w2 by jenny

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lecture2  lagerstatten =motherlode; excellentfoss ilpreserva tionatthesite  systemana turaebycarolusLinn aeus(Ki ngPhilipCame OverForGoodSe x) o  impliesan order,butdoe sn’ttellyouhowt hesespecies(spp)are related  hennigphyloge neticsyste matics“clad istics”  cladistics amethodofclassifyinganimalsa ndplantsac cordingto the proportion ofmeasurablecharact eristicsth eyhaveinco mmon  assumptionsofcladistics: o  iforganismsarerelatedbydescent,thecloserdescent=the morerecent commonancestry o  becauseorganismschange throughevolutionary time,theypa ssontheir changestotheiroffspring o  therefore,organ ismsthatare mostcloselyrelat edshouldsharethe most derived characters,a cquiredbye volution,thatare notpresent inother groups o  thesearchforthesesetsoftraitsshouldgiveus thebest approximationofthe evolutionarysequence(aphylogeny)  branchlengthusual lyrefers tomoleculardata suchthatlongerbra nches=longe r timeperiods  clade=agroupt hatincludesMR CAandallco mmondescendents  parsimony=use thefewe ststepsnecessarytoconstructalegitimatetree make it assimpleaspossible o  lookforthesimp lestpossible explana tionsoyouonlyhave afew traits uniquetoataxa o  thisisahypothesisaboutevolution  crown-baseddefinitionunifyyourclad esoitincorporateseve rythingdescen ded fromtheMRCA  trait-base ddefinitionincludeeve ryanimaltha thadfur,oranop posablethumb o  worksforexta nttaxa,butnotalwaysfore xtincttaxa  monophyleticgroupincludesthe MRCAa ndallitsde scendants  paRaphyleticgroupincludestheM RCA,butdoes n’tincludeallofi tsdesce ndants  pOlyphyleticgroupn OMRCAoralldescendents;2mono-orpara-phyleticclades incorrectlygroupedtogether  systematicsthescientificstudyofthedivers ityoforganisms,which reveals the evolutionaryre lationshipsbe tweenorganis ms  taxonomy(as ubdivisionofsy stematics) thetheoryandp racticeofclas sifying organisms  rejectingthe sppapproa chsomepplsu ggesttrashinga llnamesand usingaunique numbertoIDe verythingonea rthorper hapsusingcytochrome 1todefi neaspp  phylogenyahypothesisthat describesthehistoryofdescent ofagroupof organismsfromt heircommonanc estor  bifurcatingtree aline ageisre presented asabranchingtre e,inwhiche achsplitor noderepresentsaspeciationevent  o  branchinginto2,notmany o  doesn’tne cessarilyhapp enthatway ,butit’sthemostpa rsimonious explanation

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8/7/2019 W2 by Jenny

http://slidepdf.com/reader/full/w2-by-jenny 1/4

lecture2

•  lagerstatten=motherlode;excellentfossilpreservationatthesite

•  systemanaturaebycarolusLinnaeus(KingPhilipCameOverForGoodSex)

o  impliesanorder,butdoesn’ttellyouhowthesespecies(spp)arerelated•  hennig–phylogeneticsystematics–“cladistics”

•  cladistics–amethodofclassifyinganimalsandplantsaccordingtotheproportionofmeasurablecharacteristicstheyhaveincommon

•  assumptionsofcladistics:o  iforganismsarerelatedbydescent,thecloserdescent=themorerecent

commonancestry

o  becauseorganismschangethroughevolutionarytime,theypassontheir

changestotheiroffspring

o  therefore,organismsthataremostcloselyrelatedshouldsharethemost

derivedcharacters,acquiredbyevolution,thatarenotpresentinother

groupso

 thesearchforthesesetsoftraitsshouldgiveusthebestapproximationofthe

evolutionarysequence(aphylogeny)•  branchlengthusuallyreferstomoleculardatasuchthatlongerbranches=longer

timeperiods

•  clade=agroupthatincludesMRCAandallcommondescendents•  parsimony=usethefeweststepsnecessarytoconstructalegitimatetree–makeit

assimpleaspossible

o  lookforthesimplestpossibleexplanationsoyouonlyhaveafewtraitsuniquetoataxa

o  thisisahypothesisaboutevolution

•  crown-baseddefinition–unifyyourcladesoitincorporateseverythingdescended

fromtheMRCA

•  trait-baseddefinition–includeeveryanimalthathadfur,oranopposablethumbo  worksforextanttaxa,butnotalwaysforextincttaxa

•  monophyleticgroup–includestheMRCAandallitsdescendants

•  paRaphyleticgroup–includestheMRCA,butdoesn’tincludeallofitsdescendants

•  pOlyphyleticgroup–nOMRCAoralldescendents;2mono-orpara-phyleticclades

incorrectlygroupedtogether

•  systematics–thescientificstudyofthediversityoforganisms,whichrevealsthe

evolutionaryrelationshipsbetweenorganisms

•  taxonomy–(asubdivisionofsystematics)–thetheoryandpracticeofclassifyingorganisms

•  rejectingthesppapproach–somepplsuggesttrashingallnamesandusingaunique

numbertoIDeverythingonearth–orperhapsusingcytochrome1todefineaspp•  phylogeny–ahypothesisthatdescribesthehistoryofdescent ofagroupoforganismsfromtheircommonancestor

•  bifurcatingtree–alineageisrepresentedasabranchingtree,inwhicheachsplitor

noderepresentsaspeciationevent 

o  branchinginto2,notmany

o  doesn’tnecessarilyhappenthatway,butit’sthemostparsimonious

explanation

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•  starburstphylogenies

•  phylogenetictreesareconstructedbyanalyzingevolutionarychangesinthetraitsoforganisms

•  cladogenesis;node;stem;crowngroup•  ancestraltraits–inheritedfromacommonancestor(hair,milkformarsupials,

monotremesandeutheria)•  derivedtraits–atraitthatdiffersfromitsancestralform(placentaforeutheria

(non-marsupial/monotrememammals),hairandmilkformammaliaasagroup)

•  homologous–anyfeatures(DNAsequences,behavioral,anatomical)sharedby2+

sppthatbothinheritedthetraitfromacommonancestor

o  thesimilarityisduetotheirsharedancestry

o  thevertebralcolumninvertebrates

o  bonesinbird/batwings

•  analogous–performingasimilarfunctionbuthavingadifferentevolutionaryorigin

(wings/flightinpterosaurs,batsandbirds)•

 convergentevolution–whenindependentlyevolvedfeaturessubjectedtosimilar

selectivepressuresbecomesuperficiallysimilaro  convergenttraitsareindependentlyderived

•  evolutionaryreversal–whenacharacterrevertsfromaderivedstatebacktoan

ancestralstate•  homoplasies–traitsthataresimilarforsomereasonotherthaninheritancefroma

commonancestor

o  lossofeyesincaecilians=evolutionaryreversalandahomoplasy(they’reamphibiansthatlooklikeeyelesssnakes)

•  outgroup–alineagethatiscloselyrelatedtoaningroup(thelineageofinterest)buthasbranchedofffromtheingroupbelowitsbaseontheevolutionarytree

o  ancestraltraitsshouldbefoundintheoutgroupandtheingroup;derived

traitsarefoundonlyintheingroup•  toreconstructaphylogeny:

o  1.selectagroupoforganismstoclassify(theingroup)andanappropriate

outgroup

o  2.choosethecharactersthatwillbeusedintheanalysisandidentifythe

possibleforms(traits)ofthecharacter

o  determinetheancestralandderivedtraits

o  distinguishhomologousfromhomoplastictraits

•  systematistsusephysiological,behavioral,molecular,andstructuralcharactersoflivingandfossilorganismstoreconstructphylogenies

•  richness=numberofdifferenttaxa(alpha,betadiversity,etc)•  diversity=numberofdifferenttaxaandtheirrelativeabundances

evolutionarythoughtlecture

•  homology–anysimilaritybetweencharactersthatareduetotheirsharedancestry

•  homoplasy–whencharactersaresimilar,butnotderivedfromacommonancestor

•  synapomorphy–charactersthataresharedbyagroupoforganisms,butarenotfoundintheancestorsofthatgroup

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•  parsimony–adoptionofthesimplestassumptionintheformulationofatheoryor

intheinterpretationofdata

•  Lamarckianevolution–sppcouldchangethroughtheuseordisuseofstructures

duringtheindividualorganism’slifetimeo  theenvironmentpresentsconditionswhichproduce“needs”inorganisms

o  needsleadtomodificationoforgansandbehaviorso  usedpartsincreaseinsize,disusedpartsdecreaseinsize

o  modificationoccursduringthelifespanoftheorganismandarepassedonto

theorganism’soffspring

•  uniformatarianism–thesamenaturallawsandprocessesthatoperateintheuniversenow,havealwaysoperatedintheuniverseinthepastandapply

everywhereintheuniverse

•  Malthus–humansreproduceatahigherratethantheirresourcescansustainthem;

theonlythingkeepingthemincheckisdeathanddisease(competition)

•  darwin’sargumentfornaturalselectiono

 1.organismshaveagreatpotentialfertility,permittingexponentialgrowthof

popns(Malthus)o  2.naturalpopnsdon’tnormallyincreaseexponentially,butremainfairly

constantinsize(Darwin)

o  3.naturalresourcesarelimited(Malthus)o  4.astruggleforexistenceoccursamongorganismsinapopulation

(competition,Malthus)

o  5.variationoccursamongorganismswithinpopns(darwin’spigeons,barnacles)

o  6.variationisheritable(animalbreeding)o  7.varyingorganismsshowdifferentialsurvivalandreproduction,favoring

advantageoustraits=naturalselection(Darwin)

o  8.naturalselection,actingovermanygenerations,graduallyproducesnewadaptationsandnewspecies

•  Wallace

•  thejointletter–hookerandlyell

•  themodernsynthesis–Mendel;THmorgan–genesonchromosomes(theflylab);

mutationssupplytherawmatlfornaturalselectiontoactupon

o  JBSHaldane,SewallWright,andRonaldFisher

o  mathematicscanexplainhowselectionandmutationcanaffectgene

frequencieswithinpopulationso  populationgeneticsyieldsagenotypeexpressedinthephenotype

o  populationsaregenepoolswithvariationingeneticfrequencieso  geneticvariationisaresultofrandomsubstitutionsofbasepairsinthe

genotypebymutation,andinsexualorganisms,thevariationcanalsobe

introducedwithrecombinationo  thevariationingenotyperesultsinavariationinthephenotype,whichis

thenactedonbynaturalselection

o  naturalselectioncullsthe‘lessfit’oftherandomvariants

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o  thus,evolutionhasno‘direction,’excepttowardsincreasingfitness,whichin

anextremeformmeansthateveryfeatureinanorganismishonedbynaturalselection(panselectionism)

ReadingsforWeekTwoNote:Ioutlinedthepapersinthe1,2,3formMattsuggested,where1)isthequestionposed

inthepaper,2)isthemethodology,and3)isthemainconclusion.Hippopotamusandwhalephylogeny,Thewissenetal.2007

1.  Arehipposandcetacea(whales,dolphins,porpoises)placedcorrectlyontheirmammalianphylogeny?

2.  GeislerandTheodorperformedanewanalysisusingweightedcharacters

(homoplasiesweredown-weighted)tocreatewhattheythoughtwasthemostparsimonioustree.Theyemphasizetheimportanceofincludingmoleculardataas

well.

3.  Theauthorsofthebriefcommentary(GeislerandTheodor)thinkhippposarethemostcloselyrelatedextanttaxontoCetacea.Theauthorsoftheinitialpaper

(Thewissenetal.)respondedbysayingthatGeislerandTheodorupheldtheirmainfinding,whichwasthatraoellids(nowextinct)arethesistergrouptocetaceans.

“Herbivorousecomorphologyandspecializationpatternsintheropoddinosaurevolution,”

ZannoandMakovicky20111.  Howdoweusemorphologicalfactorstodetermineifsomethingwascarnivorousor

herbivorous?

2.  TheauthorsidentifiedlotsofcharactersreliablefortheindicationofherbivoryusingthreestatisticalanalysesandthentestedthesetraitsinCoelurosauria.

3.  Usingstats,youcanhavesomefairconfidenceaboutthefeedingbehaviorsoflong-extinctanimals.WithinCoelurosauria,thereisevidenceofincreasingspecialization

toherbivoryinsomeclades,indicatingthatthecoelurosaurianbeakwaslikely

correlatedwithanherbivorousdiet.Thisisanexampleofecomorphology,whichisthestudyoftherelationshipbetweentheecologicalroleofanindividualandits

morphologicaladaptations(structureandfunction).

Evolutionofanatomyandgenecontrol,Koentges20081.  Shouldtherebemorecommunicationbetweenthesubdisciplinesofbiology?

2.  Koentgesdidadetailedliteratureviewandprovidedalotofhisownanalysisandsuggestionsforpotentialcollaborations.

3.  Weneedtointegratedifferentdisciplines(e.g.,paleontology,evodevo,taxonomy,

genomics,molecularbio,etc.)toanswerthebigquestionsingenomicsystemsbiology.Wealsoneedalargecentralizeddatabasetointegratedataacross

subdisciplines.

StudyingFunctionandBehaviorintheFossilRecord,Benton20101.  Howdopaleobiologistsuseformtostudyfunction?

2.  Bentondiscusses/analyzesthethreemaintechniquesusedtoinferfunctionandbehaviorfromfossils:empiricalevidence,comparisonwithmodernanalogs,and

biomechanicalmodeling.Hedoesn’treallycritiquethesemethods,though.

3.  Usingthesethreemaintechniques,paleobiologistscamakeevidence-basedreconstructionsandreallyinferfunctionfromform.