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Releaseof cltochrome c from mitochondria can be detected as early as 6 hours after exposure of a population of cells to UV light, and it continuesto increase for more than 10 hours thereafter. Does thismean that individual cells slowly release theircltochrome c over this time period? Or, alternatively, doindividual cells release their cytochrome c rapidly butwith different cells being triggered over the longer timeperiod?To answerthis fundamental question,you have fused thegene for green fluorescent protein (GFP) to the gene forc1'tochromec, so that you can observethe behavior of individual cells by confocal fluorescencemicroscopy.In cellsthat are expressingthe cytochrome c-GFP fusion, fluorescence shows the punctate pattern typical of mitochondrialproteins.
You then irradiate these cells with W light andobserveindividual cellsfor changesin the punctate pattern.TWo such cells (outlined in white) are shovrn in FigureQfB-2A and B. Releaseof cytochrome c-GFPis detectedasa change from a punctate to a diffuse pattern of fluorescence.Times after W exposureare indicated as hours:minutes below the individual panels.Which model for cytochrome c releasedo these observationssupport?Explainyour reasoning.REFERENCESAdamsJM,HuangDC,StrasserA et al (2005)Subversionofthe Bcl2life/deathswitchin cancerdevelopmentandtherapyColdSpringHarbSympQuantBiol7A.46977KM& SalvesenBoatrightGS(2003)Mechanismsof caspaseactivation,CurrOpinCellBiol15:725731DanialNN& KorsmeyerSJ(2004)Celldeath:criticalcontrolpointsCel/116:205-219) echanismE l l i sR EY, u a nJ Y& H o r v i tRz A( 1 9 9 1Ma sn df u n c t i o nosf c e l ldeath AnnuRevCellBiol7:663-698givingphosphatidylserineFadokVA& HensonPM(2003)Apoptosis:recognitionan assist-witharwisrCurrBial13:R655-R657GalonekHL& HardwicklM (2006)Upgradingthe BCL-2networkNatureCellBiolB:131/- 1319ten minutesto dead Cel/GreenDR(2005)Apoptoticpathways:1 2 1.
6 7 1 , 6 7 4HorvitzHR(2003)Worms,life,and death(Nobellecture)Chembiochem-7 114'697Hyun-EuiK,FengheD,FangM & WangX (2005)Formationofis initiatedby cytochromec-induceddATPhydrolysisapoptosomeand subsequentnucleotideexchangeon Apaf-1ProcNatlAcadSciUSA102:175451755aMD,WeilM & RaffMC (1997)ProgrammedcelldeathinJacobsonanimalde',elnnme^raoll88347-354,apoptosisAnnuRevJiangX & WangX (2004)CytochromeC-mediatedBiochem/3.87-106KerrJF,WyllieAH & CurrieAR(1972)Apoptosis:a basicblologicalphenomenonin tissuekineticswith wide-ranglnglmplicationsBJ Cancer26.239-25710:09l 0 : 1517il017:18analysismicroscopicvideo,fluorescenceFigureQ18-2 Time-lapseof individualcellsfrom mitochondriaof cytochromec-GFPrelease(Problem18-7).(A)Cellsobservedfor 8 minutes,10 hoursafter(B)Cellsobservedfor 6 minutes,17 hoursafterUV irradiation.UV irradiation.One cell in (A)and one in (B),eachoutlinedin white,theircytochromec-GFPduringthe time frameof thehavereleasedbeloweachpanel.which is shownas hours:minutesobservation,(FromJ.C.Goldsteinet al.,Ndt CellBiol.2:156-'162,2000.Ltd.)from MacmillanPublishersWith permissioncelldeath CellDeathfunctionin programmedCaspaseKumarS (2007)Differ14:3243J Cel/PH(2005)DeathreceptorsignalingLavrikl,GolksA & KrammerSci118,265-267tumoursuppressionLoweSW,CeperoE & EvanG (2004)IntrinsicNature432307 315.ofLumJJ,BauerDE,KongM et al (2005)GrowthfactorregulationCel/ofapoptosis,in the absenceandcellsurvivalautophagy120,23748ofMcCallK & Stellerl, (1997)Facingdeathin the fly:geneticanalysisTrendsGenet13222-226apoptosisin Drosophila.AnnuRevGenetapoptosisNagataS (1999)Fasligand-induced33.2955and programmedin developmentNagataS (2005)DNAdegradationcelldeath,AnnuRevlmmunol23.853-875TheapoptosomeGS(2006)S & SalvesenPopC,TimmerJ,SperandioMolCell22:269-275by dimerizationcaspase-9activatesRaffMC (1999)Cellsuicidefor beginnersNature396:119-122inof apoptosisRathmell.JC& ThompsonCB(2002)PathwaysanddiseaseCel/homeostasis,development,lymphocyte109.s97-107celldeathof programmedTittelJN& StellerH (2000)A comparisonbetween speciesGenomeBiol 1AM,CoulsonEJ& VauxDL (2001)Inhibitorof apoptosisVerhagenBiolGenomeproteinsandtheirrelatives:lAPsand otherBIRPs-3009102:3009.1VousdenKH(2005)Apoptosisp53and PUMA:a deadlyduo Science685-1686309:1prot'oinshow BH3-onlyWillisSN& AdamsJM (2005)Lifein the balance:induceapoptosisCurrapin CellBiol17:617-625CELLSIN THEIRSOCIALCONTEXT[€ttE6rrtlvMttS)INVtd tHlslnssu_r^rlf tNNo)sLLLIVWINVIO XIUIVWUilNII]fVUIX] ]HINOISSHCV6 9 rtV9LL85t I,xtulvy\j-tt3fCNVSNIU9]I.NIVNIY\JV-IIVSVB]HIVlVI^IS]COt/\JSVIdCNVSNOIIfNNTdv9 :ttl) or tttfWOUISAVM]DVSSVdvnlHltdlIO NOIIVZINV9UO]HI09r r cNV sNotlf Nnf 1H9U_€€IINOtStHCVIl)-ttl)CNVSNIU]HCV)laldeq) sFlI ul'elrs uorsaqpP ol alrs uorseqpPruo{ 'lle3 qJBe Jorra}ureq} ssoJJEsessaJlsJoSuplpusueD 'paJoqJup eJe slueruelg 1erala>lsol,(JeJeqm 'suorsaqpe llef,-lleo,{q ,{.1lcar1preqlo qrea ol peqoupu ere sllec eql 'urnlaqtrda aqt ul{tl6'leeqsaql Jo aJeJ auo Surr(lrapun'(auatqutara luautasaq rc) aurruDl 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