How adaptation currents change threshold, gain and variability of neuronal spiking

Many types of neurons exhibit spike rate adaptation, mediated by intrinsic slow $\mathrm{K}^+$-currents, which effectively inhibit neuronal responses. How these adaptation currents change the relationship between in-vivo like fluctuating synaptic input, spike rate output and the spike train statistics, however, is not well understood. In this computational study we show that an adaptation current which primarily depends on the subthreshold membrane voltage changes the neuronal input-output relationship (I-O curve) subtractively, thereby increasing the response threshold. A spike-dependent adaptation current alters the I-O curve divisively, thus reducing the response gain. Both types of adaptation currents naturally increase the mean inter-spike interval (ISI), but they can affect ISI variability in opposite ways. A subthreshold current always causes an increase of variability while a spike-triggered current decreases high variability caused by fluctuation-dominated inputs and increases low variability when the average input is large. The effects on I-O curves match those caused by synaptic inhibition in networks with asynchronous irregular activity, for which we find subtractive and divisive changes caused by external and recurrent inhibition, respectively. Synaptic inhibition, however, always increases the ISI variability. We analytically derive expressions for the I-O curve and ISI variability, which demonstrate the robustness of our results. Furthermore, we show how the biophysical parameters of slow $\mathrm{K}^+$-conductances contribute to the two different types of adaptation currents and find that $\mathrm{Ca}^{2+}$-activated $\mathrm{K}^+$-currents are effectively captured by a simple spike-dependent description, while muscarine-sensitive or $\mathrm{Na}^+$-activated $\mathrm{K}^+$-currents show a dominant subthreshold component.Comment: 20 pages, 8 figures; Journal of Neurophysiology (in press

Predictive Coding as a Model of Biased Competition in Visual Attention

Attention acts, through cortical feedback pathways, to enhance the response of cells encoding expected or predicted information. Such observations are inconsistent with the predictive coding theory of cortical function which proposes that feedback acts to suppress information predicted by higher-level cortical regions. Despite this discrepancy, this article demonstrates that the predictive coding model can be used to simulate a number of the effects of attention. This is achieved via a simple mathematical rearrangement of the predictive coding model, which allows it to be interpreted as a form of biased competition model. Nonlinear extensions to the model are proposed that enable it to explain a wider range of data

Allocation of Computational Resources in the Nervous System.

The nervous system integrates past information together with predictions about the future in order to produce rewarding actions for the organism. This dissertation focuses on the resources underlying these computations, and the task-dependent allocation of these resources. We present evidence that principles from optimal coding and optimal estimation account for overt and covert orienting phenomena, as observed from both behavioral experiments and neuronal recordings. First, we review behavioral measurements related to selective attention and discuss models that account for these data. We show that reallocation of resources emerges as a natural property of systems that encode their inputs efficiently under non-uniform constraints. We continue by discussing the attentional modulation of neuronal activity, and showthat: (1) Modulation of coding strategies does not require special mechanisms: it is possible to obtain dramatic modulation even when signals informing the system about fidelity requirements enter the system in a fashion indistinguishable from sensory signals. (2) Optimal coding under non-uniform fidelity requirements is sufficient to account for the firing rate modulation observed during selective attention experiments. (3) The response of a single neuron cannot bewell characterized by measurements of attentional modulation of only a single sensory stimulus. (4) The magnitude of the activity modulation depends on the capacity of the neural circuit. A later chapter discusses the neural mechanisms for resource allocation, and the relation between attentional mechanisms and receptive field formation. The remainder of the dissertation focuses on overt orienting phenomena and active perception. We present a theoretical analysis of the allocation of resources during state estimation of multiple targets with different uncertainties, together with eye-tracking experiments that confirm our predictions. We finish by discussing the implications of these results to our current understanding of orienting phenomena and the neural code