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Here we present a new biophysicochemical model of a somatosensory cortical layer 4 to layer 2\/3 synapse to study the role of astrocytes in spike-timing-dependent long-term depression (t-LTD) <jats:italic>in vivo<\/jats:italic>. By applying the synapse model and electrophysiological data recorded from rodent somatosensory cortex, we show that a signal from a postsynaptic neuron, orchestrated by endocannabinoids, astrocytic calcium signaling, and presynaptic N-methyl-D-aspartate receptors coupled with calcineurin signaling, induces t-LTD which is sensitive to the temporal difference between post- and presynaptic firing. We predict for the first time the dynamics of astrocyte-mediated molecular mechanisms underlying t-LTD and link complex biochemical networks at presynaptic, postsynaptic, and astrocytic sites to the time window of t-LTD induction. During t-LTD a single astrocyte acts as a delay factor for fast neuronal activity and integrates fast neuronal sensory processing with slow non-neuronal processing to modulate synaptic properties in the brain. Our results suggest that astrocytes play a critical role in synaptic computation during postnatal development and are of paramount importance in guiding the development of brain circuit functions, learning and memory.<\/jats:p>","DOI":"10.1371\/journal.pcbi.1008360","type":"journal-article","created":{"date-parts":[[2020,11,10]],"date-time":"2020-11-10T18:49:23Z","timestamp":1605034163000},"page":"e1008360","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":31,"title":["Astrocyte-mediated spike-timing-dependent long-term depression modulates synaptic properties in the developing 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