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Chapter 1 The nature of cells(第2页)

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Microtubules(Mt)—dynamiccytoskeletalpos,whistantlyassembledaiditywithioplasm,andactas‘rails’foriransport.

Mitoi)—arethesiteseionforallcellularauu)—sthe‘blueprint’forallcellactivity,storediheDNA,whiecessitatestaiooplas

Nuvelope(hedoublemembraingnud.Theoutermembrainuouswiththeendoplasmicreticulu

Nuclearpores(NP)—thousandsofelsinthehattroltherapidexgesbetweennud.

Nuo)—siteoftratedRNAandribosomeprodu.

Plasmamembrane(Pm)—alipidbilayerihproteihecell,withspecializedsitesforattaduniwithneighbcells.

Ribosomes(R)—numerous(millionsperaesthatassembleproteins.

Vadvesicles(V)—avarietyofmembraments,whichfulfllspecializedfunparticularcells

speitodria,forexample,produergyforallofthecell’sactivitiesfromthebreakdownoffoodmoleanimalcells.Plantiquelyhavechloroplasts,whivertsunlightandtosugarsasaneoitodria.Bothmitodriaandaythemselveshavebeenfree-livinganismsearlyiion,beforebealyier,plexcell.Everycellhasa‘blueprint’foritsowioheDNAofitsgenes.Inanyparti,theDNAinformatiohesameiype,whethertheyarebrain,gut,orskiatictaintwocopiesofeaAmolecule(theyaresaidtobediploid),exceptthegermlinecells(eggsandsperm)inwhiAexistsasasihesegermlinecellsarehaploid).ermfusetoproducethefirstbryo(thezygote),twocopiesoftheDored.HowtheDofhaplcellsisreducedwillbedisChapter4.

IheDNAiscirdlieshients,butinplantandaisedwithinananelleknownasthenuLatin nucula,‘littleisfoldedintoes.Cellswithermedeukaryotes(aGreekword,meaning‘trueker’),rokaryreek,‘beforenuts’)suchasbacteriahaverelativelylittlespecializatioentsiernalanelles.

Allcellsreproducethemselvesbysplittingintotwo.Setoiheirtsrapidlyenoughtodivisionbyaproasbinaryfissioninaslittleas20mihemuchlargereukaryotecellsmay takethebestpartofadaytodoubletheirsizebeforedivision.Cellsasmaeshaveanunparalleledeffianddiversityofpohebasigblocksofcellareproteinmoledeverycellhastensofthousandsofdifferentproteins,inmillionsofcopies.Aumbersofthevariousmoleaeukaryoticcellareextremelydifficulttoquantify,butestimatesdoexistforbacteria,whichhave40%oftheirvolumemadeupofaroundonemillionmoleculesofsolubleproteins.Avarietyoffivemillionsmallmoleculesatfor3%,thenDNA2%.Thecellmembraerbacterialcell20%,andtheremaihetsaremadeupofthemolecularmaeededtosyeins,ing2000ribosomes.Thesefiguresberoughlyscaledup1000-foldfortheinvolumeofanimalandplantcells,whitainhundredsofindividualanellessuchasmitodria,andaroundtenmillionribosomes.Ribosomesaresmallmolecularmaesthatprovidefortheassemblyofein,acrucialrequiremeeheprovisioeinpriortocelldivision.

Itishardtofindman-mademaerythatmatchesthewsecell.Perhapsthebiggestsuperputerohwouldeclose,butitwouldalsoytophysicallyreproduceitselfwithfasterandfasterprocessors.Whilethismaysouatement,itbeorereasonablewhenyousiderthatcellshavehadaroundfourbilliettheiracttogether,tlydriveingpressuresofion.Ierms,ioifacelladaptstoitse,feedsandreproduitsurvives,butfailuremeahisprocesshasproducedaself-propagatiainingandrepairifunswithalevelofefficyrarelyapproachedbyman-mademaery.Muchoftheeofnanoteology—theengineeringoffunalsystemsatamolecularscale—isaimedatrepliolecularreaswiththeefficylevelsfoundinlivingcells.Aswellastheirhighlyeffitmetabolism(life-sustaiivity),cellsarealsocapableuuralrigidity,asfouhatgivewood,palms,andbamboovaluesofmeiaareexalinas(skystheFarEastoftenuselocalbambooratherthansteelsg).

Tissuesaion

Themaiicofeukaryoticcellsistheirabilitytoaltertheirshape,pos,aabolismtofulfllaparticulartask—todifferentiate—afacilitywhitoetogetherandformmulticellulartissues,tobiissuesintans,andthenformaireanismsuchasahumanbeing.Weareformedfromaround200differenttypesofcellsthefourmaihelia(surfaectivetissue(blood,bone,andcartilage),muservoustissue.Cellsareproducedatwidelydifferentrates,fromsperm(atarateof1000iimeasaheartbeat)toamaysurvivealifetime.Someofourbloodcellssurvivefhthourswhereastheredcellscirculateforabout120days.

Asanexampleofdifferentiation,webrieflysiderthosecellsthatformourbarriertotheehecellsatthesurfaceofourskinhaveundergoneamajor‘remodelliion)tobeeeocytes,whicharefiattenedpolygonalplatesmademaihesameproteinthatmakesnails,hair,ahersinbirds).Eaeodsaboutadayatthesurface,befshedaheohatwepresentanewlayerofskiheworldeverydayofourlives.Theshedcellsarereplacedbydivisionofuhatdiffereheypassupwardsthroughthe24celllayersthatmakeuptheoverallthianskihetoplayerofeocyteseffitlyremovesaccumulateddebris,alongwiththe7.5millionbacteriapersquaretimetreandthevariousfungalgrowthsthattlyattempttoizeouroutersurfaces.Thereare1000eocytespersquaremillimetre,andouroverallskinsurfaceareaisjustuwosquaremetres,leadingtoadailyloss(a)ofaroundtwothousandmillioncellseachday.

Theshedskincellsmakeupabout60%ofhouseholddust,andthoseweloseinbedprovidethedailybreadofthemillionorsodustmitesthatliveinourmattresses.Notallpartsofourbodiesarereplacedatthisrate,buttheskin,an,providesagoodexampleofthebasicpropertiesofcells—repli,divisioioasafunalpartofatissue,andultimatelydeath.

Aresingle-ssimple?

Instarktrasttothesinglepurposeiaskincellaspiresto,weshouldsiderwhatlifeislikefle-celledeukaryaheyaresmall,theycouldbesidered‘simple’and‘primitive’.Tosurvive,however,theyfood,mailees,reprodudavoidbeiheranisms.Protozoaarethelargestgroupofsingled-imalserhapsthemostfamiliar.AmoebaehavebeeopursueandcatchotherprotozoasuchasParamecium—perhapstheunicellularequivalentofaliongazebra.Possiblyeveillareagroupofprotozoaoria.Theseunissudroetesparadoxusdoheirfoodbutiachthemselvestovarioussurfadtheirtentadwaitforsomeuozoan,suchasaParamecloseby.Iftheteouched,the‘prey’isinstantlyparalysed,asofitsbodyaresuthetehebodyofthesu,redugthepreytoashrivelledhuskinamatterofmimethodforthetransferofpreytsisunknown,butitisdriveaculararraysoficstructurescalledmicrotubulesioriaacles(see Figure 6c;microtubuleswillbedesChapter2).Havinghadagoodmeal,suayate’,aprtheattaodifiedteoeachhwhichtheyexuclei(theyhaveone‘maudthreesmallones).Ifbingeingaeeshasanfacility.Deeslivesattachedtothegillplatesof Gammarus,afreshwatershrimp. Gammarusmularly,sothatDeesrisksbeibehiyshell,losiflowofwaterillplatesahisbrings.However,Deesreizestheearlieststagesofmoulting(possiblybyrespondingtomoultinghormone),aamorphosesintstruownasciliawhichallowsitto‘upstidfiofgillplates(ciliawillbedesChapter2).Thus,‘simple’single-shavejustasplexalifestyleasthatfoundinmanymultis.

Althoughprotozoa(andsis)mustihtheireosurvive,theirperislargelylimitedtophysidchemiterasattheirmembrawithsfeatures.Insissuchasthegreenalgae onas,a’isvisiblewithinthechloroplastumicroscope.Theeyespotisaplexsandwibrahrowsofgraaround200differentproteins,ingthesamerhodopsinsfouinaofourowneye.Signalsfromthephotoreceptiveeyespotcausetheflagella(whip-liketailsonthesurfaceofthealgae)tobeatiways,sothealgaeswimthterlight,butawayfromlightthatistht.Whilsttheeyespotmightbethoughtofasarudimehereisnoimaginginvolved,orrequired,astheeyespotsuppliesalltheinformatioheanism’stoight()rhythms,andoptimizephotosyivity.

Tissueculture

Anenormousamountofinformationaboutmammaliancellshas invitrostudies;cellsgrowninaglassorplasticflask,maintainedihat37degreesinanatmosphereof5%dioxide,repligthesinthebodyascloselyaspossible.Plantalsobecultured,oftenwiththeircelledoff.Growthofdtissuesoutsidethebodygoesbackasfarasthelate19thtury,ahodologyfortissueaurewasestablishedbyRossHarrisoninBaltimore,USAaround1910.Tissuessugfromlaboratorymiceembryosusuallygrowwellassingleing‘primarycultures’,buttheyalmostalwayshavealimitedlifespan,dyingoffafter50orsoroundsofcelldivisioareisolatedfromsolidtissuesrequireattattothesurfaceoftheplasticgrowthvesseltodivideandgrow,butbloodcells,whiallyexistinaliquid,doadgroension’culture.Cellsareusuallyculturedasasimixedcellculturessuchasthosegrownfromisolatedbonemarrowwillbothsurviveandmaintainthecellulariionsthatothebody.ieceofsolidtissueisexdputintocultures,thecellsthatgrowthemostreadilyarethosethatrespondtowoundinginthelivinganimal.Theseownasfibroblasts,arethecellsthatmaintaiissuethatformsthestructuralframeworkofthebody,suchasligamentsandtendons.Afterinjury,fibroblastspulltheedgesofwouher,ahethatformsscartissue.Iionstheyarelongandthin,withaleadifansoutatthefrontofthecellasitmrowingsurface(Figure 3c).Withinafewdays,uousroundsofcelldivisiohesurfaceoftheflasktobeeinglycrowded,ahereisnomoreroomfornewlydividedcellstoreattachtsurface,divisiohisissity-depeinhibitionofgrowth,andischaracteristialcells,butnotthosecellsderivedfromtumours.Atthispointthecellsaresub-culturedintonewflasksataredusity.Becauseprimaryculturesdieoffafterabout50divisions,thisputsatimelimitonanyseriesofexperiments,so‘perma’es(usuallyderivedfromtumours)whichdivideielyareoftenpreferredbyresearchers.

&ypesandtheirshape.(a)Haemopoieticell(spherical),(b)epithelial(polygonal)cell,(c)fibroblast,(d)adipocyteorfatcell.Allthesecellshavebeengrowniure.Althoughthesurfaceofthefatcellissmooth,theothercellsaremoretypical,withmembranefolds,ruffles,andfiensions(microvilli)overtheirsurfaces.Thedifferensizebetweedanormalcellisshownbytherouhebottomleft.(e)DeesparadoxusfeedingonaParameciumattachedtoitstentacles

HeLacells,thefirstpermahumane

Althoughpermaesfromhumansaregenerallyderivedfromtumours,tumourbiopsiesarenotoriouslytrickytoestablishihabout1issuccessful.Thefirsttimethathumancellsweregrownincultureasapermaewasin1951,byGeeyinBaltimore,USA.ThesewereHeLacells,socalledbecauseoftheirsource—acervicaltumourbiopsyfromaladyriettaLarietta’swasdiagnosedinJohnsHopkiheveryfewhospitalsintheUSAin1951thatwouldtreatamemberoftheblackpopulationwithouthealthihetime,althoughourbiopsieseretlyputintocultures,nonewerthanafewdays.Hea’scells,hantodoublewithindays,sadlyiheextremelyaggressiveandfastgrowingtumthroughoutherbody,whichkilledherwithinmonths.Thea’scellsveryquicklyspreadaroundtheworldofcellbiology.Iheworldwidedemapermahumane,theTuskegeeInstitutebeganmassprodugthecells,shipping20,000tubesor6trillioncellseveryweek.Thismeansthateveryfewmonths,thesamevolumeofcellsthathadbeeoformHeaherselflefttheprodue.HeLacellswereexperimentedonineveryceivableway,noneofwhichossibleusing‘whole’humans.Theywereexposedtthatmightbetoxiours,everytypeofradiationpossible,aoxinsahemostparttheydidinasignifitlydifferentwaytodifferentspealiaitortanttoshowthathumaurecellsdidnotbehaveinanyuniqueways.Ashercellshavenowbeengroideforoverhalfatury,thereisprobablyenoughofHeheflecells)aroundtheworldforhertopopulateherowntown.Infaehters,afterseeingthefilmJurassicPark,wasbrieflycedthattherewere‘otherinLondon,whereshehadreadthatmuchresearchhadbeeonHeLarietta’swouldprobablyneverhaveknownabouttheirmother’scellsexceptthattheyweretrabyJohnsHopkinsresearcherswhowaoparetheDNAfromHeLacellswithitsanmatch.Thisiedthemtothemultimilliondollarindustrythathadbeeheirmother’stumour,and,notunreasoheirlawyers)triedtoextrathesituation.Uhem,in1951nhtstopatieed,aeofiahassubseque‘aperson’sdiscardedtissuesarenottheirpropertyandercialized’.Evea’sarestilluoaffordhealthiheUSA.Despite60yearsofintensivereseargreAanalysis,theactualreasonsfortheprolifiatureofHeLacellgrowtharestillunknown.HeLaostbesideredthe‘weeds’oftissueculture,havingtakeheresbyatalination,afaetolightwhenculturedcellsbegaerizedbytheirDNAsome2o.

Fordecades,researedonvitrohasbeeioojusthowrepreseis,becauseallyinathree-dimensionaleinamanismatoavarietyofstressesandfaotexperiencedbyasinglelayerofcellsgrolasticdish.Remarkably,inoverhalfaturyoftratedresearch,culturehaveproducedverylittleinthewayofmisleadinginformation,andwithoutthisaccumulatedkisuhetpotentialofstemantherapywouldexistatall.

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