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首页 权力、性、自杀:线粒体与生命的意义

权力、性、自杀:线粒体与生命的意义.pdf

权力、性、自杀:线粒体与生命的意义

九月虺
2009-06-22 0人阅读 举报 0 0 暂无简介

简介:本文档为《权力、性、自杀:线粒体与生命的意义pdf》,可适用于人文社科领域

Power,Sex,SuicideMitochondriaandtheMeaningofLifeThispageintentionallyleftblankPower,Sex,SuicideMitochondriaandtheMeaningofLifeNICKLANEGreatClarendonStreet,OxfordoxdpOxfordUniversityPressisadepartmentoftheUniversityofOxfordItfurtherstheUniversity’sobjectiveofexcellenceinresearch,scholarship,andeducationbypublishingworldwideinOxfordNewYorkAucklandCapeTownDaresSalaamHongKongKarachiKualaLumpurMadridMelbourneMexicoCityNairobiNewDelhiShanghaiTaipeiTorontoWithofficesinArgentinaAustriaBrazilChileCzechRepublicFranceGreeceGuatemalaHungaryItalyJapanPolandPortugalSingaporeSouthKoreaSwitzerlandThailandTurkeyUkraineVietnamOxfordisaregisteredtrademarkofOxfordUniversityPressintheUKandincertainothercountriesPublishedintheUnitedStatesbyOxfordUniversityPressInc,NewYork©NickLaneThemoralrightsoftheauthorhavebeenassertedDatabaserightOxfordUniversityPress(maker)FirstpublishedAllrightsreservedNopartofthispublicationmaybereproduced,storedinaretrievalsystem,ortransmitted,inanyformorbyanymeans,withoutthepriorpermissioninwritingofOxfordUniversityPress,orasexpresslypermittedbylaw,orundertermsagreedwiththeappropriatereprographicsrightsorganizationEnquiriesconcerningreproductionoutsidethescopeoftheaboveshouldbesenttotheRightsDepartment,OxfordUniversityPress,attheaddressaboveYoumustnotcirculatethisbookinanyotherbindingorcoverandyoumustimposethesameconditiononanyacquirerBritishLibraryCataloguinginPublicationDataDataavailableLibraryofCongressCataloginginPublicationDataDataavailableTypesetbyFootnoteGraphicsLimitedPrintedinGreatBritainonacidfreepaperbyClaysLtd,StIvesplcISBN–––––––ForAnaAndforEnekoBorn,appropriatelyenough,inPartThispageintentionallyleftblankContentsListofIllustrationsixAcknowledgementsxiIntroductionMitochondria:ClandestineRulersoftheWorldPartHopefulMonster:TheOriginoftheEukaryoticCellTheDeepestEvolutionaryChasmQuestforaProgenitorTheHydrogenHypothesisPartTheVitalForce:ProtonPowerandtheOriginofLifeTheMeaningofRespirationProtonPowerTheOriginofLifePartInsiderDeal:TheFoundationsofComplexityWhyBacteriaareSimpleWhyMitochondriaMakeComplexityPossiblePartPowerLaws:SizeandtheRampofAscendingComplexityThePowerLawsofBiologyTheWarmBloodedRevolutionPartMurderorSuicide:TheTroubledBirthoftheIndividualConflictintheBodyFoundationsoftheIndividualPartBattleoftheSexes:HumanPreHistoryandtheNatureofGenderTheAsymmetryofSexWhatHumanPrehistorySaysAbouttheSexesWhyThereAreTwoSexesPartClockofLife:WhyMitochondriaKillusintheEndTheMitochondrialTheoryofAgeingDemiseoftheSelfCorrectingMachineACureforOldAgeEpilogueGlossaryFurtherReadingIndexviiiContentsListofIllustrationsSchematicstructureofamitochondrion,showingcristaeandmembranesSchematicillustrationsofabacterialcellcomparedwithaeukaryoticcellHydrogenosomesinteractingwithmethanogensCourtesyofProfessorBlandFinlay,FRS,CentreforEcologyandHydrology,WinfrithTechnologyCentre,DorsetSchematicshowingthestepsofthehydrogenhypothesisAdaptedfromMartinetalAnoverviewofendosymbioticmodelsfortheoriginsofeukaryotes,theirATPproducingorganelles(mitochondriaandhydrogenosomes)andtheirheterotrophiclifestyle,BiologicalChemistry:–Therespiratorychain,showingcomplexesThe‘elementaryparticlesoflife’ATPaseinthemitochondrialmembraneFromGogol,EP,Aggeler,R,Sagerman,MCapaldi,RA,‘CryoelectronmicroscopyofEscherichiacoliFadenosinetriphosphatasedecoratedwithmonoclonalantibodiestoindividualsubunitsofthecomplex’Biochemistry,(),–©()AmericanChemicalSociety,reprintedwithpermissionTherespiratorychain,showingthepumpingofprotonsPrimordialcellswithironsulphurmembranesFromMartin,W,andRussell,MJ,‘Ontheoriginsofcells’,PhilosophicalTransactionsoftheRoyalSocietyB(),–Merezhkovskii’sinvertedtreeoflife,showingfusionofbranchesFromMereschkowsky,C,‘TheoriederzweiPlasmaartenalsGrundlagederSymbiogenesis,einerneuenLehrevonderEntstehungderOrganismen’BiolCentralbl(),–,–,–,–InternalmembranesofNitrosomonas,givingita‘eukaryotic’look©YuichiSuwaTherespiratorychain,showingthecodingofsubunitsGraphshowingthescalingofrestingmetabolicrateversusbodymassFromMackenzie,DScience:,withpermissionMitochondrialnetworkwithinacellFromGriparic,LvanderBliek,AM,‘Themanyshapesofmitochondrialmembranes’Traffic(),–©MunksgaardBlackwellPublishingGraphshowinglifespanagainstbodyweightinbirdsandmammalsFromPerezCampoetal,‘Therateoffreeradicalproductionasadeterminant’,JournalofComparativePhysiologyB(),–BykindpermissionofSpringerScienceandBusinessMediaChapterheadingillustrations©InaSchuppeKoistenenThepublishersapologizeforanyerrorsoromissionsintheabovelistIfcontactedtheywillbepleasedtorectifytheseattheearliestopportunityxListofIllustrationsAcknowledgementsWritingabooksometimesfeelslikealonelyjourneyintotheinfinite,butthatisnotforlackofsupport,atleastnotinmycaseIamprivilegedtohavereceivedthehelpofnumerouspeople,fromacademicspecialists,whomIcontactedoutofthebluebyemail,tofriendsandfamily,whoreadchapters,orindeedthewholebook,orhelpedsustainsanityatcriticalmomentsAnumberofspecialistshavereadvariouschaptersofthebookandprovideddetailedcommentsandsuggestedrevisionsThreeinparticularhavereadlargepartsofthemanuscript,andtheirenthusiasticresponseshavekeptmegoingthroughthemoredifficulttimesBillMartin,ProfessorofBotanyattheHeinrichHeineUniversityinDüsseldorf,hashadsomeextraordinaryinsightsintoevolutionthatarematchedonlybyhisaboundingenthusiasmTalkingwithBillisthescientificequivalentofbeinghitbyabusIcanonlyhopethatIhavedonehisideassomejusticeFrankHarold,emeritusProfessorofMicrobiologyatColoradoStateUniversity,isaveteranoftheOxPhoswarsHewasoneofthefirsttograspthefullmeaningandimplicationsofPeterMitchell’schemiosmotichypothesis,andhisownexperimentaland(beautifully)writtencontributionsarewellknowninthefieldIknowofnobodywhocanmatchhisinsightintothespatialorganizationofthecell,andthelimitsofanoverlygeneticapproachtobiologyLastbutnotleast,IwanttothankJohnHancock,ReaderinMolecularBiologyattheUniversityoftheWestofEnglandJohnhasawonderfullywideranging,eclecticknowledgeofbiology,andhiscommentsoftentookmebysurpriseTheymademerethinktheworkabilityofsomeoftheideasIputforward,andhavingdonesotohissatisfaction(Ithink)IamnowmoreconfidentthatmitochondriareallydoholdwithinthemthemeaningoflifeOtherspecialistshavereadchaptersrelatingtotheirownfieldofexpertise,anditisapleasuretorecordmythanksWhenrangingsowidelyoverdifferentfields,itishardtobesureaboutone’sgraspofsignificantdetail,andwithouttheirgenerousresponsetomyemails,naggingdoubtswouldstillbesetmeAsitis,Iamhopefulthattheloomingquestionsreflectnotjustmyownignorance,butalsothatofwholefields,fortheyarethequestionsthatdriveascientist’scuriosityInthisregard,Iwanttothank:JohnAllen,ProfessorofBiochemistry,QueenMaryCollege,UniversityofLondonGustavoBarja,ProfessorofAnimalPhysiology,ComplutenseUniversity,MadridAlbertBennett,ProfessorofEvolutionaryPhysiologyattheUniversityofCalifornia,IrvineDrNeilBlackstone,AssociateProfessorofEvolutionaryBiologyatNorthernIllinoisUniversityDrMartinBrand,MRCDunnHumanNutritionUnit,CambridgeDrJimCummins,AssociateProfessorofAnatomy,MurdochUniversityChrisLeaver,ProfessorofPlantSciences,OxfordUniversityGottfriedSchatz,ProfessorofBiochemistry,UniversityofBaselAloysiusTielens,ProfessorofBiochemistry,UniversityofUtrechtDrJonTurney,ScienceCommunicationGroup,ImperialCollege,LondonDrTiborVellai,InstituteofZoology,FribourgUniversityandAlanWright,ProfessorofGenetics,MRCHumanGeneticsUnit,EdinburghUniversityIamverygratefultoDrMichaelRodgers,formerlyofOUP,whocommissionedthisbookasoneofhisfinalactsbeforeretiringIamhonouredthatheretainedanactiveinterestinprogress,andhecasthiseagleeyeoverthefirstdraftmanuscript,providingextremelyhelpfulcriticalcommentsThebookismuchimprovedasaresultInthesamebreathImustthankLathaMenon,SeniorCommissioningEditoratOUP,whoinheritedthebookfromMichael,andinvesteditwithherlegendaryenthusiasmandappreciationofdetailaswellasthelargerpictureManythankstootoDrMarkRidleyatOxford,authorofMendel’sDemon,whoreadtheentiremanuscriptandprovidedinvaluablecommentsIcan’tthinkofanyonebetterabletoevaluatesomanydisparateaspectsofevolutionarybiology,withsuchagenerousmindI’mproudhefounditastimulatingreadAnumberoffriendsandfamilymembershavealsoreadchaptersandgivenmeagoodindicationofwhatthegeneralreaderispreparedtotolerateIwanttothankinparticularAllysonJones,whoseunfeignedenthusiasmandhelpfulcommentshaveperiodicallysentmyspiritssoaringMikeCarter,whohasbeenfriendenoughtotellmefranklythatsomeearlydraftsweretoodifficult(andthatlateronesweremuchbetter)PaulAsbury,whoisfullofthoughtsandabsorbingconversation,especiallyinwildcornersofthecountrywheretalkisunconstrainedIanAmbrose,alwayswillingtolistenandadvise,especiallyoverapintDrJohnEmsley,fullofguidanceandinspirationProfessorBarryFuller,bestofcolleagues,alwaysreadytotalkoverideasinthelab,thepub,oreventhesquashcourtandmyfather,TomLane,whohasreadmostofthebookandbeengenerousinhispraiseandgentleinpointingoutmystylisticinfelicities,whileworkingtotightdeadlinesonhisownbooksMymotherJeanandbrotherMaxhavebeenunstintingintheirsupport,asindeedhavemySpanishfamily,andIthankthemallThefrontispieceillustrationsarebyDrInaSchuppeKoistenen,aresearcherinbiomedicalsciencesinStockholmandnotedwatercolorist,whoismakinganameinscientificartTheserieswasspeciallycommissionedforthisbook,andinspiredbythethemesofthechaptersI’mverygratefultoher,asIthinktheybringtolifethemysteryofourmicroscopicuniverse,andgivethebookauniqueflavourSpecialthankstoAna,mywife,whohaslivedthisbookwithme,throughxiiAcknowledgementstimesbestdescribedastestingShehasbeenmyconstantsparringcompanion,bouncingideasbackandforth,contributingmorethanafew,andreadingeveryword,well,morethanonceShehasbeentheultimatearbiterofstyle,ideas,andmeaningMydebttoherisbeyondwordsFinally,anotetoEneko:heisantitheticaltowritingbooks,preferringtoeatthem,butisabundleofjoy,andaneducationinhimselfAcknowledgementsxiiiThispageintentionallyleftblankINTRODUCTIONMitochondriaClandestineRulersoftheWorldMitochondriaaretinyorganellesinsidecellsthatgeneratealmostallourenergyintheformofATPOnaveragethereare–ineverycell,givingtenmillionbillioninthehumanbodyEssentiallyallcomplexcellscontainmitochondriaTheylooklikebacteria,andappearancesarenotdeceptive:theywereoncefreelivingbacteria,whichadaptedtolifeinsidelargercellssometwobillionyearsagoTheyretainafragmentofagenomeasabadgeofformerindependenceTheirtortuousrelationswiththeirhostcellshaveshapedthewholefabricoflife,fromenergy,sex,andfertility,tocellsuicide,ageing,anddeathAmitochondriononeofmanytinypowerhouseswithincellsthatcontrolourlivesinsurprisingwaysThispageintentionallyleftblankMitochondriaareabadlykeptsecretManypeoplehaveheardofthemforonereasonoranotherInnewspapersandsometextbooks,theyaresummarilydescribedasthe‘powerhouses’oflifetinypowergeneratorsinsidelivingcellsthatproducevirtuallyalltheenergyweneedtoliveThereareusuallyhundredsorthousandsoftheminasinglecell,wheretheyuseoxygentoburnupfoodTheyaresosmallthatonebillionofthemwouldfitcomfortablyinagrainofsandTheevolutionofmitochondriafittedlifewithaturbochargedengine,revvedupandreadyforuseatanytimeAllanimals,themostslothfulincluded,containatleastsomemitochondriaEvensessileplantsandalgaeusethemtoaugmentthequiethumofsolarenergyinphotosynthesisSomepeoplearemorefamiliarwiththeexpression‘MitochondrialEve’shewassupposedlythemostrecentancestorcommontoallthepeopleslivingtoday,ifwetraceourgeneticinheritancebackupthematernalline,fromchildtomother,tomaternalgrandmother,andsoon,backintothedeepmistsoftimeMitochondrialEve,themotherofallmothers,isthoughttohavelivedinAfrica,perhapsyearsago,andisalsoknownas‘AfricanEve’Wecantraceourgeneticancestryinthiswaybecauseallmitochondriahaveretainedasmallquotaoftheirowngenes,whichareusuallypassedontothenextgenerationonlyintheeggcell,notinthespermThismeansthatmitochondrialgenesactlikeafemalesurname,whichenablesustotraceourancestrydownthefemalelineinthesamewaythatsomefamiliestrytotracetheirdescentdownthemalelinefromWilliamtheConqueror,orNoah,orMohammedRecently,someofthesetenetshavebeenchallenged,butbyandlargethetheorystandsOfcourse,thetechniquenotonlygivesanideaofourancestry,butitalsohelpsclarifywhowerenotourancestorsAccordingtomitochondrialgeneanalysis,Neanderthalmandidn’tinterbreedwithmodernHomosapiens,butwasdriventoextinctionatthemarginsofEuropeMitochondriahavealsomadetheheadlinesfortheiruseinforensics,toestablishthetrueidentityofpeopleorcorpses,includingseveralcelebratedcasesAgain,thetechniquedrawsontheirsmallquotaofgenesTheidentityofthelastRussianTzar,NicholasII,wasverifiedbycomparinghismitochondrialgeneswiththoseofrelativesAyearoldgirlrescuedfromariverinBerlinattheendoftheFirstWorldWarclaimedtobetheTzar’slostdaughterAnastasia,andwascommittedtoamentalinstitutionAfteryearsofdispute,herclaimwasfinallydisprovedbymitochondrialanalysisfollowingherdeathinMorerecently,theunrecognizableremainsofmanyvictimsoftheWorldTradeCentercarnagewereidentifiedbymeansoftheirmitochondrialgenesDistinguishingthe‘real’SaddamHusseinfromoneofhismanydoubleswasachievedbythesametechniqueThereasonthatthemitochondrialgenesaresousefulrelatespartlytotheirabundanceEverymitochondrioncontainstocopiesofitsgenesBecausethereareusuallyhundredsofmitochondriaineverycell,therearemanythousandsofcopiesofthesamegenesineachcell,whereasthereareonlytwocopiesofthegenesinthenucleus(thecontrolcentreofthecell)Accordingly,itisrarenottobeabletoextractanymitochondrialgenesatallOnceextracted,thefactthatallofussharethesamemitochondrialgeneswithourmothersandmaternalrelativesmeansthatitisusuallypossibletoconfirmordisprovepostulatedrelationshipsThenthereisthe‘mitochondrialtheoryofageing’,whichcontendsthatageingandmanyofthediseasesthatgowithitarecausedbyreactivemoleculescalledfreeradicalsleakingfrommitochondriaduringnormalcellularrespirationThemitochondriaarenotcompletely‘sparkproof’Astheyburnupfoodusingoxygen,thefreeradicalsparksescapetodamageadjacentstructures,includingthemitochondrialgenesthemselves,andmoredistantgenesinthecellnucleusThegenesinourcellsareattackedbyfreeradicalsasoftenastotimesaday,practicallyanabuseeverysecondMuchofthedamageisputrightwithoutmoreado,butoccasionalattackscauseirreversiblemutationsenduringalterationsingenesequenceandthesecanbuildupoveralifetimeThemoreseriouslycompromisedcellsdie,andthesteadywastageunderpinsbothageinganddegenerativediseasesManycruelinheritedconditions,too,arelinkedwithmutationscausedbyfreeradicalsattackingmitochondrialgenesThesediseasesoftenhavebizarreinheritancepatterns,andfluctuateinseverityfromgenerationtogeneration,butingeneraltheyallprogressinexorablywithageMitochondrialdiseasestypicallyaffectmetabolicallyactivetissuessuchasthemuscleandbrain,producingseizures,somemovementdisorders,blindness,deafness,andmusculardegenerationMitochondriaarefamiliartoothersasacontroversialfertilitytreatment,inwhichthemitochondriaaretakenfromaneggcell(oocyte)ofahealthyfemaledonor,andtransferredintotheeggcellofaninfertilewomanatechniqueknownas‘ooplasmictransfer’Whenitfirsthitthenews,oneBritishnewspaperranthestoryunderthecolourfulheading‘Babiesbornwithtwomothersandonefather’Thischaracteristicallyvividproductofthepressisnottotallywrongwhileallthegenesinthenucleuscamefromthe‘real’mother,someofthemitochondrialgenescamefromthe‘donor’mother,sothebabiesdidindeedreceivesomegenesfromtwodifferentmothersDespitethebirthofmorethanapparentlyhealthybabiesbythistechnique,bothethicalandpracticalconcernslaterhaditoutlawedinBritainandtheUSClandestineRulersoftheWorldMitochondriaevenmadeitintoaStarWarsmovie,totheangerofsomeaficionados,asaspuriouslyscientificexplanationofthefamousforcethatmaybewithyouThiswasconceivedasspiritual,ifnotreligious,inthefirstfilms,butwasexplainedasaproductof‘midichlorians’inalaterfilmMidichlorians,saidahelpfulJediKnight,are‘microscopiclifeformsthatresideinalllivingcellsWearesymbiontswiththem,livingtogetherformutualadvantageWithoutmidichlorians,lifecouldnotexistandwewouldhavenoknowledgeoftheforce’Theresemblancetomitochondriainbothnameanddeedwasunmistakeable,andintentionalMitochondria,too,haveabacterialancestryandlivewithinourcellsassymbionts(organismsthatshareamutuallybeneficialassociationwithotherorganisms)Likemidichlorians,mitochondriahavemanymysteriousproperties,andcanevenformintobranchingnetworks,communicatingamongthemselvesLynnMargulismadethisoncecontroversialthesisfamousinthes,andthebacterialancestry

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权力、性、自杀:线粒体与生命的意义

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