發(fā)育與進化（Evolutionarydevelopmentalbiology，常簡稱Evo-Devo）王文中德MaxPlanck青年科學家小組遺傳資源與進化國家重點實驗室中國科學院昆明動物研究所發(fā)育與進化內(nèi)容：1、進化發(fā)育生物學簡介2、我們的思考和研究做科學要有好的科學問題：科學體系發(fā)展中不可回避、必須回答的問題可以操作的問題TheArtoffeasible!內(nèi)容：做科學要有好的科學問題：TheArtoffeas背景：遺傳、發(fā)育和進化是生命科學中的本質(zhì)問題，我們看到的各種各樣的生物組織、器官和性狀無不是千百萬年來遺傳變異經(jīng)自然選擇固定并最后通過個體發(fā)育展現(xiàn)而來。長期以來科學家們一直在試圖把這三方面聯(lián)系起來。隨著分子生物學、進化基因組學和發(fā)育生物學等現(xiàn)代生命科學和技術的迅速發(fā)展和結合，這種努力首次變得富有希望。20多年前興起的進化發(fā)育生物學（Evolutionarydevelopmentalbiology，常簡稱Evo-Devo）是這種希望的一個開端，該學科試圖從基因表達在發(fā)育過程中的變化理解不同物種形態(tài)發(fā)生和功能的進化(Hall，1992）。背景：遺傳、發(fā)育和進化是生命科學中的本質(zhì)問題，我們看到的各種Haeckel‘stheory:

Ontogenyrecapitulatesphylogeny（1874）thefetaldevelopmentofanindividual(ontogeny)isaspeeded-upreplayofmillionsofyearsofspeciesevolution(phylogeny)Haeckel‘stheory:

OntogenyrecThomasHuntMorgan,“EmbryologyandGenetics”(1934)第一次提出了胚胎發(fā)育的可能細胞分化機制：基因表達的時空分化在過去的30年中，我們已經(jīng)發(fā)現(xiàn)了各種動物均使用的發(fā)育調(diào)控基因基本的細胞過程，形成了“進化發(fā)育生物學”這一分支學科。進化發(fā)育生物學（evolutionarydevelopmentalbiology）：旨在闡明生物進化中發(fā)育過程和機制怎樣改變以及這些改變?nèi)绾卧炀瓦^去和現(xiàn)在的生物多樣性的一門學科。（BagunaandGarcia-Fernandez,2003).物種之間發(fā)育基因的比較（SeanCarrol，2005） 常簡稱“EvoDevo”ThomasHuntMorgan,“Embryolog"Evo-devo"isoneofthenewestareasofbiology.Itseekstoexplainevolutionaryeventsthroughthemechanismsofdevelopmentalbiology.Itsmajorquestionsareoftenthequestionsofthelate1800swhichhadbeenleftwaitingforothergenerationsofbiologiststounravel.Wearethosegenerations.Thesequestionsinclude:Howdidwingsevolve?Howdidlimbsevolve?Howdidvertebratesarisefrominvertebrates?Howdidvertebratesformtheirhead?TheModernSynthesisofthe1930sand1940swasthesynthesisofevolutionarybiologyandpopulationgenetics.Itisoneofthegreatestexplanatorysystemsinscience,anditexplainsincrediblywelltheorigin,maintenance,andevolutionofvariationswithinpopulationsofspecies.However,itdidnotidentifythegenesinvolvedinthesechanges,nordiditexplaintheoriginofhighertaxa.Evolutionarydevelopmentalbiologyseeksto“complete”thesynthesisbyaddingdevelopmentalgeneticstopopulationgenetics.TraditionalEvo-devolooksattheregulatoryregionsofgenesratherthantheircodingsequences.Itemphasizesthearrivalofthefittestratherthanthesurvivalofthefittest."Evo-devo"isoneofthenewesPolymorphismofgenetoyandsphinxinaworldwideandalocalDrosophilamelanogasterpopulationWang,W.,Thornton,K.,Berry,A.andLong,M.2002,Science,295:134-137.群體遺傳學案例（基因在群體中的變異情況）PolymorphismofgenetoyandsNocrossingoverSubjecttorelentlessselectivesweepandbackgroundselectioninpopulationsDrosophilaChromosomeNocrossingoverDrosophilaVariationsin18generegionswereinvestigatedWang,W.,Thornton,K.,Berry,A.andLong,M.2002,Science,295:134-137.Variationsin18generegions(1)Phylogeny(系統(tǒng)發(fā)育)Evo-Devo中的重要概念:

(1)Phylogeny(系統(tǒng)發(fā)育)Evo-Devo中進化和發(fā)育-生物化學與細胞生物學研究所課件Evo-Devo中的重要概念:

(2)Bodyplan胚胎前后軸（A－P）、背腹軸（D－V）、左右軸（L－R）等決定Evo-Devo中的重要概念:

(2)BodyplanDeRobertis,2008)DeRobertis,2008)BodyplanBodyEvo-Devo中的重要概念:(3)Toolkitgenes:mastergenesthatgoverntheformationandpatterningoftheirBodiesandbodypartsofallkindsofanimals.Despitebeingmorphologicallyverydiverse,multicellularorganismsaremadebyaveryconservedsetofregulatorygenes(toolkitgenes)playingcomparabledevelopmentalroles.Thismostunexpectedfindingrepresentedapowerfulmolecularproofofevolutionas‘descentwithmodification’(Darwin,1859).最近：Asetofgenesthatisrequiredfordevelopmentandissharedwidelyamongspecies(Canestroetal,NRG,2007)Evo-Devo中的重要概念:傳統(tǒng)的發(fā)育進化生物學研究范式：經(jīng)典的進化發(fā)育生物學(EvoDevo)認為所有動物的發(fā)育由同一套工具箱基因（toolkitgenes）控制傳統(tǒng)的發(fā)育進化生物學研究范式：經(jīng)典的進化發(fā)育生物學(Evo果蠅蜈蚣在果蠅，ubx抑制腹部dsl基因的表達的足產(chǎn)生，但在蜈蚣中其的C端磷酸化位點不存在，導致不抑制腹部的足產(chǎn)生不同動物眼睛的發(fā)育控制由一套同源的基因（Pax6）控制果蠅蜈蚣在果蠅，ubx抑制腹部dsl基因的表達的足產(chǎn)生，但在Nonetheless,itbringsabigparadox:ifdevelopmentalgenesarethesame,howaredifferencesindevelopmentandinthefinalmorphologyindifferentorganismstobeaccountedfor?Nonetheless,itbringsabigp(Canesttroetal.,NRG,2007)(Canesttroetal.,NRG,2007)(Canesttroetal.,NRG,2007)(Canesttroetal.,NRG,2007)最近，SeanCarroll等認為發(fā)育進化主要由調(diào)控序列的改變導致最近，SeanCarroll等認為發(fā)育進化主要由調(diào)控序列的LossofanEnhancerinSticklebackFishEvolutionleadstodramaticmorphologicaldifference

LossofanEnhancerinStickleButterflyWingPatternEvolutionIsAssociatedwithChangesinaNotch/Distal-lessTemporalPatternCurrentbiology2004,13:1159-1166ButterflyWingPatternEvoluti進化和發(fā)育-生物化學與細胞生物學研究所課件進化和發(fā)育-生物化學與細胞生物學研究所課件(4)Homologyvs.convergent,parallelandhomoplasmySortingoutparallelismfromhomologyinlandmarkissuessuchastheorigin(s)ofsegmentation,theorigin(s)oftheeye(Hodin,2000)warrantsfurtherstudiesofadditionalphyla,andadeepunderstandingofplasticityandconstraintsingeneanddevelopmentalevolution.Whenastructurewhichishomologousbetweencloselyrelatedorganismsisbuiltusingdifferentgenes,itshouldbeconsideredconvergentratherthanhomologous.Homologousgenesmayberesponsiblefornon-homologousmorphologies.Genesandpathwaysareutilizedonmanyseparateoccasionsduringdevelopment.Asexamples,theNotch-signallingpathwayisbroadlyusedduringDrosophilaandvertebratedevelopment(Simpson,1997;Robey,1997),andhedgehog,TGF-βandWntfamilymembersareusedoverandoveragainduringdevelopment.Evolutionmaywellhaveworkedby“genetictinkering”(Jacob,1977)or“bricolage”ofgenenetworks(DubouleandWilkins,1998).paralogues,orthologuesParalogues:源自基因重復；共同祖先是基因。Orthologues：源自祖先物種，共同祖先是物種。(4)Homologyvs.convergent,pMacroevolutionvs.microevolutionAsDobzhansky(1937)firstlypointedout,themainissueinthemacrovs.microevolutiondebateiswhethermutationsresultinginrealevolutionarynoveltiesareofthesamekindasthoseoccurringdailyorwhetherweshouldexpectspecial,raremutationsonlyoccurringongeologicaltimescales.Macroevolutionvs.microevolutEvoDevo使用的主要方法1、比較基因組學2、各種生物胚胎發(fā)育中基因的表達圖式3、同源基因的替換或轉基因EvoDevo使用的主要方法1、比較基因組學Thefuturetaskformacro-Evo-Devowillbetounravel:i)preexistingdevelopmentalpotential;inotherwords,whatwasbeforetheCambrian,whichmeansanalyzingthedevelopmentaltoolkitcomponentoftheclosestsistergroups,relativesofeubilaterians,theacoelomorphflatworms(Ruiz-Trilloetal.,1999;2002);ii)theextentandqualityofbricolageofthisbasalbilateriantoolkitcomparedtohigherbilaterians;andiii)tryingtolinktheknown,andthosewhichremaintobediscovered,fossilgroupsOngoingdebatesorunsolvedquestionsinevolutionarydevelopmentalbiology

OriginofBilateria

OriginofVertebrata

OriginofBirds

O