Synaptic Correlates of Working Memory Capacity

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Synaptic Correlates of Working Memory Capacity Yuanyuan Mi, Mikhail Katkov, Misha Tsodyks  Neuron  Volume 93, Issue 2, Pages 323-330 (January 2017) DOI: 10.1016/j.neuron.2016.12.004 Copyright © 2017 Elsevier Inc. Terms and Conditions

Figure 1 STP-Based WM Network Model (A) Network architecture: a number of recurrent excitatory neural clusters, shown in different colors, reciprocally connected to an inhibitory neuron pool, shown in black. (B) Model of a synaptic connection with STP. In response to a presynaptic spike train (lower panel), the neurotransmitter release probability u increases and the fraction of available neurotransmitter x decreases (middle panel), representing, synaptic facilitation and depression, respectively. The effective synaptic efficacy is proportional to ux (upper panel). (C) Network simulation with five loaded memory items. Upper panel: firing rates of different clusters. Five clusters are sequentially stimulated by brief external excitation (shaded colored rectangles). Different colors correspond to different clusters as in (A). Following the stimulation, four clusters continue sequential activation in the form of PSs while the remaining item fades away. Lower panel: the instantaneous synaptic efficacy JEEux for stimulated neural clusters during loading and subsequent reactivations. (D) Probability of retaining an item in WM as a function of its serial position in a train of 16 loaded items, computed with 450 simulated trials with presentation frequencies between 16 and 66 items/s. (E) Red: same as (D); computed with 450 simulated trials with presentation frequencies between 0.25 and 2 items/s. Blue: analytical calculation of the probability, as explained in the text and Supplemental Information. The parameters are as follows: JEE=8, τf=1.5s, τd=0.3s, U=0.3, τ=8ms, JIE=1.75, JEI=1.1, α=1.5, P=16, and Ib=3.0Hz in (C) and Ib=8Hz in (D) and (E). Neuron 2017 93, 323-330DOI: (10.1016/j.neuron.2016.12.004) Copyright © 2017 Elsevier Inc. Terms and Conditions

Figure 2 WM Capacity: Calculations and Numerical Results (A) Activity of two consecutively activated excitatory clusters, μ and μ + 1, and the inhibitory cluster. The cluster μ elicits a PS at time t0 (magenta trace), which triggers the response of the inhibitory neural pool (black trace), the latter generating negative synaptic current at the cluster μ + 1 (green trace). The cluster μ + 1 then recovers from inhibition until its current reaches the threshold point where next PS is emitted. (B) The function F(h) that determines the simplified one-dimensional dynamics of synaptic current according to Equation 7. Synaptic current recovers from initial hyperpolarized level h0 to the slowest point of the flow at hmin, after which the flow speed is increasing and a PS is generated. The speed of the synaptic current flow is illustrated with the red arrows. (C) The WM capacity as a function of τ, obtained with numerical simulations of the model. τd and τf as in Figure 1. (D) Same as (C); analytically estimated with Equations 5, 6, and 9. In Equation 9, we neglected the dependence of C, h0, and Icrit on synaptic parameters of the model and replaced them by the constant values C = 4, h0=−200Hz, and Icrit=2.45Hz. See Supplemental Information for details. τd and τf as in Figure 1. (E) The WM capacity as a function of τf and τd obtained with numerical simulations of the model. (F) Same as (E); analytically estimated as in (D). The parameters JEI, JIE, JEE, U, α, P, and Ib are the same as Figure 1D. Neuron 2017 93, 323-330DOI: (10.1016/j.neuron.2016.12.004) Copyright © 2017 Elsevier Inc. Terms and Conditions

Figure 3 Spiking Neural Networks (A) An example of network simulation, including spontaneous activity and WM triggered by loading ten stimuli. Spikes of 8,000 excitatory neurons are shown as dots; neurons are arranged in order such that the first 6,400 neurons are encoding ten patterns stored in the network. Out of those, eight items are loaded sequentially into the network after 5 s of spontaneous activity. Parameters are as follows: τ=20ms, τf=3s, τd=0.6s, and U=0.2. (B) The WM capacity as a function of τ, obtained with multiple simulations of the same network and averaging the results. Error bars: SEM. (C) The WM capacity as a function of τf and τd. Parameters of the spiking network model are given in the Supplemental Information. Neuron 2017 93, 323-330DOI: (10.1016/j.neuron.2016.12.004) Copyright © 2017 Elsevier Inc. Terms and Conditions

Figure 4 Segmenting a Stream of Inputs into Chunks via Modulation of Background Excitation (A) Two groups of inputs of four items each are loaded sequentially with a pause between them (shaded color rectangles). Upper panel: activity of each cluster is shown in a different color. Lower panel: background input. After the fourth’s input is presented, all four representations from the first chunk are active until the background input is reduced. When the background input is increased again and the next four items are presented, the second chunk is active after the eighth’s memory is loaded. (B) Same as (A), but with a shorter pause between two groups of inputs. The segmentation fails because when the background input is increased after the pause, the memories from the first chunk are reactivated and mix in with the second chunk. All other parameters as in Figure 1D. Neuron 2017 93, 323-330DOI: (10.1016/j.neuron.2016.12.004) Copyright © 2017 Elsevier Inc. Terms and Conditions