Tc.h. Bittethnol.

l99l, 52, 149-156

Microbial HydrocarbonDegradation-Bioremediation
of
Oil Spills

RonaldM. Atlas
Depanmenlof Biology.UniveGityof Louisville.Louisville.Kentucky40292,USA

(Receivedt8 Dcccmber1990:acccptcd29 Jatruaryl99l )

ABSTRACT

Riorcnedietion has bc(on,e a najor nctho.l enplorcd in rcstotdtioh of

fi n a rer'cval syslem, o.
holaopyin8, Ecording, or
xrcly otCh.micattndustry,
st€red wnn rh. Copyrisht
tnal or i cm.l u*. or fo,
. condirionrhar thc copier
n pa8c or e.ch artictefo.
iSht Las Tbc appropriare
ra3raph.ro th. Copyrighr
ode apFa^ in an ai'cle.
nut b€ obr.in.d diErlv
tins, snch a tor eEncal
ne rcw coll4i!. worts.
)*ner for such copli.g.
oil-polh e.lenrironnents tlmt nmkesuseoJnolunl n iuobial biodeqrulativc
uditities. Riorcnediation oJ petoleunt pollutants ovt(ones the fattors
lihtitinltrutesof ntiuobial h\dto&rbon bio.leatulation.Olien thisi^)olrcs
usinsthe en:l],llJtit Mpabilitiesof the ikdigenous htdto.arbondesrutlitlo
nioobial population eul nlodifi inq endrunnental fa&)^. pa i&larlr
rcn.entrutions of noleelar oxrsen.Ji\el fo,ns of nitosen. andphospharc
b dthierc enhancedruter ol hldrocathon hiodegradation.Biodegrada!ion
of oih sludgcsand hioremediationol oil-ontaninated siteshas heen
a.hieftd h.t oxtaen ddrlition-(.9.. ht tilling soilsin lan{arming and hr
uddntahfdro.ten peruxide ot pvtfins o\_tseninn oile.l uquifersalons yith
a.Uilioti ofhitrcgen and phosphorous-contuining fe ili:ers. The sucrcssoI
r?eding oil spils vith niuobial prcparurions is anhiguous. Succes{ul
hiorcn?diatiol of a nnjor narine oil lpill has been uthieued basedupon
odtlitionol mtru,Jcn I pho:rhorust;' tli:u\.
Ke-f Bords: oil bioremediation.hydrocarbon degradation- marine
contamination,soil contamination.groundwatercontamination.

I MICROBIAL DEGRADATION OF PETROLEU]II HYDROCARBONS

lhorlHiU b. 6led b
po$ible di*ninarion of
The spill of mor€ than 20O000 barrclsof crude oil from tbe oil lank€r firotr
Voldezin Pr;nceWilliam Sound,Alaska on 24 March I9E9.as well as smaller
or oama8eto Frsors or spillsin Texas,RhodeIslandand the DelawareBay,haslocusedattenrionon rhe
l.any us or op.rarion oi problemof hydrocarboncootaminationin marine and estuarine€nvironments
Lodaftls.inclusior in dis
and the role of microbialhydrocarbondegradationin thc removalof p€troleum
or valu€of such producr
149

J.(hcn. Tah. aior..hrol.0163-:5759l'So3 50 a l99l SCI. Pnntd i. Orcat Brnai.

pollurrnk.' An .sruddion of hydrocrrhons ht ndrnl popuL(ions .f hrw b€cn$me (pori\ of los noletulnr wci[hr rtunatics beinsrnNkRl6dr'
tr,icroortrnGms.cpwnu do. orIh. timcy me.h!ni\ b) |'.tolcum !n'Jr .\in rhc JlrrlA I oslr 4 , r '.po'(J 'l'i rhc m
nnd othdi hydtuc bonpdluranGrc.liminakn frun rh..nvnonmcnr.r' "hich R 6 1"'"''*.-** u,t.^-u;. .rL b. d'r'ncr rrom 'hu{ ih3' lrr(r rl'nhri'
ofb'odcsrudarionundn onrmrr ruhoBrory$ndiriols htrv. bcn rcpo(cd io bc ti,-",t-' v" +"t a"rJk' 'td drh^ h'rdl'!rn' llnnrr sroq on
f t h i e h r s 2 5 0 0l u r 0 O 0 s b ' d r y ' lRd.5),undurin{iruudndnionrll.kolcuD ,i"",::r' i". -a ,.r. r.T. thu.. 'hc d.sud, ' on / d'fiHcn' rr_- n'
i,,.t,*rt"^ -,r rourh J'ncrcnr n'nurrr'otr' Jl

"'iii"'lil1ffi,
.*. ,ia i"*-g ..'-". -,.t
t FA(-ToRS DIITERI'ININC x,rTEs OF PETROLf,L,M isubsrr dcr hvJ;\.trFoh ^r h'or.sirdniun nredc\ hrrn!h'J htdrtu'hn\'
BroDt:cl{iDAlIoN '""r, ntr",l"* dd s,bnnui; polvn$har romdicr' $ch ii cr
"' *-rrv connnurc !tr inmdsins pDltdiion 'r rhd rsLdutl
thc otca of p.tolcuo hydrelrbon bidl.Andatnr in rh! cnvmnf,,cir !'r "r,-*'rti"*.
i,.r.*l'm .r-.--t,.'*'-E' ths indudrion or ci,lmcs tor poltcv'ric
ddsmincd by rhc populurionsdl ln,irs.no$ hydn)!xrbon ncsndrg micro. r,vd-..r.' dclmdrton dcFndr u|'on n$er dol'cullr scIshr
d.8uan' rhe phlsiologicrl flllbilirics ol tho\c uopuldnrns. rnd rrndu\ "i,,rl.
,;,-.,'- *." * " ,l'c rind nc '-' 'ne '_4ne' r'r Jes ihnr i
,hioric frcroG rhrr innucnli srD*rh 'ar6 or rhd* hydntid.n.,lcsmdinr i.".i -.. a""..\'\-i","
; '-" n.a'G' hin\ l' r n' i m'rurqi br rr\"ub{' rc i{ir
" 'ifrtnr ,$al'ncr 'r rhes lomnod'r' I
'nrrob!'l populLtri.n\' 1hcp$li.nccof ndiolcum polluratrNdq')lndr onrt's ,, ..-'.,1', "i'.'., i . r -rt .
qudriry d qullirl oI rhL hydrocrrbonmnlurc unJ .n rh. prop.n'cs .f rh. N'cr;bi!r rrrr!r ^lihoush dts. " ttl ri sl'r 'rrcs, rl$c rNpoudds arc
hrdRHrbon\ dn 0.rxr clmrurll\ cdnFicd 6 clrlnJn diude ' srrs ht nncr)or$tr'sn\
Ftolcln,
itrdsllnir.ly.*hsca5 und$ rnothcr \d olcdfldtrionr rlrc$mc hldocrrbon5 !rn hrr! hc.n '{'lrkd rhir dcst lr +nn! Findd'rr lt'mrnc
MtcDbdiN
| . t , o n . nd c h h i . . J + ' - d . dE r h i n . " c L .r , r l r l c q \ ' a n . . , 1 . D . .
P.i.. crposur of a microbialco muniry ro hydrocr.brtrs's 'mpdnr n "t '.i 1. t'-'.'*. t-' '-.. '. ' ' ' . ' "- .r .. . ' t ' . r ' - " a F r r ' ' n . l 1 ' i r F P ^ H ' d ' ' 0 n 'c!' r' '
Jdflm'mne ho* Bpidlr {'h*qucnr hrdi..rrbon lrpurs lri tt htudqrudd.' n;m $il' incndcr snnr.f,rpl.r *JrE
Cosmunirics rrposed ri hydroatrbonsh$om xltrptd. lrl,ibirins kLcdi!. Ikntu d dl r r fLNndrhtrr(hc mirtub'rl dllrl'dlion ofu\phdtnd rnd c$i!'
r6ulrin! 0 insrlul pn'p.dnn\ ol qhirh hrn D'crbunv b({ osld$ld .clrlivd} rc(lcirrrtrr h hkrdcurlddion'
hydn,.rrbon-dcsrdins d.ir: -.,' *,r""f . --"i."-" Romrn' rr /rrrllr n'trnd clidend ftncoordr(nrn
crhboL'( se.cs. h' unpdll!td .trr n)inrtrr\ hrdn,crrboi 'irllrdss $n.'rLl 'i.{hr.rln+ rr{ nL mmf"nnd\ in n'ncJ
:ni"r_.r, t''h tr
coninut bss rhln l% of thc .ricrobirl co mutrily, $hcrs in oil I'ollurcd un{'11'fl$rn-" +JrJn$ll
dosyn.ns hydNa.botr dcsnd.^ ofro .ct'r* I l0'- dt trrictuDryJnilN' "..t',.r.rr','.
sine rp(ed frtrobirl q)mnu',irics hrrc h'shtr phpoiril,ns of hrni\xrrbdn r.-.*.'i",no',r'"'r*.t.i{obirlhvJrutrftof, dcsHJinsrdtrr'c!hv
dc8mdes. th.y $n refond b rhc tNdnlc ol lrtdnurhdtr nolltrrrtrr\ $irh I lr rn;1on thc phlticrlnrtrrc rntl dEmictl (nrposi )i 'lrh' oLllprdL!urrn)
houc.Inrhecrf orrhc/lR.4, :rnd ?rn&sp'll\rlon!ihcrunof ariurtry. r. .,*,". -"i i".t,lt" nn nncn'hi c.knriarnD lnd rh( h!-drturbitr\
Fhtrc.. f(r eiumplo.rhc idaptd htdtu($oniulri'ns p.puhdN 'n*.tr*d '"'"1"i"" - 1", -i{obi.l mdibohc rulclrrnd the rati of
by $!!rrl ord$ olnusnirud. rnd biodqr rion o(ui.rl N fx{ or tu'{c. rhri lNdrod;onndlb;lism "r"' "*t".i hv mico.rsrnisms r! ^i1o* r.nrpemrtrcslhc r\cosrv
clapo hdn in rhc dlyl l lo{in* ihcsn'lll 'i rh. .il 'ncru$. lrt voLiril',Iion or ro$ rhon thr in L'lkrrurtr rcdu!d rnd
ol
Thc phr:icul nrk oI ll.kolcun hldrccrrhon! hs J mrrtcd crsr on rlw rho' r'!t* $lub'l t h ''{u$d. Jdrrns rhc on$r dr h'od'lhdrii'n Rrts
r*' *t'rdci'LI(qr1 f'r-!nrr1npc"!n 'htr
b'odc$ldrlor Ar r*y lo{ nrclnrd.trs hr_dtulrrhun\iru *irhh ,n *d$. . 4,,'--".*'.-r, !'
bur mon oil spiliincidcn( rcler$ I'ctolun, hldnurbDn! in.onentrion\tuf ."-r'*.r *.,'*.,.i'-.'.r'i J('."'rI'r" r c n / ! t r ' r t r' ' ' r r r " '
in.rcc* oIrh. $lubil,ry limirs lh. dcBrcuotstrcrd'ns ddsninA r rJn rhc d . . . . f l d r t" l F h ( k n E ' r r u ' n u ( r r r h cr . i " o f h t J n J h n t r ' e r ' ' h { r mh
strfrc! rrer of oil rvai[hh for mi!rubid coLonizxrnrb) h]&ocrrbon-tlc8rador :;*'.". ;jr.-,,, .. 'h' '.-s
mic'odlg!nisns:'ir,tu!r'csj{dn\.1h.oiltr.rh!ll!ltr ls.I$miig!lhinslicr ''.." '--,".*.1'' t *..rc. crl'( n'oph'lR,lr'n:.i'l/nrl"i' 1
"^-.
M'cmbid Jqradarion dfoil hrs b.rn nx)\n to drru. by rri3cr .n ,lipnrk ';cludinsobnsahhydr&atbnnutlvcis $ft dfstuch hatnr! tuen k'lrrcd lr''n
r lishr iNnlri. fh.lidir ofrhr oil. *ih high.mdle$hr{crAhr rrrm!rics. r' N. (ui {{ rhum rcnr rcsionsrr ''
(
rnd r\pMlt.nA consid. 'l lo b. rc$lcitlot .r crhih tns orl! vsr low sr$ rL i1. _ 'r'.'cr. n 'h.-o'.rsi 'JJ'" u h) lo''6 n' b- f"e'L J''i ni'
hiod{bdarion, ilrhdtrgh q'me iud'6 hnr rqxtrrd rhff rmoerl ' ccn.nllr.
n hr: bc.n considcrcdrhrr ntuisfir chrin rlrrne! nru crsxr r' d.sridr r rhcrc 'iq"i'"J' e-.r',. -*ri,.* -r. ihnelot nesrl., rrr out'orm'c'ob'!l
"
o r d t o n . f h y d o k r b o n s i n r h c e r ! n o n m c dr h e i ! r i t r b i r y r r . r ' l c r r i s o r t r .
rcmd lot oily *rn.s.rrr' fftro hrlc hccnsonc npo \ oi nobilwaion ol dil
\dnio$. und iquitc K oltcn timiri'rFJnLldspen,lsi .n r ryp..f$t!n,l
inrortu$ilcoLumd. bur ir mo$ !ascsthn. hash-.n lirllc cridene ld {sn'lilrnr
dosnwrd le!ch,.s ofoil. wlng rnd Brnhr:' rentlv nudi.d rhe cfet or
by,nicroorunism\..cuB,. rrhenn$orrnNrobt!hyJnf!6,)nbioJcstudxron, hbEm.d ion on rc\idtr.sdlL'cl spill\ 'n $ill::rcmrcm
r ofidr lu'l hrrtin8
hd{cr(.tr'c !sl' L()f,rnrl irr r.dLr)sicrlrirn'ticinccnlpcrr\ rj hc ni,rrL! dlcsl Dil, resp(d rclJ) l'cnnrou rnd l(riciry of rhc i ndcr*d Ln
oi1r
s.n'al i'lcnisro^ hrlc rcponqt rhrr cnlrntrrhonr.r Irihbtc nirrolco rh. olld orjct fucl<hcahs oil<disl o'l BnncmcdHnonkcrrmcnr sonsrv
x(r li]cphrls in hnririn! b m,cftnrrt lrJ(llodfhoi iry dincrr{Jn'crcbirldivivrscosnlEd
Jc$.rr.dlurlNBnrenc!rnnb
d!!.rddi.n.rr sd'nc invsr-srtrn hos!!cr. hi!c rc&lu nr opr.$t n) conrminrrcd brr trnrRxtd vr1. Btrtrcmcdiitun tc
. i ^ n ^ ' ! n . , c . r h . r t r r ' u r . i . ' n J p h . n p t , , { n r . r , c n nhr. n i , n ! ! n \ r $ , i c , ! t h c soill{onkhn$rd soil\ in ,1 6 *tcks t' r &ER thrr .an \uppon planr .ove'
J'rl*.nt. - T.,,lh . n."J.^.c.' ,.ntt.,Ft,J.. ',' Fr F.,r.t,, t'-...di ion I rcarn$r ltmoi complctlr.liminrtul
w-""r,,r " '""*tt,t'"
rdrrnrn ot lrydtulrrr)n\ *ihji ntr o rtL.t rr P A l l . \ i ' , l l * $ i \ w i r h o u l b k n u m c d i r L nr)l n
5. r 2 5 " / ' . l r h c l r i e h { * c i ! h r P A H
rhc biod+r arnr of n)tubtc hldrft,trhm\. whco cotrldcntrr n trbts {rs n'll prsmr rr ll *.cls ?\ftcr{brrorial niri muirt.nicirv rnd rricnv
rnd rh.\ntNrt'\ rru pR$nhjr nor h ni[! j,ct r|[ rhc ()nrrminxi. so,l rIlnnDchrd rhc hrrksround levcl otnnconrrdn!tdl \oil
!'LubilLry.f rhc hylounrN n nr Lliw r\ trr f,cr trJr $rrht,rhtrriL if m L'ftsr!l sclt of bitY.n.dirri,)i Dctr li.r(ion s.tconrplclc ii 20 weels Song
Inrdis'rh cd'iJcrins rhc rrr. rr h,s t.|c| d.r:'brncnc 'Icd ltrtxEror! \onl\or[rnn'nL.jcr rud huriry oil. dic\uldil.
rn'l btrnr.r( o,ldn solr\ condodin!(hrr sirh hnftm.di bn rtrrnDnr n should
hc p.\$hlc h rcduc. lrt rocrrhonsh insisniricrnrh!cLs n (rrrminlki soilt

r BtoRtrMl]rn4T()\Or.O . ( O\'rr]\trNt rr:Dsot L ,|\D I aloRlMEDrAno\ or lrlanlNt oll, sPlll,s

CROI NDB^ I'[R
'1n"d h b'Ff.J-r"rr\"nt' n e d i : In ' r d
Rc$fn'rnr rhrr hrer ol r'!'lnutrh.n hk .Br.,Jir6n .ncn {( rntrcu nr ,hr l,rxN..rnhir'on rlrnFllrc'Fdr ral'n l'!\ " ll'. 1 ,lrfflorl'r"l-r
cl$nur or thc o'l rpill{d fom rhc 6\\d rrritu: {s Pht!.!l wsh'n8 of o 'd
rr rd.rr nh urdr riLtrr \hordi,$ !trh lt'ah trcsurc sdcr wrscxprnslrcrnd clcrned\hidines hccrmc
lrunrd rhc u\c df nnrnulrnfl\ rc.dt(l nrcn! .cclclnir3 B'orcn,cdidbn.thercinc.lrr .onnds.d rs i mcthod
bkrumcd'rril)ncDrLci$ h)!tn'crrh.d\ r) r r\r |h ,$ $ru.,nd .rrhon ro rusmcnr.rhc clcrnup prKdurcs Ths USlil'A trntl truon.nr.'ed mro rn
!ctrurl rfnri'rlr.\ ri hor!nrsti .l rsrs rnt ldi'ily to !rtldc (hc fcasibllir'.r u$ns bi(ncm.dirrnr Ihr roj'cr
mdd'nflri(n,.\uch i\ rhnD!h Lir'lr/cinrrl..ik{ rtrd ns i(tr, rid n{drotr.l rNu\ld on dctminins {hdha nut'cnr ruEmrnrlr'ctr nruld ninulrt s of
rdrl.d hrtho(rhotr desh $ h) \c.dnr! h 't!!.idr on ThrB ryFoInurnsnr sut nrnkrn,n *ca !\'rdcrsl: $!r.r
In so'lttrrd si{Nrlwrl$ i\u$ 'r. rsDtri rns riJ rhe tr\r orrLLLins. r,ftut \olLLhhl2r:l N P sitlNi icnilt.r ntmtr!rtlon) sl!w icl.r!e lnobtrrvlcn*liurcr
rc'r'on_ rM hldn)sd' p$o\tdc 1r.[n.'1 !rn hL tr\cf"] r, r\w,nrina thc riLr or.trcn!\dlrr.d N1 l')r rnd dlcdrhilic (lnip.l liAP 12=ohic !d'1. urci LrurvL
rimidrn .ru*d hj to\' 'rn.Nhr o\lsur ir.,it.,b,tir:r r! thc !$ of ttrlit r6tr{ 'n hborrro.Y nnttrhrn,nsatui 'n lietd
rhosnhler. lirch fcdrli/s *
iinnlcngrri on tjr\ hcsr u\cd !r !trfLt, ! rhN to \hd* rhc srlicNY
htridcsndlri(,n of !r!nN Mon grtrndtrJcl \,!!tic\ con\idcrrrns rar rin, sivenn) p.rc!r'!iad!(* dolL,ticd dlsri prii!ulrrl,
conrrinr nuoninl nDpuhrtrD(nrhtr.roit brxlcs,rnJnnr I t,)\c\ar. unrdrkn\ $trdphicJion dre n) als,'l blin)msrnd kricry b n\h and ,nrcn brurc' Thc
| m s n i r r r r. ' n d m c l n h o L \ norf . i l r { 1 l l ) I r ! L L m
lirs Rn)nntrl r/.rr frNfd rppliunion of th. olcoth'l'c fciiliz{ pt)'luccd rsydtnm ic ault. {'mtrlarinr
r,{.cd rftrn,n inJ trinr!{ rnJ rho\nr$,u\ Mdi otr dduL,t\ sniricrN r rcdu( bi.'icsirdlri,lr {!ch rhrt rhuturri.es .f rhc oil'hlr.kcncd 'ootl rlr{ \lrrulitrc
ql er$rn'. fr.tr' r conrrnro.trrdsn)undrrr.r rurndi rhirc rnd r.( !5*nhlly o .ftc *nhin lodlyt rficr rrudmc.i rhc ur
rcsc otr Ihq d .n ,niumiutfrrc.r!)i.sodnn th.\thnr.xnrtd nrJnm uf lnlDol rid r n.w rkasc frnilt* *r\ rfpnNqi ft' shorchrurie mtn( !
pho\phrrr n' crnni'r{onrrNturnl sturundslcr l r u\ rt{, pumpcd o r|tr
{as u\ud r\ tr mrjor lln orrh! cleriup ctroi ^ jo | .n trsEPA sL e oI
lnrnd*rtr $ih.' rallconrnn$or Iha rcdnrN incrc,N,l rh. trumE(rd ihl|(r monirorin!cllon rollo*cdrhuclrcdiEnc\r ot rl'. bDrcmd'n'on rflimcn(
hydtucnrhin urililis micn!4rinds. rnd I $r\ c{inrxrd rhnr r {)hrr.cLs.t lhich trrs cimrrcd tu incRr\c rhc rrrc5.I bk esr rrion ri lcri rhrccJdld
!r{rtr,. rcrc rcmovd h\ ( nuLrrc d+ tdir)n owlrs 10ilr dlcclvor s bfiLmcdLrrnr bsr g LhemrjLn tu{mcn( mdhod nt
kd rt'. fcrsih'trr d{ u\i,rr trndrlmin! rrtr rcn.! 'nB dn n. uknr fr.n rh. imFcr.d lh.rtliG of l''nr willir sodnd
 Ano(h.rapprouchlor rcdlne oilsfiUs lhd htr! b.m coftid.rcd bur n.rr ticld dlt 1l.d iqdrlfl\ :lcng wrh rJ
'"'r r'ob!l
rcscd inrolvd *cdlng *i(h s.ndically .nsinccrudni.roo{aiismr. Thc poknrial ',.i0 *n ."r t r" sRu-PrrhIs|lr nrcuBfon
" ". r.". v"n" &/u spill huLrhc $1"'s or rhn trcrthcnr n
Ior cr.arns,lh.ough gcndic mrnipul.{ior, nicrobial nrains rblc b dqrud. r l',, l'"a . ,"*,r"
vrridy oI dircrcnr ryp.s ol hydrearbols hus b..n d.nonsrat d. Th. s.ncri.
innrmation ror ar L!$ sohc cnzync inlolvcd in alkaD. d simplc roha!. .,r,r"i"" . * a-nt' t'. t -s." dnd Dhu'nhv-u' r'rdfl ru uqrrJc oil n
idnlci.d t'l rhc r939Ah\krD dil tpill l,oft m'!irnon h!rh{om'
hrdrmrbon rGnsfom,rion @uF on DhsnidJrrr and rn u*ofnrminwnh rd|.y.horclms
rhrr maks
cnl'Ercd d.smids a5an inGulun dunnsRdin!*ould pcltrdc rhc phbl.hs ,, '.i". -.r',*l ..int.y"a . -""raiion or.ilrrnrurcd cn'nonmnt
rsEred sirh snpchon bd*ei nrais in ! nired culrur. Hoy.vcr. rh.n n{ ot nrrunl m'($iil btudctrdan$ t! rcs
rx .onsidcr!blc.ontokiiy :u(oundin! rhc (l.r* of luch scn lcdly.nsin..rcd
m'c'oo'sannml ,nro rh. cn!i'onmcnr rnd h(l.l k{,rs ol rhcv .'-!rntsm\ 1u.
rhcrcrdo bc dclrycd u.ril rh. is!trs of s {ry, ooililnmenr, lid p.ktrliol ror Rtrff,RllN(lis
cc.losiclldamryc lrc rcsolrqdI A hydmcnr6ondesrrdiDgpsuaomonnd las
cn8inc.rcdbyChlkrlhriy rrd sasrh. orulnirm rhlrrh. Sup€s.(l)un orrhc
r)n{.d shks. in ! lindha.l dcnion. rulcd could b. par?nr.d rhc .r3rnnm : r i " , p. n , i , i " ' . v"..
.ngin cred by chrrnhrd! i\ drrb|t old.!hdrn! I numtlr.f lor nrclddr'
*.th r aroDric hydt Nn.bons. bu' dr\ no' dcend. ' hc h ishc' mokc ur.tr *cignr . i.,i.:.r *.r-,
r.rsisrcnrFlrnudcrr riondis and rhus h)\ odr hocncdnsidcrod tir ui in rh.
hioRm.dirloo of orl spilk. Gird rh. cu(cnr rcsularoryfrrmc*ork lor rh. ,,'. h,
-.'i,i*',
dslibcrare(lcrs ofscnc(ica[ycngrn.i.d nr(dorsinfm\. n f untitcLy(hr( my ,;1,;; R ^^,".
suchors$*n *ouLdgrin rh. d.lcsiry r.g!h(ryupprorrlin rimc h hc ornuch
us?in rrearin! rn oll sDill "i r* .i i. .i. t g' 1. " .' 'r'
sedinp *nb non8.ndicdly cnsinercd hdcri, wls u$d fororshorc oilxpill.d
".
'rs
Biocmed' on df (hc ,rr.4u 4,., \pdl .lt ihc
coN conrntd ofrftlyinra sdctrlrurc $nhn fcrcr caralynprodu(i hl Alphl .iii.l,rnur".r.r.
co4orarionro rh.o'lai \.a r' Th. mriond. ln rhn rrlllrd.h rd redrn! n rhi,
ind4.noN D,.robr.l to puhrionsmry.or tE (prbl. old?sftdin! rhc rids 6n!?
- i,,-ll, n r"r"ir 'ion({{ r( rhsftqsr
ofpor.nrid subnnks prckm r suchc.mptci mnrurc\ rs p.trdhum H.wNc.,
th. premisa rhrr rh. nicroorsrnishr nlunlly pre$nl ln rn rnsn,nmcnr
l,liir;r m"".nt""" ^ -',t" .t
M,.; i br|+'D,'q'\
:ubrlcted(oconhindion*irh oil would hclnoiNbbor.ratrsi!0ly,i08nd0,s ' r-...nr
pcrrolcun aid lhrr !dd!d microo4rnismr would bc ablc 10 do u sup(rortob
Ded ro b..ramircd ar.fully clrins w{. nrdc rhlr rhc \cedinstclrm.nr oI
th. ,n.r, &r, snill *ri sucas{ul !r .ompl.rcly rnoling rhd oil. bu( rhr
crcdivcncs ofrhl Alph, \adine ro nimuldc htultcFad jon h$ nor brcn Kriri.d. J;,.r,,..,.,, .i ,,, ."".*i".' ..'
n.rh.srh..ff6tiv.n6solrhcculrurbEn$.rnmsd bv rhc USnPA in hbthra^ r-{ | \"i- R
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.n..*ltr'n*":',lt't..*
l:nlnonm.nral rd$ ol nydrnarbor dclrrdlrion rr. hmir.d by rhc .o7]m!r.
dpabiliti6 or rhc Indi*cnoushyduarhon-tlclrdin! microbi p.pul,rions rnd rqr F r! 1
bt cnu.onmsnrrl rr.tns. prdicuhny co.!r.rhrionr of molsul.. ornrn. tued ij.,,+j..i,..J r ii u^ x--,l-n'N.hdr " N- \-'
,' r",, j i r.rtr- -.r 'h-
t m{ oI nnog . d phosphr. B'orcmcdi tu. or oir norrtrhnr inr)lr$ \d;'. d!/
orcrcomins rhc rNc lrsnh! f!d.B Btr{.gr.drlion or .ilt rludrcr rnd n s",,r.r, o rl w*".c o a
bioRmcdirrbn ol oil-co.tamimlcd :ir$ h$ bcen rchiLwd hl ory3cn *.i.-* lr"..r,.' ' t.c",n
trddnon cs.. by lilling $il\ in lindr!mir13 rnd by ddlns htdogcn nc'ord. !rnl'ri 5 5 l r e N e i : 3 161
Ureas. Immobilircd on Chitosrn Membranc. Inactiration
by Hesry Mclrl lons

Birhrrr Krtucwskn

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1' x srs ptu$lr i rlDr rh. inh'hitr,n or!ruric
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