Generation of Diverse Biological Forms through Combinatorial Interactions between Tissue Polarity and Growth 英文参考文献.docVIP

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Generation of Diverse Biological Forms through Combinatorial Interactions between Tissue Polarity and Growth 英文参考文献.doc

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Generation of Diverse Biological Forms through Combinatorial Interactions between Tissue Polarity and Growth 英文参考文献

GenerationofDiverseBiologicalFormsthrough CombinatorialInteractionsbetweenTissuePolarityand Growth RichardKennaway1,EnricoCoen2,AmeliaGreen2,AndrewBangham1* 1SchoolofComputingSciences,UniversityofEastAnglia,Norwich,UnitedKingdom,2JohnInnesCentre,Norwich,UnitedKingdom Abstract A major problem in biology is to understand how complex tissue shapes may arise through growth. In many cases this process involves preferential growth along particular orientations raising the question of how these orientations are specified. One view is that orientations are specified through stresses in the tissue (axiality-based system). Another possibility is that orientations can be specified independently of stresses through molecular signalling (polarity-based system). The axiality-based system has recently been explored through computational modelling. Here we develop and applyapolarity-basedsystemwhichwecalltheGrowingPolarisedTissue(GPT)framework.Tissueistreatedasacontinuous materialwithinwhichregionallyexpressedfactorsundergeneticcontrolmayinteractandpropagate.Polarityisestablished bysignalsthatpropagatethroughthetissueandisanchoredinregionstermedtissuepolarityorganisersthatarealsounder geneticcontrol.Ratesofgrowthparallelorperpendiculartothelocalpolaritymaythenbespecifiedthrougharegulatory network.Theresultinggrowthdependsonhowspecifiedgrowthpatternsinteractwithintheconstraintsofmechanically connectedtissue.Thisconstraintleadstotheemergenceoffeaturessuchascurvaturethatwerenotdirectlyspecifiedby the regulatory networks. Resultant growth feeds back to influence spatial arrangements and local orientations of tissue, allowingcomplexshapestoemergefromsimplerules.Moreover,asymmetriesmayemergethroughinteractionsbetween polarityfields.WeillustratethevalueoftheGPT-frameworkforunderstandingmorphogenesisbyapplyingittoagrowing Snapdragonflowerandindicatehowtheunderlyinghypothesesmaybetestedbycomputationalsimulation.Wepropose thatcombinatorialintractionsbetweenorientationsandratesofgrowth,whicha