Mitoi)—arethesiteseionforallcellularauu)—sthe‘blueprint’forallcellactivity,storediheDNA,whiecessitatestaiooplasm。
Nuvelope(hedoublemembraingnud。Theoutermembrainuouswiththeendoplasmicreticulum。
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。