{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,2,27]],"date-time":"2026-02-27T06:14:54Z","timestamp":1772172894934,"version":"3.50.1"},"update-to":[{"DOI":"10.1371\/journal.pcbi.1008996","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2021,6,11]],"date-time":"2021-06-11T00:00:00Z","timestamp":1623369600000}}],"reference-count":85,"publisher":"Public Library of Science (PLoS)","issue":"6","license":[{"start":{"date-parts":[[2021,6,1]],"date-time":"2021-06-01T00:00:00Z","timestamp":1622505600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["www.ploscompbiol.org"],"crossmark-restriction":false},"short-container-title":["PLoS Comput Biol"],"abstract":"<jats:p>Several homeostatic mechanisms enable the brain to maintain desired levels of neuronal activity. One of these, homeostatic structural plasticity, has been reported to restore activity in networks disrupted by peripheral lesions by altering their neuronal connectivity. While multiple lesion experiments have studied the changes in neurite morphology that underlie modifications of synapses in these networks, the underlying mechanisms that drive these changes are yet to be explained. Evidence suggests that neuronal activity modulates neurite morphology and may stimulate neurites to selective sprout or retract to restore network activity levels. We developed a new spiking network model of peripheral lesioning and accurately reproduced the characteristics of network repair after deafferentation that are reported in experiments to study the activity dependent growth regimes of neurites. To ensure that our simulations closely resemble the behaviour of networks in the brain, we model deafferentation in a biologically realistic balanced network model that exhibits low frequency Asynchronous Irregular (AI) activity as observed in cerebral cortex. Our simulation results indicate that the re-establishment of activity in neurons both within and outside the deprived region, the Lesion Projection Zone (LPZ), requires opposite activity dependent growth rules for excitatory and inhibitory post-synaptic elements. Analysis of these growth regimes indicates that they also contribute to the maintenance of activity levels in individual neurons. Furthermore, in our model, the directional formation of synapses that is observed in experiments requires that pre-synaptic excitatory and inhibitory elements also follow opposite growth rules. Lastly, we observe that our proposed structural plasticity growth rules and the inhibitory synaptic plasticity mechanism that also balances our AI network both contribute to the restoration of the network to pre-deafferentation stable activity levels.<\/jats:p>","DOI":"10.1371\/journal.pcbi.1008996","type":"journal-article","created":{"date-parts":[[2021,6,1]],"date-time":"2021-06-01T16:27:08Z","timestamp":1622564828000},"page":"e1008996","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":2,"title":["Growth rules for the repair of Asynchronous Irregular neuronal networks after peripheral 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Turrigiano","year":"1999","journal-title":"Trends in neurosciences"},{"issue":"5874","key":"pcbi.1008996.ref059","doi-asserted-by":"crossref","first-page":"385","DOI":"10.1126\/science.1150516","article-title":"The antidepressant fluoxetine restores plasticity in the adult visual cortex","volume":"320","author":"JFM Vetencourt","year":"2008","journal-title":"Science"},{"key":"pcbi.1008996.ref060","doi-asserted-by":"crossref","first-page":"100","DOI":"10.1016\/S0006-8993(03)02717-3","article-title":"Extracellular GABA concentrations in area 17 of cat visual cortex during topographic map reorganization following binocular central retinal lesioning","volume":"976","author":"A Massie","year":"2003","journal-title":"Brain research"},{"key":"pcbi.1008996.ref061","doi-asserted-by":"crossref","first-page":"347","DOI":"10.3109\/08990229109144757","article-title":"Injury-induced reorganization of somatosensory cortex is accompanied by reductions in GABA staining","volume":"8","author":"PE Garraghty","year":"1991","journal-title":"Somatosensory & motor research"},{"issue":"11","key":"pcbi.1008996.ref062","doi-asserted-by":"crossref","first-page":"877","DOI":"10.1038\/nrn1787","article-title":"Critical period plasticity in local cortical circuits","volume":"6","author":"TK Hensch","year":"2005","journal-title":"Nature Reviews Neuroscience"},{"issue":"6774","key":"pcbi.1008996.ref063","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1038\/35004582","article-title":"Inhibitory threshold for critical-period activation in primary visual cortex","volume":"404","author":"M Fagiolini","year":"2000","journal-title":"Nature"},{"issue":"2","key":"pcbi.1008996.ref064","doi-asserted-by":"crossref","first-page":"374","DOI":"10.1016\/j.neuron.2012.03.015","article-title":"Elimination of inhibitory synapses is a major component of adult ocular dominance plasticity","volume":"74","author":"D van 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Available from: https:\/\/doi.org\/10.5281\/zenodo.2605422."},{"key":"pcbi.1008996.ref070","doi-asserted-by":"crossref","DOI":"10.3389\/fnana.2016.00057","article-title":"Automatic generation of connectivity for large-scale neuronal network models through structural plasticity","volume":"10","author":"S Diaz-Pier","year":"2016","journal-title":"Frontiers in neuroanatomy"},{"key":"pcbi.1008996.ref071","doi-asserted-by":"crossref","first-page":"32","DOI":"10.3389\/fninf.2018.00032","article-title":"Toward Rigorous Parameterization of Underconstrained Neural Network Models Through Interactive Visualization and Steering of Connectivity Generation","volume":"12","author":"C Nowke","year":"2018","journal-title":"Frontiers in Neuroinformatics"},{"issue":"3","key":"pcbi.1008996.ref072","doi-asserted-by":"crossref","first-page":"287","DOI":"10.1007\/s12021-012-9146-1","article-title":"The connection-set algebra\u2014a novel formalism for the representation of connectivity structure in neuronal network models","volume":"10","author":"M Djurfeldt","year":"2012","journal-title":"Neuroinformatics"},{"issue":"1","key":"pcbi.1008996.ref073","doi-asserted-by":"crossref","DOI":"10.1038\/srep18854","article-title":"GeNN: a code generation framework for accelerated brain simulations","volume":"6","author":"E Yavuz","year":"2016","journal-title":"Scientific Reports"},{"issue":"1","key":"pcbi.1008996.ref074","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1016\/j.jneumeth.2012.03.001","article-title":"Power-efficient simulation of detailed cortical microcircuits on SpiNNaker","volume":"210","author":"T Sharp","year":"2012","journal-title":"Journal of Neuroscience Methods"},{"key":"pcbi.1008996.ref075","unstructured":"Billaudelle S, Cramer B, Petrovici MA, Schreiber K, Kappel D, Schemmel J, et al. Structural plasticity on an accelerated analog neuromorphic hardware system."},{"issue":"5","key":"pcbi.1008996.ref076","doi-asserted-by":"crossref","first-page":"964","DOI":"10.1016\/j.neuron.2017.10.013","article-title":"A Commitment to Open Source in Neuroscience","volume":"96","author":"P Gleeson","year":"2017","journal-title":"Neuron"},{"key":"pcbi.1008996.ref077","doi-asserted-by":"crossref","DOI":"10.3389\/neuro.11.012.2008","article-title":"PyNEST: a convenient interface to the NEST simulator","volume":"2","author":"JM Eppler","year":"2008","journal-title":"Frontiers in neuroinformatics"},{"key":"pcbi.1008996.ref078","unstructured":"Kunkel S, Morrison A, Weidel P, Eppler JM, Sinha A, Schenck W, et al. NEST 2.12.0; 2017."},{"key":"pcbi.1008996.ref079","unstructured":"Denker M, Yegenoglu A, Gr\u00fcn S. Collaborative HPC-enabled workflows on the HBP Collaboratory using the Elephant framework. In: Neuroinformatics 2018;. p. P19. Available from: https:\/\/abstracts.g-node.org\/conference\/NI2018\/abstracts#\/uuid\/023bec4e-0c35-4563-81ce-2c6fac282abd."},{"issue":"3","key":"pcbi.1008996.ref080","doi-asserted-by":"crossref","first-page":"542","DOI":"10.1016\/j.neuron.2011.06.017","article-title":"Broad inhibition sharpens orientation selectivity by expanding input dynamic range in mouse simple cells","volume":"71","author":"Bh Liu","year":"2011","journal-title":"Neuron"},{"issue":"7430","key":"pcbi.1008996.ref081","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1038\/nature11665","article-title":"Inhibition dominates sensory responses in the awake cortex","volume":"493","author":"B Haider","year":"2013","journal-title":"Nature"},{"issue":"9","key":"pcbi.1008996.ref082","doi-asserted-by":"crossref","first-page":"3555","DOI":"10.1073\/pnas.0810390106","article-title":"The fractions of short-and long-range connections in the visual cortex","volume":"106","author":"A Stepanyants","year":"2009","journal-title":"Proceedings of the National Academy of Sciences"},{"key":"pcbi.1008996.ref083","doi-asserted-by":"crossref","first-page":"2848","DOI":"10.1073\/pnas.0138051100","article-title":"Mexican hats and pinwheels in visual cortex","volume":"100","author":"K Kang","year":"2003","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"issue":"1","key":"pcbi.1008996.ref084","doi-asserted-by":"crossref","first-page":"P21","DOI":"10.1186\/1471-2202-14-S1-P21","article-title":"Developing orientation maps using realistic patterns of lateral connectivity","volume":"14","author":"P Rudiger","year":"2013","journal-title":"BMC Neuroscience"},{"issue":"9","key":"pcbi.1008996.ref085","doi-asserted-by":"crossref","first-page":"1117","DOI":"10.1038\/nn1747","article-title":"Spine growth precedes synapse formation in the adult neocortex in vivo","volume":"9","author":"GW Knott","year":"2006","journal-title":"Nature neuroscience"}],"updated-by":[{"DOI":"10.1371\/journal.pcbi.1008996","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2021,6,11]],"date-time":"2021-06-11T00:00:00Z","timestamp":1623369600000}}],"container-title":["PLOS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1008996","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,6,11]],"date-time":"2021-06-11T13:47:34Z","timestamp":1623419254000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1008996"}},"subtitle":[],"editor":[{"given":"Abigail","family":"Morrison","sequence":"first","affiliation":[],"role":[{"role":"editor","vocabulary":"crossref"}]}],"short-title":[],"issued":{"date-parts":[[2021,6,1]]},"references-count":85,"journal-issue":{"issue":"6","published-online":{"date-parts":[[2021,6,1]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1008996","relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/810846","asserted-by":"object"}]},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,6,1]]}}}