscompononsCandDohoman-FT8asarocopor,
buhocyoplasmiccompononAEisnoroquirodorhisuncion.Thoboundlacococcin
Ariggorspormoabili.aionohomombrano,causingloakagooinracollularcomponons.
?009.Man-FT8ishokoypahwayormannosoand
glucosoupakoinbacoria.Foconly,hooxracollular
loopohomombrano-locaodman-FT8wasshown
oborosponsibloorspociicargorocogniionby
classabacoriocinsK|osoal.,?010.
Classbwo-popidobacoriocinsconsiso?di-
oronunmodiiodpopidos,bohowhichmusbo
prosoninapproximaolyoqualamounsinordoro
oxoranibioic aciviyCooroal.,?005b.
LacococcinGG plusGandlacococcinO
O plusOaroclassbbacoriocinsisolaod
rom//ac//s.Fopidosandshowahighlovolo
homology,wihonly6and3amino-aciddioroncosin
mauropopidos,rospocivoly.Thohybridcombina-
ionsohosopopidossuchasG plusO orO
plusG showodanibacorialaciviy,indicaing
hosobacoriocinsobooxchangoabloZondooal.,
?006.Thusar,pormoabili.aionohoargocoll
mombranoisconsidorodashomodooaciono
classbbacoriocinsOppogardoal.,?007.Thoy
inducoholoakagooinornalsubsancossuchas
monovaloncaions,phosphao,andATFAbooo
al.,1994,Molloal.,1996,Molloal.,1999,Cuo..o
oal.,?003.8ocondary srucuralsudios o
lacococcinG,planaricinE/F,andplanaricinJ/K
havorovoalodhamombrano-likooniiosinduco
amphiphilic-holicosobohpopidos,andaddi-
ionalsrucuringwasobainodwhoncomplomonary
popidosworooxposodsimulanoouslyohomom-
branooniiosHaugooal.,1998,Haugooal.,
1999.Alhoughinograodmombranoprooinsaro
spoculaodoboarocopor,rocogniionargosinho
classbbacoriocinsysomhavoyooboidoniiod.
8imilaro mosbacoriocins,class c cyclic
bacoriocinspormoabili.ohomombranoosuscopi-
blocolls,rosulinginholoakagooions,dissipaion
ohomombranopoonial,andulimaolycolldoah
vanEolkum oal.,?011.norosingly,onorocin
A8-48,gassoricinA,subilosinA,andcarnocyclinA
canoxorhoiraciviywihouanyrocoporGalvo.
oal.,1991,Kawaioal.,?004,Thonnarasuoal.,
?005,Gongoal.,?009.EnorocinA8-48ormsnon-
solocivoporos,whichcausoholoakagoolowmo-
locularwoighcompoundsGalvo.oal.,1991.On
hoohorhand,carnocyclinAormsanion-solocivo
ionchannols,whichrosulinvolago-dopondonporo
ormaionhaisabolishoduponmombranodopolari-
.aionGongoal.,?009.
Classdnon-caogori.odbacoriocinsarodi-
vorsoinhoirsrucurosaswollasmodosoacion.
LacococcinAandlacococcinEarosuggosodo
uili.ohoman-FT8asarocopor,pormoabili.oho
cyoplasmicmombrano,andcausoloakagoosoluos
acrosshomombrano.LacococcinsAandEdono
sharosoquoncosimilariyooachohororwihclass
abacoriocins.Unlikoclassabacoriocins,which
roquiro a paroho C prooin oman-FT8,
lacococcinsAandEroquirobohCandDprooins
orman-FT8argoingDiopoal.,?007Fig.?C.
LacicinO isaloadorlossclassdbacoriocin.
FrovioussudiosroporodhalacicinOdoosnoro-
quiroarocopororismombrano-pormoabili.ingac-
iviy and acs via a oroidal-poro mochanism
Yonoyamaoal.,?009a,Yonoyamaoal.,?009b.
Lacococcin97?Lcn97?washoirsidoniiod
non-lanibioicbacoriocinhaspociicallyinoracs
wihlipid.Lcn97?aciviywasanagoni.odbylipid
/^/c.co-procipiaoswihlipidmicollosandin-
hibishoaciviyo?on.ymos,FEF?andFomX,
whichusolipidasasubsrao.nhocasoonisin,
ho-orminallanhioninoringsaroinvolvodinlipid
binding.Lcn97?lacksalanhioninoring,andhonco
mighdisplayanovollipid-bindingmoiMarino.
oal.,?008.
Resistancetobacteriocins
isinAandpodiocinFA-1/AcHarohusarho
onlybacoriocinscommorciallyusodasoodpro-
sorvaivos.Thoomorgoncoobacoriocin-rosisan
srainsshouldbocomploolyavoidod,horooro,un-
dorsandinghomochanismobacoriocinrosisanco
isovialimporanco.
Onoimporanmodiaorobacorialsrossro-
sponsoishoalornaivosigmaacor8igE.8igEhas
boon shown o conribuo posiivoly o /
c^cc,/c^solorancoohobacoriocinsnisin
andlacicin3147Eogloyoal.,?006.Tho/
c^cc,/c^swo-compononsignalransducion
sysom,LisFK,playsasigniicanroloinacid,oha-
nol,andoxidaivosrossandinmurinovirulonco.
wasproposodhaLisFKalsodoorminoshosonsi-
iviyo/c^cc,/c^sonisinCooroal.,
?00?.VirF8isawo-compononsysomharogu-
laos ho oxprossion oJ//^ and o/ in /
c^cc,/c^sandinluoncoshobacorium'ssus-
copibiliyocaionicanibacorialpopidosMandin
oal.,?005.Thodoloiono//ando/isound
orosulin3?-and16-oldroducionsinrosisancoo
nisinCollinsoal.,?010a.Thoponicillin-binding
prooingonoo/oGravosonoal.,?001,hoollurio
rosisancogono//^Collinsoal.,?010b,hogono
oxprossinghopormoasocompononoanATF-
bindingcassooAECransporor,a^/Collinso
al.,?010a,andhogluamaodocarboxylasogono
aJEogloyoal.,?010aroalsoroporodoplaya
roloinhoinnaorosisancoo/c^cc,/c^so
nisin.
Thomochanismorosisancoonisinandclassa
K.SNMTFTAL.
bacoriocinsisrolaodocollmombranoluidiyand
coll-suraco chargoVadyvaloo o al., ?00?,
Vadyvaloooal.,?004.wasroconlydomonsraod
hahoroaro?ma|orrosisancomochanismsoman-
FT8-argoinglacococcinA in/c^cc,/c^s
and//ac//sK|osoal.,?010.Thoirsinvolvos
downrogulaionoman-FT8gonooxprossion,andho
socondinvolvosnormaloxprossionohoman-FT8
sysom,buhoundorlyingmochanismorosisanco
orhosocollsisunknown.Howovor,ihasboonsug-
gosodhacollsuracochangoshaaochoinor-
acionbowoonhobacoriocinandismombrano-
locaodrocopormighboinvolvod.
8udiosonbacoriocinrosisancohavoocusod
onlyonspociicbacoriocinssuchasnisinandsomo
man-FT8-argoingpopidos.Exonsivousooavari-
oyobacoriocinswouldroquiropainsakingro-
soarchoavoidhopossiblodovolopmonowido-
sproadbacoriocinrosisanco.
iosyntheticmechanismbiosyntheticen-
zymes
Eacoriocinsarousuallyribosomallysynhosi.od
asprocursorpopidoshaconsisoan-orminal
loadorpopidoandaC-orminalpro-rogion.Tho
loadorpopidokoopshobacoriocinsinacivoandis
spoculaodoproochoproducingorganismsrom
boingkillodbyisownbacoriocin.Tholoadorpop-
ido,whichishoughoboascaoldorhobiosyn-
hoicon.ymos,iscloavodo,andhoroloasodpro-
rogionbocomosamaurobacoriocinCooroal.,
?005b,Duouroal.,?007,Oppogardoal.,?007,
WilloyandvandorDonk,?007.
Lanibioicsroquiropos-ranslaionalmodiicaions
oinroducounusualaminoacidsinohopro-rogion.
Classlanibioicssuchasnisin,subilin,andFop5
aromodiiodby?disincon.ymos,LanEandLanC.
LanEdohydraoshosorinoandhrooninorosiduoso
hopro-rogion,andLanCcycli.osdohydraodrosi-
duoswihcysoino.LanTisanAECranspororha
ransporshomodiiodprocursorpopidos.Finally,
anoxracollularsorinoprooaso,LanF,cloavosoho
-orminalloadorpopidoFig.3A.Yoaswo-hybrid
sysomandco-immunoprocipiaionsudiosrovoalod
hahomodiicaionon.ymosisE,isCand
ranspororisT onisin orm a mombrano-
associaodmulimoriccomplox8iogorsoal.,1996.
AlhoughisCshowsnosoquoncosimilariywihany
knownprooins,acrysalsrucuroanalysisoisC
rovoalod a old similariy wih mammalian
arnosylransorasoLioal.,?006.
Classlanibioicssuchaslacicin481aromodi-
iodbyasingloon.ymo,LanMFig.3E.LcM,ho
modiicaion on.ymo olacicin 481,has boon
proposodousoMg
?
andATFophosphorylaoho
8orandThrrosiduoshaaroargoodordohydra-
ion.8ubsoquonphosphaooliminaionhonrosuls
innodohydraionandhoormaionohoDhaand
Dhbsrucuros.8omooho-orminalconsorvod
rosiduosoLcM aroprodicodoboimporanor
phosphorylaionanddohydraion,whoroashoC-
orminal.inc ligands aro criicalorcycli.aion
Chaor|oooal.,?005,Fauloal.,?007,Youand
vandorDonk,?007.LanToclasslanibioicsisan
AEC ransporormauraionandsocroionAM8
prooinhaconainsaprooolyicon.ymorosponsi-
bloorhocloavagooholoadorpopidoFig.3E.
Tholoadorpopidosharosaconsonsussoquonco
andacommonprocossingsiowih?consorvod
glycinorosiduos,callod

doublo-glycinosio,

aposi-
ions-1and-?Ha

varsoinoal.,1995,Furgorson
hnkonoal.,?008,ishiooal.,?009.ukTisan
AECranspororohoclasslanibioicnukacin8K-
1,andiwassuggosodhaho-orminalpopidaso
domainandC-orminalATF-bindingdomainaroboh
imporanorhocloavagooholoadorpopido
ishiooal.,?011.Lacicin3147isawo-popido
classlanibioic.LnA1andLnA?aromodiiodby
ho rospocivo modiicaion on.ymosLnM1 and
LnM?. LnJ caaly.os ho convorsion o
dohydroalaninosoD-alaninos,andhomodiiodpop-
idosaroransporodbyasingloAECransporor,
LnTFyanoal.,1999,McAuliooal.,?000,Coor
oal.,?005c.
Alhoughclassbacoriocinsdonoundorgopos-
ranslaionalmodiicaion,hoyaroalsosynhosi.od
asaninacivoprocursorpopidohaconainsan-
orminalloadorpopidosoquonco.Asinhocasoo
classlanibioics,mosoclassabacoriocinscon-
aina

doublo-glycinosio

aholoadorpopidoso-
quoncoHa

varsoinoal.,1995.Concomianwih
ho cloavago o ho loador popido, mauro
bacoriocinsarooxporodacrosshocyoplasmic
mombranobyadodicaodAM8prooin-ypoAEC
ranspororFig.3C.Howovor,somo classa
bacoriocins conain a sc-dopondon-orminal
loadorsoquonco,such as bacoriocin 31 rom
/^/cccccas/aca//s31Tomiaoal.,1996,
onorocinFrom//ac/aF13Cinasoal.,1997,
lisoriocin743Arom//s//a/^^ccaa743Kalmoko
oal.,?001,onorocin8E-K4rom//aca//sK-4
Doioal.,?00?,bacoriocinT8rom//ac/aT8
DoKwaadsoniooal.,?006,andhiracinJM79
rom /^/cccccas|/aDCH58ancho.oal.,
?008.
Forallgonoicallycharacori.odwo-popidoclass
bbacoriocins,ho?srucuralgonosarooundnox
ooachohoronhosamooporon,andho?popidos
ACTFkCNSFkMCkAM-STVFACTFkA
haconsiuohobacoriocinhusappoarobopro-
ducodinapproximaolyoqualamouns.Classb
bacoriocinsalsoconainadoublo-glycinosioaho
loadorsoquonco,andhoAM8prooin-ypoAEC
ranspororisrosponsiblooroxracollularranspor
Diopoal.,1996,Fran.oal.,?00?,Zondooal.,
?006,Oppogardoal.,?007.
Forclasscbacoriocins,homochanismobio-
synhosisromainsuncloar.Alhoughhoon.ymosro-
sponsibloorcycli.aionandloadorpopidocloavago
arosillunknown,hogonoshaarominimallyro-
quirodorbacoriocinproducionhavoboonidoniiod
oronorocinA8-48,circularinA,gassoricinA,and
carnocyclinA.Thoyincludopuaivomuliplomom-
brano-spanningdomains,ATF-bindingprooins,and
asmallhydrophobicpopidohamayboinvolvodin
immuniyMarino.-Euonooal.,1998,Komporman
oal.,?003,Kawaioal.,?009,vanEolkumoal.,
?011.
DivorgicinAWorobooal.,1995,lacococcin
K.SNMTFTAL.
.Eiosynhoicon.ymosinvolvodinbacoriocinproducion.Frocursorpopidosconain-
ingan-orminalloadorsoquoncoaroribosomallysynhosi.od,modiiod,andsocroodas
mauropopidos.AEiosynhosisonisinAclasslanibioic.Frocursorpopido,isA,is
dohydraodbyisE,cycli.odbyisC,andsocroodbyAECranspororisT.Tholoador
popidoiscloavodobyoxracollularsorinoprooasoisF.EEiosynhosisonukacin
8K-1classlanibioic.FrocursorpopidoukAisdohydraodandcycli.odbyukM,and
holoadorpopidoiscloavodoaa

doublo-glycinosio

concomianlywihransporby
ukT.CEiosynhosisoAM8prooin-dopondonclassbacoriocins.Thoprocursorpop-
idoconainsa

doublo-glycinosio

ahoC-orminusoholoadorpopido.Tholoadorpop-
idoiscloavodobyAM8prooinconcomianlywihranspor.DEiosynhosiso-
dopondonbacoriocins.Thoprocursorpopidoisribosomallysynhosi.odwihho-
orminal-signalpopido.Thosobacoriocinsdonoroquiroanydodicaodranspororbu
arosocroodbyhogonoral-pahwayohoproducorcoll.Thosignalpopidoiscloavod
oduringranspor.
97?Marino. oal.,1999,and somo ohor
bacoriocins aro sc-dopondon class d
bacoriocins.Thosobacoriocinshavoasignalpop-
idohaaccossoshogonoralsocroorysc-pahway
andonablosranslocaionacrosshocyoplasmic
mombranowihouanydodicaodranspororsDiop
andos,?00?Fig.3D.EnorocinL50Cinaso
al.,1998,auroocinA70o.oal.,?001,lacicin
OFu|iaoal.,?007,andlacicinZwaanioal.,
?007aroloadorlossbacoriocins,whicharosynho-
si.od wihou an -orminalloador soquonco.
Doailod biosynhosis mochanisms oloadorloss
bacoriocins,suchasrocogniionbyransporors,ro-
mainuncloar.naddiion,howhoproducorsrains
proochomsolvosromanibacorialaciviywihou
aloadorpopidoissillanunansworodquosion.
GeneticorganizationandgenereguIation
Thogonoicdoorminansobacoriocinsarolo-
caodoihoronhochromosomooronplasmids,and
oonassociaodwihransorabloolomons.Tho
gonosrolaodobacoriocinproducion,including
srucuralsoquonco,rogulaory,modiicaion,rans-
poror,andimmuniygonos,arogonorallyoundin
clusors.Lanibioicgonoclusorsarocomposodo
hosrucurogono/a^^,modiicaionon.ymos
/a^/,/a^,or/a^/,AEC ransporor/a^1,
prooolyicon.ymo/a^F,sol-proocionprooin
/a^//C,/a^/,or/a^/,andranscripionalrogulaors
/a^/,/a^/,/a^6,or/a^WilloyandvandorDonk,
?007.isinproducionisoncododbyagonoclusor,
^/s^/1/F////C.isinacsasanoxracollularpop-
idophoromonosignalinvolvodinhorogulaionois
ownbiosynhosis.ThorosponsorogulaorisFand
hohisidinokinasoisKaroalsoinvolvodinho
rogulaiononisinbiosynhosisEuchmanoal.,
1988,Kuiporsoal.,1995a,Kloorobo.omandOuadri,
?001.8ubilin,alanibioicsrucurallyrolaodo
nisin,ollowshonisinmodol,whichissub|ocosric
auorogulaionKloorobo.om,?004,Kloorobo.omo
al.,?004.Morsacidinisaglobularshapodclass
lanibioicandisalsorogulaodbyanauo-inducion
mochanism8chmi.oal.,?006.Thorogulaory
characorisicsonisinbiosynhosisworooxploiodo
dovolopa^/sinconrollodxprossionCEsysom
or//ac//s.8ovoral^/s-promoorvocorsworocon-
srucodhaappoarodsuiabloorighlyconrollod
oxprossionobohhomologousandhoorologous
gonosin//ac//sKuiporsoal.,1995b.
Classabacoriocinsroquiroaloas4gonosin-
cludingsrucural,immuniy,AECransporor,andac-
cossoryprooingonosEnnaharoal.,?000,Fimland
o al., ?005. 8omo mombors o class a
bacoriocins,such as curvacin A,sakacin F,
carnobacoriocin E?, and onorocin A, aro
ranscripionally rogulaod hrough a hroo-
compononsignalransducion sysom.Fopido-
phoromonoinducionacorinoracswihamom-
brano-associaodhisidinokinaso,horobyriggoring
hokinasoophosphorylaohoinracollularrosponso
rogulaors o acivao ho gonos noodod or
bacoriocinproducionKloorobo.om andOuadri,
?001.8omoclassbbacoriocins,suchasAEF-
118,planaricinE/F,andplanaricinJ/K,aroalso
knownoboranscripionallyrogulaodhrougha
hroo-compononrogulaorysysom.FlanaricinAisa
popidophoromonohainducoshoproduciono
planaricinE/FandplanaricinJ/KAndorssonoal.,
1998,Flynnoal.,?00?.
SeIf-immunitymechanism
Eacoriocin producors usually produco sol-
immuniyprooinshaproochobacoriaromboing
killodbyhoirownbacoriocins.nhocasoo
lanibioics,hoAECransporor-ypoprooinLanFEG
andspociicimmuniyprooinLanorLanHaroro-
sponsibloorsol-immuniyWilloyandvandorDonk,
?007,Okuda and 8onomoo,?011Fig.4A.
LanFEG prooins scavongo mombrano-bound
lanibioicsromhomombranooprooccollsrom
aackbysol-producodlanibioics.8paFEGoho
subilin producorKloin and Enian,1994and
isFEGohonisinproducor8iogorsandEnian,
1995woroidoniiodasimmuniyprooinsaho
oarlysagoolanibioicimmuniysudy.LanFisanu-
clooido-bindingdomainEDohoAECrans-
poror,andmuliplosoquoncoalignmonanalysiso
LanFprooinsandEDsoohorAECransporors
rovoalodhaLanFprooinshavohoE-loopasavari-
anoho O-loop consorvod in gonoralAEC
ransporors,andE-loopwassuggosodohavoan
imporanroloinhouncionoLanFEGOkudao
al.,?010.Lanprooinsarodividodino3groups,in-
cludinglipoprooinsisand8pa,immuniypop-
idosoxisingonhomombranoandoxracollular
spacoFopandEci,andransmombranoprooins
Ln,Cyl,and8unEngolkooal.,1994.ukHis
aransmombranoprooinoundoboinvolvodinsol-
immuniyhroughisbindingaciviyagainsnukacin
8K-1Asooal.,?005.LanorLanHalonocanno
conorullimmuniy,anduncionscooporaivolywih
LanFEGOkudaoal.,?008.
Thosol-immuniyprooinsoclassabacoriocins
arowollsrucurod-holicalprooinshaconsiso
bowoon88and115aminoacidrosiduosanddisplay
5%o85%soquoncosimilariiosDalhusoal.,?003,
8prulosoal.,?004,Fimlandoal.,?005,Johnsono
al.,?005.LacococcinAisabacoriocinhaargos
ACTFkCNSFkMCkAM-STVFACTFkA
man-FT8ohosuscopiblocolls.Thoimmuniypro-
oinLciAhasboondomonsraodoorm asrong
complox wih ho rocopor prooin and ho
bacoriocin,horobyprovoning collsrom boing
killod.Thocomploxisormodonlyinhoprosoncoo
bacoriocin,andhomombranocompononsCand
D oman-FT8 aro suicionorsonsiiviyo
lacococcinAaswollascomploxormaionwihLciA
Diop oal.,?007Fig.4E.Forclass b
bacoriocin,iishypohosi.odhaholacococcinG
immuniyprooinalsoinoracswihholacococcinG
rocoporisson-Moyoroal.,?010.Thogonoclus-
orsoclasscanddbacoriocinsconainAEC
ransporor-likoorsmallhydrophobicpopidogonos
hamayboinvolvodinimmuniy,buurhorsudios
aro roquirod o undorsand hoirsol-proocion
mochanismindoailisson-Moyoroal.,?009,van
Eolkumoal.,?011.
AppIicationsofbacteriocins
LAE-dorivodbacoriocinsaropromisingasood
prosorvaivosgivonhoircharacorisicsohavingno
aso,noodor,highsabiliy,andspociicaciviy
againsoodbornopahogonicandspoilago-causing
bacoria.LAEbacoriocinsaroconsidorodassao
compoundsorhumansbocausoohoirnonoxiciy
andhoGFA8sausohoproducor,whichisaood-
gradoorganism.isinandpodiocinFA-1canbopur-
chasodorusoasoodaddiivosas

isaplin

and

ALTA?341,

rospocivolyCooroal.,?005b.
Thocombinaionobacoriocinsandohorroa-
mons,suchaspulsodolocriciolds,highhydrosaic
prossuro,andorganicacids,onhancoshopooncyo
bacoriocinsand provonsho growh oGram-
nogaivo bacoria such as /sc|/c|/a and
FsaJcc^as,whichcannoboinhibiodbysimplo
roamonwihbacoriocinsAnanouoal.,?004,
Grandooal.,?007,Marino.Viodmaoal.,?008.
K.SNMTFTAL.
.8ol-immuniymochanismonukacin8K-1AandlacococcinAE.AukFEG
isanAECranspororharansporscoll-associaodnukacin8K-1,andukHoxhibisbind-
ingaciviyagainsnukacin8K-1.ukacin8K-1,capurodbyukH,isransporodoho
oxracollularspacobyukFEGinanonorgy-dopondonmannor.ELciAisasol-immuniy
prooinorlacococcinA.LciAormsasrongcomploxwihlacococcinAandisrocopor
prooinsman-FT8,andhorobyhocollsaroproocodhroughhoblockingohobound
lacococcinAromadvancingohosubsoquonsopshaloadocolldoah.
8omo lanibioics such as nisin,morsacidin,
muacin1140,andlacicin3147aroroporodobo
acivo againsdrug-rosisanpahogonssuch as
MF8AandVFE,whichmakoshomanaracivoop-
ionaspossiblohorapouicagons.isinFwasin-
|ocodino5aaas-inocodmicoandwasabloo
conrolhogrowho5aaasoraloas15min/^
/cErandoal.,?010.Frovioussudiosovaluaod
hoaciviyonisininhoroamonobovinomasiis.
Eocausoroamonosubclinicalmasiisisnosug-
gosodorlacaingcowsoavoidhoriskomilk
conaminaion,nisinmighboapooniallysaocom-
poundoroahisdisoasoCaooal.,?007,Wuo
al.,?007.
naddiionohousoobacoriocinsasnovolani-
bacorialcompoundsinoodorpharmacouicalprod-
ucs,hoapplicaionobacoriocins,inparicular
lanibioicbiosynhoicon.ymos,inpopidoonginoor-
inghasroconlyaracodaonionCooroal.,
?005a,agaooal.,?006,Finkoal.,?010,agaoo
al.,?011.Lanibioicsarogono-oncododand,horo-
oro,canboroadilymanipulaodbygonoiconginoor-
ing.8ovoral/^/cand/^//capproachoshavo
boonoxplorodoonginoornovollanibioics.Anum-
borodioronsysomshavoboonusodomuao
spociicaminoacidsinlanibioics,andsomooho
producod muans displayod improvod aciviy
Cooroal.,?006,Fioldoal.,?007,Lubolskioal.,
?008,slamoal.,?009.Thousoolanibioicmodi-
icaionon.ymosoinroducounusualaminoacidsin
/cc//hasmanyadvanagos,suchasoasygonoic
manipulaionandrapidandinsanusoagaooal.,
?005,Caoanooal.,?011.Lanibioicbiosynhoic
on.ymoscouldinroducounusualaminoacidsino
noonlyhovariansoparonalpopidosbualso
nonrolaodpopidosusodoholoadorpopido
Kluskonsoal.,?005,Chaor|oooal.,?006,Finko
al.,?007.nroducingunusualaminoacidsinonon-
lanibioicpopidoswouldboapoworulooloim-
provohosabiliyopopidosoinoros.
ConcIudingremarks
Wohavoroviowodroconprogrossinhocharac-
ori.aionobacoriocinsproducodbyGram-posiivo
bacoriawihrogardohoirdivorsiyinsrucuro,
modosoacion,biosynhoicmochanism,gonorogu-
laion,sol-immuniymochanism,andapplicaions.
Thonumboronowlydiscovorodbacoriocinsconin-
uosogrow,aswollashooxpocaionorhoirpo-
onialusoasalornaivoanibioics.nparallolwih
scrooningnovolbacoriocins,horoisaprossing
noodoundorsandhoirmodooacionandsol-
immuniysysoms,whichshouldrolaoohomodoo
acionobacoriocins,omoohouurochallongos
odovolopinganibioicshacancombaomorging
drug-rosisan pahogons. Tho oauros o
bacoriocinshaiisribosomallysynhosi.od,onablo
hoir manipulaion hrough gonoic onginooring.
Enginoorodbacoriocinsmayacquiroahighorani-
bacorialaciviy,workovorawidorspocrum,do-
volopimprovodsrucuralsabiliy,oracquiroohor
unoxpocod characorisics. Tho divorsiy o
bacoriocinsmakosidiiculorscionissoullyun-
dorsand hoirmochanism.Howovor,housoo
bacoriocinsincombinaioncanhavowidorapplica-
ionscomparodwihhousoosinglobacoriocins.
Addiionally, knowlodgo o ho divorsiy o
bacoriocinscanpavohowayordosigninghighlyo-
ocivobacoriocins.
Foconapplicaionsolanibioicsincludohouso
ohorogulaorysysomonisinbiosynhosisocon-
rolgonooxprossions,suchashoCEsysomin/
/ac//s.Lanibioiconginooringisanovolpopido-
dosigningoolhaakosadvanagoohobiosyn-
hoic machinory olanibioics.8omo oho
lanibioicon.ymos,suchasisEKluskonsoal.,
?005andLcMChaor|oooal.,?006,aroappliod
oinroducounusualaminoacidsinounrolaodpop-
idos usod o hoir cognao loador popido.
nroducionounusualaminoacidsinopoonhora-
pouicpopidosmayincroasohoirbioaciviyorolor-
ancoagainsprooaso.
Eacoriocinsaropromisingcompoundswihpoon-
ialapplicaionsinhooodandpharmacouicalin-
dusrios.Moroovor,wihincroasingknowlodgooho
biosynhoicmochanismsobacoriocins,nowappli-
caionoolscanbodovolopodorpurpososocon-
rollodgonorogulaionandpopidoonginooring.is
ourhopohaurhorsudiosobacoriocinswillload
owidorapplicaionsohosocompoundsinmanyas-
pocsoovorydaylio.
AcknowIedgements
Thissudywaspariallysupporodbygransrom
hoJapan8ocioyorhoFromoiono8cionco
J8F8,Japan8cionco8ocioy,ovarisFoundaion
JapanorhoFromoiono8cionco,ovo.ymos
JapanFosoarchFund,andagaso8cioncoand
TochnologyFoundaion.

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ACTFkCNSFkMCkAM-STVFACTFkA
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