Reverse Evolution Driving Forces Behind the Loss of Acquired Photosynthetic Traits 英文参考文献.docVIP

ReverseEvolution:DrivingForcesBehindtheLossof AcquiredPhotosyntheticTraits FranciscodeCastro1*,UrsulaGaedke1,JensBoenigk2 1DepartmentofEcologyandEcosystemModelling,UniversityofPotsdam,Potsdam,Germany,2InstituteforLimnology,AustrianAcademyofSciences,Mondsee,Austria Abstract Background:Thelossofphotosynthesishasoccurredoftenineukaryoticevolution,evenmorethanitsacquisition,which occurredatleastninetimesindependentlyandwhichgeneratedtheevolutionofthesupergroupsArchaeplastida,Rhizaria, Chromalveolata and Excavata. This secondary loss of autotrophic capability is essential to explain the evolution of eukaryotes and the high diversity of protists, which has been severely underestimated until recently. However, the ecologicalandevolutionaryscenariosbehindthisevolutionary‘‘stepback’’arestilllargelyunknown. Methodology/PrincipalFindings:Usingadynamicmodelofheterotrophicandmixotrophicflagellatesandtwotypesof prey,largebacteriaandultramicrobacteria,weexaminetheinfluenceofDOCconcentration,mixotroph’sphotosynthetic growthrate,andexternallimitationsofphotosynthesisonthecoexistenceofbothtypesofflagellates.Ourkeypremisesare: large bacteria grow faster than small ones at high DOC concentrations, and vice versa; and heterotrophic flagellates are moreefficientthanthemixotrophsgrazingsmallbacteria(bothempiricallysupported).Weshowthatdifferentialefficiency inbacteriagrazing,whichstronglydependsoncellsize,isakeyfactortoexplainthelossofphotosynthesisinmixotrophs (which combine photosynthesis and bacterivory) leading to purely heterotrophic lineages. Further, we show in what conditionsanheterotrophmutantcancoexist,orevenout-compete,itsmixotrophicancestor,suggestingthatbacterivory andcellsizereductionmayhavebeenmajortriggersforthediversificationofeukaryotes. Conclusions/Significance:Ourresultssuggestthat,providedthemixotroph’sphotosyntheticadvantageisnottoolarge, the (small) heterotroph will also dominate in nutrient-poor environments and will readily invade a community of mixotrophsandbac