11# Lecture 4D: Short-Term Plasticity
22
3- In this lesson, we will study how short-term plasticity (STP) dynamics
4- using one of ngc-learn's in-built synapses, the ` STPDenseSynapse ` .
3+ In this lesson, we will study how short-term plasticity (STP) <b >[ 1] </b > dynamics
4+ -- where synaptic efficacy is cast in terms of the history of presynaptic activity --
5+ using ngc-learn's in-built ` STPDenseSynapse ` .
56Specifically, we will study how a dynamic synapse may be constructed and
67examine what short-term depression (STD) and short-term facilitation
78(STF) dominated configurations of an STP synapse look like.
@@ -22,21 +23,31 @@ to emulate dynamics from a STD-dominated perspective.
2223
2324### Starting with Facilitation-Dominated Dynamics
2425
25- One experiment goal with using a "dynamic synapse" is often to computationally
26+ One experiment goal with using a "dynamic synapse" < b > [ 1 ] </ b > is often to computationally
2627model the fact that synaptic efficacy (strength/conductance magnitude) is
27- not a fixed quantity (even in cases where long-term adaptation/learning is
28- absent) and instead a time-varying property that depends on a fixed
29- quantity of biophysical resources, e.g., neurotransmitter chemicals. This
30- means, in the context of spiking cells, when a pre-synaptic neuron emits a
31- pulse, this will affect the relative magnitude of the synapse's efficacy;
28+ not a fixed quantity -- even in cases where long-term adaptation/learning is
29+ absent -- and instead a time-varying property that depends on a fixed
30+ quantity of biophysical resources. Specifically, biological neuronal networks,
31+ synaptic signaling (or communication of information across synaptic connection
32+ pathways) consumes some quantity of neurotransmitters -- STF results from an
33+ influx of calcium into an axon terminal of a pre-synaptic neuron (after
34+ emission of a spike pulse) whereas STD occurs after a depletion of
35+ neurotransmitters that is consumed by the act of synaptic signaling at the axon
36+ terminal of a pre-synaptic neuron. Studies of cortical neuronal regions have
37+ empirically found that some areas are STD-dominated, STF-dominated, or exhibit
38+ some mixture of the two.
39+
40+ Ultimately, the above means that, in the context of spiking cells, when a
41+ pre-synaptic neuron emits a pulse, this act will affect the relative magnitude
42+ of the synapse's efficacy;
3243in some cases, this will result in an increase (facilitation) and, in others,
3344this will result in a decrease (depression) that lasts over a short period
34- of time (several milliseconds in many instances). Considering the fact
35- synapses have a dynamic nature to them, both over short and long time-scales,
36- means that plasticity can be thought of as a stimulus and resource-dependent
37- quantity, reflecting an important biophysical aspect that affects how
38- neuronal systems adapt and generalize given different kinds of sensory
39- stimuli.
45+ of time (several hundreds to thousands of milliseconds in many instances).
46+ As a result of considering synapses to have a dynamic nature to them, both over
47+ short and long time-scales, plasticity can now be thought of as a stimulus and
48+ resource-dependent quantity, reflecting an important biophysical aspect that
49+ affects how neuronal systems adapt and generalize given different kinds of
50+ sensory stimuli.
4051
4152Writing our STP dynamic synapse can be done by importing
4253[ STPDenseSynapse] ( ngclearn.components.synapses.STPDenseSynapse )
@@ -106,9 +117,37 @@ want to set to obtain different desired behavior from this in-built dynamic
106117synapse -- setting $\tau_f > \tau_d$ will result in STF-dominated behavior
107118whereas setting $\tauf < \tau_d$ will produce STD-dominated behavior. Note
108119that setting $\tau_d = 0$ will result in short-term depression being turned off
109- completely ($\tau_f 0$ disables STF).
120+ completely ($\tau_f 0$ disables STF).
110121
111- We can then write the simulated input Poisson spike train as follows:
122+ Formally, given the time constants above the dynamics of the ` STPDenseSynapse `
123+ operate according to the following coupled ordinary differential equations (ODEs):
124+
125+ $$
126+ \tau_f \frac{\partial u_j(t)}{\partial t} &= -u_j(t) + N_R (1 - u_j(t)) s_j(t) \\
127+ \tau_d \frac{\partial x_j}{\partial t} &= (1 - x_j(t)) - u_j(t + \Delta t) x_j(t) s_j(t) \\
128+ $$
129+
130+ and the resulting (short-term) synaptic efficacy:
131+
132+ $$
133+ W^{dyn}(t + \Delta t) = \Big( W^{max}_{ij} u_j(t + \Delta t) x_j(t) s_j(t) \Big)
134+ + W^{dyn}_{ij} (1 - s_j(t))
135+ $$
136+
137+ where $N_R$ represents an increment produced by a pre-synaptic spike (and
138+ in essence, the neurotransmitter resources available to yield facilitation),
139+ $W^{max}_ {ij}$ denotes the absolute synaptic efficacy (or maximum response
140+ amplitude of this synapse in the case of a complete release of all
141+ neurotransmitters; $x_j(t) = u_j(t) = 1$) of the connection between pre-synaptic
142+ neuron $j$ and post-synaptic neuron $i$, and $W^{dyn}_ {ij}(t)$ is the value
143+ of the dynamic synapse's efficacy at time ` t ` .
144+
145+ ### Simulating and Visualizing STF
146+
147+ Now that we understand the basics of how an ngc-learn STP works, we can next
148+ try it out on a simple pre-synaptic Poisson spike train. Writing out the
149+ simulated input Poisson spike train and our STP model's processing of this
150+ data can be done as follows:
112151
113152``` python
114153t_vals = []
@@ -143,7 +182,6 @@ x_vals = jnp.squeeze(jnp.asarray(x_vals))
143182t_vals = jnp.squeeze(jnp.asarray(t_vals))
144183```
145184
146-
147185We may then plot out the result of the STF-dominated dynamics we
148186simulate above with the following code:
149187
@@ -225,4 +263,15 @@ Now, modify your meta-parameters one last time to use a higher-frequency
225263input spike train, i.e., ` firing_rate_e = 20 ## Hz ` , to obtain a plot similar
226264to the one below:
227265
228- <img src =" ../../images/tutorials/neurocog/20Hz_stp_std_dom.jpg " width =" 500 " />
266+ <img src =" ../../images/tutorials/neurocog/20Hz_stp_std_dom.jpg " width =" 500 " />
267+
268+ You have now successfully simulated a dynamic synapse in ngc-learn across
269+ several different Poisson input train frequencies under both STF and
270+ STD-dominated regimes. In more complex biophysical models, it could prove useful
271+ to consider combining STP with other forms of long-term experience-dependent
272+ forms of synaptic adaptation, such as spike-timing-dependent plasticity.
273+
274+ ## References
275+
276+ <b >[ 1] </b > Tsodyks, Misha, Klaus Pawelzik, and Henry Markram. "Neural networks
277+ with dynamic synapses." Neural computation 10.4 (1998): 821-835.
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