Chapter26

TheTreeofLife:
AnIntroductiontoBiologicalDiversity

LectureOutline

Overview:ChangingLifeonaChangingEarth

Lifeisacontinuumextendingfromtheearliestorganismstothegreatvarietyofforms
alivetoday.
Organismsinteractwiththeirenvironments.
Geologicaleventsthatalterenvironmentschangethecourseofbiologicalhistory.
Whenglaciersrecedeandthelandrebounds,marinecreaturescanbetrappedinwhat
graduallybecomefreshwaterlakes.
Populationsoforganismstrappedintheselakesareisolatedfromparentpopulations,
andmayevolveintonewspecies.
Lifechangestheplanetitinhabits.
Theevolutionofphotosyntheticorganismsreleasedoxygenintotheair,witha
dramaticeffectonEarthsatmosphere.
TheemergenceofHomosapienshaschangedtheland,water,andairatanunprecedented
rate.
Historicalstudyofanysortisaninexactdisciplinethatdependsonthepreservation,
reliability,andinterpretationofrecords.
Thefossilrecordofpastlifeisgenerallylessandlesscompletethefurtherintothepast
wedelve.
Fortunately,eachorganismalivetodaycarriestracesofitsevolutionaryhistoryinits
molecules,metabolism,andanatomy.
Still,theevolutionaryepisodesofgreatestantiquityaregenerallythemostobscure.

Concept26.1ConditionsonearlyEarthmadetheoriginoflifepossible
Mostbiologistsnowthinkthatitiscrediblethatchemicalandphysicalprocesseson
Earthproducedsimplecells.
Accordingtoonehypotheticalscenario,therewerefourmainstagesinthisprocess:
1. Theabioticsynthesisofsmallorganicmolecules(monomers).
2. Thejoiningofmonomersintopolymers.
3. Thepackagingofthesemoleculesintoprotobionts,dropletswithmembranesthat
maintainedadistinctinternalchemistry.

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 4. Theoriginofselfreplicatingmoleculesthateventuallymadeinheritance
possible.
Thescenarioisspeculativebutdoesleadtopredictionsthatcanbetestedinlaboratory
experiments.
Earthandtheotherplanetsinthesolarsystemformedabout4.6billionyearsago,
condensingfromavastcloudofdustandrockssurroundingtheyoungsun.
Itisunlikelythatlifecouldhaveoriginatedorsurvivedinthefirstfewhundredmillion
yearsaftertheEarthsformation.
Theplanetwasbombardedbyhugebodiesofrockandiceleftoverfromtheformationof
thesolarsystem.
Thesecollisionsgeneratedenoughheattovaporizeallavailablewaterandpreventthe
formationoftheseas.
TheoldestrocksontheEarthssurface,locatedatasitecalledIsuainGreenland,are3.8
billionyearsold.
Itisnotclearwhethertheserocksshowtracesoflife.
ThefirstcellsmayhaveoriginatedbychemicalevolutiononayoungEarth.
ItiscrediblethatchemicalandphysicalprocessesonearlyEarthproducedthefirstcells.
Accordingtoonehypothesis,therewerefourmainstagestothisprocess:
1. Abioticprocessessynthesizedsmallorganicmolecules,suchasaminoacidsand
nucleotides.
2. Thesemonomerswerejoinedintopolymers,includingproteinsandnucleicacids.
3. Polymerswerepackagedintoprotobionts,dropletswithmembranesthat
maintainedaninternalchemistrydistinctfromtheirsurroundings.
4. Selfreplicatingmoleculesarose,makinginheritancepossible.
Abioticsynthesisoforganicmonomersisatestablehypothesis.
AsthebombardmentofearlyEarthslowed,conditionsontheplanetwereverydifferent
fromtoday.
Thefirstatmospheremayhavebeenareducingatmospherethickwithwatervapor,along
withnitrogenanditsoxides,carbondioxide,methane,ammonia,hydrogen,andhydrogen
sulfide.
Similarcompoundsarereleasedfromvolcaniceruptionstoday.
AsEarthcooled,thewatervaporcondensedintotheoceansandmuchofthehydrogen
waslostintospace.
Inthe1920s,RussianchemistA.I.OparinandBritishscientistJ.B.S.Haldane
independentlypostulatedthatconditionsonearlyEarthfavoredthesynthesisoforganic
compoundsfrominorganicprecursors.
Theyreasonedthatthiscouldnothappentodaybecausehighlevelsofoxygeninthe
atmosphereattackchemicalbonds.
Areducingenvironmentintheearlyatmospherewouldhavepromotedthejoiningof
simplemoleculestoformmorecomplexones.
Theconsiderableenergyrequiredtomakeorganicmoleculescouldbeprovidedby
lightningandtheintenseUVradiationthatpenetratedtheprimitiveatmosphere.

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 YoungsunsemitmoreUVradiation.Thelackofanozonelayerintheearlyatmosphere
wouldhaveallowedthisradiationtoreachEarth.
Haldanesuggestedthattheearlyoceanswereasolutionoforganicmolecules,a
primitivesoupfromwhichlifearose.
In1953,StanleyMillerandHaroldUreytestedtheOparinHaldanehypothesisby
creating,inthelaboratory,theconditionsthathadbeenpostulatedforearlyEarth.
Theydischargedsparksinanatmosphereofgasesandwatervapor.
TheMillerUreyexperimentsproducedavarietyofaminoacidsandotherorganic
molecules.
OtherattemptstoreproducetheMillerUreyexperimentwithothergasmixtureshave
alsoproducedorganicmolecules,althoughinsmallerquantities.
Itisunclearwhethertheatmospherecontainedenoughmethaneandammoniatobe
reducing.
Thereisgrowingevidencethattheearlyatmospherewasmadeupprimarilyofnitrogen
andcarbondioxide.
MillerUreytypeexperimentswithsuchatmosphereshavenotproducedorganic
molecules.
Itislikelythatsmallpocketsoftheearlyatmospherenearvolcanicopeningswere
reducing.
Alternatesitesproposedforthesynthesisoforganicmoleculesincludesubmerged
volcanoesanddeepseaventswherehotwaterandmineralsgushintothedeepocean.
Theseregionsarerichininorganicsulfurandironcompounds,whichareimportantin
ATPsynthesisbypresentdayorganisms.
SomeoftheorganiccompoundsfromwhichthefirstlifeonEartharosemayhavecome
fromspace.
ResearchersarelookingoutsideofEarthforcluesabouttheoriginoflife.
EvidenceisgrowingthatMarswasrelativelywarmforabriefperiod,withliquidwater
andanatmosphererichincarbondioxide.
Duringthatperiod,prebioticchemistrysimilartothatonearlyEarthmayhaveoccurred
onMars.
DidlifeevolveonMarsandthendieout,ordiddroppingtemperaturesandathinning
atmosphereterminateprebioticchemistrybeforelifeevolved?
LiquidwaterliesbeneaththeicecoveredsurfaceofEuropa,oneofJupitersmoons,
raisingthepossibilitythatEuropashiddenoceanmayharborlife.
Detectionoffreeoxygenintheatmosphereofanyplanetsoutsideoursolarsystemwould
bestronglysuggestiveofoxygenicphotosynthesis.
LaboratorysimulationsofearlyEarthconditionshaveproducedorganicpolymers.
Theabioticoriginhypothesispredictsthatmonomersshouldlinktoformpolymers
withoutenzymesandothercellularequipment.
Researchershaveproducedpolymers,includingpolypeptides,afterdrippingsolutionsof
monomersontohotsand,clay,orrock.
SimilarconditionslikelyexistedonearlyEarthatdeepseaventsorwhendilutesolutions
ofmonomerssplashedontofreshlava.

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Protobiontscanformbyselfassembly.
Lifeisdefinedbytwoproperties:accuratereplicationandmetabolism.
Neitherpropertycanexistwithouttheother.
DNAmoleculescarrygeneticinformation,includingtheinformationneededforaccurate
replication.
ThereplicationofDNArequireselaborateenzymaticmachinery,alongwithacopious
supplyofnucleotidebuildingblocksprovidedbycellmetabolism.
AlthoughMillerUreyexperimentshaveyieldedsomeofthenitrogenousbasesofDNA
andRNA,theyhavenotproducedanythinglikenucleotides.
Thus,nucleotideswerelikelynotpartoftheearlyorganicsoup.
Selfreplicatingmoleculesandametabolismlikesourceofthebuildingblocksmusthave
appearedtogether.
Thenecessaryconditionsmayhavebeenprovidedbyprotobionts,aggregatesof
abioticallyproducedmoleculessurroundedbyamembraneormembranelikestructure.
Protobiontsexhibitsomeofthepropertiesassociatedwithlife,includingreproduction
andmetabolism,andcanmaintainaninternalchemicalenvironmentdifferentfromtheir
surroundings.
Laboratoryexperimentsshowthespontaneousformationofprotobiontsfromabiotically
producedorganiccompounds.
Forexample,dropletsofabioticallyproducedorganiccompoundscalledliposomesform
whenlipidsandotherorganicmoleculesareaddedtowater.
Thelipidsformamolecularbilayeratthedropletsurface,muchlikethelipidbilayerofa
membrane.
Thesedropletscanundergoosmoticswellingorshrinkingindifferentsaltconcentrations.
Someliposomesstoreenergyintheformofamembranepotential.
Liposomesbehavedynamically,growingbyengulfingsmallerliposomesorgiving
birthtosmallerliposomes.
IfsimilardropletsforminginpondsonearlyEarthincorporaterandompolymersof
linkedaminoacidsintotheirmembranes,andifsomeofthesepolymersmadethemembranes
permeabletomolecules,thenthosedropletscouldhaveselectivelytakenuporganic
moleculesfromtheirenvironment.
RNAmayhavebeenthefirstgeneticmaterial.
ThefirstgeneticmaterialwasprobablyRNA,notDNA.
ThomasCechandSidneyAltmanfoundthatRNAmoleculesnotonlyplayacentralrole
inproteinsynthesis,butalsoareimportantcatalystsinmoderncells.
RNAcatalysts,calledribozymes,removetheirownintronsandmodifytRNAmolecules
tomakethemfullyfunctional.
RibozymesalsohelpcatalyzethesynthesisofnewRNApolymers.
Ribozymecatalyzedreactionsareslow,buttheproteinsnormallyassociatedwith
ribozymescanincreasethereactionratemorethanathousandfold.
LaboratoryexperimentshavedemonstratedthatRNAsequencescanevolveunderabiotic
conditions.

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 UnlikedoublestrandedDNA,singlestrandedRNAmoleculescanassumeavarietyof3
Dshapesspecifiedbytheirnucleotidesequences.
RNAmoleculeshavebothagenotype(nucleotidesequence)andaphenotype(three
dimensionalshape)thatinteractswithsurroundingmolecules.
Underparticularconditions,someRNAsequencesaremorestableandreplicatefaster
andwithfewererrorsthanothersequences.
Occasionalcopyingerrorscreatemutations;selectionscreensthesemutationsforthe
moststableorthebestatselfreplication.
BeginningwithadiversityofRNAmoleculesthatmustcompeteformonomersto
replicate,thesequencebestsuitedtothetemperature,saltconcentration,andother
featuresofthesurroundingenvironmentandhavingthegreatestautocatalyticactivity
willincreaseinfrequency.
ItsdescendentswillbeafamilyofcloselyrelatedRNAsequences,differingdueto
copyingerrors.
Somecopyingerrorswillresultinmoleculesthataremorestableormorecapableofself
replication.
SimilarselectioneventsmayhaveoccurredonearlyEarth.
ModernmolecularbiologymayhavebeenprecededbyanRNAworld.
Naturalselectioncouldrefineprotobiontscontaininghereditaryinformation.
ThefirstRNAmoleculesmayhavebeenshort,viruslikesequences,aidedintheir
replicationbyaminoacidpolymerswithrudimentarycatalyticcapabilities.
Thisearlyreplicationmayhavetakenplaceinsideprotobionts.
RNAdirectedproteinsynthesismayhavebegunasweakbindingofspecificaminoacids
tobasesalongRNAmolecules,whichfunctionedassimpletemplatesholdingafew
aminoacidstogetherlongenoughforthemtobelinked.
ThisisonefunctionofrRNAtodayinribosomes.
SomeRNAmoleculesmayhavesynthesizedshortpolypeptidesthatbehavedasenzymes
helpingRNAreplication.
Earlychemicaldynamicswouldincludemolecularcooperationaswellascompetition.
OtherRNAsequencesmighthavebecomeembeddedintheprotobiontmembrane,
allowingittousehighenergyinorganicmoleculessuchashydrogensulfidetocarryout
organicreactions.
Aprotobiontwithselfreplicating,catalyticRNAwoulddifferfromotherswithoutRNA
orwithRNAwithfewercapabilities.
Ifthatprotobiontcouldgrow,split,andpassitsRNAmoleculestoitsdaughters,the
daughterswouldhavesomeofthepropertiesoftheirparent.
Thefirstprotobiontsmusthavehadlimitedamountsofgeneticinformation,specifying
onlyafewproperties.
Becausetheirpropertieswereheritable,theycouldbeactedonbynaturalselection.
Themostsuccessfuloftheseprotobiontswouldhaveincreasedinnumbers,becausethey
couldexploitavailableresourcesandproduceanumberofsimilardaughterprotobionts.
OnceRNAsequencesthatcarriedgeneticinformationappearedinprotobionts,many
furtherchangeswerepossible.

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 OnerefinementwasthereplacementofRNAastherepositoryofgeneticinformationby
DNA.
DoublestrandedDNAisamorestablemolecule,anditcanbereplicatedmore
accurately.
OnceDNAappeared,RNAmoleculeswouldhavebeguntotakeontheirmodernrolesas
intermediatesintranslationofgeneticprograms.
TheRNAworldgavewaytoaDNAworld.

Concept26.2ThefossilrecordchronicleslifeonEarth
Radiometricdatinggivesabsolutedatesforsomerockstrata.
Therelativesequenceoffossilsinrockstratatellsustheorderinwhichthefossilswere
formed,butitdoesnottellustheirages.
Geologistshavedevelopedmethodsforobtainingabsolutedatesforfossils.
Oneofthemostcommontechniquesisradiometricdating,whichisbasedonthedecay
ofradioactiveisotopes.
Anisotopeshalflife,thenumberofyearsittakesfor50%oftheoriginalsampleto
decay,isunaffectedbytemperature,pressure,orotherenvironmentalvariables.
Fossilscontainisotopesofelementsthataccumulatedwhiletheorganismswerealive.
Forexample,thecarboninalivingorganismcontainsthemostcommoncarbonisotope,
carbon12,aswellasaradioactiveisotope,carbon14.
Whenanorganismdies,itstopsaccumulatingcarbon,andthecarbon14thatitcontained
atthetimeofdeathslowlydecaystonitrogen14.
Bymeasuringtheratioofcarbon14tototalcarbonortonitrogen14inafossil,wecan
determinethefossilsage.
Withahalflifeof5,730years,carbon13isusefulfordatingfossilsuptoabout
75,000yearsold.
Fossilsolderthanthatcontaintoolittlecarbon14tobedetectedbycurrent
techniques.
Radioactiveisotopeswithlongerhalflivesareusedtodateolderfossils.
Paleontologistscandeterminetheageoffossilssandwichedbetweenlayersofvolcanic
rocksbymeasuringtheamountofpotassium40inthoselayers.
Potassium40decaystothechemicallyunreactivegasargon40,whichistrappedinthe
rock.
Whentherockisheatedduringavolcaniceruption,theargonisdrivenout,butthe
potassiumremains.
Thisresetstheclockforpotassium40tozero.
Thecurrentratioofpotassium40toargon40inalayerofvolcanicrockgivesan
estimateofwhenthatlayerwasformed.
Magnetismofrockscanalsobeusedtodatethem.
Whenvolcanicorsedimentaryrockforms,ironparticlesintherockalignthemselves
withEarthsmagneticfield.
Whentherockhardens,theirorientationisfrozenintime.

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 GeologistshavedeterminedthatEarthsnorthandsouthmagneticpoleshave
reversedrepeatedlyinthepast.
Thesemagneticreversalshavelefttheirrecordonrocksthroughouttheworld.
Patternsofmagneticreversalcanbematchedwithcorresponding
patternselsewhere,allowingrockstobedatedwhenothermethodsarenot
available.
GeologistshaveestablishedageologicrecordofEarthshistory.
Bystudyingrocksandfossilsatmanydifferentsites,geologistshaveestablisheda
geologicrecordofthehistoryoflifeonEarth,whichisdividedintothreeeons.
ThefirsttwoeonstheArchaeanandtheProterozoiclastedapproximatelyfourbillion
years.
ThesetwoeonsarereferredtoasthePrecambrian.
ThePhanerozoiceoncoversthelasthalfbillionyearsandencompassesmuchofthetime
thatmulticellulareukaryoticlifehasexistedonEarth.
Itisdividedintothreeeras:Paleozoic,Mesozoic,andCenozoic.
EachagerepresentsadistinctageinthehistoryofEarthandlifeonEarth.
Theboundariesbetweenerascorrespondtotimesofmassextinction,whenmanyforms
oflifedisappeared.
MassextinctionshavedestroyedthemajorityofspeciesonEarth.
Aspeciesmaybecomeextinctformanyreasons.
Itshabitatmayhavebeendestroyed,oritsenvironmentmayhavechangedinadirection
unfavorabletothespecies.
Biologicalfactorsmaychange,asevolutionarychangesinonespeciesimpactothers.
Onanumberofoccasions,globalenvironmentalchangesweresorapidandmajorthat
themajorityofspecieswentextinct.
Suchmassextinctionsareknownprimarilyfromthelossofshallowwater,marine,hard
bodiedanimals,theorganismsforwhichthefossilrecordismostcomplete.
ThePermianmassextinctiondefinestheboundarybetweenthePaleozoicandMesozoic
eras.
Ninetysixpercentofmarineanimalspecieswentextinctinlessthan5millionyears.
Terrestriallifewasalsoaffected.
TheCretaceousextinctionof65millionyearsmarkstheboundarybetweentheMesozoic
andCenozoiceras.
Morethanhalfofallmarinespeciesandmanyfamiliesofterrestrialplantsandanimals,
includingthedinosaurs,wentextinct.
ThePermianmassextinctionhappenedatatimeofenormousvolcaniceruptionsinwhat
isnowinSiberia.
Theseeruptionsmayhaveproducedenoughcarbondioxidetowarmtheglobalclimate.
Reducedtemperaturedifferencesbetweentheequatorandthepoleswouldhaveslowed
themixingofoceanwater.
TheresultingoxygendeficitintheoceansmayhaveplayedalargeroleinthePermian
extinction.

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 AcluetotheCretaceousmassextinctionisathinlayerofclayenrichediniridiumthat
separatessedimentsfromtheMesozoicandCenozoic.
IridiumisaveryrareelementonEarththatiscommoninmeteoritesandotherobjects
thatfalltoEarth.
WalterandLuisAlvarezandtheircolleaguesattheUniversityofCaliforniaproposed
thatthisclayisfalloutfromahugecloudofdebristhatwasthrownintotheatmosphere
whenanasteroidoralargecometcollidedwithEarth.
Thecloudwouldhaveblockedsunlightanddisruptedtheglobalclimateforseveral
months.
A65millionyearoldcraterscarhasbeenlocatedbeneathsedimentsontheYucatn
coastofMexico.
At180kmindiameter,itistherightsizetohavebeencausedbyanobjectwitha
diameterof10km.
Muchremainstobelearnedaboutthecausesofmassextinctions.
Itisclearthattheyprovidedlifewithopportunitiesforadaptiveradiationsintonewly
vacatedecologicalniches.

Concept26.3Asprokaryotesevolved,theyexploitedandchangedyoungEarth
Theoldestknownfossilsare3.5billionyearoldstromatolites,rocklikestructures
composedoflayersofcyanobacteriaandsediment.
Ifbacterialcommunitiesexisted3.5billionyearsago,itseemsreasonablethatlife
originatedmuchearlier,perhaps3.9billionyearsago,whenEarthfirstcooledtoa
temperaturewhereliquidwatercouldexist.
Prokaryotesdominatedevolutionaryhistoryfrom3.5to2.0billionyearsago.
Theearlyprotobiontsmusthaveusedmoleculespresentintheprimitivesoupfortheir
growthandreplication.
Eventually,organismsthatcouldproducealltheirneededcompoundsfrommoleculesin
theirenvironmentreplacedtheseprotobionts.
Arichvarietyofautotrophsemerged,someofwhichcoulduselightenergy.
Thediversificationofautotrophsallowedtheemergenceofheterotrophs,whichcould
liveonmoleculesproducedbytheautotrophs.
ProkaryoteswereEarthssoleinhabitantsfrom3.5to2.0billionyearsago.
Theseorganismstransformedthebiosphereoftheplanet.
Relativelyearly,prokaryotesdivergedintotwomainevolutionarybranches,thebacteria
andthearchaea.
Representativesfrombothgroupsthriveinvariousenvironmentstoday.
Metabolismevolvedinprokaryotes.
ThechemiosmoticmechanismofATPsynthesisiscommontoallthreedomains
Bacteria,Archaea,andEukarya.
Thisisevidenceofarelativelyearlyoriginofchemiosmosis.

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 TransmembraneprotonpumpsmayhavefunctionedoriginallytoexpelH +that
accumulatedwhenfermentationproducedorganicacidsaswasteproducts.
ThecellwouldhavetospendalargeportionofitsATPtoregulateinternalpHbydriving
H+pumps.
Thefirstelectrontransportpumpsmayhavecoupledtheoxidationoforganicacidstothe
transportofH+outofthecell.
Finally,insomeprokaryotes,electrontransportsystemsefficientenoughtoexpelmore
H+thannecessarytoregulatepHevolved.
ThesecellscouldusetheinwardgradientofH+toreversetheH+pump,whichnow
generatedATPinsteadofconsumingit.
Suchanaerobicrespirationpersistsinsomepresentdayprokaryotes.
Photosynthesisprobablyevolvedveryearlyinprokaryotichistory.
Themetabolismofearlyversionsofphotosynthesisdidnotsplitwaterandliberate
oxygen.
Somelivingprokaryotesdisplaysuchnonoxygenicphotosynthesis.
TheonlylivingphotosyntheticprokaryotesthatgenerateO2arecyanobacteria.
Mostatmosphericoxygenisofbiologicalorigin,fromthewatersplittingstepof
photosynthesis.
Whenoxygenicphotosynthesisfirstevolved,thefreeoxygenitproducedlikelydissolved
inthesurroundingwateruntiltheseasandlakesbecamesaturatedwithO 2.
AdditionalO2thenreactedwithdissolvedirontoformtheprecipitateironoxide.
Thesemarinesedimentswerethesourceofbandedironformations,redlayersofrock
containingironoxidethatareavaluablesourceofironoretoday.
About2.7billionyearsago,oxygenbeganaccumulatingintheatmosphereandterrestrial
rockswithoxidizedironformed.
Whileoxygenaccumulationwasgradualbetween2.7and2.2billionyearsago,itshotup
to10%ofcurrentvaluesshortlyafterward.
Thisoxygenrevolutionhadanenormousimpactonlife.
Initsfreemolecularandionizedformsandincompoundssuchashydrogenperoxide,
oxygenattackschemicalbonds,inhibitsenzymes,anddamagescells.
Theincreaseinatmosphericoxygenlikelydoomedmanyprokaryotegroups.
Somespeciessurvivedinhabitatsthatremainedanaerobic,wheretheirdescendents
surviveasobligateanaerobes.
OtherspeciesevolvedmechanismstouseO2incellularrespiration,whichusesoxygento
helpharvesttheenergystoredinorganicmolecules.

Concept26.4Eukaryoticcellsarosefromsymbiosesandgeneticexchangesbetween
prokaryotes
Eukaryoticcellsdifferinmanyrespectsfromthesmallercellsofbacteriaandarchaea.
Eventhesimplestsinglecelledeukaryoteisfarmorecomplexinstructurethanany
prokaryote.

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 Whilethereissomeevidenceofearliereukaryoticfossils,thefirstclearlyidentified
eukaryoteappearedabout2.1billionyearsago.
Otherfossilsthatresemblesimple,singlecelledalgaeareslightlyolder(2.2billionyears)
butmaynotbeeukaryotic.
Tracesofmoleculessimilartocholesterolarefoundinrocksdatingback2.7billion
years.
Suchmoleculesarefoundonlybyaerobicallyrespiringeukaryoticcells.
Ifconfirmed,thiswouldplacetheearliesteukaryotesatthesametimeastheoxygen
revolutionthatchangedtheEarthsenvironmentsodramatically.
Prokaryoteslackinternalstructuressuchasthenuclearenvelope,endoplasmicreticulum,
andGolgiapparatus.
Theyhavenocytoskeletonandareunabletochangecellshape.
Eukaryoticcellshaveacytoskeletonandcanchangeshape,enablingthemtosurround
andengulfothercells.
Thefirsteukaryotesmayhavebeenpredatorsofothercells.
Acytoskeletonenablesaeukaryoticcelltomovestructureswithinthecellandfacilitates
themovementofchromosomesinmeiosisandmitosis.
Mitosismadeitpossibletoreproducethelargeeukaryoticgenome.
Meiosisallowedsexualrecombinationofgenes.
Howdidthecomplexorganizationoftheeukaryoticcellevolvefromthesimpler
prokaryoticcondition?
Aprocesscalledendosymbiosisprobablyledtomitochondriaandplastids(thegeneral
termforchloroplastsandrelatedorganelles).
Theendosymbiotictheorysuggeststhatmitochondriaandplastidswereformerlysmall
prokaryoteslivingwithinlargercells.
Thetermendosymbiontisusedforacellthatliveswithinahostcell.
Theproposedancestorsofmitochondriawereaerobicheterotrophicprokaryotes.
Theproposedancestorsofplastidswerephotosyntheticprokaryotes.
Theprokaryoticancestorsofmitochondriaandplastidsprobablygainedentrytothehost
cellasundigestedpreyorinternalparasites.
Thesymbiosisbecamemutuallybeneficial.
Aheterotrophichostcouldusenutrientsreleasedfromphotosynthesis.
Ananaerobichostwouldhavebenefitedfromanaerobicendosymbiont.
Astheybecameincreasinglyinterdependent,thehostandendosymbiontsbecameasingle
organism.
Alleukaryoteshavemitochondriaortheirgeneticremnants.
Thetheoryofserialendosymbiosissupposesthatmitochondriaevolvedbeforeplastids.
Overwhelmingevidencesupportsanendosymbioticoriginofplastidsandmitochondria.
Theinnermembranesofbothorganelleshaveenzymesandtransportsystemsthatare
homologoustothoseintheplasmamembranesofmodernprokaryotes.
Mitochondriaandplastidsreplicatebyasplittingprocesssimilartoprokaryoticbinary
fission.

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 Likeprokaryotes,eachorganellehasasingle,circularDNAmoleculethatisnot
associatedwithhistone.
TheseorganellescontaintRNAs,ribosomes,andothermoleculesneededtotranscribe
andtranslatetheirDNAintoprotein.
Ribosomesofmitochondriaandplastidsaresimilartoprokaryoticribosomesintermsof
size,nucleotidesequence,andsensitivitytoantibiotics.
Whichprokaryoticlineagesgaverisetomitochondriaandplastids?
ComparisonsofsmallsubunitribosomalRNAfrommitochondria,plastids,andvarious
livingprokaryotessuggestthatagroupofbacteriacalledthealphaproteobacteriaarethe
closestrelativestomitochondriaandthatcyanobacteriaaretheclosestrelativesto
plastids.
Overtime,geneshavebeentransferredfrommitochondriaandplastidstothenucleus.
Thisprocessmayhavebeenaccomplishedbytransposableelements.
SomemitochondrialandplasticproteinsareencodedbytheorganellesDNA,while
othersareencodedbynucleargenes.
Someproteinsarecombinationsofpolypeptidesencodedbygenesinbothlocations.
Theoriginsofotheraspectsofeukaryoticcellsareunclear.
Someresearchershaveproposedthatthenucleusitselfevolvedfromanendosymbiont.
Nucleargeneswithcloserelativesinbothbacteriaandarchaeahavebeenfound.
Thegenomeofeukaryoticcellsmaybetheproductofgeneticannealing,inwhich
horizontalgenetransfersoccurredbetweenmanydifferentbacterialandarchaeallineages.
Thesetransfersmayhavetakenplaceduringtheearlyevolutionoflife,ormayhave
happenedrepeatedlyuntilthepresentday.
Theoriginofothereukaryoticstructuresisalsothesubjectofactiveresearch.
TheGolgiapparatusandtheendoplasmicreticulummayhaveoriginatedfrominfoldings
oftheplasmamembrane.
Thecytoskeletalproteinsactinandtubulinhavebeenfoundinbacteria,wheretheyare
involvedinpinchingoffbacterialcellsduringcelldivision.
Thesebacterialproteinsmayprovideinformationabouttheoriginoftheeukaryotic
cytoskeleton.
Someinvestigatorshavesuggestedthateukaryoticflagellaandciliaevolvedfrom
symbioticbacteria.
However,the9+2microtubuleapparatusofeukaryoticflagellaandciliahasnotbeen
foundinanyprokaryotes.

Concept26.5Multicellularityevolvedseveraltimesineukaryotes
Agreatrangeofeukaryoticunicellularformsevolvedasthediversityofpresentday
protists.
Molecularclockssuggestthatthecommonancestorofmulticellulareukaryoteslived1.5
billionyearsago.
Theoldestknownfossilsofmulticellulareukaryotesare1.2billionyearsold.
RecentfossilfindsfromChinahaveproducedadiversityofalgaeandanimalsfrom570
millionyearsago,includingbeautifullypreservedembryos.

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 Whyweremulticellulareukaryotessolimitedinsize,diversity,anddistributionuntilthe
lateProterozoic?
GeologicevidencesuggeststhatasevereiceagegrippedEarthfrom750to570million
yearsago.
AccordingtothesnowballEarthhypothesis,lifewouldhavebeenconfinedtodeepsea
ventsandhotspringsorthosefewlocationswhereenoughicemeltedforsunlightto
penetratethesurfacewatersofthesea.
Thefirstmajordiversificationofmulticellulareukaryoticorganismscorrespondstothe
timeofthethawingofsnowballEarth.
Thefirstmulticellularorganismswerecolonies.
Somecellsinthecoloniesbecamespecializedfordifferentfunctions.
Suchspecializationcanbeseeninsomeprokaryotes.
Forexample,certaincellsofthefilamentouscyanobacteriumNostocdifferentiateinto
nitrogenfixingcellscalledheterocysts,whichcannotreplicate.
Theevolutionofcolonieswithcellularspecializationwascarriedmuchfurtherin
eukaryotes.
Amulticellulareukaryotegenerallydevelopsfromasinglecell,usuallyazygote.
Celldivisionandcelldifferentiationhelptransformthesinglecellintoamulticellular
organismwithmanytypesofspecializedcells.
Withincreasingcellspecialization,specificgroupsofcellsspecializedinobtaining
nutrients,sensingtheenvironment,etc.
Thisdivisionoffunctioneventuallyledtotheevolutionoftissues,organs,andorgan
systems.
Multicellularityevolvedseveraltimesamongearlyeukaryotes.
AnimaldiversityexplodedduringtheearlyCambrianperiod.
Mostofthemajorphylaofanimalsappearsuddenlyinthefossilrecordintheadaptive
radiationknownastheCambrianexplosion.
Cnidarians(thephylumthatincludesjellies)andporiferans(sponges)werealready
presentinthelatePrecambrian.
However,mostofthemajorgroups(phyla)ofanimalsmaketheirfirstfossilappearances
duringtherelativelyshortspanoftheCambrianperiodsfirst20millionyears.
Molecularevidencesuggeststhatanimalphylaoriginatedandbegantodivergebetween1
billionand700millionyearsago.
AtthebeginningoftheCambrian,thesephylasuddenlyandsimultaneouslyincreasedin
diversityandsize.
Plants,fungi,andanimalscolonizedthelandabout500millionyearsago.
Thecolonizationoflandwasoneofthepivotalmilestonesinthehistoryoflife.
Thereisfossilevidencethatcyanobacteriaandotherphotosyntheticprokaryotescoated
dampterrestrialsurfaceswellmorethanabillionyearsago.
However,macroscopiclifeintheformofplants,fungi,andanimalsdidnotcolonizeland
untilabout500millionyearsago,duringtheearlyPaleozoicera.

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 Thegradualevolutionfromaquatictoterrestrialhabitatswasassociatedwithadaptations
thatallowedorganismstopreventdehydrationandtoreproduceonland.
Forexample,plantsevolvedawaterproofcoatingofwaxontheirphotosyntheticsurfaces
toslowthelossofwater.
Plantscolonizedlandinassociationwithfungi.
Inthemodernworld,therootsofmostplantsareassociatedwithfungithataidinthe
absorptionofwaterandnutrientsfromthesoil.
Thefungiobtainorganicnutrientsfromtheplant.
Thisancientsymbioticassociationisevidentinsomeoftheoldestfossilizedroots.
Plantscreatednewopportunitiesforalllife,includingherbivorous(planteating)animals
andtheirpredators.
Themostwidespreadanddiverseterrestrialanimalsarearthropods(includinginsectsand
spiders)andvertebrates(includingamphibians,reptiles,birds,andmammals).
Terrestrialvertebrates,whichincludehumans,arecalledtetrapodsbecauseoftheirfour
limbs.
Earthscontinentsdriftacrosstheplanetssurfaceongreatplatesofcrust.
Earthscontinentsdriftacrosstheplanetssurfaceongreatplatesofcrustthatfloatonthe
hot,underlyingmantle.
Platesmayslidealongtheboundaryofotherplates,pullingapartorpushingagainsteach
other.
Continentalplatesmoveslowly,buttheircumulativeeffectsaredramatic.
Mountainsandislandsarebuiltatplateboundariesoratweakpointsontheplates.
Platemovementshavehadamajorinfluenceonlife.
About250millionyearsago,neartheendofthePaleozoicera,allthecontinental
landmassescametogetherintoasupercontinentcalledPangaea.
Oceanbasinsdeepened,sealevellowered,andshallowcoastalseasdrained.
Manymarinespecieslivinginshallowwatersweredrivenextinctbythelossof
habitat.
Theinteriorofthesupercontinentwassevere,cold,anddry,leadingtomuchterrestrial
extinction.
DuringtheMesozoicera,180millionyearsago,Pangaeabegantobreakup.
Asthecontinentsdriftedapart,eachbecameaseparateevolutionaryarenawith
lineagesofplantsandanimalsthatdivergedfromthoseonothercontinents.
Continentaldriftexplainsmuchabouttheformerandcurrentdistributionoforganisms.
Australianfloraandfaunacontrastsharplyfromthatoftherestoftheworld.
MarsupialmammalsfillecologicalrolesinAustraliaanalogoustothosefilledby
placentalmammalsonothercontinents.
MarsupialsprobablyevolvedfirstinwhatisnowNorthAmericaandreachedAustralia
viaSouthAmericaandAntarcticawhilethecontinentswerestilljoined.
ThebreakupofthesoutherncontinentssetAustraliaadrift.
InAustralia,marsupialsdiversifiedandthefewearlyeutheriansbecameextinct.
Onothercontinents,marsupialsbecameextinctandeutheriansdiversified.

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Concept26.6Newinformationhasrevisedourunderstandingofthetreeoflife
Inrecentdecades,moleculardatahaveprovidednewinsightsintotheevolutionary
relationshipsoflifesdiverseforms.
Thefirsttaxonomicschemesdividedorganismsintoplantandanimalkingdoms.
In1969,R.H.Whittakerarguedforafivekingdomsystem:Monera,Protista,Plantae,
Fungi,andAnimalia.
Thefivekingdomsystemrecognizedthattherearetwofundamentallydifferenttypesof
cells:prokaryotic(thekingdomMonera)andeukaryotic(theotherfourkingdoms).
Threekingdomsofmulticellulareukaryotesweredistinguishedbynutrition,inpart.
Plantsareautotrophic,makingorganicfoodbyphotosynthesis.
Mostfungiaredecomposerswithextracellulardigestionandabsorptivenutrition.
Mostanimalsingestfoodanddigestitwithinspecializedcavities.
InWhittakerssystem,Protistaincludedalleukaryotesthatdidnotfitthedefinitionof
plants,fungi,oranimals.
Mostprotistsareunicellular.
However,somemulticellularorganisms,suchasseaweeds,wereincludedinProtista
becauseoftheirrelationshipstospecificunicellularprotists.
Thefivekingdomsystemprevailedinbiologyformorethan20years.
Duringthepastthreedecades,systematistsappliedcladisticanalysistotaxonomy,
constructingcladogramsbasedonmoleculardata.
ThesedataledtothethreedomainsystemofBacteria,Archaea,andEukaryaas
superkingdoms.
BacteriadifferfromArchaeainmanykeystructural,biochemical,andphysiological
characteristics.
Manymicrobiologistshavedividedthetwoprokaryoticdomainsintomultiplekingdoms
basedoncladisticanalysisofmoleculardata.
Asecondchallengetothefivekingdomsystemcomesfromsystematistswhoaresorting
outthephylogenyoftheformermembersofthekingdomProtista.
MolecularsystematicsandcladisticshaveshownthattheProtistaisnotmonophyletic.
Someoftheseorganismshavebeensplitamongfiveormorenewkingdoms.
OthershavebeenassignedtothePlantae,Fungi,orAnimalia.
Clearly,taxonomyatthehighestlevelisaworkinprogress.
Therewillbemuchmoreresearchbeforethereisanythingclosetoanewconsensusfor
howthethreedomainsoflifearerelatedandhowmanykingdomsshouldbeincludedineach
domain.
Newdata,includingthediscoveryofnewgroups,willleadtofurthertaxonomic
remodeling.
Keepinmindthatphylogenetictreesandtaxonomicgroupingsarehypothesesthatfitthe
bestavailabledata.

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