A Local Circuit Model of Learned Straitial and Dopamine Cell Responses under Probabilistic Schedules of Reward

Before choosing, it helps to know both the expected value signaled by a predictive cue and the associated uncertainty that the reward will be forthcoming. Recently, Fiorillo et al. (2003) found the dopamine (DA) neurons of the SNc exhibit sustained responses related to the uncertainty that a cure will be followed by reward, in addition to phasic responses related to reward prediction errors (RPEs). This suggests that cue-dependent anticipations of the timing, magnitude, and uncertainty of rewards are learned and reflected in components of the DA signals broadcast by SNc neurons. What is the minimal local circuit model that can explain such multifaceted reward-related learning? A new computational model shows how learned uncertainty responses emerge robustly on single trial along with phasic RPE responses, such that both types of DA responses exhibit the empirically observed dependence on conditional probability, expected value of reward, and time since onset of the reward-predicting cue. The model includes three major pathways for computing: immediate expected values of cures, timed predictions of reward magnitudes (and RPEs), and the uncertainty associated with these predictions. The first two model pathways refine those previously modeled by Brown et al. (1999). A third, newly modeled, pathway is formed by medium spiny projection neurons (MSPNs) of the matrix compartment of the striatum, whose axons co-release GABA and a neuropeptide, substance P, both at synapses with GABAergic neurons in the SNr and with the dendrites (in SNr) of DA neurons whose somas are in ventral SNc. Co-release enables efficient computation of sustained DA uncertainty responses that are a non-monotonic function of the conditonal probability that a reward will follow the cue. The new model's incorporation of a striatal microcircuit allowed it to reveals that variability in striatal cholinergic transmission can explain observed difference, between monkeys, in the amplitutude of the non-monotonic uncertainty function. Involvement of matriceal MSPNs and striatal cholinergic transmission implpies a relation between uncertainty in the cue-reward contigency and action-selection functions of the basal ganglia. The model synthesizes anatomical, electrophysiological and behavioral data regarding the midbrain DA system in a novel way, by relating the ability to compute uncertainty, in parallel with other aspects of reward contingencies, to the unique distribution of SP inputs in ventral SN.

CONFIGR: A Vision-Based Model for Long-Range Figure Completion

CONFIGR (CONtour FIgure GRound) is a computational model based on principles of biological vision that completes sparse and noisy image figures. Within an integrated vision/recognition system, CONFIGR posits an initial recognition stage which identifies figure pixels from spatially local input information. The resulting, and typically incomplete, figure is fed back to the “early vision” stage for long-range completion via filling-in. The reconstructed image is then re-presented to the recognition system for global functions such as object recognition. In the CONFIGR algorithm, the smallest independent image unit is the visible pixel, whose size defines a computational spatial scale. Once pixel size is fixed, the entire algorithm is fully determined, with no additional parameter choices. Multi-scale simulations illustrate the vision/recognition system. Open-source CONFIGR code is available online, but all examples can be derived analytically, and the design principles applied at each step are transparent. The model balances filling-in as figure against complementary filling-in as ground, which blocks spurious figure completions. Lobe computations occur on a subpixel spatial scale. Originally designed to fill-in missing contours in an incomplete image such as a dashed line, the same CONFIGR system connects and segments sparse dots, and unifies occluded objects from pieces locally identified as figure in the initial recognition stage. The model self-scales its completion distances, filling-in across gaps of any length, where unimpeded, while limiting connections among dense image-figure pixel groups that already have intrinsic form. Long-range image completion promises to play an important role in adaptive processors that reconstruct images from highly compressed video and still camera images.

A Unified Model of Spatiotemporal Processing in the Retina

A computational model of visual processing in the vertebrate retina provides a unified explanation of a range of data previously treated by disparate models. Three results are reported here: the model proposes a functional explanation for the primary feed-forward retinal circuit found in vertebrate retinae, it shows how this retinal circuit combines nonlinear adaptation with the desirable properties of linear processing, and it accounts for the origin of parallel transient (nonlinear) and sustained (linear) visual processing streams as simple variants of the same retinal circuit. The retina, owing to its accessibility and to its fundamental role in the initial transduction of light into neural signals, is among the most extensively studied neural structures in the nervous system. Since the pioneering anatomical work by Ramón y Cajal at the turn of the last century[1], technological advances have abetted detailed descriptions of the physiological, pharmacological, and functional properties of many types of retinal cells. However, the relationship between structure and function in the retina is still poorly understood. This article outlines a computational model developed to address fundamental constraints of biological visual systems. Neurons that process nonnegative input signals-such as retinal illuminance-are subject to an inescapable tradeoff between accurate processing in the spatial and temporal domains. Accurate processing in both domains can be achieved with a model that combines nonlinear mechanisms for temporal and spatial adaptation within three layers of feed-forward processing. The resulting architecture is structurally similar to the feed-forward retinal circuit connecting photoreceptors to retinal ganglion cells through bipolar cells. This similarity suggests that the three-layer structure observed in all vertebrate retinae[2] is a required minimal anatomy for accurate spatiotemporal visual processing. This hypothesis is supported through computer simulations showing that the model's output layer accounts for many properties of retinal ganglion cells[3],[4],[5],[6]. Moreover, the model shows how the retina can extend its dynamic range through nonlinear adaptation while exhibiting seemingly linear behavior in response to a variety of spatiotemporal input stimuli. This property is the basis for the prediction that the same retinal circuit can account for both sustained (X) and transient (Y) cat ganglion cells[7] by simple morphological changes. The ability to generate distinct functional behaviors by simple changes in cell morphology suggests that different functional pathways originating in the retina may have evolved from a unified anatomy designed to cope with the constraints of low-level biological vision.

KInNeSS: A Modular Framework for Computational Neuroscience

Making use of very detailed neurophysiological, anatomical, and behavioral data to build biological-realistic computational models of animal behavior is often a difficult task. Until recently, many software packages have tried to resolve this mismatched granularity with different approaches. This paper presents KInNeSS, the KDE Integrated NeuroSimulation Software environment, as an alternative solution to bridge the gap between data and model behavior. This open source neural simulation software package provides an expandable framework incorporating features such as ease of use, scalabiltiy, an XML based schema, and multiple levels of granularity within a modern object oriented programming design. KInNeSS is best suited to simulate networks of hundreds to thousands of branched multu-compartmental neurons with biophysical properties such as membrane potential, voltage-gated and ligand-gated channels, the presence of gap junctions of ionic diffusion, neuromodulation channel gating, the mechanism for habituative or depressive synapses, axonal delays, and synaptic plasticity. KInNeSS outputs include compartment membrane voltage, spikes, local-field potentials, and current source densities, as well as visualization of the behavior of a simulated agent. An explanation of the modeling philosophy and plug-in development is also presented. Further developement of KInNeSS is ongoing with the ultimate goal of creating a modular framework that will help researchers across different disciplines to effecitively collaborate using a modern neural simulation platform.

KInNeSS: A Modular Framework for Computational Neuroscience

Making use of very detailed neurophysiological, anatomical, and behavioral data to build biologically-realistic computational models of animal behavior is often a difficult task. Until recently, many software packages have tried to resolve this mismatched granularity with different approaches. This paper presents KInNeSS, the KDE Integrated NeuroSimulation Software environment, as an alternative solution to bridge the gap between data and model behavior. This open source neural simulation software package provides an expandable framework incorporating features such as ease of use, scalability, an XML based schema, and multiple levels of granularity within a modern object oriented programming design. KInNeSS is best suited to simulate networks of hundreds to thousands of branched multi-compartmental neurons with biophysical properties such as membrane potential, voltage-gated and ligand-gated channels, the presence of gap junctions or ionic diffusion, neuromodulation channel gating, the mechanism for habituative or depressive synapses, axonal delays, and synaptic plasticity. KInNeSS outputs include compartment membrane voltage, spikes, local-field potentials, and current source densities, as well as visualization of the behavior of a simulated agent. An explanation of the modeling philosophy and plug-in development is also presented. Further development of KInNeSS is ongoing with the ultimate goal of creating a modular framework that will help researchers across different disciplines to effectively collaborate using a modern neural simulation platform.

Simulating Effects of Learning and Lesions with a Model of Intrinsic and Synaptically Gated Responses of Striatal Cholinergic Interneurons

The giant cholinergic interneurons of the striatum are tonically active neurons (TANs) that respond with characteristic pauses to novel events and to appetitive and aversive conditioned stimuli. Fluctuations in acetylcholine release by TANs modulate performance- and learning-related dynamics in the striatum. Whereas tonic activity emerges from intrinsic properties of these neurons, glutamatergic inputs from thalamic centromedian-parafascicular nuclei, and dopaminergic inputs from midbrain, are required for the generation of pause responses. No prior computational models encompass both intrinsic and synaptically-gated dynamics. We present a mathematical model that robustly accounts for behavior-related electrophysiological properties of TANs in terms of their intrinsic physiological properties and known afferents. In the model, balanced intrinsic hyperpolarizing and depolarizing currents engender tonic firing, and glutamatergic inputs from thalamus (and cortex) both directly excite and indirectly inhibit TANs. If the latter inhibition, presumably mediated by GABAergic interneurons, exceeds a threshold, its effect is amplified by a KIR current to generate a prolonged pause. In the model, the intrinsic mechanisms and external inputs are both modulated by learning-dependent dopamine (DA) signals and our simulations revealed that many learning-dependent behaviors of TANs are explicable without recourse to learning-dependent changes in synapses onto TANs. The "teaching signal" that modulates reinforcement learning at cortico-striatal synapses may be a sequence composed of an adaptively scaled DA burst, a brief ACh burst, and a scaled ACh pause. Such an interpretation is consistent with recent data on cholinergic control of LTD of cortical synapses onto striatal spiny projection neurons.