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1、Vol466j26August2010jdoi:10.1038/nature09205ANALYSISTheevolutionofeusociality112MartinA.Nowak,CorinaE.Tarnita&EdwardO.WilsonEusociality,inwhichsomeindividualsreducetheirownlifetimereproductivepotentialtoraisetheoffspringofothers,underliesthemostadvancedformsofsocial
2、organizationandtheecologicallydominantroleofsocialinsectsandhumans.Forthepastfourdecadeskinselectiontheory,basedontheconceptofinclusivefitness,hasbeenthemajortheoreticalattempttoexplaintheevolutionofeusociality.Hereweshowthelimitationsofthisapproach.Wearguethatstan
3、dardnaturalselectiontheoryinthecontextofprecisemodelsofpopulationstructurerepresentsasimplerandsuperiorapproach,allowstheevaluationofmultiplecompetinghypotheses,andprovidesanexactframeworkforinterpretingempiricalobservations.ormostofthepasthalfcentury,muchofsociobi
4、ologicalgreaterthantwotimesthecosttothealtruist(R51/2)oreighttimestheoryhasfocusedonthephenomenoncalledeusociality,inthecaseofafirstcousin(R51/8).whereadultmembersaredividedintoreproductiveand(par-Duetoitsoriginalityandseemingexplanatorypower,kinselectionFtially)no
5、n-reproductivecastesandthelattercareforthecametobewidelyacceptedasacornerstoneofsociobiologicaltheory.young.HowcangeneticallyprescribedselflessbehaviourarisebyYetitwasnottheconceptitselfinitsabstractformthatfirstearnednaturalselection,whichisseeminglyitsantithesis?
6、Thisproblemfavour,buttheconsequencesuggestedbyHamiltonthatcametohasvexedbiologistssinceDarwin,whoinTheOriginofSpeciesbecalledthe‘‘haplodiploidhypothesis.’’Haplodiploidyisthesex-declaredtheparadox—inparticulardisplayedbyants—tobethedeterminingmechanisminwhichfertili
7、zedeggsbecomefemales,andmostimportantchallengetohistheory.Thesolutionofferedbytheunfertilizedeggsmales.Asaresult,sistersaremorecloselyrelatedtomasternaturalistwastoregardthesterileworkercasteasa‘‘well-oneanother(R53/4)thandaughtersaretotheirmothers(R51/2).flavoured
8、vegetable’’,andthequeenastheplantthatproducedit.HaplodiploidyhappenstobethemethodofsexdeterminationintheThus,hesaid,thewholecolonyistheunitofsele