Intrinsic Stabilization of Output Rates by Spike-Based Hebbian Learning

We study analytically a model of long-term synaptic plasticity where synaptic changes are triggered by presynaptic spikes, postsynaptic spikes, and the time differences between pre- and postsynaptic spikes. We show that plasticity can lead to an intrinsic stabilization of the mean firing rate of the postsynaptic neuron. Subtractive normalization of the synaptic weights (summed over all presynaptic inputs converging on a postsynaptic neuron) follows if, in addition, the mean input rates and the mean input correlations are identical at all synapses. If the integral over the learning window is positive, firing-rate stabilization requires a non-Hebbian component, whereas such a component is not needed, if the integral of the learning window is negative. A negative integral corresponds to `anti-Hebbian' learning in a model with slowly varying firing rates. For spike-based learning, a strict distinction between Hebbian and `anti-Hebbian' rules is questionable since learning is driven by correlations on the time scale of the learning window. The correlations between presynaptic and postsynaptic firing are evaluated for a piecewise-linear Poisson model and for a noisy spiking neuron model with refractoriness. Whereas a negative integral over the learning window leads to intrinsic rate stabilization, the positive part of the learning window picks up spatial and temporal correlations in the input