We introduce the notion of Morton-style factorial coding and illustrate how it may help understand information integration and perceptual coding inthe brain. We show that by focusing on average responses one may miss the existence of factorial coding mechanisms that become only apparent when analyzing spike count histograms. We show evidence suggesting that the classical/nonclassical receptive field organization in the cortex effectively enforces the development of Morton-style factorial codes. This may provide some cues to help understand perceptual coding inthe brain and to develop new unsupervised learning algorithms. While methods like ICA (Bell & Sejnowski, 1997) develop independent codes, in Morton-style coding the goal is to make two or more external aspects of the world become independent when conditioning on internal representations.

A population of neurons typically exhibits a broad diversity of responses to sensory inputs. The intuitive notion of functional classification is that cells can be clustered so that most of the diversity is captured by the identity ofthe clusters rather than by individuals within clusters. We show how this intuition can be made precise using information theory, without anyneed to introduce a metric on the space of stimuli or responses. Applied to the retinal ganglion cells of the salamander, this approach recovers classicalresults, but also provides clear evidence for subclasses beyond those identified previously. Further, we find that each of the ganglion cellsis functionally unique, and that even within the same subclass only a few spikes are needed to reliably distinguish between cells.

Here we derive optimal gain functions for minimum mean square reconstruction fromneural rate responses subjected to Poisson noise. The shape of these functions strongly depends on the length T of the time window within which spikes are counted in order to estimate the underlying firingrate. A phase transition towards pure binary encoding occurs if the maximum mean spike count becomes smaller than approximately three provided the minimum firing rate is zero. For a particular function class, we were able to prove the existence of a second-order phase transition analytically.The critical decoding time window length obtained from the analytical derivation is in precise agreement with the numerical results. We conclude that under most circumstances relevant to information processingin the brain, rate coding can be better ascribed to a binary (low-entropy) code than to the other extreme ofrich analog coding. 1 Optimal neuronal gain functions for short decoding time windows The use of action potentials (spikes) as a means of communication is the striking feature of neurons in the central nervous system.

Single unit activity in the striatum of awake monkeys shows a marked dependence on the expected reward that a behavior will elicit. We present a computational model of spiny neurons, the principal neurons of the striatum, to assess the hypothesis that direct neuromodulatoryeffects of dopamine through the activation of D1 receptors mediate the reward dependency of spiny neuron activity. Dopamine release results in the amplification of key ion currents, leading to the emergence of bistability, which not only modulates the peak firing rate but also introduces a temporal and state dependence of the model's response, thus improving the detectability oftemporally correlated inputs. 1 Introduction The classic notion of the basal ganglia as being involved in purely motor processing has expanded over the years to include sensory and cognitive functions. A surprising newfinding is that much of this activity shows a motivational component. For instance, striatal activity related to visual stimuli is dependent on the type of reinforcement (primary vs secondary) that a behavior will elicit [1].

Experimental and theoretical studies suggest that these different forms of variability play different behavioral, neural and computational roles, and may be reported by different (notably neuromodulatory) systems. Here, we refine ourprevious theory of acetylcholine's role in cortical inference in the (oxymoronic) terms of expected uncertainty, and advocate a theory for norepinephrine in terms of unexpected uncertainty. We suggest that norepinephrine reports the radical divergence of bottom-up inputs from prevailing top-down interpretations, to influence inference and plasticity. We illustrate this proposal using an adaptive factor analysis model.

We consider a statistical framework for learning in a class of networks ofspiking neurons. Our aim is to show how optimal local learning rules can be readily derived once the neural dynamics and desired functionality of the neural assembly have been specified, in contrast to other models which assume (sub-optimal) learning rules. Within this framework we derive local rules for learning temporal sequencesin a model of spiking neurons and demonstrate its superior performance to correlation (Hebbian) based approaches. We further show how to include mechanisms such as synaptic depression andoutline how the framework is readily extensible to learning in networks of highly complex spiking neurons. A stochastic quantalvesicle release mechanism is considered and implications on the complexity of learning discussed.

We show that two important properties of the primary visual cortex emerge when the principle of temporal coherence is applied to natural image sequences. The properties are simple-cell-like receptive fields and complex-cell-like pooling of simple cell outputs, which emerge when we apply two different approaches to temporal coherence. In the first approach we extract receptive fields whose outputs are as temporally coherent aspossible. This approach yields simple-cell-like receptive fields (oriented, localized, multiscale). Thus, temporal coherence is an alternative tosparse coding in modeling the emergence of simple cell receptive fields. The second approach is based on a two-layer statistical generative model of natural image sequences. In addition to modeling the temporal coherence of individual simple cells, this model includes inter-cell temporal dependencies.Estimation of this model from natural data yields both simple-cell-like receptive fields, and complex-cell-like pooling of simple cell outputs. In this completely unsupervised learning, both layers ofthe generative model are estimated simultaneously from scratch. This is a significant improvement on earlier statistical models of early vision, where only one layer has been learned, and others have been fixed a priori.

Inner-product operators, often referred to as kernels in statistical learning, define amapping from some input space into a feature space. The focus of this paper is the construction of biologically-motivated kernels for cortical activities. Thekernels we derive, termed Spikernels, map spike count sequences into an abstract vector space in which we can perform various prediction tasks. We discuss in detail the derivation of Spikernels and describe an efficient algorithm forcomputing their value on any two sequences of neural population spike counts. We demonstrate the merits of our modeling approach using the Spikernel and various standard kernels for the task of predicting hand movement velocitiesfrom cortical recordings. In all of our experiments all the kernels we tested outperform the standard scalar product used in regression with the Spikernel consistently achieving the best performance.

By comparison to some other sensory cortices, the functional properties ofcells in the primary auditory cortex are not yet well understood. Recent attempts to obtain a generalized description of auditory cortical responses have often relied upon characterization of the spectrotemporal receptivefield (STRF), which amounts to a model of the stimulusresponse function(SRF) that is linear in the spectrogram of the stimulus.

Cortical neurons have been reported to use both rate and temporal codes. Here we describe a novel mode in which each neuron generates exactly 0 or 1 action potentials, but not more, in response to a stimulus. We used cell-attached recording, which ensured single-unit isolation, to record responses in rat auditory cortex to brief tone pips. Surprisingly, the majority of neurons exhibited binary behavior with few multi-spike responses; several dramatic examples consisted of exactly one spike on 100% of trials, with no trial-to-trial variability in spike count. Many neurons were tuned to stimulus frequency. Since individual trials yielded at most one spike for most neurons, the information about stimulus frequency was encoded in the population, and would not have been accessible to later stages of processing that only had access to the activity of a single unit. These binary units allow a more efficient population code than is possible with conventional rate coding units, and are consistent with a model of cortical processing in which synchronous packets of spikes propagate stably from one neuronal population to the next.