Mutual Repression Enhances the Steepness and Precision of Gene Expression Boundaries 英文参考文献.docVIP

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Mutual Repression Enhances the Steepness and Precision of Gene Expression Boundaries 英文参考文献.doc

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Mutual Repression Enhances the Steepness and Precision of Gene Expression Boundaries 英文参考文献

MutualRepressionEnhancestheSteepnessand PrecisionofGeneExpressionBoundaries ThomasR.Sokolowski1,ThorstenErdmann2,PieterReintenWolde1* 1FOMInstituteAMOLF,Amsterdam,TheNetherlands,2UniversityofHeidelberg,InstituteforTheoreticalPhysics,Heidelberg,Germany Abstract Embryonicdevelopmentisdrivenbyspatialpatternsofgeneexpressionthatdeterminethefateofeachcellintheembryo. While gene expression is often highly erratic, embryonic development is usually exceedingly precise. In particular, gene expression boundaries are robust not only against intra-embryonic fluctuations such as noise in gene expression and protein diffusion, but also against embryo-to-embryo variations in the morphogen gradients, which provide positional information to the differentiating cells. How development is robust against intra- and inter-embryonic variations is not understood.Acommonmotifinthegeneregulationnetworksthatcontrolembryonicdevelopmentismutualrepression betweenpairsofgenes.Toassesstheroleofmutualrepressionintherobustformationofgeneexpressionpatterns,we haveperformedlarge-scalestochasticsimulationsofaminimalmodeloftwomutuallyrepressinggapgenesinDrosophila, hunchback(hb)andknirps(kni).Ourmodelincludesnotonlymutualrepressionbetweenhbandkni,butalsothestochastic andcooperativeactivationofhbbytheanteriormorphogenBicoid(Bcd)andofknibytheposteriormorphogenCaudal (Cad),aswellasthediffusionofHbandKnibetweenneighboringnuclei.Ouranalysisrevealsthatmutualrepressioncan markedlyincreasethesteepnessandprecisionofthegapgeneexpressionboundaries. Incontrasttoothermechanisms suchasspatialaveragingandcooperativegeneactivation,mutualrepressionthusallowsforgene-expressionboundaries thatarebothsteepandprecise.Moreover,mutualrepressiondramaticallyenhancestheirrobustnessagainstembryo-to- embryovariationsinthemorphogenlevels.Finally,oursimulationsrevealthatdiffusionofthegapproteinsplaysacritical rolenotonlyinreducingthewidthofthegapgeneexpressionboundariesviathemechanismofspatialaveraging,butalso inrepairingpatt