蝴蝶翅膀的进化

Evo-DevoofButterflyWingPatterns2002.11.29高锋演化发育生物学家们关注的一个焦点是：蝴蝶翅膀的颜色模式多样性是怎样通过自然选择形成的以及特定模式的形成过程。演化发育生物学中一个正在蓬勃发展的领域，试图探索在发育过程与自然选择的分选作用下的遗传变异与表型之间的一种联系。蝴蝶翅膀模式成为了一种为数不多的在生物组织不同水平来研究形态学差异的成功例子。1.蝴蝶翅膀的颜色模式无论在种间还是在种内都有着非常壮观的差异。2.蝴蝶翅膀模式是一个完整研究塑造形态的演化和发育过程相互关系的理想系统。3.蝴蝶翅膀模式在发育过程中的可塑性。4.蝴蝶翅平面模式图。5.Wing-pattern突变体及eyespot的形成Figure1|Examplesofmodelbutterflyspecies.Evo–devostudiesofbutterflywingpatternshaveconcentratedonafewmodelspecies,whichprovidedifferentopportunitiesforresearch.a|Heliconiusmelpomemeandb|Papiliodardanusbutterfliesareclassicalexamplesofspeciesinvolvedinpolymorphicanti-predatormimicrysystems.c|Pontiaoccidentalishasbeenusedtostudyphenotypicplasticity,which,inthisspecies,hasaknownecologicalrelevance.d|Preciscoeniahasbeentheclassicalmodelforthedevelopmentaldissectionofwing-patternformation.e|Papiliobutterflies,includingP.glaucus,haveacharacteristicpigmentsystemandhavebeenafavouritemodelforcharacterizingthepigmentbiosynthesispathways.f|Bicyclusanynanahasbeensuccessfullyestablishedinthelaboratoryand,duringthepast12years,hasbecomethebutterfly‘lab-rat’,asitisamenabletocontrolledpopulationandorganism-levelapproachestothestudyofwing-patternvariationanddevelopment.H.melpomenecourtesyofJimMallet,UniversityCollegeLondon,UK;P.dardanuscourtesyofFredNijhout,DukeUniversity,NorthCarolina,USA;P.occidentaliscourtesyofJoelKingsolver,UniversityofNorthCarolina,USA;P.coeniacourtesyofScottWeatherbeeandSeanCarroll,UniversityofWisconsin,Madison,USA;P.glaucuscourtesyofRichardffrench-Constant,UniversityofBath,UK.1.Comparativemethod2.Experimentalcrosses3.Artificialselection4.Surgicalmanipulationsofpupalwings5.Biochemistry6.Physiology7.Gene-expressionpatterns8.Genemapping9.GeneticmanipulationVariousmethodshavebeenusedtoanalysethetwo-dimensionalpatternsonbutterflywings.1.在幼虫的发育过程中，表型可随着外界环境的不同而变化。如，温度、光周期等。2.目前对可塑性的遗传学基础还不清楚，包括是否有可塑性的特定调节基因。3。在一些蝴蝶种类中,脱皮甾酮激素(EC)能调节非遗传多型性。Figure2|The‘nymphalidgroundplan’.Despitethegreatdiversityinbutterflywingpatterns,arecognizableidealized‘groundplan’hasbeenproposedtodescribemostwingpatternvariants6.Accordingtothisgroundplan,thetypesofpatternelement(forexample,eyespots,chevronsandbands)areorganizedinparallelseries:themarginalandsubmarginalbands(whichmightformchevronelements),theborderocellioreyespots,andthecentralandbasalbands.Homologouspatternelementsareseriallyrepeatedinindividualwingcompartmentsthatareborderedbyveins(thickenedwingveinsontheforewingillustratetheboundariesofsuchacompartment,knownasawingcell).Patternelementsinindividualwingcellsarerepeatedalongtheposterior–anterioraxisandcan,toalargeextent,changeindependentlyofeachother.Thefigureshowsaforewing(top)andahindwing(bottom),bothorientatedwiththeanteriorparttothetopanddistalmargintotheright.ReproducedwithpermissionfromREF.33©(2001)Wiley.Figure3|Wing-patternmutants..Allphotosareoftheventralsurfaceoftheforewing(top)andhindwing(bottom)ofB.anynanafemales.a|Wildtypephenotypewithtwoeyespotsontheforewingandsevenonthehindwing,withcharacterisicrelativesizes;b|Bigeyemutantwithoverallenlargedeyespots46;c|Spottymutantwithextraeyespotsontheforewing46;andd|Goldeneyemutant,inwhichthecharacteristicallyblackscalesofthecentralringoftheeyespotsarealmostcompletelyreplacedbygoldscales56.Theindividualshownindis,infact,aBigeye–Goldeneyedoublemutant.Othermutantsnotillustratedhereincludecometandcyclops,whichhavealteredeyespotshape46,andmutant3–4,whichhasfewereyespotsonitshindwing9,38.一些在eyespot形成过程中表达的基因：Apterous:翅膀的背面表达Wingless:在wingmargin中表达Distal-less:在wingmargin中表达Engrailed:翅的后部cubitusinterruptus:翅的前部表达中有重部署的现象：Distal-less,engrailedandspalt：不仅在眼点中心中共同表达，在周边的颜色区域也分别表达。cubitusinterruptus,patchedandhedgehog：在特定的时间表达（在眼点中心或周围）不在眼点中心表达，但是能发挥影响的有：decapentaplegicandwingless：在眼点的定位中起作用Ultrabithorax：参与控制眼点的大小中a|Typicaladulteyespotmorphologywithacentralwhitepupil,ablackringandanoutergoldring.b,c|Expressionpatternof

Distal-lessinthecentresofthepresumptiveeyespotsin