ADVISORYE DITORS H.W . BEAMS AUDREYM UGGLETON-HARRIS HOWARDA .B ERN DONALDG .M URPHY DEANB OK ANDREASO KSCHE GARYG .B ORISY MURIELJ .O RD C PIETB ORST VLADIMIRR. P ANTI STANLEYC OHEN W. J.P EACOCK RENEC OUTEAUX DARRYLC .R EANNEY MARIEA .D IBERARDINO LIONEIL. R EBHUN BERNDTE HRNGER JEAN-PAURLE VEL CHARLESJ .F LICKINGER L.E VA NSR OTH NICHOLAGSI LLHAM JOANS MITH-SONNEBORN YUKIOH IRAMOTO WILFREDS TEIN YUKINORHII ROTA RALPHM .S TEINMAN MARK HOGARTH HEWSONS WIFT K.K UROSUMI K.TANAKA ARNOLDM ITTELMAN DENNILS. T AYLOR KEITHE .M OSTOV TADASHUITAKOJI ALEXANDERY UDIN INTERNATIONAL Revieowf C ytology Editor-in-Chief G.H.BOURNE StG.e orgUen'isv erSscihtooyofM l e dicine StG.e ogre'Gsr,e nada WesItn dies Editors K.W .J EON MARTINFRI EDLANDER DepartmoefZn oto logy JulSetse EiyneI nstitute UniverofsT ientyn essee UCLAS chooofMl e diince KnxovilTlenen,e ssee LoAs ngelCeaslo,ifrn ia SUPPLEME1N7T CytolaongdCy e lPlh ysiology FourtEdhi tion Editebdy G.H.BOURNE StG.e orgUen'isv erSscihtooyofM l e dicine StG.e ogre'Gsr,e nada WesItn dies ACADEMIPCR ESSI,N C. HarcoBurratJc oev anoPuvbilcihs,h ers SanD iegoN ewY orkB erkelBeoys ton LondoSny dneTyo kyoT oronto COPYRIGH©T 1 897B Y ACADEMIPCR ESSI,N C. ALLR IGHTRSE SERVED. NOP ARTO FT HIS PUBLICATIBOENR EMPARYO DUCEODR TRANSMITTIENAD N YF ORMO RB YA NYM EANSE,L ECTRONIC ORM ECHANICAILN,C LUDIPNHGO TOOCPYR,E CORDINOGR, ANYI NFORMATISOTNO RAGAEN DR ETRIEVSAYLS TEMW,I THOUT PERMIISOSNI NW RITINFGR OMT HEP UBLISHER. ACADEMIPCR ESS,I NC. SanD iegCoa,l ifo9r2n1i0a1 UnitKeidn gdoEmd itipounb lisbhye d ACADEMIPCR ESISN C.( LONDONL)TD . OvaRlo adL,on doNnW ?DX 24-28 I LIBRAROYF C ONGRESCSA TALOGC ARDN UMBER:5 2-5203 ISBN0 -123-64378-3 PRINTEIDN THUNEI TESDT ATEOSF A MERICA 87 88 89 90 98 7654 321 Foreword Whent hfier setd itoifoC nyt oloagnydC elPlh ysioalpopegayr iend1 492i,t covered fatihrselt yao tfew eklnlo wlecdeglbeil o olfao tgt yh at itnai pmper ox­ imat4e0l0pyr inptaegde As s.e coenddi taipopne airne1 d59 1at 5 2p4a egsT.h e thierddi tairorni ivne1 d69 4w it7h8 p0a geasn ndo w,i n1 98,a7 ftaenar v alanche ofr eseairncc ehlb li ol,ot ghyfoeu rtehd itiiseo vne lna r.g er Thes tuodfym acrolmeocuolfbe iso losgiicganli fiicsapa anrcotefc eblilo lo­ gya ndl eaidnse vittaoib tTl.hy e t er"mm olecubliaorl owgaysfi" r suts ebdy WilliAasmt buirn1y 49 5a ndF reifelidnhe irbs o oskya s",W heintw asf irst decidteobd r eaokp ean l ivicnegla ln ds tudiyti sn nweork rin,gm solecular biolowgaysb orn." Sinmcoel ecubliaorl aongdcy e lbilo ldoegaywl i tche lwlhsai ts" cyto"l?og y In1 790N oviksoaffit dh awth iclyet oliostg hyse t udya ltolyf p esc eolbfly as l l typeosf t echniaqnudte hsae n alyosfc iesla clt iviintt eireomsfs m icroisccoaplly visisbtrluec etsuc,re lbli oloagniasltyscz eela lc tiviintt ieersmos f mcoulle.e s Thiryteya argsoc ytolwoagsmy o roer l esssy nonymwoiutcshe lbli ology. Thed egrteowe h ichh awveae d dedo utkron owleodfcg eel lasc einnt aunrdy ah alifsa mazi.Bn agcikn 1 65R5o beHrootk ew itahp enk nifea,m agnifying gla,sa sndap ieocfec orska wt hatl atthwteae srm aduep moyfr iaodfs minute compartmoercn etlsIl ns1 .76 4L eeuwenhfooeucnkdfr ee-licveinligln sv,i sible tot he nakefldo uiersyheii,nn sg t agnwaantteS rc.h leiadb eontn,ai sitn1, 3 88 andS chwaanz no,o gliositn1, 38 9s tattehdaa tlp ll anatnsda nimwaelrsme a de upo fc elTlhsu.ts h "e cetlhle o"r wyas bSocrnh. leainddeS nc hwabnenl ieved that ccoeullbeld sf ormfreod mm ateriawla snt ohctae tl lubluaVtri ,r chionw 1585c laimienhd i fsa muosa phor"iOsmm nicse lluel acee lltuhlaaatl, l" cells came frootmh ceerl ls. 1381B rowhna dd iscovtehrneeu dcu lseF,l emmidnigsv ceoremdio tsiisn In 1880a,n di n1 980W aldeydeirs covcehrreodm oessoM.me ndeforlm ulattheed lawosfh eredbiatsye d oenx pehriism esnttuadlii n1e s68 5b,u tth lei mitaotfi ons cytolaottg hyte i mper evenstceide nfrtoimsu tnsd erstatnhcdeei lnlgmu elcahra ­ nismisn volMvieeds.c ihseorl antuecdla eciicfrd o mp usi n1 87a1nd n ameidt "Nuciln"e. Hertiwg obsfeerrtlviiezda itni1 o78n5. These twheneri en eteceenntthhu irgyh liingt hhdtees v elopomfoe unkrtn owl­ edgoefc ytolaongdcy e lplh ysiology. Int he etawrelnyt iceetnhtu trhyep r ogsrsoe fo ur knowolfet dhgece e ll gathesrteed.a B myt hbee ginonfi ntcghe en tMuernyd 'esslt udwieersae c cepted when whoirswk a sr ediscobvyeD reeV dr ieasn dh icso llesa.Tg hueec ellular mecahnisfmoM re ned'lsfni dinwgassp roviwdheedSn u ttion1n 0 92s uggested xiii xiv FOREWORD th"ec hromostohmeeoo rfhy e rediitnwy h"i chhei ndictahtatethdg e e netsh at carrtihehede rediftcaatrowyre sr per esiennt thc eh romosoTmheess t.u doife s Morgaann dh icso lleapgruoevsie dxepde rimpernotooafftl h iasn dab eginning wasm adoen t hmea ppionftg h dei stribouftg ieonnea sl ontghl ee ngotfth h e chromosDoumrei.tn hgfe i rhsatlo fft hcee ntcuerlbyli olowgeirsietn st ribgyu ed thnea tuorfge e neasn dt hewraes a genebrealli tehfat th ewye rper otein in natuarlet,h ovuagrhi ous studtiehsaD tN Aim nidgihbctaep t leady air nogl ien genaec tivIin1t 59y3.W atsaonndC ricdke monstrtahtaDetNd A h ada d ouble helisctarlu catnudtr hea ittp roviad meedc hanbiysw mh icghe necso ulbde duplicaantdte rda nsfefrrormeo dn ec eltloa nothNeirr.e nbaenrOdgc howae re ablteo" breatkh"ge e nectoidcae n sdh owheodw c ombinaotfbi aosndeses c id­ edw hicahm inaoc iwdesr teob ei ncorpoirnatstope edc pirfioct eWionrsk.ei rns molecular hgaevsneue btsiecqsu pernotvliuyds we idt ahs tonishing results lead­ intgor ecombiDnNaA(n gte neetnigci neearniadn loglfi) t ssc ienatnidpfo itce n­ timaeld icaaplp lications. Duritnhgi ssa mpee ritohde wreer mea joard vanicneo su rk nowleodfg e cellmuelmabrr asnteasr wtiitnthgh w eo rokfO vertwohnop ropoasl eidpl iady er ast hien terbfeatcweet ehnee n vironamnedtn htce e lGlo;r taenrdG rendienl 1297s howtehdle i pliady meurs bte b imolecularH.a rDvaeanyni,dDe alvlsio,n showetdh aittm ushta vper otein awsistoihcta i lastToeh.de yb uiulptt he concepotfit ohnep aucimoleculoaftr h emne amtburraaenn edl ateelre ctron microsgcaovpaeym orpholobgaistcioats lh meo deNlo.w f iyfy tealrast tehre bioloofgm ye mbrainsse tsi dlolm inabtyDe adn ielcloint'rsi butions. Thes tuodfyt heen zymoefts h cee lplr oceerdaepdid dulryi tnhgte w entieth centuoruyt;s tannadmienisgnt hifise lidn clWuadreb u(rWga rbuyregl'lso w enzymKee)i,l (icny tochcryotmoec,h rooxmied aasnedK) r,e b(st ricarboxylic acicdy clAet)t .hs ea mtei mteh meo rpholobgaisocifais lm porctealnltlu olcair int hcee ltlh,me i tochownedrrbeie ai,en xgp lobryPe adl aadnedb yP ortaenrd otherMso.r phologaincdea nlz ymset udioens e ndoplarsemtiicc ulum, ribosolmyesso,s o(mDeeDs u vep)e,r oxisGoomlegasip, p aratthunesu ,c lear membrannuetr,i toifoc ne llcsh,a ngienss e nescecnhcaen,gi enrs e spotnos e drugasn,dm anmyo rfea tro nou mertooum se ntihoanvb ee edne veloapnidn g havmea dnee cesass aerryis eusca hst hIen ternatRieovnioaeflCw y tol.og y Thed evelopomfoe unkrtn owleodftg hece e lrlap na rawliltethlh dee velop­ menotfn ewt echnolsougcaihsme isc rochecmyitsotcrhye,m tiisstcsruuyle,t ure, differecnetnituarglia fti(ocnefr lalc tionastpieocnt)r osUcVo mpiyc,r oscopy, cytophototmreatrnys,m iasnssdic oann neilnegc trmoinc roscXo-pryaa,yn alysis, electropmhiocrreosbiiesra,rm a diaantdai uotno,r adiography. Toc ovaedre quaitnae nlryye asondaebtlaeia lm otuhonefti nformaotnit ohne celtlh aitsc urreanvtaliyl waobullrede quail reen gstehryio efvs o lumSeisn.c e wea rper oducoinnlogyn ev oluwmeem usotm isto maes peocftc se lbli ology. Weh avgei vtehnoe p porttuoan igtryoo feu xpp etrott asla kb otuhte sipre cialties FOREWORD xv inc elbli olaongdwy e b elitehvaet what ttosh aeyy bwheioa lfvil en tetrote hset heterogegnreoououpfsr eadwehrosw istho b ritnhge mseulpvt eods a tien certaino fta hrfieee alsd . Thev olume begainnas c cwoiuotnfhht o wl iving sayrsoetr egmasn ized starwtiitnvhgi ruasnedts h per okaryFortoemts h.e wreem ovet oe ukaryotes witahd escriopftt hireoe ns uolbttsa iwniettdhh sec anneilnegc tmriocnr oscope, followbeyda serioefcs h aptoenro sr ganeslulcaehss t hmei tochoannddr ia plasttihGdeos l,ga ip parattheuens d,o plarsemtiicc ulum, rliybsoossoommeess,, andt hset rucatnudfr uen ctoifto hnce e lmle mbrane. Mechanifosrcm esml olv emeonrfot r m ovimnagt eriinca oln twaictcthe lalrse discunsesxeTtdh. en uclenuusc,lm eeamrb rannuec,l eoalnuhdse ,t erochromatin ardee awlitt ihnfo urc haptTehrestn.h cee lilt seiltcfsy, c lietr,se pasiern,e s­ cencaen,dd eatahrr ee viewTehdev. o lumiesc oncluwdietadhn a rtiocnl e growfatcht or. Ia mg reaitnldye bttote hdae u thoofrt sh vea rious fchotarhp eteiexrrc se p­ tionpaalt iednucreia nv ge reyx tended gpeesrtifoaortdt i hoveno luamned t o AcadePmriecfo srst hepirro fessiontahlepiiasrtm i,e annctdeh ,eh ierli pnf inally putttihnivgso lutmoeg ether. INTERNATIONREAVLIE W OF CYTOLOGY, SUPPLEMENT1 7 OrigionfC elClo mponentSst,r uctuarned,F unction LAURLAI NHGs u Officofe Cso ntinuSitnuigde sR,i cUen iver,sH iotyust,oT nexa7s7 251 I.I ntroduction Lifea,s w ek nowi ti,sb aseodnc elluunlaiertn sc lobsyes de mipermeable membranTehsep. r incmiapcarlo molceocmuploanroe nfct esla lrste h per oteins andn uclaeciicdO sn.eo ft hneu claeciicbd a s(easd eniinnt eh)fe o romf a de­ nositnrei phosipsth hameta ej osro urocfeb iochemeincearlgA yl.lo ft hese materairacelo so rdinsatrtuecdt uarnadlf luyn ctioinnatsloel lyf -perpetuating, evolvalbilvesi,yn sgt eImnss e.a rcfohrit nhgoe r igionft( hsce)e loln,me u sfitr st askw haptr ebiolporgoicceasclso euslh da vper odubcieodg ecnoimcp ounds whiccho uhladv eev olvteobd e comtehf eri slti visnygs tems. Opar(i1 n9 3w8a)so neof t hfier sstc ienttopi rsotpsot sheai tnt, h aeb seoncfe bioloagcitciavlti htoeir eisg,i lni wfeoa fsv isat epwsiysnet haensades ss emblies ofp arMtosl.e cuelvaorl utipornoacreybs asseoesnd u nivelraswaoslfc hemistry andp hysiacrssee eans s ourcoefis n creascionmgpllbyei xo gemnoilce cules ("biogeinsui scei"dn t hairs titcodl ees crmiabtee roirca olnsd itwihoinclshe ad tot hmea terainadcl osn ditoiflo infesO) p.a railns sotr esstehdne e efodr c el­ lulaforrit thyee m ergeonflc iefeA .l thosuogmhde e taoiftl hsih syp othheasvies requiurpedda taison ugkr n owleedxgpea ntdhsce,e ntprrailn coifpOl peasr in's propohsaavrlee maisnceide ntisfiocuanTldhl.eye ssensttieaiplnts h oer igoifn lievf icah emiecvaoll uatriveoi ns ualaisfz oeldl ows: -elmeomleenc-utlse s macromol-eccuellelsu laorriitgoyif ln -i fe. II.B ioelements Att hmeo sbta sic chemiwcean lo ttelh eaavtlee lll ,e mecnotnaslt itoufe nts protopalrarese ma dily aivnta hiieln aabnliwemo artlaedr ouuns(d s eTea bIl)e. Ther elaatbiuvned aonftc hee eslee meinntssia dnedo utsoifdl ei visnygs tems areh,o wevneoret,q uivaTlheionsbt s.e rvatiotnh wasthu igwlgeeec satensx pect thien aniemnavtier ontmope rnotv aildoleft hnee ceseslaermye inntgarle dients ofl ifei,it se vidtehnactte rtealienm eanrtbese tstueirt teobd e cotmheme a terials ofl itfeh aont heTrhsec. o ncoefp" tfi tneisfuss n"d amenttota hlue n derstanding ofn atusrealle catnideo vno luitnib oino logaiswc ealalls p rebiolsoygsitceamls (Wal1d649,, 1 790). Copyri©g h1t9 8b7y A cademPreiscs ,I nc. All righotfrs e producitnia onnyfo nn reserved. 2 LAURAL INGH SU TABLEI ESSENTBIIAOLE LEMEINNPT RSO TOPLASM ARRANGEIDNO RDEORF A BUNADNCE0 Bioelements Traceel ements Hydrogen Manganese Oxygen Iron Carbon Cobalt Nitrogen Copper Sodium Zinc Calcium Phosphorus Sulfur Poatssium Chlorine Magnesium 0Somoet heerl ememnatysbe essential for speciofrigicas nms. IIIA.b iologiMcoalle culEavro lution Ino rdteoru ndershtoawnc dh emiecvaoll uctoiuolnhd a vtea kepnl aocne prebiolEoagritichati,l sn ecestsoaf riyre sstt abalsic slho,s aeslp yo ssiab le, descriopfte iaornEl ayr tcho nditIinofonrsm.a tfrioomnv ariosuosu rcheass helpiendv estitgoaa rtroiravstae c omposciotnec epStuicoihnn .fo rmaitiso n derifrvoemd btoetrhr easntedix atlr atesroruerscAtelista.hl o tuhgpehi ctubrye is nom eancso mplseotmege,e neprraolp ecratbnie er se liaatbtlryi tbopu rtiemdo r­ diaEla rth. A.C ONDITOINOP NR IMORDEIAARLT H Thea geo ft hpel anEeatr tihse stimaatta ebdo u4t. b5i lliona nydte haer s oldest mkincorwonfo sasrdieal tsae tda bo3ut. b5i llion( Byaeragrhsao nodm Schop1f69,6 E;n geelta l.,1 968R)e.m aionfps o ssilbilffeeo rmosrb iological materoifsa ulcsah n tiqaurigete yn erpaololrpylr ye searnvader dde i fuflittcod ate ori dentpirfeyc isely. Geologsitcuadloi nem si nerdaelp osoiltdeteshr a anb ou3t. b5i llyieoanr s sholwo woexri dasttiaottnhe astn h ofrsoem n ewesrt raFtoaer.x ampsluel,f ides ofi rolne,a adn,dz inhca vbee efno unidno xidation sctoautelhsda vew hich exisotneldiy ft haet mosphceornidci twieornneos n oxid(iHzoilnlga1 n9d6,2 ). Materdiaatllesad t tehrat nh atti mpee risohdo hwi ghsetra otfeo sx idactoimo­n patiwbilteih n crefraeseeo dx ygteenn siinot nh een vironAm ecnotr.r elation ORIGIONF C ELLC OMPONENTS 3 betwegeeno logaincdpa all eontodlaotgiain cdailct ahtametos l ecuolxayrg en appeairnae pdp recaimaobulnaetf stt ehrae p pearoafmn iccer ofosrseislesm bling photosynotrhgeatniiTcsh mussfr.,e eo xygewnh,i cihsam ajcoorm ponoeftn hte Eartaht'mso spthoedraeiy sg, e nerbaellliye tvobe eda p roduocflt i fen,o at prereqfourii stiostr ei gIinfan c.t fr,e eo xygceanbn e d estrutcout nipvreo tected biomolecules. Thec onceopfat n a naeropbriicm iattimvoes pihsefu rret hseurp porbtye d analyosfev so latmialtee rwihailcshh a vbee etnr appedt hbEeya rtchr'uss t presumaattb hlteyi moefE artcho'nss olidatioonfs. u cmAhan atleyrfrsioeamsl s varivooulsc aonuitcg ashsaivncego sn sisstheonwtinln ys igniafimcoaunnottf s freeo xygeTnh.em ajocro mponenotfas l vlo lcagnaisces sof asrt udaireed primawraitlveyar p aonrsd e condcaarribldoyin o xiOdteh.ce orn stistuuceahns t s onimae,t hane, hydrohgyednr osgeuetlnchfi,.ad vebe,e eind entified in amm smaller aamnodiu nvn atrsy pinrgo porfrtoimoo nnsev olcatnoao n other (Fox andD ose1,9 77). Thefo llowisncge nairsri eoc onstwriutcthth ebede satv aildaabtlfraeo m varisoouusr scuecsah sg eochemiassttrroyn,o amnydt, h eorecthiecmailsA ttr y. thtei moeft hEea rtfohr'msa titohnhe,i gdhe groefge e ological atnudr bulence temperhaatddur riev oeffmn u cohf t hlei ghgtaessentso ,t ahbyldyr ogheenl,i um, ando thneorb lgea seAss.t hEea rtbhe gatnoc ooaln ds olyi,dp fockoetfts h e remainvionlga mtaitleer bieaclasme en trappuendd etrhE ea rtchr'ussT th.e se pockeertrsu ppteerdi odiincl aalrlgtyeh lseya mmea nnaesrt hedyot odaMyu.c h oft hveo lcaonuitcg aswsaisrn egt aiwnietdht ihnge r avitaptuiloolfan c aolo ling ands olidiEfayrittnohfog r ma s econdaatrmyo sphTehrece o.m posiotfti hoen secondaanlrdoy n g-laatsmtoisnpgwh aesarm ei xtoufrm ei ldrleyd ucgiansgae ss represbeynv toeldc anic oouftt ogda(asRysu ebse1 y69,4 R;e vel1l69e5,)T .h e composiotfti hoaent mosphhaersre em aiensesde ntsitaalbellxyec ,et phtafr te e oxygaepnp ealraetdae sra by-proodfup chto tosynatchteitviiActn ioetsh.e r schoooftl h ougvhite wtsh per imorEdairatalht mospahser rees embmloirneg clostehlceyo mposiotfmi aotne rgieanlesr foaulnldiy nt huen ive(rOspea rin, 1398M;i llaenrUd r ey1,9 59T)ha.it st ,h per imcaormyp onweonutlshd a vbee en hydroagnedhn e liHuimg.h rleyd ucceodm ponesnutcashs onimae,t hane, amm andh ydrocgyeann wioduel hda vbee epnr esienng tr eaatmeoru ntthsac na rbon monoxiadnedc arbon dioxide. Whethtehrpe r imorEdairatalht 'mso spwhaesri,enfa cth,i gholrmy o derately reduciisan ngi ntereqsuteisntpgie orsn e A.s w ek nown owh,o wevtehri,s problheamts u rneodu tto b ea r athaecra deomniewc i trhe spteotc htoe r igoifn biogemnaitce rials. uEsxipvneagrri imceeodnm tbsi naotfci oonndsi triaonngsi ng fromh ightlomy i ldrleyd uchianvgpe r oduecqeudi vatlyepnaetns da mounotfs aminaoc iadnsdn umerous other organic1 59m6o1,l59 e7cK;ue lrers, (Abelson, 1890)O.fp articulairst hifena tctethr ahetys dtr ogen acc yoamnmiodpner ,e cur­ sotro a minaoc iadnsd p uriniesfos r,m ewdi tehi threerd ucoirno gx idizing 4 LAURAL INGH SU reactmiioxnt u(rCehso ugh1ul9e8y4T,)h .ef acttha avt a rioefbtiyo gemnoilce ­ culceasen a sialryifr soem p rebicootnidci tsiuopnpsot rhtceso ncoefpo tr igoifn livfei aab iologicale vmoolluetciuolna.r B.C OSMCIHCE MISTRY Thacth emiecvaoll uitsai ogne nuenriavle,pr hseanlo meinsfo unr tshuebrs tan­ tiabtyew dh aitsk nowanb ocuot smcihce misEtrlye.m eenvtosl frvoem h ydro­ genu ndears tronocmoincdailt oifto enmsp eraptruerses,au nrdee ,n ergetics. Atomfiiscs iaonnfudss ioanrspe a thwtaofy osr nme we lemeTnhtosse.el ements whicgho t ofo rmo rganciocm pouanrdaesm ontgh meo satb undcaonstm ic chemic(aBlusha ln dP onnamper1u79m1a)T,.a b lIeIl isctosm pouwnhdisc h havbee eind entiinfo iuetdse pra (cCea mer1o79n0,D; o se1,89 4)T.h em ost abundmaonlte cuslpaerca ipepse tarobs ew ateforl,l owbeydc ompoucnodns­ taincianrgb Tohne.p redomionfaw nacteae nrcd arbcoonm pousnudgsg etshtast thper evaulneinvte crhseamli issotr ryg a(nOirc1o ,90 8)a ntdh laitef w,h ereivte r maya riisnte h uen ivewrosuelb,de b asoendt hsea mbei oeleamnebdni tosm ole­ cules. C.M ETEORITES A direscotu rocfei nformatiinso unp poorftt h aeb iolofogrimcaatli on of biogenic miosfol uencdiu nlm eest eor(iOtreo1s,9 7P2o;n namper1u9m7a2,) . Meteorwihtiecsch o ntacirny staclalribnoen acceoomupso uncdasl,lc eadr ­ bonacecohuosn drhiatveyesi ,e ladnei dm presasrirvoaefyo rgamnoilce cules incluadmiinngao c idWsh.e thtehre asmei naoc iadrste r uolfyn onbiological origoirdn e rifrvoemdb iologciocnatla miniasat piroonb lwehmi ccho ncerns researcThheecr rsi.t emroisoutns efiunld istinguaimsihnaiocn igdfr so m abiolosgoiucravclee rsst uhsoo sfbe i oloogriicgiaistln h p er eseanncrdee latively higphr oporotfti hoseno -canlolnepdr oteivnaarcieeootufais me isn aoc iadss comparteotd h onsoer mafolulnydi np roteinsa.r aeTb hoeu1rt08e s pecies of aminaoc ikdnso wtnoo cciunrn atuornel;2y 0o ft heasrece o mmonfoluyn di n natuprraolt eFiunrst.h eromnolroyen ,e( Lfo rmso)f t htew op ossisbtleer ­ eochemiiscoamle forrimcs andL formsi)sfo undi nb iologaicatliplvryeo ­ (o tein(sA.n alogobuisollyo,g iaccatlislvuyeg aarrsep redominoafnt thley o seri)Ae msi.naoc ifodrsm efrdo mn onbiolporgoicceasolsc ecsui rnr acemic mixtuarnedis n clsuidgen ifaimcoaunnottf ns o nproteisntrauccetovuuarsra ila ­ tioTnhsi.fs ca th abse edne monstrbaylt aebdo rastyonrtyha ensfrdeo sm a nalyses onc arbonaccheoonudsr Iitct aebnse .r elicaobnlcyl utdheamdto lecuelvaorl u­ tiocna ann ddo eosc ciunrt haeb seonfcle i visnygs tuenmdsle arb oractoonrdyi ­ tioanssw elalse xtraterrwehsetrrtieha mele ltye oroirtiegsi nate.