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1、Vol466j26August2010jdoi:10.1038/nature09205ANALYSISTheevolutionofeusociality112MartinA.Nowak,CorinaE.Tarnita&EdwardO.WilsonEusociality,inwhichsomeindividualsreducetheirownlifetimereproductivepotentialtoraisetheoffspringofothers,underliesthemostadvancedformsofsocia
2、lorganizationandtheecologicallydominantroleofsocialinsectsandhumans.Forthepastfourdecadeskinselectiontheory,basedontheconceptofinclusivefitness,hasbeenthemajortheoreticalattempttoexplaintheevolutionofeusociality.Hereweshowthelimitationsofthisapproach.Wearguethatst
3、andardnaturalselectiontheoryinthecontextofprecisemodelsofpopulationstructurerepresentsasimplerandsuperiorapproach,allowstheevaluationofmultiplecompetinghypotheses,andprovidesanexactframeworkforinterpretingempiricalobservations.ormostofthepasthalfcentury,muchofsoci
4、obiologicalgreaterthantwotimesthecosttothealtruist(R51/2)oreighttimestheoryhasfocusedonthephenomenoncalledeusociality,inthecaseofafirstcousin(R51/8).whereadultmembersaredividedintoreproductiveand(par-Duetoitsoriginalityandseemingexplanatorypower,kinselectionFtiall
5、y)non-reproductivecastesandthelattercareforthecametobewidelyacceptedasacornerstoneofsociobiologicaltheory.young.HowcangeneticallyprescribedselflessbehaviourarisebyYetitwasnottheconceptitselfinitsabstractformthatfirstearnednaturalselection,whichisseeminglyitsantith
6、esis?Thisproblemfavour,buttheconsequencesuggestedbyHamiltonthatcametohasvexedbiologistssinceDarwin,whoinTheOriginofSpeciesbecalledthe‘‘haplodiploidhypothesis.’’Haplodiploidyisthesex-declaredtheparadox—inparticulardisplayedbyants—tobethedeterminingmechanisminwhichf
7、ertilizedeggsbecomefemales,andmostimportantchallengetohistheory.Thesolutionofferedbytheunfertilizedeggsmales.Asaresult,sistersaremorecloselyrelatedtomasternaturalistwastoregardthesterileworkercasteasa‘‘well-oneanother(R53/4)thandaughtersaretotheirmothers(R51/2).fl
8、avouredvegetable’’,andthequeenastheplantthatproducedit.HaplodiploidyhappenstobethemethodofsexdeterminationintheThus,hesaid,thewholecolonyistheunitofsele
