Cytology and Cell Physiology. International Review of Cytology: Supplement 17

858 Pages·1987·28.423 MB·English

by  Geoffrey H Bourne

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
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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.