Perturbing Dynamin Reveals Potent Effects on the Drosophila Circadian Clock 英文参考文献.docVIP

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Perturbing Dynamin Reveals Potent Effects on the Drosophila Circadian Clock 英文参考文献.doc

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Perturbing Dynamin Reveals Potent Effects on the Drosophila Circadian Clock 英文参考文献

PerturbingDynaminRevealsPotentEffectsonthe DrosophilaCircadianClock ValerieL.Kilman1,2,LuoyingZhang1,Rose-AnneMeissner1,ElyssaBurg1¤,RaviAllada1,2* 1Department of Neurobiology and Physiology, Northwestern University, Evanston, Illinois, United States of America, 2Center for Sleep and Circadian Biology, NorthwesternUniversity,Evanston,Illinois,UnitedStatesofAmerica Abstract Background:Transcriptionalfeedbackloopsarecentraltocircadianclockfunction.However,theroleofneuralactivityand membrane events in molecular rhythms in the fruit fly Drosophila is unclear. To address this question, we expressed a temperature-sensitive,dominantnegativealleleoftheflyhomologofdynamincalledshibirets1(shits1),anactivecomponent inmembranevesiclescission. PrincipalFindings:Broadexpressioninclockcellsresultedinunexpectedlylong,robustperiods(.28hours)comparableto perturbation of core clock components, suggesting an unappreciated role of membrane dynamics in setting period. Expressioninthepacemakerlateralventralneurons(LNv)wasnecessaryandsufficientforthiseffect.Manipulationofother endocyticcomponentsexacerbatedshits1’sbehavioraleffects,suggestingitsmechanismisspecifictoendocyticregulation. PKA overexpression rescued period effects suggesting shits1 may downregulate PKA pathways. Levels of the clock componentPERIODwerereducedintheshits1-expressingpacemakersmallLNvoffliesheldatafullyrestrictivetemperature (29uC). Less restrictive conditions (25uC) delayed cycling proportional to observed behavioral changes. Levels of the neuropeptidePIGMENT-DISPERSINGFACTOR(PDF),theonlyknownLNvneurotransmitter,werealsoreduced,butPERIOD cyclingwasstilldelayedinflieslackingPDF,implicatingaPDF-independentprocess.Further,shits1expressionintheeyealso results in reduced PER protein and per and vri transcript levels, suggesting that shibire-dependent signaling extends to peripheralclocks.ThelevelofnuclearCLK,transcriptionalactivatorofmanycoreclockgenes,isalsoreducedinshits1flies, andClkoverexpressionsuppressestheperi