蝴蝶翅膀的進化

Evo-DevoofButterflyWingPatterns2002.11.29高鋒演化發育生物學家們關注的一個焦點是：蝴蝶翅膀的顏色模式多樣性是怎樣通過自然選擇形成的以及特定模式的形成過程。演化發育生物學中一個正在蓬勃發展的領域，試圖探索在發育過程與自然選擇的分選作用下的遺傳變異與表型之間的一種聯系。蝴蝶翅膀模式成為了一種為數不多的在生物組織不同水平來研究形態學差異的成功例子。1.蝴蝶翅膀的顏色模式無論在種間還是在種內都有著非常壯觀的差異。2.蝴蝶翅膀模式是一個完整研究塑造形態的演化和發育過程相互關系的理想系統。3.蝴蝶翅膀模式在發育過程中的可塑性。4.蝴蝶翅平面模式圖。5.Wing-pattern突變體及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.在幼蟲的發育過程中，表型可隨著外界環境的不同而變化。如，溫度、光周期等。2.目前對可塑性的遺傳學基礎還不清楚，包括是否有可塑性的特定調節基因。3。在一些蝴蝶種類中,脫皮甾酮激素(EC)能調節非遺傳多型性。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形成過程中表達的基因：Apterous:翅膀的背面表達Wingless:在wingmargin中表達Distal-less:在wingmargin中表達Engrailed:翅的后部cubitusinterruptus:翅的前部表達中有重部署的現象：Distal-less,engrailedandspalt：不僅在眼點中心中共同表達，在周邊的顏色區域也分別表達。cubitusinterruptus,patchedandhedgehog：在特定的時間表達（在眼點中心或周圍）不在眼點中心表達，但是能發揮影響的有：decapentaplegicandwingless：在眼點的定位中起作用Ultrabithorax：參與控制眼點的大小中a|Typicaladulteyespotmorphologywithacentralwhitepupil,ablackringandanoutergoldring.b,c|Expressionpatternof

Distal-lessinthecentresofthepresumptiveeyespotsin