Background Synaptic Activity in Rat Entorhinal Cortex Shows a Progressively Greater Dominance of Inhibition over Excitation from Deep to Superficial Layers.docVIP

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Background Synaptic Activity in Rat Entorhinal Cortex Shows a Progressively Greater Dominance of Inhibition over Excitation from Deep to Superficial Layers.doc

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Background Synaptic Activity in Rat Entorhinal Cortex Shows a Progressively Greater Dominance of Inhibition over Excitation from Deep to Superficial Layers

BackgroundSynapticActivityinRatEntorhinalCortex ShowsaProgressivelyGreaterDominanceofInhibition overExcitationfromDeeptoSuperficialLayers StuartDavidGreenhill1¤a,SophieElizabethLynChamberlain1¤b,AlexLench1,PeterVernonMassey1, KathrynHeatherYuill3,GavinLawrenceWoodhall2,RolandSpencerGwynneJones1* 1DepartmentofPharmacyandPharmacology,UniversityofBath,ClavertonDown,Bath,UnitedKingdom,2AstonBrainCentre,SchoolofLifeandHealthSciences,Aston University, Birmingham,United Kingdom, 3School of Biomedical Healthcare Sciences, Plymouth University Peninsula Schools of Medicine and Dentistry, Plymouth, UnitedKingdom Abstract Theentorhinalcortex(EC)controlshippocampalinputandoutput,playingmajorrolesinmemoryandspatialnavigation. Different layers of the EC subserve different functions and a number of studies have compared properties of neurones across layers. We have studied synaptic inhibition and excitation in EC neurones, and we have previously compared spontaneous synaptic release of glutamate and GABA using patch clamp recordings of synaptic currents in principal neurones of layers II (L2) and V (L5). Here, we add comparative studies in layer III (L3). Such studies essentially look at neuronal activity from a presynaptic viewpoint. To correlate this with the postsynaptic consequences of spontaneous transmitter release, we have determined global postsynaptic conductances mediated by the two transmitters, using a methodtoestimateconductancesfrommembranepotentialfluctuations.Wehavepreviouslypresentedsomeofthisdata forL3andnowextendtoL2andL5.Inhibitiondominatesexcitationinalllayersbuttheratiofollowsaclearrankorder (highesttolowest)ofL2.L3.L5.Thevarianceofthebackgroundconductanceswasmarkedlyhigherforexcitationand inhibition in L2 compared to L3 or L5. We also show that induction of synchronized network epileptiform activity by blockade of GABA inhibition reveals a relative reluctance of L2 to participate in such activity. This was associated with maintenanceofadominantbackgroundin