Visual Feature Encoding Ganglion Cell Response Transience Is Determined by the Summation of Converging Parallel Signals

Visual Feature Encoding Ganglion Cell Response Transience Is Determined by the Summation of Converging Parallel Signals | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Visual Feature Encoding Ganglion Cell Response Transience Is Determined by the Summation of Converging Parallel Signals Alma Ganczer, Gergely Szarka, Márton Balogh, Ádám Jonatán Tengölics, and 2 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-477066/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Retinal ganglion cells (RGCs) summate inputs across their receptive fields and forward a corresponding spike train code to the brain. Considering the many visual aspects carried by this code the comprehension of RGC firing kinetics and the underlying mechanisms is markedly important. RGCs can generate a maintained spiking (sustained) or a quickly decaying brief burst of spikes (transient) upon ON- and/or OFF-set of prolonged light stimuli. Our results here challenge the classical view that claims an outer retinal origin for RGC response transience and explains the observed response dichotomy with the dissimilar glutamate receptor kinetics in postsynaptic bipolar cell dendritic surfaces. We find that activation of the same glutamate receptor subtype can result in transient, sustained, and intermediate RGC responses. Moreover, even signaling via a single bipolar cell subtype can result in RGC responses with a variety of transience values. Contrary, a change in the dominance of inputs delivered by converging retinal pathways can alter RGC response transience considerably. Such response component fine-tuning occurs via inner retinal GABAergic inhibitory and gap junction mediated excitatory interactions. The above data thus indicate that RGC light response temporal characteristics are determined by inner retinal microcircuits and fine-tuned in a context dependent manner. Neurobiology of Disease Cellular & Molecular Neuroscience gap junction electrical synapse ganglion cell inner plexiform layer ganglion cell layer retina parallel signaling Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Full Text Due to technical limitations, full-text HTML conversion of this manuscript could not be completed. However, the manuscript can be downloaded and accessed as a PDF. Additional Declarations No competing interests reported. Supplementary Files Supplementalfigure1.eps Supplemental Figure 1 Supplementalfigure2.eps Supplemental Figure 2 Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-477066","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":24144340,"identity":"b1c6c220-3718-4b47-b313-5f45faf8a143","order_by":0,"name":"Alma Ganczer","email":"","orcid":"","institution":"University of Pécs, Szentágothai Research Centre","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Alma","middleName":"","lastName":"Ganczer","suffix":""},{"id":24144341,"identity":"74fd21df-13c0-4502-8f93-e06e8f8d58db","order_by":1,"name":"Gergely Szarka","email":"","orcid":"","institution":"University of Pécs, Szentágothai Research Centre","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Gergely","middleName":"","lastName":"Szarka","suffix":""},{"id":24144342,"identity":"7d93621d-2c69-4f9d-a788-6746a00e8bc1","order_by":2,"name":"Márton Balogh","email":"","orcid":"","institution":"University of Pécs, Szentágothai Research Centre","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Márton","middleName":"","lastName":"Balogh","suffix":""},{"id":24144343,"identity":"e9739e6b-3afb-4987-9909-20d001f6dae5","order_by":3,"name":"Ádám Jonatán Tengölics","email":"","orcid":"","institution":"University of Pécs, Szentágothai Research Centre","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Ádám","middleName":"Jonatán","lastName":"Tengölics","suffix":""},{"id":24144344,"identity":"e2f46110-7d79-4aa7-92a9-56c9a9f7ccf3","order_by":4,"name":"Tamás Kovács-Öller","email":"","orcid":"","institution":"University of Pécs, Szentágothai Research Centre","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Tamás","middleName":"","lastName":"Kovács-Öller","suffix":""},{"id":24144345,"identity":"d1f531f7-3be2-472a-b1b8-d89b4b10c97f","order_by":5,"name":"Béla Völgyi","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAABEklEQVRIiWNgGAWjYFAC5gYGBgMJHn64AA+ESsCthRGkxUZGsgHCleABaTlAUAtDmo3BAWK1GBxvbH7NU3CYx/j84qebbtQw1NnzHGB7/KGCIQ9mCIaWMwfbrHkMDvOY3XhmdjvnGNAW3gZ2gwNnGIpxaZGckdhmDNFyAKiFjQEUEGwSB9sYEjcQ0mI84/i32zn/YFr+4dbCL5HY/JjHII3HgL/H7HZuG9hhQC0NeLTwHGxjnGNgwyNxg6fsdm6fhGTPmYPtBmeOSSTOxKGFjb358Ic3fyTs+fuPb7ud882Gn70n+diDihqbxD4cWkC6pMDxLZEAJoGYsQ3KwAmYP/4AOxFhKBs+5aNgFIyCUTDyAAB1VV54OrnnCAAAAABJRU5ErkJggg==","orcid":"","institution":"University of Pécs, Szentágothai Research Centre","correspondingAuthor":true,"submittingAuthor":false,"prefix":"","firstName":"Béla","middleName":"","lastName":"Völgyi","suffix":""}],"badges":[],"createdAt":"2021-04-29 07:59:07","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-477066/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-477066/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":8885616,"identity":"83092974-bbfa-4088-811a-c15ddb0d9912","added_by":"auto","created_at":"2021-05-06 23:28:02","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":87746,"visible":true,"origin":"","legend":"A Great Variety of Response Transience Across the RGC Population. a. Representative perievent raster diagrams show that individual RGCs provide light-evoked spiking responses upon full-field illumination that are rather similar across trials (four consecutive trials for each recorded cell). However, RGCs display a great variety in terms of their response length (or decay – expressed as the PSTHτ value in this work) for both the ON (cells 1 and 2) and OFF (cells 3 and 4) subpopulations. The white bar below the recordings represents the timing of the on- and offset of the stimulus in this and in all other figure panels of this paper. b. Frequency histogram shows the distribution of PSTHτ values for the ON (white) and OFF (black) RGC subpopulations. Clearly, the distribution of PSTHτ values appears unimodal and does not allow for the clear separation of transient and sustained RGC responses. This unimodality as well as the rather wide range are features shared by both the ON and OFF RGC subpopulations.","description":"","filename":"Fig1.png","url":"https://assets-eu.researchsquare.com/files/rs-477066/v1/fa117569c0cbb355e48eb6e9.png"},{"id":8885741,"identity":"808f343d-4fbb-4406-be14-495a5569b2cd","added_by":"auto","created_at":"2021-05-06 23:31:02","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":70327,"visible":true,"origin":"","legend":"Scotopic RGC Response Transience Values are Just as Diverse as Photopic Responses. a. Representative ON RGC light responses (raster to the left and PSTH to the right for the same RGC) were evoked first by scotopic (top) and then photopic (bottom) full-field light stimuli. This change in the stimulus strength induced a clear change in response amplitude but the overall shape (response delay and decay) of the response remained largely unchanged. b. Similar to the ON RGC is panel a this representative OFF RGC display light responses that, besides subtle changes in response amplitude, overall remain unchanged in scotopic (top) and photopic (bottom) stimulating conditions. c. Diagram displays PSTHτ value pairs for individual RGC light responses in scotopic (left) and photopic (right) light stimulations. Clearly, the change in stimulus strength induced the alteration of PSTHτ values for many examined RGCs, however, the range of response transience values for the entire RGC population were comparable in scotopic conditions (if not even wider) to those obtained with photopic stimulation paradigms. d. Floating bar graphs show that the variety of scotopic RGC response PSTHτ values is just as great as for photopic responses for the same RGCs.","description":"","filename":"Fig2.png","url":"https://assets-eu.researchsquare.com/files/rs-477066/v1/746ce9af7eb7d756fe2602e3.png"},{"id":8885769,"identity":"aa48fc7c-05e9-43c1-a772-5b52e0e83a62","added_by":"auto","created_at":"2021-05-06 23:34:03","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":110100,"visible":true,"origin":"","legend":"Changes in Response Transience can be the Result of Signal Interference through Parallel Retinal Pathways. a. Representative ON RGC light-evoked rasters (left column) and PSTH cohorts (right column) recorded as a response to stimuli of various strength (intensity values are reflected in the right top corner of each -panel). PSTHτ values of this RGC changed non-monotonously during this experiment when the stimulus intensity was gradually increased (see also panel c). PSTHs clearly show a peak of very sensitive signal component (red arrow) evoked by weak, scotopic stimuli. This sensitive response component appears relative delayed when it is compared to the less sensitive but brisk signal component (light blue arrow). These two signal components differ in their delays but appear similar in response decay, therefore PSTHτ values are shifted towards the sustained range when the two signals are summated (mesopic conditions - 2nd, 3rd and 4th panels), whereas remain transient when only one signal is present (scotopic condition – 1st panel) or dominates over the other component (photopic conditions – 5th. 6th and 7th panels; see also panel c). b. Representative OFF RGC light-evoked rasters (left column) and PSTHs (right column) recorded as a response to varying intensity stimuli (intensity values are reflected in the right top corner of each -panel). PSTHτ values of this RGC clearly changed during this experiment as the stimulus intensity was gradually increased. Similar to the cell in panel a this OFF RGC showed a very sensitive but rather delated response peak (red arrow) and a faster but less sensitive (light blue arrow) peak. The two signal components differed in their delays and sensitivities and a slight alteration in PSTHτ values occurred as a result of the summation of components (mostly in mesopic conditions – middle panels). While the distinction of response components can clearly be differentiated for the ON RGC in a, this OFF cell (and most examined RGCs) showed a less obvious and less separable summation of incoming signals. c and d. Diagrams show that, similar to cells shown in panels a and b (values of these cells appear in black and red in the diagrams), most recorded RGCs displayed stimulus strength driven changes of PSTHτ values (grey curves). e. Diagram shows minimum/maximum PSTHτ value pairs for the recorded cells during the course of the stimulus intensity recording paradigm. The examined RGCs showed ~18-73% PSTHτ changes during the course of this experiment.","description":"","filename":"Fig3.png","url":"https://assets-eu.researchsquare.com/files/rs-477066/v1/0228a25c26900f504c881b38.png"},{"id":8885768,"identity":"df44e781-5ed0-4ab0-80c5-37b498447386","added_by":"auto","created_at":"2021-05-06 23:34:03","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":288961,"visible":true,"origin":"","legend":"GABA Receptor Blockade Induces Abrupt Changes in RGC Response Transience. a and b. Representative ON (a) and OFF (b) RGC light-driven responses (rasters on the top and corresponding PSTHs below) are clearly altered when the nonspecific GABA receptor blocker PTX was applied (bottom panels). The observed changes include both the disappearance (e.g. the sustained response shoulder for the OFF cell) and the unmasking (OFF pathway driven spiking for the ON cell and ON pathway driven spiking of the OFF cell; transient ON inhibition of for the ON cell) of various response components.","description":"","filename":"Fig4.png","url":"https://assets-eu.researchsquare.com/files/rs-477066/v1/b3a016baaec40d54a4240e5a.png"},{"id":8885618,"identity":"58283d35-7268-43b9-87dd-03b87b156049","added_by":"auto","created_at":"2021-05-06 23:28:02","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":85357,"visible":true,"origin":"","legend":"GABA Receptor Blockade Induces a General Decrease of RGC PSTHτ values. a and b. Apart from a handful of cells that showed no considerable change, the GABA receptor blockade induced an overall decrease of PSTHτ values for most examined ON (left) and OFF (right) RGCs. This PTX mediated decrease is also reflected in a drop of mean and median PSTHτ values (b) for both ON (left) and OFF (right) RGC subpopulations.","description":"","filename":"Fig5.png","url":"https://assets-eu.researchsquare.com/files/rs-477066/v1/506fd04d3a195a8e4afd07b7.png"},{"id":8885740,"identity":"5bd37cb2-75eb-4f51-ad00-5573959e54c2","added_by":"auto","created_at":"2021-05-06 23:31:02","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":224915,"visible":true,"origin":"","legend":"Gap Junction Blockade Induces Alterations in RGC Response Transience. a and b. Representative ON (left) and OFF (right) RGC light-evoked responses (rasters on the top and PSTHs on the bottom) are altered when the nonspecific gap junction blocker MFA was applied. These changes are less obvious when compared to the PTX induced changes, but a clear transience reduction can be observed for both the ON (left) and OFF (right) cells presented here.","description":"","filename":"Fig6.png","url":"https://assets-eu.researchsquare.com/files/rs-477066/v1/62759df6fa969e98e49a7584.png"},{"id":8885744,"identity":"72457598-9194-49bc-b933-ef9be4907c2c","added_by":"auto","created_at":"2021-05-06 23:31:03","extension":"png","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":121968,"visible":true,"origin":"","legend":"Gap Junction Blockade Induces an Overall Decrease of RGC PSTHτ values. a and b. Although a number of RGCs showed no change or increase of PSTHτ values as a response of a pharmacological gap junction blockade via the application of MFA, most RGCs responded with a decrease of their response decays for both the examined ON (left) and OFF (right) RGCs. The observed MFA mediated decrease is also depicted by the decreased mean and median PSTHτ values (b) for both ON (left) and OFF (right) RGC subpopulations.","description":"","filename":"Fig7.png","url":"https://assets-eu.researchsquare.com/files/rs-477066/v1/c08b15809e0600ace7369b31.png"},{"id":8885742,"identity":"48c2fc61-264f-4237-a93f-8c8e68567894","added_by":"auto","created_at":"2021-05-06 23:31:03","extension":"png","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":217930,"visible":true,"origin":"","legend":"Summary Drawing of Potential Signal Summation Mechanisms that Affecting RGC Response Transience. a. Two bipolar cells of different subtypes provide transient inputs to the same RGC (light blue EPSC curves). These two inputs have dissimilar delays (due to differential bipolar cell signaling and/or different location of synapses over the RGC dendritic arbor) and therefore the summation of the responses results in an intermediate or sustained RGC spiking response. b. This RGC receives excitatory inputs from two sources, from a transient bipolar cell (light blue EPSC) and from a gap junction coupled amacrine cell (purple depolarization). If the dynamics of these two inputs differ their summation will induce intermediate and/or sustained RGC spiking. c. This RGC receives excitation from a bipolar cell (light blue EPSC) and delayed inhibition (red IPSC) from an amacrine cell resulting in a transient RGC response. d. An RGC that receives excitation from a bipolar cell (light blue EPSC) and inhibition (red IPSC) from an amacrine cell. In this scenario the two inputs have about the same delays therefore the excitation will be truncated and the RGC output is an intermediate/sustained spiking.","description":"","filename":"Fig8.png","url":"https://assets-eu.researchsquare.com/files/rs-477066/v1/6e049a307289d1f709ddafd3.png"},{"id":13629186,"identity":"662628d6-ca1b-458c-9215-befdaa140f8f","added_by":"auto","created_at":"2021-09-17 08:05:09","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2112351,"visible":true,"origin":"","legend":"","description":"","filename":"Ganczer202120210429withfiguresinserted.pdf","url":"https://assets-eu.researchsquare.com/files/rs-477066/v1_covered.pdf"},{"id":10057551,"identity":"6904387b-7e72-4c92-9789-4d0411c8fba0","added_by":"auto","created_at":"2021-06-07 09:59:33","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2108854,"visible":true,"origin":"","legend":"","description":"","filename":"Ganczer202120210429withfiguresinserted.pdf","url":"https://assets-eu.researchsquare.com/files/rs-477066/v1_covered.pdf"},{"id":8885842,"identity":"1b331380-9908-4fc4-bcd6-7f130dc55382","added_by":"auto","created_at":"2021-05-06 23:37:08","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1772839,"visible":true,"origin":"","legend":"","description":"","filename":"Ganczer202120210429withfiguresinserted.pdf","url":"https://assets-eu.researchsquare.com/files/rs-477066/v1_stamped.pdf"},{"id":8885625,"identity":"5c2ed526-38ac-41f9-bad7-018e9595a791","added_by":"auto","created_at":"2021-05-06 23:28:03","extension":"eps","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":3161214,"visible":true,"origin":"","legend":"Supplemental Figure 1","description":"","filename":"Supplementalfigure1.eps","url":"https://assets-eu.researchsquare.com/files/rs-477066/v1/152ea371dc1a2697c00060a3.eps"},{"id":8885621,"identity":"322d57a9-ead5-445b-94c7-9476cd548ec3","added_by":"auto","created_at":"2021-05-06 23:28:03","extension":"eps","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":2626706,"visible":true,"origin":"","legend":"Supplemental Figure 2","description":"","filename":"Supplementalfigure2.eps","url":"https://assets-eu.researchsquare.com/files/rs-477066/v1/d8e08de4b58737402a9c5902.eps"}],"financialInterests":"No competing interests reported.","formattedTitle":"\u003cp\u003eVisual Feature Encoding Ganglion Cell Response Transience Is Determined by the Summation of Converging Parallel Signals\u003c/p\u003e","fulltext":[{"header":"Full Text","content":"\u003cp\u003eDue to technical limitations, full-text HTML conversion of this manuscript could not be completed. However, the manuscript can be downloaded and accessed as a PDF.\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":false,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":true,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"gap junction, electrical synapse, ganglion cell, inner plexiform layer, ganglion cell layer, retina, parallel signaling","lastPublishedDoi":"10.21203/rs.3.rs-477066/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-477066/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"Retinal ganglion cells (RGCs) summate inputs across their receptive fields and forward a corresponding spike train code to the brain. Considering the many visual aspects carried by this code the comprehension of RGC firing kinetics and the underlying mechanisms is markedly important. RGCs can generate a maintained spiking (sustained) or a quickly decaying brief burst of spikes (transient) upon ON- and/or OFF-set of prolonged light stimuli. Our results here challenge the classical view that claims an outer retinal origin for RGC response transience and explains the observed response dichotomy with the dissimilar glutamate receptor kinetics in postsynaptic bipolar cell dendritic surfaces. We find that activation of the same glutamate receptor subtype can result in transient, sustained, and intermediate RGC responses. Moreover, even signaling via a single bipolar cell subtype can result in RGC responses with a variety of transience values. Contrary, a change in the dominance of inputs delivered by converging retinal pathways can alter RGC response transience considerably. Such response component fine-tuning occurs via inner retinal GABAergic inhibitory and gap junction mediated excitatory interactions. The above data thus indicate that RGC light response temporal characteristics are determined by inner retinal microcircuits and fine-tuned in a context dependent manner.","manuscriptTitle":"Visual Feature Encoding Ganglion Cell Response Transience Is Determined by the Summation of Converging Parallel Signals","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2021-05-06 23:28:00","doi":"10.21203/rs.3.rs-477066/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"4f5cbe10-ed77-4541-889f-1593bb746ad3","owner":[],"postedDate":"May 6th, 2021","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":4111435,"name":"Neurobiology of Disease"},{"id":4111436,"name":"Cellular \u0026 Molecular Neuroscience"}],"tags":[],"updatedAt":"2021-06-07T09:59:16+00:00","versionOfRecord":[],"versionCreatedAt":"2021-05-06 23:28:00","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-477066","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-477066","identity":"rs-477066","version":["v1"]},"buildId":"7rjqhiLT3MXkJMwkYKINL","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}