Secondary Metabolites From Marine Bacteria

8/2/2019 Secondary Metabolites From Marine Bacteria

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Annu Rev Micbol. / 44Copyrght by Ann Reews Inc A rghts reseed

STRATEGIES FOR THEDISCOVRY OF SECONDARY

METABOLITS FROM MARINE

BACTERI: cological

Perspect yes

Pu R. Jnn nd W FnScpps Instition of Oeanography, La Jolla Calfoa 9090

Y WORDS: bacterial ecology, biological activity marine natural products chmitry

ONTENTS

INTODUON . . . 560

ECOLOGCAL CONSDRAOS ....... . . . . . . . . 561Ptc Dv . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3e Dtut Batea te Sea .. . ... ... 64Te at Batea t lat ad Iveebate . 66T R St Bacta 67t Ma Mtalte p t Dtut Bactea 69e Aptat Bata t Dee Ma 570 7Te Dtt Bt Ma ad Teetal Batea 572

BOLOGCA ACVS AND NOV ORANC SBSANS....... 574

ONUDN RMARKS 76

T

Marine microorganisms have become an imporan poin of sudy in he search

for novel microbial producs. Today, both academic and indusrial ineres inmarine mcroorgaism i on he rse in par becaue of he growing numberof unique biologically acive econdary meabolie repored from marine

baceria abiliy o asses he biosynheic poenial of marine baceriaoeer i ieitably coped to or baic dertandig of their bioogy.Ti eie mmize the chemica dicoerie ad biologica aciiie

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ENSEN & FENC

repote from marne batera an fouses upon several mrobologal opsas hey elae o aal odc dscovey, cld he dsbos, dvesity ad clrably of arie bacea as wel as he ole of syboic

batera n the prouon of substances prevously ascrbe o other sources.

INRDUIN

Hstorally the stuy of natural prouts has fouse upon the elaborae

bosynhec paways of eestral plants and mcooanss I he lae1960 howeer the seach for novel metaboltes took a new recton as thereal of exploaion expae to nclue plans an anals n the sea Ths

avene of esearch was naed paly by acaecas and faclaed byhe develope of sba, whh ovded a eecve eans o olle shallow-wate aine orgasms Today as the esult of extensve eseach thestuy of mane natural prouts is eognze as both an negral omponen

of atual pos hesy a a sgnfiat esoue fo ug scovey aton the suesses of mare natural poucs chemsty have helpeue he own scplne of ane hecal eoloy an aea of eseachthat has otrbute sgncantly to our unerstanng of the ecologal rolesof marne seonary metaboltes (see 142)

he scovery hat mane ogasms repesent a rh soure of prevoslyescrbe seonary metaboltes s not entely suprsng onseng thatay alal an nveebae phyla rese exclsvey he sea n hndsht

this disovery sees only loial ven that many soboe aine oa

ss lakin obvos suctural efenses podce toxc seconday etabolitesas a means of hemal efense Altough our unerstanng of hemcal

efense an ohe eooga oles of ae seonay eaboes has jus

ben chemal suies of ane oanss have been extensve, yen

thousans of unique metaboltes feaurn new stuual lasses fntioalgrops an a relatvely hgh ncence of halogenaton Desrptons of thechemal iscoveries i this fel can be foun i peioically upae reviews

fo 977 to ate (36 ad prevos evews ced there)As the checal polfcacy of ane oanss becae appaent so

ther ustal potental an marne metboltes are now ben sreene

extensvely for pharmaologal ativity The resls of hese eots have le

to seveal sfcant boecal dscovees These ncle he clical eval

atio of at least two aine natual pocs as antcancer agetsbyosta

43, 6 whch s cuetly n lcal ials n both Grea Bta and heUted Saes an e B 50 whch ecenly oplete phase laltals n he Ue States (see 77 descios of he boloal aviesof varos marne plant an nvetebrate taxa (151) as well as etale revews

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SECONDR MEBOLIES ROM MRINE BCTERI 56

of the bomedal potental and pharmaologal popetes of mane metaboltes, a numbe of evews ae avalable 8

Contnung efforts to obtan hemally unque marne organsms have led

to the use of new and nnovatve methods of sample ollecton. These effortsnlude travel to far reahes of te gobe to olet prevousy unstudedspees the use of deepdvng submesbles to aess habtats that le beondthe depths of conventonal scuba, and the colleton of ogansms that weepevously overlooed owng to dultes n obtanng adequate bomass forchemcal evaluaton These efforts have aompaned a growng nterest n thesudy of arne batera, a logcal and potentally productve avenue of re-seach gven the hstocal sgnfane of seonday metaboltes fom teestal bactea and the enewable natue of ultuable esoues

Durng the past deade studes of seonday metaboltes from arne batera have begun n laboraores throughout the world. n the most smpledsense these studes use standard dluton and platng tehnques to solatebatera from marne samples. Isolated batera ae cultured n shaefasson aga plates, o n fementos, and he oganc components of these cultuesae then xtrated, usually wth ogan solvents. The extracts are tested fobologal atvtes, eg standard antbot ds dffuson assays, and analyzedfor the presene of unusua hea onsttuents Organc materals of nterestare purfed and the stutues analyed, a omplex process that geneally

eques sophstcated NMR ethodologesBeause of gowng nteest n the study of seonday metaboltes fom

marne batea, we would le to address n ths revew several tops nmobal eology that ae elevant to natual produt dscovey. s these topsae boad n sope and fundamental to mane mcobolog, ou goal s tofous upon e aspets at pean to natural produt dsovery. Rathe thandsuss n detal hemal dscoveres from marne batera, we refer the readeo recent revews orented toward ths objetve (5 4087 11) and ofera sumary of the bologa atvtes and novel seondary metaboltes e

poed to dae fom arne bcea All of the followng dscussons of marnebter elude yanobatera (unless otherws ndated) whh are none

theess an mpotant esoure for unque seonday metaboltes (36 and pe-vous evews ted theen) n addton, one should note that novel seondaymetaboltes hae been repoed fom oher taxonom groups of marne mcoogansms, ncludng fung and mroalgae; however, these goups ae notdsussed here.

EOLOGIL ONSIDERTIONS etendng the seah fo novel bal eaboltes to the sea a fundaental queston ust be addressed, .e do arne batra produce prevously

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56 NSEN & FENC

Novel ogaic molecules ted from marine bacteria

Prcin san Soue Bomedia ativity

Acomycete

haa rea Sedient Antbotc, aicancrremyes ejmaress Sedimet Atbiotcremyes rses Sedimet Atibiotcremyes syaess Sediment AticancerMadromycete Sedimet Atibioticremyes sp. Sediment AntiboticActinomycte Sedimet AtibioticActinomycete Sediment No activity epoedActinomycete Gorgonia sot cora Atcancer atibotcremyes sp Jllysh Atiioti, aammatoyremyces sp Spoge AtibioticNoactoycete

semas rms (Aerm Searass Atibioticas sp.)

hrmaerm Aermas sp Seawater AtibioticPsedomonad Tide poo seawaer AnibioicBeeea aes Vbr Salt s edmet No ativy epoedAermas ra Not reported BohodlaoUnidetified Seawater o activity eporteUnidentified ram-positive scies Deepsea sedment Atcacr atviaAerma sp Spoe AtcacAermas haas Deep-sea sedimet AticacerVbr agarm Fi No activty repotedAermas sp Seawater Ezyme btioAemnas sp Sng AtmoiaPseudmas sp. Anticanceredmna n Ada AmcobaUnidentiied am-stive scies Deep-sea sedment AaeTems Hyotemal AntunBas Deep-sea sedimet AticacerAermas eaea Seawater No atvty epoed

Referenes

7 20, 7 77

18 10

059

527

desed seoday meaoles ad so, ae evomeal aos se o e oea eed o e odo o ese ssas Ts eso

s eme o oe dameal o e develome o mae moologys snf dsp: h s do epes uue spso meey eetral stas aaed o he odo o he sea

I esose o the atte eso, he exsee o se mae aeaas domeed ealy 50 yeas ao see 08. Te aly o mae aeao ode evosy udesed seoday meaoes has also ee ade-uately etaihed ae ) oeig the odtio eea o theodo o seoda meaoles mae aea,he solao o a ovelmeaole does o sel ld osess o s same ssae

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SECONDR METBOLIES ROM MRINE BCERI 56

b a nonman san. Howv hs sbsanc s podcd on whncondiions pcic o h main nvionmn a povidd .g. sawaincasd hdosaic pssu o a man nun h liihod o is

subsanc bing pducd as a nonan nain podc is lw Moo i hs sbsanc posssss as a obsvd wih mais omsa baca sch as h psnc o halogn aoms h agmn os uniq man nau s uh snghnd.

Fo a naual poucs pspci, an acia ain a laiunplod soc o nol sconda mabos. uh nvsmn inhs aa fom boh an acaic an nusial pspciv wi ulimamin h uli of his souc To maim h civnss o uusach a b ndsandng of man bacal coog s ssnial Ths

unsanding can oand hogh conind sdis in o ampl hdvs dsbion and clai o man aca and h adapaionsof bacia o suvival in oh nomal and m main nvionmns.

Prokaotic Diversi

aca anmc an abolic is a unamnall inaa f pan naua poduc isc In ain sss hw- a nl bginnng o unsan h u n of paoic

s hs fac s aqual monsad b h iscovs o abndanpochloophs 22 canobaca 89 and mos cnl acha a 28 42nhabng h wol's opn ocans. Ths dscovs n pa accound o h appcaon o chnqus n molcula olog o h sd o mcoacolog confm pvious blifs ha acal ivs n main ssms sa ga han can b accond o b h cdpnn chnqspcall s o scb bacal communis s sis o pokaoics coninu h knowlg gain can b us assss h dg och s bs f h a paoc couni can oband culu an ncu n h sac o nol cobia pocs

bsa a aca s ka ha o bn gnaua cosao n cical suis f an bacia is supp bh lal fw gna fo which no oculs ha bn poabl 1) hs h onl aa ha podc nq mabols o a hspl h nl aa si o a? os sach in hs ld sms o havccnad upon maindd acincs and gamngaiv hopic bacia capabl f gwh n nuinich mdia. This appachs logcal as h solaon and cl cnqu sd o hs bacia alavl simpl and man sains can oband wh minmm o.oo bcaus sus f an baca ha n bn nsi hsas sola ll ll b n naua pducs sach. This appoach

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54 JENSEN ENC

oeve eglets te osslty tat ote vese taxa oue ovel seoy metotes.

ig tu oucts stues to iclue ivese ctel gous eg

aaeoes) eues sec moologcl exetse a esets te oo-uity o ollaoatos etee atual outs cemsts a moolo-gi amla t tese metodologes. Te alleges of estalsg amcoil sceeg pogm eseclly oe tt clues vese cteltxe ell ko to te pamceutcl iusty, te ppoces useo mee tese calleges ve ee esce e.g 12,1,2,14 Altoug coeion etee xonomic ad ioytetc vesty as yet to edefined fo mne micoognim, e incuion of diee cei x intul poucts eseac stll epesets atol ppoc to ovel meto-

lte iscovey.

The Dsrbuon of Ber n he Se

Te actos cotollg te stiuto o te te se e comlex vstly eet om tose epeece y teestal ate Te most po-found of ee diffence my we e queou ilieu ief wic poves eectve meum o ceil disesl oseg tt seteycy contns 104 o 06 actera m-1 4 nd ocen curent cn trnorthee onim conidebe dince mine cei my encoune a f

gee iey oat nd enionmen condiion t teestal ate- Te alty o mae cte to suvve utle te esouces oui tese vious itts il ultimtely ve sgiict eect up teistutio te sea

Te atats tat mae atea eoute lue ufce ot nimendmate. I con o eei em mine uce e coniuallyate y assemlage o ctei Te suitaility o tese vious suceso coloto may sgctly ect ctel stutos me -. Aou e nume nd ype of cei coonizin mine ufceve no een dequey documened e coonion poce nd e fcoectg coloiato ve ee iscusse etl 4 41101 11 17

Fom te esetve o nu oduc dicoey a geeal undendingo te istiutio o maie cte a te ctos ectg tese istiutos is eecl I speciic txoomic goup o cte s cose ocemicl stuy iomtio esciig te istiutio o tt goup illcltte te colleto o ote samles I g eeal,tese smples maynude ewe edimen a ot nime nd innime ojec [Seu11) as evee te ssotos o te t mae smples. Coseg tt te stiuto o cte s ot uiom tougout te mieevomet ulttve escptos o tese stutios ca e use tomxime e nume of dee cei tax otae in ultue.

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SECONDRY MET BOlTES FROM MRINE BCRI 565

In cour o iolaing arin bacria or cical valuaion w avurvyd diribuion o on cicaly igniican group acinoyc T b ar ourc o acioyc idid u ar ar di

n ro wic hir ioaion i w docund ( 5 44 47 ) T raiv nubr o acinoyc in arn dna low owv ad lciv iolaion od (g 4 54

wic includ a and anibioic ran ar uul. Following iolaion od uiabl o cuur and cical vauaion o arinacoyc cud o ud ioically by paracuical iduyor oi acinoyc wi o coon dnc bing incorpoaion o aar or pcc an u no ow du (.

)

vau o udying acinoyc diribuion in din i donad by obvaio a hir ubr can vary dpndi upon dp o wic ap ar cocd ( ) or xapl a udyo nar-or an dn tpoyc wr ncound nrqunly

pa ubliora on wl acnoplan w iolad n rar nuba diac o o crad () T oao no ony uu

n arging pcic aa or olaion bu can b ud o a indig

ou au o acoyc olaa pota codao a ogiad acvy o acoyc olad o a ouc a b uod

53 56 137 quo a vad v a acoyc a oabudan ral oil a in ain dn () ow varygdgr o al olranc ( 4 7 77) and poduc por a undoubdly wa lag nubr o or ino a w y ay avabl o a udind pod o

Aoug quon abou ndignou arn nau o acnoyc

a dicul o addr paly o pog a bn ad n gard o xapl w av obvd a dgr o c adivdacoyc uir alin condiio o grow vai dpdig upon

aooc up udd () I th apl vually all ddvdacoplan urd aa o go and oud gar nuba dac o o cad T obvao u a aci

opla ly adapd o h an vionn and rprnoblga a baca On o and ub o poycdcrad a dianc ro o incad bacra could grow n aoa diu wic ndica a rpoyc a aculaivlya ad pobly o al oi. Radl o oi o axooc

aliaion all o acinoyc iolad grw luurianly n a awarbad diu ad or av poal o no only b abolicallyacv n arin viro bu o poduc uu aboli wn

povdd pcc a go aco

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ESE & FE

The Associations of Bacteria with Plant and Invetebate

The dsbuon of bacea on vng mane surfaces can reveal e extent to

which tese ognisms e consistntly ssited wi scic plns ndnimls. Docuenting te spcicity of ese ssiations is an intgral butnot in itelf conclusive par o dening bacteal symbioses e quest fornew ntu poduct, thee ston e pculrly sgncnt becusehe potenal ole of bacteal symbionts in he pduction of metabolites

peviously ascbed to other oganims is open to debte ee secon onbacteil symbioses) see 3 39 Without evidence fo e consistent ssociation of bactei with a gven pan o inveebate one cannot disngush

ymbionts om nsient ssocites nd te role o symboc bacte n

metabolte poducton becomes obscuedepite increasng documentaon o he coextence beten bacte ad

ine invetebtes (see 6 conclusive evidence of symboses s oenlackng (4. Among the nvetebates frequently collcted for natual producttudy ponge-bactel ocation e pobably e mot thoughly descibed [s eview by Wikinson (94]. ese stdes showd t spongeassociated bactea can be disnct from bactera n e surroundng seawter5 9 95 Also phenotypically sila bactea wee isolated fom 9 o sponge species collected om two distinct geogpicl egions (95

hese esults sggest that the bacteial counites associated with spongesy be highly pecic Howeve hi pecicty may not apply to ll spongeseg.

In the case of coelenterate another goup of organms typicaly collectedo cheical study the mucus of indivduls within he Odes Zonhdae andGogonce can hbo dense bacteal counites (33 55 83 Despitethee obsevations le s knon about how e communties va ithin obetween spies. In ddition bctei ssocited wih lge cn be disnctom bactea in the suounding seawater(3 84 2 6 but te sccityo hese ssociation i not clea Given the lmted numbe o bactel associtions descbed eltve to te numbe o divese mne orgnisms vlbleo study this subject emains a elatvely uneploed aea fo futue eseach

lthough much remin unkown bout bctel on th maneplants nd invetebate many highly pecic associatons hve been docu

ented These eamples include the association o bioluinescent Vb

fscher with he light organ o cephalopods (9 dniogen-xng bcteiawih both te gstointestinl tct o sea uchins (5 nd te gland ofehye in shpwoms (88 ubcutcula bacte i echnodem 5) and the spiochete Crsspr p with the cysalline style o molluscs96) The boad spectum of ese associatons suggests that scc bacterialinvetebate symbioses e common in he sea. Future studes may even ho

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ONDARY MTABOLIT FROM MARIN BATRIA

ha he majory of marine oganisms have specific baceria populaionsassociaed ih heir exeal surfaces and ineal issues

A recen observaon in our aboraory ndcaes ha unusual and prevously

undocumened bacerial assocaions occur ih marne agae During a sudyof baceria in he coral reef environmens of Belize Cenal Ameica we fondha a large percenage of he culrable baceria associaed wih agal sufacesere gram posiive More specificaly 30% of he baceria associaed wih hegreen alga Hameda sp and he bron aga Lobopho areaa ere gramposve (3) Ths observaon conradics he eary beef ha % of marnebaceria are gram negaive (20). Alhough gramposiive baceria have beenrepored in greaer abndance in sedimens () he relaive abundance ofhese baceria in mos marne habas has no been adequaely assessed

We have included unicelular gramposiive marine isolaes in our sdiesof bacerial secondary meabolie producion Of approimaely 00 srainsseened hus far promising insecicidal aciviy enzyme prodcion cyo-oxicy and a eas one novel molecule (7) have been deeced Alhoughmarne gramposiive bacera do no appear o rva he acnomycees in heirabily o produce arge quanes o srucurally compex secondary meabo-lies addiiona sudes of hese bacera are cleary araned As nvesga-ions ino he disribion of gramposive and oher baceria on he srfaces

of marine plans and nverebraes coninue previously undocumened associaions may be revealed These associaions may eec baceial suvivasraegies ha nclude he producion of unusual secondary meabolies as ellas provde poran nomaon reang o he bacerial producon o meabolies prevously ascribed o oher sources

Rl f ymbc Bacra cda Mablrdc

Hsorcally he search for marne naural producs has cenered around chemi-

cal sudes of mane plans and soboded nverebraes onsderng haally all organsms colleced or chemcal sudy nclude assocaed mcroorganisms quesons abou he rue biosynheic origin of moecules isolaedfrom plans and inveebraes mus be addressed In some cases circumsanialeidence leads o he suspicon ha meaboles are of bacerial oigin hisedence can nclude exremely lo (eg l% by eigh) and oen variablemeabole yields he solaon of he same meabole from diverse invere-brae axa consisen microbial assocaes in he producing organism and hesmilary of he molecue n queson o a previousy described microbia

produc Addressing qesions of biosynhec origin are paricuarly appropriae hen one consders ha n cerain marine inverebraes eg spongesassocaed bacera can accoun or up o 40% of he celluar volume (4)

Researchers nvesgang he bosynhec orgns of marne meaboies

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56 JNN FNI

hav takn svra prmnta approachs to addrss ths quston. Th moscommon approach has bn h solaton and cultur of bactra from hproducng organsm followd by chcal analyss of h culur or h

mabol of nrst. n most cass, ths fforts hav bn unsuccssful,whch s not supsng whn on consds th lkhood of obtanng thproducng stran n cultur and provdng ths stran wh h culur condonsncssary for mtabot producon. f th bactral producton of a mtabolcanno b dmonstrad n culur th documnaon of a spcc bactralpopulaon assocatd wth th producng organsm n an abundanc ha canaccount for th onntaton of th mtabolt n th hot an pd mpo

an daa n suppot of baral b oynhss.obably h most notabl amp of th bactra pducton of a mta

bol orgnaly ascrbd o anothr organsm nvovs th potnt marnnuroon, trooton (X), bs known for h lnss and dah cannct folowng th consumpton of tantd raw pufffsh Fuu spp.). n-vstgatons no th orgn of TTX (4 rvald ha X s prodcd as afrmntaton product of both a b sp oatd from th nttn of aanhdcrab Atergatis foridus (2), and a Pseudomonas s solad rom ad calcaous alga,Jaa s () Mo cn suds hav shown ha TTXs producd by a vary of aonomcally dvrs marn bacra ( 4200 ncludng acnomycs (0 Ths nformaon has ld o h proposaltha bacra ar h sol orgn of X whch may subsquntly accumulan nvrtbrats hrough th food wb (

n addton o X bacra hav bn found o roduc a las two ohrmarn ons. On ampl coms from h dbl mollusc abya japca (4) h consumpon of whch causd at last on outbrak of foodposonng n Japan apca tocty was tracd o h prsnc of nosuugaon and posurugaton boh o whch wr solatd om th dgstvgland of h toc mollusc. Mcrobologcal suds rvald ha a corynform

bacrum,also solad from h dgstv gland of japca could roduchs ons. hrd ampl of bactral ton producton s saon (STX)on of th major causatv agns n parayc shlsh posonng. STX hasbn soatd from dnoaglats, .g. yau ama (5whch wr prsumd to b h sourc of shllfsh tocty. Howvr,shshtocy s not corrlatd wh th prsnc of P amaess (2 Rcnsuds rvald hat Maxea s. solatd rom P. amaess producs

STX ( and ay b th ultma orgn of ths mtabot.In at last wo addtonal studs, th bacral producon of mabots

rv ously ascrbd o nonbacral sourcs has bn rpord Th rst (5dmosrad ha a Mcrococcu s solad from h spong Teaa roducd a srs of dtopprazns prvously rportd from th spongIn h sond study (3) bomnad dphnyl thrs orgnally solatd from

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SECONDR MEBOLIES ROM MRINE BCERI 56

he spone Dsdea sp wee obane as femenaon pros fom wosans o Vibrio sp epoe o be symbons o ysa. Despe s evene o baeal o al ea o ese papes leaves oom o

speulaon abou e e bosyne on of e spone meaboles (7

Oe aemps o eermne e bosyne on of mane meabolesinlue e pysal sepaaon o baea fom os ssues,an e subsequenexamiaio o ese ells o e pesee o e meabole quesio emos suessfl emonsaon of s enqe nvolve e use of flowyomey o sepaae yanobaeal symbons (Oscllatora spogla)ome spone Dsdea hebaea 8). o analyss o e pre yano-baea ese aos prove e s evene fo e loalzaon n

pokayoi symbo ells o a seonay meabole pevosly asbe oan nveebae os Oe eos o eeme e ellla oao o baeameaboles lue e se of moleula pobes; oweve o e bes of onowlee ese meos ave no ye me w sess

Baeal assoaons w plans an anmas pose allenn quesonsin e el o mae aual pous emsy enly e quesons oeaes ees volve e o e of baea e bosyess o meabolespevously asibe o oe oasms Pa of s nees s puely eoloal,bu s subje beomes espealy mpoa wen e meabole qeso

sows pomsn bomeal applaons One s ase s byosan 1, eanane aen solae fom e byozoan Bugula nena n e ase ofbyosan , e evene o baeal on nlues e low yiels (-06%by we) n w e meabolie s poe an e pesene of snpopulaos o baeal symbons e byooan ssues We one osesa B. as o available fom naue quanes a an effevelyfalae bomea evelopmen e possby of pon byosan byemeao beomes a poy

Eet ine eLie upn the Diiuin Beiease of e lak of expemenal evene we an ony spelae aboe effes of plan an nveebae meaboles pon e sbon of baea n e sa. ese ees may be epesse in seveal ways inl eeeee o uesiable baea o e aao o baea a epesefavoable epbons. Alo laboaoybase assays ave nae a baea an espon o mane meaboles e.. 020)lle eoloally elevanaa ae avalable o sues ese sbsaes ave a effe upo sae

olonzn baea see fo a eview of eoloal aspes affein mobal emoa beavoEay evdee suesi e volveme of mae meaboles n e

oo of epbaea dsbuos ame fom sudes of mane alae. In wo

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NSN FNIAL

separae repors invesigaors proposed ha anibioic alga phenos wereresponsib or rdcd nmbrs o sac bacria and oling organisms onhe maine algae agassum naans (6) and scphyum ndsum (5)

Sbsqny, a corraion bween redced eves o epbiosis and he pro-dcion o animicrobial sbsances by marne organisms was prsend asevidence or he ecological role o hese meabolies in anioIng eg ,7,5 74 8 86). Unornaly,h dgr o which animicrobial aciviis

ranslae o in si eecs pon poenia sracecoonizing baceria has nobeen adeqaely demonsraed

Alhoh h in si microbiological cs o marn naral podcsremain lagey ndocmented wo ndings sgges hey may inence hedistribion o bacria on living marin sraces On amp comes rom

h sdy o wo olonia ascidians ha possessd remarkably dieren nm-bers o epibioic baceria (84) Laboraory selemen assays comparing heseascidians demonsrad ha xracs o h sparsly colonid Cysoys aus were highly deeen o bacerial aachmen whie eracs o hedensely colonized ycnum anum signiicanly enhanced he aachmenprocss Ths rsls ss ha h aion o bacal aachmn byhos mabolies may b an imporan acor in he conrol o bacera ponl n marin sracs

A second eample demonsraing he eecs o secondary meabolies ponhe disribion o microorganisms involves he inhibi ion o pahogenic ngiby baceria associad wih he mbryos o wo marin crsacans aaenmacdacyus and maus amecanus n separa sdies,rsearchers obsrvd ha embryos o boh mroylus and H mrin us wr covrdby dense assemblages o wha appeared o be argey nibaceial popaions50,5) Baceria consisenly isoaed rom he embryos o macdacyusand H amecanus prodcd he previosly described meabolis isain andyrosol respecivly boh o which show aningal eecs oward he crs-

acan pahon,gnium llnts Fhr xprimns wih P mro-tylus rald ha baceriaree embryos rapidly sccmb o ngalinecion whras simar embryos reinocaed wih baceria srvived. heseresls ld o h conclsion ha symbioic baceria chemicaly deend macdacyus embryos agains ngal incion Bacerial associaions o hisnare add a new level o complexiy o h mechanisms by which secondarymeabois innce he disribions and specic associaions o marinemicroorganisms

The Adion of Bceri o Divere Mrine Hbi

Thir mabolic capaciy alows proaryoes o srvive viraly across hspcrm o nronmna coniions on on his plan Assming hameabolic and biosynheic di ersiy are correlaed he adapaions o baceria

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SECONDR METBOES ROM MRINE BCERI 571

o divrs mrin bitts provid sminly limitlss voluionry opportuniis or t proution o uniqu sonry mtbolits. Altou briadapt to divrs a viomts av apparntly rivd littl at

ion in sr for novl sonry mtbolits n impotn industrilppliio s o from t sudy of on su roup i.. tmopilir (Achabacta)ssoid i dps yrorml vnts O)s btri produ rmosbl DNA polymrs nzyms, i vbn isoltd, lond, nd mrt for biotnoloil ppliions 7)tr vin of btril adaptions to xtrm mn vironmts in-luds t ronitio tt n mp from rmniv btiumisold fom depths of the Suu Sea is expressed only at increased hydro-stati prssu (9). This disovry suggsts th existn of a gnti pogam

toupls prssur ssin to proti sytsis

Bacterial Culturabili

Minii bri in ulur is rquisi o fiv sudy of sondrymtbolit prodution, bus dqu quntiis of bril biomss nsldom b obtid from saps olld i he fild. Ufouly, tmajory of bacera observed n rain marin sampls do not form olonison the nuren-rch agar radtionay used for he soaion of marin bri1,92). In fac mcroscopic couns perormed direcly on seawaer sampes,usually omplisd i forochrome sans 4 14) typially yildbtril ouns r ordrs of mnitud ir n os obind itpang tiqus . 13) s rsults havel to t rally aptolusio tat mjority o mri batr are uulurabl,a olusiotat raiss srious ons for rsarrs intrstd i maximiin t um-br of divrs isols srnd for uniqu sondry mbolis

Svrl topis dsrv losr srutiny i disussion of bril uluraby For o mos studis av fous upo t bria i str

smpls old fr rom shore Ths my b impornt onsidrin oganc conen of openocean seawaer s generay ow 11),ad t sr bari have evoved surviva sraeges 14), iludi rsposs tosrvtion (7 7) t may drmatially rdu tir biliy to fom oloison nutrintri r .. Difo ) urtmor, disrpny btnvibl d dir ous, for so satr smpls y rsult from loplatin iinis, i. . only a f viabl indiviuals in populatio rpbl of formi olonis on r da (47)

t sould lso b notd t t prt of ulturbl bri lultd

for mri sampls n b ir 0.1 % typilly rportd for str.For xmpl 4 to 11% of t btria obsrvd in ssoition i srospon Caa chr ulurbl 1). Iflivin mrinurac provde nurench envonmens a eavey lar prnt of

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IAL

the cte sscted wh these suces my m clnes when ncultedn nunh wh md the st u nwlede, he ulull-ty cte ssced wth nuench mne htts hs nt een d-

dessed n del. One t th wuld e del such sudes s the mnenveee dgesve sysem whch cn exensve cel cmmun-es (61)

Anhe cncept h must e kept n mnd n dscussns celcululy s he nhen slectvy ll sln meds. Becuse nly lmed g ce wll gw wen ny ne sln med s used,t s n supsng lge numes cte ppe e uncultule.T deemne e ecenge cultule cte n ny cmmunty newuld hve t ttemt multple sln methds nd detemne the sum

ll unqu stns esuln m hs ds hs s n esle hene hepecentge cultule cte epted ny ne smple must e cnsd-eed n the cnex he sln mehds used

We cn ls sk uestns u the cmpstn the nncultulempnn the el cmmuny hs psn nnulule stns evusly desced seces, d ey eesen new seces gene ecen nlyses smll suun (S ctel RNA sequencese sed lg n s se F emle, s een swn ce-lnkn cmmunes cnss dly dvese ctel ssemlgeswthn whc undesced ylgenec gu e cmmn (4 As we ecme we nvel ctel ppulns, cused ets cn e mde ssess the cululty these ce.

Apently, txnmclly dvese mne cte t e eectvelyncpted nt the sech new mcl ducts new cultue mehdsneed t e develped ptcully metds th tke nt cunt the envnmenl pmees ssced wh he hs smpled. hese mehds mynclde te se mnedeved nens, wc ve een swn ndcee dcn nvel meles n les ne ep ( n hs sdy,e ddn ulvezed wn lg ku ch (ana ra s hegw medum nduced e dcn new nc, SS228 Y edevelmen mnedeved nuens my mve u ly culuemne e nd esn ddnl tunts h dsvy nvlmeles

The Ditinction Between Marine and Terretria Bacteria

Eslshng he exsence specc mne ce ws undmentl the

develpment mne mclgy s scentc dscplne. ly studesled he psl h mnc ctc cn c dstngushd m eestlsns sed n e eqemen sewe gw Susequendes dcmened edly deecle sdm equemen gw, wc

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SNDAR MABITS FRM MARIN BATRIA 57

is rary obsr for trrtria strains (0 Athough marin batria akan ofia finion thi monraion of a pi oium rquirmnt forgrowth ha bom wiy apt a a mtho to istinguish main from

most trrra batra [s riw by Mao (08 09]Mor rnty h phyioogia ro of oium wa intigat in gratr

tai or many marin batra,th tranport of arou sout into th i soium pnnt ( 75 96 h nrgy rquir for thi tranport ignat b a piationpnnt soium pump (5 his pump xtrusodium ions,gnating a tansmmbran soium gaint with potnta n-g that an b utii to pfom arious ua funtions inuing thgnration of AP. Oh t a 10 ha shown that th oium rquirmnt inmarin batria is orrat wih h posion of rspirationpnnt

soium ux,a th authors ha suggst that tron transpotrinsoium ux an b rgar as a ritrion for th finition of marinbatia

Athough th phioogia istintions btwn marn an trrstria batra ar ary important th raona for rtriting marin batria to onythos sains that mt a pdn st of rquirmnt i qutionab Foxamp n studis ondutd o th oast of i w obs that morthan 0 o th gampositi batria isoatd o a arity o sousquid awat o gowth n additon a sawatquiing strains

tt ha a montrab rquirmnt of oium for growth. A a rut thbatria an b n as marin ba upon traitiona proposas (0 202Ao bas on ths proposa,batra that i not rquir awatr or oiumfor growth,but wr apab of growth n awatrbas mum,ou babd a haotoant ttia tains ut,shou w ah thi onuiona pio and a a sut iminat nonsawat and/or nonoiumquingbatria from isussions of marin batra? o tak this qution on tpfuth, w an nision a numbr of marin nihs, g thos inhabit bynaua smbiont, in whih batria may n nount sawat Ifths batria do not ha a dmonstab sodium quirmnt, arnt thynonhss main W bi that b stiting man bata to inudon thos stain that mt a pdfin st of rquirmnt w run th isko orooing tan groups hat may b quay w aapt to suri onan or in th sa a w a ingnous marin batria that simpy o notmt th t of rqurmnts hosn

Furthrmor,th rquirmnt of awatr for th growth of ain batriaappars to ary btwn taxa In th as of grampoiti batria, non of

th oi tt in at at four parat tuis rquir awatr for growth8 100 7) Ao,a urrnty rib th gnu Plnococcs whihis ognid as a main gnus do not qui sawatr for growth 2 882 Maod 10 mas an intsting obration garing th gam

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5 JENSEN FENCA

pote coccu Gafa homari (now Aerococcus viridans), the pathogencbacteu that caue eptcea n obte Ctng unpuhed data, he tte

G. homr es require sewer r grw es g

e grsiie seies i e se se rereseies e iigeusr require eier sewer r siu r grw Wi e eei

the odeatey haophc cocc, eg Mrnoos Slnoos pp,porosn hlophl Moos hlobs wi gwiy i % s suis re yiy ise r sr sesse rus er yersie eires ere ers e re i wi grsiie i ise r e se requiresewer r siu r grw. s ubject cetany waant uthe netgaton

In ou opnon, to ake a dtncon beween ane and eea cteaaed upon odu equeent not a potant o natua poduc d

coey a the deonaton that actea oated o the ea peo unque

boynthee he bet exape o unque ane boynthee ay we be

the bactea poducton o haogenated antbotc (eg theui ese e eies ssess eue se eu-ere usie e re e e is resrie sei se rieiis en ei suies eri grw i ese iis rewre.

IOLOGIL TIVITES ND NOVEL ORGNISSNS

he abty o ane bactea to poduce antbacte ubance ws docuented oe than yea ago Neay yea ate, Bukhode andcowoke ( pued the t antbotc o a ane acteu, and ate e yea, Loe 7 ree e sruure is suse igyrie yrre. I e yers we ese ery iseries enuer ne rgi eues rere r rie eri s grwn ncude neay ubtance (abe hee ubance epeent deecae o checa tuctue and pode edence o the untpped boynthetc potenta o ane actea Athough the tota nue o unque oe

ues rere e reis w rxiey e ese wereesrie er reeig grwig ieres i is re reser.

he iogia iiies rere r e eues esrie i Te iue n y antbotc t ao antcance and anta actte o oeee ese actte ae a eecton o boaay accebty and the cenii obecte o the eeach goup epone o the dcoey thoughe ri sreeig eri exrs is essei ieiy ree

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SNDAR MABIES RM MARIE BAERIA

isovey (beause o o hi aes) an o eeiv e bioasayguie hemiaoaon,h boaay an bom burnom or ma rarh groupMaimiing he poenia o ug iovy hough inhoue ing i ao

iu o ma eearh goup oniering he many new an ophiiaargir rn vop in rn yar. or h aon inuriaan aaem ollaboaions oe an eeve avenue by hih o obanbroaprum rnng o baria mabo. A or o rn marnmiobia pou inae,aiiona biomeiay an inuiay impoanaviies i eainy be isovee

mnion ha nov onary meaboie hav been ioa om aimie axonomi goup o maine baeia. This may be miseaing as ohgna, inuing Moraxella (5) an Alcalgenes (6) pou bioogiay

aiv uban ha have no ben uy harariz In aiion,an ani-umo polyahaie ha been olae om a Flavbatium p (180) ananibioi aiviy a aoiae ih a purp phooynhi barumChroau purpuru (7) Hoeve, o he bes o ou noege manyaxa have no b een uie in eai o he pouion o bioogay aveonary maboie.

One genu ha i onspiuousy absen om he maine naua proueaue s Ballu, even hough isoaes ae aiy obaine rom mainseimens ( ) an unusua seonay meaboies have been epoe om

eesia sains 2) Ohe han he een isoaion in ou aboaoy o apvouy unrib y pippi (78) o h b o our knowege,ony one ohe subsanen nymeha been epoe rom a marineacllu p. (33) h ahaa wre alo aben om h man naualpous ieaue pio o he een isoaion o ne yi poysues omsains o anaeobiay uue Therococcu spp (5) These hemophii,uumeaboizing arhaa r o na marin hyrohrma yman he subsanes poue emonsae boh aniunga an anhemini v.

In aiion o he ases aeay menione,he hisoy o bioogia aiviiesepoe o maine baeia is ong Fo exampe ae he suvey o anibioiaiviy by Rosne Zobe () ame monaion ha marin baeiapo anivra (5 76) aniaga (6) aniane (80) an animiobiaav (6 7 5-7 3 4448 56 95 03 04 9) nvgaor havaso emonsae ha maine baeia poue baeioin (68) an nyminhibios (775 7) In some ases, hese aiviies ae ue o poeins oohe age moues o hih ruua uiaion anno b perorm

uinganar pra hnu. In oh a, amp o oa an

hararz h av prnp wre nv ma he een o hee aivi-es hoeve suppos uhe nvesigaion o eonary meaboi e ommaine baeia.

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6 JNN & FNIA

ONUDNG REMRKS

oay marin batra ar bing rogniz as an important rsour or

mirobia prouts. h growing intrst in this rsour is aquaty mnstrat by th numbr o novl mtaboits rnty rpot rom marinbatra an th numbr o rsarh abratois thoughut th wor nowworking in this r th hmi potntia o marin batra to bfftivy assss howvr ontinu basi rsarh in marn mirobioogy is ssntia. his rsarh is n baus w o not yt hav a arunrstaning o th typs o marn batra avaiab or hmia stuy thnihs thy oupy an th mhanisms by whih thy ar aapt to thman nvironmnt

h ous o our rsarh i marn mirobioogy is to isoat nov sonr mtabois rom marn batria a o ask oogia qustions aboutboth th prouing strains an th ros o th mtaboits prou. n thurs o this wor it sms appopat to onsir our rsuts in tms otraitiona onpts in man mirobioogy or xamp shou w istinuih marn from rrra bacria bad upon odum on rquremn or

or that mattr any spii physiooga ritria? Athough th uiationo pcc on rqurmn and h uncon o h ion n bacria grwthis cary imptant rom a physioogia prsptiv ths aaptations may

prov t b imporan, in an coogica conx han h dmonraionthat batria ar mtaboiay ativ in th marin nvironmnt Crtainy intms o natura prout isovry a marn isoat's abiity to prou s-ondary mabo ony whn provdd wh condon pcc o h marn

nvironmnt is a primary onsiration.Anothr gnra onpt in man mirobioogy that an b arss in

gratr tai spiay in ight o avaiab mo thniqus is th numbran ativity o grampositiv bata in th sa h rativ numbrs ogram-pov batria in sm man habitats may b gratr than prvoulybiv. or xamp a rnt stuy monstrat that o 6 ons obtain rom sawatr samps an xamin by sma subunit 6S) rRNqunc anaysis ustr within a group o gampositiv bata 4though this stuy was not intn to b quantitativ it suggsts that grampositiv batra may rprsnt a numriay signiant omponnt o thtota viab as w as th uturab 83) batria ommunit. ar aiona dcrpon grampov batra ar n o bttr cary thirnumbs and atvts n th sa.

o at ony th most priminary invstigations hav bn ma in thsarh or nov mtaboits rom marin batria From an aami prsptiv many ogia qustions unamnta to this rmain unanswr.Th qsts incud t ts o pan and invrbra mabo upon

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SECONDARY METABOLITES FROM MARINE BACERIA 77

h dsibuion of man bacria and h ols of hs mabos in chmcal dfns agans mcobal pahogns Oh mpoan qusons assh ol of symboc baca n poucon of mabols pvously

ascibd o nonbacal soucs Whou coninud cological sudis of ma-n baca hir ponal fo h poducon of novl sconay mabosmay nv b fully alizd

WDT

W han ofssos Raph Mch and Faoo Aam fo h cca au-aion of his manuscp n addon w han Carol wly fo hlpfuldral ommens

Any u eew chapter, as we as any arice cted in an u eew chaper,may be purchasd from he Annual Reviews Preprns and Reprn sevice.

183478007; 4152595017 ma: a@assg

Ltt Ctd

1 AI-Ogily SM, KnighJon EW. 977.Antifouling role of antibioics prodedb marin algae and brozoan Nature265:728-29

2 Anden Wlfe MS Falner DJ1974 utotoxic ntibiotic poducionby a man rmaerm. ar.B 272885

3 Aaway DH Zaboky OR d 993are Bely, Vl. , harma-eal a Bave Narl Prs. Nw Yok Pnm 500 p

4 Austn B 1988 Marine MicrobiologyNw Yok: Cambdg Univ Prss

5 Astn B . 989 Nol prmcuicmnds rm mrne J.Appl. Baceriol. 6746170

8aam RB, Ganh M etas Y no v man mgnim: ntbael ptm Hell Wss eeresr 8891

7 Bllsr M Blster M, Blih P.1977 oltion nd ctezaon of mlelr wet n red by a rEcol 3289303

8 Brin , brri Eg L 987 Enumertn st nd o yioogc propt of cmcfom awatr and dimnt. Sys. Appl.

Mirobiol 108599 Bt Wrigh M, Yaano A

vermn M 989. oltn o negated b hyosti re n adea m. Natur 34257274

10 Bll W Mithl R 972 Chotiand growth repon of mrine btrio agal xtrclar produc BBll 143:26577

1 1 Brthle M MacLo RA. 199 Thro of N' in bn tnpot nresprtion n h rne elea aesa 4 a J rl.3743339

2 etn V. 98. he hems anBl f As, Vl. NwYork li 91 pp.

13 Boinchs Leos ML Baja L. 988Puatin ynas htrorhibtel couii itd wts veslss aI lva ra inn Microb. Ecol. 534557

4 Bona DB, Win RM Colwel RR.86 Mo-nv eaion n pontil fo bionog .r l 200

5 B Ahe G Roh r Myr S 1963 nbon o yt by mrne um. J Baerl. 85132-3

16 Buc My SP, Kp KM 962Mrin btri wi n y titySee 233940

Buess Myas udo H, Mtsunga T 991 Ano produtonb th arin photynhi bteru

Chomatium purpuaum NKPB03104: localition of ciiy o tematopo rl 843016

18 Buod PR ir RM Lz P.

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57 JENEN & FENIAL

966 Pruction of a pyrrole antibioticby a maine bacterum Appl robol14:4953

9 Buhell ME Gfe U ed 989 Boate Metabolte from roorganm

Pge n Ida Mrobolog Sere Vol 7 Amterdamew YokElsevier 49 p

0 Chet Aeth Mitchell R 97eulion of bacteria fm marne urface Appl Mrobol 0 0

het I Mitchell R 976 cologicalapect of micobial chemotactic behaio Annu ob 09

. Chiholm SW Olon R Zettler ERGece R Wateruy ]B Welchmeye NA 1988 A noel ree-ling

hlohyte abundant in he oceaneuhoic one. Nature 40 Clau D Fite D ocur M 99

Genera related to the genu Ballupa, pa Pou Fa, and aopanonn Te Proote dboo o teBolog of Batera opologIsluion, Idntcatin Applicatind A Balow G Tue M DwoinW Hade K Schleife 694 New: Sprngeerag nd ed

. Cope WA 970. Attachment of maine

bacteria to olid urface n Adeonn Bologal yem, RS Manlypp 7-87 New o Academic

. Cundell AM Sleeter D Mitchell R.977. Micrbial populaton aiatedwith he uace f he bown alga A-opyllum nodoum robal Eo/489

6 Didon BS Schumcher RW 99olation and ynthei of caprlactinA and B new aolatam from amaine bactem Tetredro 4994

7 Dai AR agett NM MCnell OJoung M 989. ibioi of marineagae and enhi inveebat natuaroduct chemitry and othe mechanim inhibiting ettlement nd oergrowth. See e 8 pp 8

8 DeLong E. 2 Archaea in coatalmane envinment Pro Natl AadS USA 896889

9 Demain A Solomon NA ed 986.Manual ofIdutral Mrobology adBotenology Wahigton D Am.Soc. Microbiol 466 pp

0 Do ogue mada C Noguchi hwada Smd U. 99 eodtoxin uction of actnomcetela 'rm marn edimnt ppBaerol 068

Do ogue K Simid [990.

Identificaton of dee-eaediment bactea which duce teodotoin Appln bl 611626

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Vbro Baterol 997-7. Ducow W Mtchel R 979. Bac

teial oulation and adatation in hemucu layer on liing coral mnolOeanogr 7

. -Naeeb MA Lechevalie A 96.Selectve olation o aebc actomycete Appl robol 777

Elyao GB uzntoa Mihailo Mal tev I oinov G edoeyevSA. 99. Bminated dipheny ehefrom a marne bacterum acated whhe ponge dea perea 7

6-6 auler DJ 99 Maine natual d

uct. Nat Pro Re 097-937 . auer JD He H Unon M ewey

CA. 99 New metabolite fom maneponge ae ymbiont imoant?zz Ital 07

8. autn DG ed. 988 Bomedal Imrane f Marin Oganims Mm.al Ad S 9

9 enical W 99 Chemical udie ofmarine bactera e Re 9678

0 enical W enen PR 99. Mane

micoganim a new biomedical reoue See Ref . 49-754 letche M Marhall C 98 Are

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uhman JA Mallum Davi AA.99. Novel major archaebacteralgrou fm me planton Nature6489

. uhman JA Mallum K Davi AA.99 hylogeneic diverity o uburfce mie micbi commuiie

fm the Atlantic and Pacifc Ocean.Appl nron Mrobo!' 99

uetani N ima D Matunaga Sahimoto taga et al 987.3-Amino3-deo--glucoe an antbiotic uced by a dee-ea bacteum.peea 4464-6

4 Gauthie 976 Altermona rubrap nov a new marne antbiotcproducing bacteum. t J Syt Baterol264966

46 Gauthie MJ. 976. Mication of bactel repiration by a polyanioic antibioic poduced by a mane Aeomoa Antmrob Agent emoter966

7 Gauthier MJ Breittmayer A 979 Anew ntibiotic-pducing bacterum

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SCNDARY MABLIS FRM MARIN BACRIA 57

f sewe Alterma auratasp nv t yt ate 29:-72

48 Guhe MJ Flu GN. 1976 Anbel v ne vle-ened Alrmnas w spleeene e pdun bned punds a J. bl22161219

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