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1、GrowthtradeoffsproducecomplexmicrobialcommunitiesonasinglelimitingresourceMichaelManhart∗andEugeneI.ShakhnovichDepartmentofChemistryandChemicalBiology,HarvardUniversity,Cambridge,MA02138,USA(Dated:June1,2018)Therelationshipbetweenthedynamicsofacommunityanditsconstituentpairwiseinteractions
2、isafundamentalprobleminecology.Higher-orderecologicaleffectsbeyondpairwiseinteractionsmaybekeytocomplexecosystems,butmechanismstoproducetheseeffectsremainpoorlyunderstood.Hereweshowthathigher-ordereffectscanarisefromvariationinmultiplemicrobialgrowthtraits,suchaslagtimesandgrowthrates,onasin
3、glelimitingresourcewithnootherinteractions.Theseeffectsproducearangeofecologicalphenomena:anunlimitednumberofstrainscanexhibitmul-tistabilityandneutralcoexistence,potentiallywithasinglekeystonestrain;strainsthatcoexistinpairsdonotcoexistalltogether;andthechampionofallpairwisecompetitionsma
4、ynotdominateinamixedcommunity.Sincevariationinmultiplegrowthtraitsisubiquitousinmicrobialpopulationsduetopleiotropyandnon-geneticvariation,ourresultsindicatethesehigher-ordereffectsmayalsobewidespread,especiallyinlaboratoryecologyandevolutionexperiments.Keywords:microbialgrowth;microbialec
5、ology;higher-ordereffects;coexistence;non-transitiveselectionComplexcommunitiesofmanydistinctspeciesorbutalsonon-transitiveselection[21],inwhichpairwisestrainsaboundinbothnature[1,2]andthelabora-competitionsofstrainscanexhibit“rock-paper-scissors”tory[3,4].Oneofthemostfundamentalproblemsin
6、behaviors[22].ecologyistounderstandtherelationshipbetweenacom-Variationinmultiplegrowthtraits,suchaslagtime,munity’sbehaviorandthepairwiseinteractionsofitsexponentialgrowthrate,andyield(resourceefficiency),constituents[5–8].Inparticular,thekeyquestionisispervasiveinmicrobialpopulations[23–2
7、5].Notonlywhetherthesepairwiseinteractionslargelydeterminethearesinglemutationsknowntobepleiotropicwithrespectbehaviorofthecommunityasawhole.However,ecol-tothesetraits[26,27],butevengenetically-identicallin-ogistshavelongconsideredthepossibilityof“higher-eagesmaydem