Spikes, Synchrony, and Attentive Learning by Laminar Thalamocortical Circuits

This article develops the Synchronous Matching Adaptive Resonance Theory (SMART) neural model to explain how the brain may coordinate multiple levels of thalamocortical and corticocortical processing to rapidly learn, and stably remember, important information about a changing world. The model clarifies how bottom-up and top-down processes work together to realize this goal, notably how processes of learning, expectation, attention, resonance, and synchrony are coordinated. The model hereby clarifies, for the first time, how the following levels of brain organization coexist to realize cognitive processing properties that regulate fast learning and stable memory of brain representations: single cell properties, such as spiking dynamics, spike-timing-dependent plasticity (STDP), and acetylcholine modulation; detailed laminar thalamic and cortical circuit designs and their interactions; aggregate cell recordings, such as current-source densities and local field potentials; and single cell and large-scale inter-areal oscillations in the gamma and beta frequency domains. In particular, the model predicts how laminar circuits of multiple cortical areas interact with primary and higher-order specific thalamic nuclei and nonspecific thalamic nuclei to carry out attentive visual learning and information processing. The model simulates how synchronization of neuronal spiking occurs within and across brain regions, and triggers STDP. Matches between bottom-up adaptively filtered input patterns and learned top-down expectations cause gamma oscillations that support attention, resonance, and learning. Mismatches inhibit learning while causing beta oscillations during reset and hypothesis testing operations that are initiated in the deeper cortical layers. The generality of learned recognition codes is controlled by a vigilance process mediated by acetylcholine.

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.

Neural Dynamics Underlying Impaired Autonomic and Conditioned Responses Following Amygdala and Orbitofrontal Lesions

A neural model is presented that explains how outcome-specific learning modulates affect, decision-making and Pavlovian conditioned approach responses. The model addresses how brain regions responsible for affective learning and habit learning interact, and answers a central question: What are the relative contributions of the amygdala and orbitofrontal cortex to emotion and behavior? In the model, the amygdala calculates outcome value while the orbitofrontal cortex influences attention and conditioned responding by assigning value information to stimuli. Model simulations replicate autonomic, electrophysiological, and behavioral data associated with three tasks commonly used to assay these phenomena: Food consumption, Pavlovian conditioning, and visual discrimination. Interactions of the basal ganglia and amygdala with sensory and orbitofrontal cortices enable the model to replicate the complex pattern of spared and impaired behavioral and emotional capacities seen following lesions of the amygdala and orbitofrontal cortex.

A Neural Model of Visually Guided Steering, Obstacle Avoidance, and Route Selection

A neural model is developed to explain how humans can approach a goal object on foot while steering around obstacles to avoid collisions in a cluttered environment. The model uses optic flow from a 3D virtual reality environment to determine the position of objects based on motion discontinuities, and computes heading direction, or the direction of self-motion, from global optic flow. The cortical representation of heading interacts with the representations of a goal and obstacles such that the goal acts as an attractor of heading, while obstacles act as repellers. In addition the model maintains fixation on the goal object by generating smooth pursuit eye movements. Eye rotations can distort the optic flow field, complicating heading perception, and the model uses extraretinal signals to correct for this distortion and accurately represent heading. The model explains how motion processing mechanisms in cortical areas MT, MST, and posterior parietal cortex can be used to guide steering. The model quantitatively simulates human psychophysical data about visually-guided steering, obstacle avoidance, and route selection.

Neural Dynamics of Motion Perception, Recognition Learning, and Spatial Attention

Advanced Research Projects Agency (ONR N00014-92-J-4015); National Science Foundation (IRI-90-24877); Office of Naval Research (N00014-91-J-4100)

Integrating Symbolic and Neural Processing in a Self-Organizing Architechture for Pattern Recognition and Prediction

British Petroleum (89A-1204); Defense Advanced Research Projects Agency (N00014-92-J-4015); National Science Foundation (IRI-90-00530); Office of Naval Research (N00014-91-J-4100); Air Force Office of Scientific Research (F49620-92-J-0225)

Boundary, Brightness, and Depth Interactions During Preattentive Representation and Attentive Recognition of Figure and Ground

This article applies a recent theory of 3-D biological vision, called FACADE Theory, to explain several percepts which Kanizsa pioneered. These include 3-D pop-out of an occluding form in front of an occluded form, leading to completion and recognition of the occluded form; 3-D transparent and opaque percepts of Kanizsa squares, with and without Varin wedges; and interactions between percepts of illusory contours, brightness, and depth in response to 2-D Kanizsa images. These explanations clarify how a partially occluded object representation can be completed for purposes of object recognition, without the completed part of the representation necessarily being seen. The theory traces these percepts to neural mechanisms that compensate for measurement uncertainty and complementarity at individual cortical processing stages by using parallel and hierarchical interactions among several cortical processing stages. These interactions are modelled by a Boundary Contour System (BCS) that generates emergent boundary segmentations and a complementary Feature Contour System (FCS) that fills-in surface representations of brightness, color, and depth. The BCS and FCS interact reciprocally with an Object Recognition System (ORS) that binds BCS boundary and FCS surface representations into attentive object representations. The BCS models the parvocellular LGN→Interblob→Interstripe→V4 cortical processing stream, the FCS models the parvocellular LGN→Blob→Thin Stripe→V4 cortical processing stream, and the ORS models inferotemporal cortex.

Neural Control of Interlimb Coordination and Gait Timing in Bipeds and Quadrupeds

1) A large body of behavioral data conceming animal and human gaits and gait transitions is simulated as emergent properties of a central pattern generator (CPG) model. The CPG model incorporates neurons obeying Hodgkin-Huxley type dynamics that interact via an on-center off-surround anatomy whose excitatory signals operate on a faster time scale than their inhibitory signals. A descending cornmand or arousal signal called a GO signal activates the gaits and controL their transitions. The GO signal and the CPG model are compared with neural data from globus pallidus and spinal cord, among other brain structures. 2) Data from human bimanual finger coordination tasks are simulated in which anti-phase oscillations at low frequencies spontaneously switch to in-phase oscillations at high frequencies, in-phase oscillations can be performed both at low and high frequencies, phase fluctuations occur at the anti-phase in-phase transition, and a "seagull effect" of larger errors occurs at intermediate phases. When driven by environmental patterns with intermediate phase relationships, the model's output exhibits a tendency to slip toward purely in-phase and anti-phase relationships as observed in humans subjects. 3) Quadruped vertebrate gaits, including the amble, the walk, all three pairwise gaits (trot, pace, and gallop) and the pronk are simulated. Rapid gait transitions are simulated in the order--walk, trot, pace, and gallop--that occurs in the cat, along with the observed increase in oscillation frequency. 4) Precise control of quadruped gait switching is achieved in the model by using GO-dependent modulation of the model's inhibitory interactions. This generates a different functional connectivity in a single CPG at different arousal levels. Such task-specific modulation of functional connectivity in neural pattern generators has been experimentally reported in invertebrates. Phase-dependent modulation of reflex gain has been observed in cats. A role for state-dependent modulation is herein predicted to occur in vertebrates for precise control of phase transitions from one gait to another. 5) The primary human gaits (the walk and the run) and elephant gaits (the amble and the walk) are sirnulated. Although these two gaits are qualitatively different, they both have the same limb order and may exhibit oscillation frequencies that overlap. The CPG model simulates the walk and the run by generating oscillations which exhibit the same phase relationships. but qualitatively different waveform shapes, at different GO signal levels. The fraction of each cycle that activity is above threshold quantitatively distinguishes the two gaits, much as the duty cycles of the feet are longer in the walk than in the run. 6) A key model properly concerns the ability of a single model CPG, that obeys a fixed set of opponent processing equations to generate both in-phase and anti-phase oscillations at different arousal levels. Phase transitions from either in-phase to anti-phase oscillations, or from anti-phase to in-phase oscillations, can occur in different parameter ranges, as the GO signal increases.

An Active Pattern Recognition Architecture for Mobile Robots

An active, attentionally-modulated recognition architecture is proposed for object recognition and scene analysis. The proposed architecture forms part of navigation and trajectory planning modules for mobile robots. Key characteristics of the system include movement planning and execution based on environmental factors and internal goal definitions. Real-time implementation of the system is based on space-variant representation of the visual field, as well as an optimal visual processing scheme utilizing separate and parallel channels for the extraction of boundaries and stimulus qualities. A spatial and temporal grouping module (VWM) allows for scene scanning, multi-object segmentation, and featural/object priming. VWM is used to modulate a tn~ectory formation module capable of redirecting the focus of spatial attention. Finally, an object recognition module based on adaptive resonance theory is interfaced through VWM to the visual processing module. The system is capable of using information from different modalities to disambiguate sensory input.

Fusion Artmap: A Neural Network Architecture for Multi-Channel Data Fusion and Classification

Fusion ARTMAP is a self-organizing neural network architecture for multi-channel, or multi-sensor, data fusion. Single-channel Fusion ARTMAP is functionally equivalent to Fuzzy ART during unsupervised learning and to Fuzzy ARTMAP during supervised learning. The network has a symmetric organization such that each channel can be dynamically configured to serve as either a data input or a teaching input to the system. An ART module forms a compressed recognition code within each channel. These codes, in turn, become inputs to a single ART system that organizes the global recognition code. When a predictive error occurs, a process called paraellel match tracking simultaneously raises vigilances in multiple ART modules until reset is triggered in one of them. Parallel match tracking hereby resets only that portion of the recognition code with the poorest match, or minimum predictive confidence. This internally controlled selective reset process is a type of credit assignment that creates a parsimoniously connected learned network. Fusion ARTMAP's multi-channel coding is illustrated by simulations of the Quadruped Mammal database.

Variable Rate Working Memories for Phonetic Categorization and Invariant Speech Perception

Speech can be understood at widely varying production rates. A working memory is described for short-term storage of temporal lists of input items. The working memory is a cooperative-competitive neural network that automatically adjusts its integration rate, or gain, to generate a short-term memory code for a list that is independent of item presentation rate. Such an invariant working memory model is used to simulate data of Repp (1980) concerning the changes of phonetic category boundaries as a function of their presentation rate. Thus the variability of categorical boundaries can be traced to the temporal in variance of the working memory code.

Cerebellar Learning in an Opponent Motor Controller for Adaptive Load Compensation and Synergy Formation

This paper shows how a minimal neural network model of the cerebellum may be embedded within a sensory-neuro-muscular control system that mimics known anatomy and physiology. With this embedding, cerebellar learning promotes load compensation while also allowing both coactivation and reciprocal inhibition of sets of antagonist muscles. In particular, we show how synaptic long term depression guided by feedback from muscle stretch receptors can lead to trans-cerebellar gain changes that are load-compensating. It is argued that the same processes help to adaptively discover multi-joint synergies. Simulations of rapid single joint rotations under load illustrates design feasibility and stability.

The VITEWRITE Model of Handwriting Production

This article describes the VITEWRITE model for generating handwriting movements. The model consists of a sequential controller, or motor program, that interacts with a trajectory generator to move a hand with redundant degrees of freedom. The neural trajectory generator is the Vector Integration to Endpoint (VITE) model for synchronous variable-speed control of multijoint movements. VITE properties enable a simple control strategy to generate complex handwritten script if the hand model contains redundant degrees of freedom. The controller launches transient directional commands to independent hand synergies at times when the hand begins to move, or when a velocity peak in the outflow command to a given synergy occurs. The VITE model translates these temporally disjoint synergy commands into smooth curvilinear trajectories among temporally overlapping synergetic movements. Each synergy exhibits a unimodal velocity profile during any stroke, generates letters that are invariant under speed and size rescaling, and enables effortless connection of letter shapes into words. Speed and size rescaling are achieved by scalar GO and GRO signals that express computationally simple volitional commands. Psychophysical data such as the isochrony principle, asymmetric velocity profiles, and the two-thirds power law relating movement curvature and velocity arise as emergent properties of model interactions.

Evaluation of Speaker Normalization Methods for Vowel Recognition Using Fuzzy ARTMAP and K-NN

A procedure that uses fuzzy ARTMAP and K-Nearest Neighbor (K-NN) categorizers to evaluate intrinsic and extrinsic speaker normalization methods is described. Each classifier is trained on preprocessed, or normalized, vowel tokens from about 30% of the speakers of the Peterson-Barney database, then tested on data from the remaining speakers. Intrinsic normalization methods included one nonscaled, four psychophysical scales (bark, bark with end-correction, mel, ERB), and three log scales, each tested on four different combinations of the fundamental (Fo) and the formants (F1 , F2, F3). For each scale and frequency combination, four extrinsic speaker adaptation schemes were tested: centroid subtraction across all frequencies (CS), centroid subtraction for each frequency (CSi), linear scale (LS), and linear transformation (LT). A total of 32 intrinsic and 128 extrinsic methods were thus compared. Fuzzy ARTMAP and K-NN showed similar trends, with K-NN performing somewhat better and fuzzy ARTMAP requiring about 1/10 as much memory. The optimal intrinsic normalization method was bark scale, or bark with end-correction, using the differences between all frequencies (Diff All). The order of performance for the extrinsic methods was LT, CSi, LS, and CS, with fuzzy AHTMAP performing best using bark scale with Diff All; and K-NN choosing psychophysical measures for all except CSi.

Fuzzy ARTMAP, Slow Learning and Probability Estimation

A nonparametric probability estimation procedure using the fuzzy ARTMAP neural network is here described. Because the procedure does not make a priori assumptions about underlying probability distributions, it yields accurate estimates on a wide variety of prediction tasks. Fuzzy ARTMAP is used to perform probability estimation in two different modes. In a 'slow-learning' mode, input-output associations change slowly, with the strength of each association computing a conditional probability estimate. In 'max-nodes' mode, a fixed number of categories are coded during an initial fast learning interval, and weights are then tuned by slow learning. Simulations illustrate system performance on tasks in which various numbers of clusters in the set of input vectors mapped to a given class.