Evolution of Networks for Body Plan Patterning; Interplay of Modularity, Robustness and Evolvability 英文参考文献.docVIP

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Evolution of Networks for Body Plan Patterning; Interplay of Modularity, Robustness and Evolvability 英文参考文献.doc

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Evolution of Networks for Body Plan Patterning; Interplay of Modularity, Robustness and Evolvability 英文参考文献

EvolutionofNetworksforBodyPlanPatterning; InterplayofModularity,RobustnessandEvolvability KirstenH.tenTusscher1,2*,PaulienHogeweg1 1TheoreticalBiologyandBioinformaticsGroup,DepartmentofBiology,FacultyofScience,UtrechtUniversity,Utrecht,TheNetherlands,2ScientificComputing,Simula ResearchLaboratory,Oslo,Norway Abstract Amajorgoalofevolutionarydevelopmentalbiology(evo-devo)istounderstandhowmulticellularbodyplansofincreasing complexity have evolved, and how the corresponding developmental programs are genetically encoded. It has been repeatedly argued that key to the evolution of increased body plan complexity is the modularity of the underlying developmental gene regulatory networks (GRNs). This modularity is considered essential for network robustness and evolvability. In our opinion, these ideas, appealing as they may sound, have not been sufficiently tested. Here we use computer simulations to study the evolution of GRNs’ underlying body plan patterning. We select for body plan segmentationanddifferentiation,astheseareconsideredtobemajorinnovationsinmetazoanevolution.Toallowmodular networks to evolve, we independently select for segmentation and differentiation. We study both the occurrence and relationofrobustness,evolvabilityandmodularityofevolvednetworks.Interestingly,weobservedtwodistinctevolutionary strategies to evolve a segmented, differentiated body plan. In the first strategy, first segments and then differentiation domains evolve (SF strategy). In the second scenario segments and domains evolve simultaneously (SS strategy). We demonstratethatunderindirectselectionforrobustnesstheSFstrategybecomesdominant.Inaddition,asabyproductof this larger robustness, the SF strategy is also more evolvable. Finally, using a combined functional and architectural approach, we determine network modularity. We find that while SS networks generate segments and domains in an integratedmanner,SFnetworksuselargelyindependentmodulestoproducesegmentsanddomains.Surprisingly,wefin