Learning and Production of Movement Sequences: Behavioral, Neurophysiological, and Modeling Perspectives

A growing wave of behavioral studies, using a wide variety of paradigms that were introduced or greatly refined in recent years, has generated a new wealth of parametric observations about serial order behavior. What was a mere trickle of neurophysiological studies has grown to a more steady stream of probes of neural sites and mechanisms underlying sequential behavior. Moreover, simulation models of serial behavior generation have begun to open a channel to link cellular dynamics with cognitive and behavioral dynamics. Here we summarize the major results from prominent sequence learning and performance tasks, namely immediate serial recall, typing, 2XN, discrete sequence production, and serial reaction time. These populate a continuum from higher to lower degrees of internal control of sequential organization. The main movement classes covered are speech and keypressing, both involving small amplitude movements that are very amenable to parametric study. A brief synopsis of classes of serial order models, vis-à-vis the detailing of major effects found in the behavioral data, leads to a focus on competitive queuing (CQ) models. Recently, the many behavioral predictive successes of CQ models have been joined by successful prediction of distinctively patterend electrophysiological recordings in prefrontal cortex, wherein parallel activation dynamics of multiple neural ensembles strikingly matches the parallel dynamics predicted by CQ theory. An extended CQ simulation model-the N-STREAMS neural network model-is then examined to highlight issues in ongoing attemptes to accomodate a broader range of behavioral and neurophysiological data within a CQ-consistent theory. Important contemporary issues such as the nature of working memory representations for sequential behavior, and the development and role of chunks in hierarchial control are prominent throughout.

Self-Organizing Hierarchical Knowledge Discovery by an ARTMAP Image Fusion System

Classifying novel terrain or objects front sparse, complex data may require the resolution of conflicting information from sensors working at different times, locations, and scales, and from sources with different goals and situations. Information fusion methods can help resolve inconsistencies, as when evidence variously suggests that an object's class is car, truck, or airplane. The methods described here consider a complementary problem, supposing that information from sensors and experts is reliable though inconsistent, as when evidence suggests that an object's class is car, vehicle, and man-made. Underlying relationships among objects are assumed to be unknown to the automated system or the human user. The ARTMAP information fusion system used distributed code representations that exploit the neural network's capacity for one-to-many learning in order to produce self-organizing expert systems that discover hierarchical knowledge structures. The system infers multi-level relationships among groups of output classes, without any supervised labeling of these relationships.

Laminar Cortical Dynamics of 3D Surface Perception: Stratification, transparency, and Neon Color Spreading

How does the laminar organization of cortical circuitry in areas VI and V2 give rise to 3D percepts of stratification, transparency, and neon color spreading in response to 2D pictures and 3D scenes? Psychophysical experiments have shown that such 3D percepts are sensitive to whether contiguous image regions have the same relative contrast polarity (dark-light or lightdark), yet long-range perceptual grouping is known to pool over opposite contrast polarities. The ocularity of contiguous regions is also critical for neon color spreading: Having different ocularity despite the contrast relationship that favors neon spreading blocks the spread. In addition, half visible points in a stereogram can induce near-depth transparency if the contrast relationship favors transparency in the half visible areas. It thus seems critical to have the whole contrast relationship in a monocular configuration, since splitting it between two stereogram images cancels the effect. What adaptive functions of perceptual grouping enable it to both preserve sensitivity to monocular contrast and also to pool over opposite contrasts? Aspects of cortical development, grouping, attention, perceptual learning, stereopsis and 3D planar surface perception have previously been analyzed using a 3D LAMINART model of cortical areas VI, V2, and V4. The present work consistently extends this model to show how like-polarity competition between VI simple cells in layer 4 may be combined with other LAMINART grouping mechanisms, such as cooperative pooling of opposite polarities at layer 2/3 complex cells. The model also explains how the Metelli Rules can lead to transparent percepts, how bistable transparency percepts can arise in which either surface can be perceived as transparent, and how such a transparency reversal can be facilitated by an attention shift. The like-polarity inhibition prediction is consistent with lateral masking experiments in which two f1anking Gabor patches with the same contrast polarity as the target increase the target detection threshold when they approach the target. It is also consistent with LAMINART simulations of cortical development. Other model explanations and testable predictions will also be presented.

A Neuromorphic Model for Achromatic and Chromatic Surface Representation of Natural Images

This study develops a neuromorphic model of human lightness perception that is inspired by how the mammalian visual system is designed for this function. It is known that biological visual representations can adapt to a billion-fold change in luminance. How such a system determines absolute lightness under varying illumination conditions to generate a consistent interpretation of surface lightness remains an unsolved problem. Such a process, called "anchoring" of lightness, has properties including articulation, insulation, configuration, and area effects. The model quantitatively simulates such psychophysical lightness data, as well as other data such as discounting the illuminant, the double brilliant illusion, and lightness constancy and contrast effects. The model retina embodies gain control at retinal photoreceptors, and spatial contrast adaptation at the negative feedback circuit between mechanisms that model the inner segment of photoreceptors and interacting horizontal cells. The model can thereby adjust its sensitivity to input intensities ranging from dim moonlight to dazzling sunlight. A new anchoring mechanism, called the Blurred-Highest-Luminance-As-White (BHLAW) rule, helps simulate how surface lightness becomes sensitive to the spatial scale of objects in a scene. The model is also able to process natural color images under variable lighting conditions, and is compared with the popular RETINEX model.

From Parallel Sequence Representations to Calligraphic Control: A Conspiracy of Neural Circuits

Calligraphic writing presents a rich set of challenges to the human movement control system. These challenges include: initial learning, and recall from memory, of prescribed stroke sequences; critical timing of stroke onsets and durations; fine control of grip and contact forces; and letter-form invariance under voluntary size scaling, which entails fine control of stroke direction and amplitude during recruitment and derecruitment of musculoskeletal degrees of freedom. Experimental and computational studies in behavioral neuroscience have made rapid progress toward explaining the learning, planning and contTOl exercised in tasks that share features with calligraphic writing and drawing. This article summarizes computational neuroscience models and related neurobiological data that reveal critical operations spanning from parallel sequence representations to fine force control. Part one addresses stroke sequencing. It treats competitive queuing (CQ) models of sequence representation, performance, learning, and recall. Part two addresses letter size scaling and motor equivalence. It treats cursive handwriting models together with models in which sensory-motor tmnsformations are performed by circuits that learn inverse differential kinematic mappings. Part three addresses fine-grained control of timing and transient forces, by treating circuit models that learn to solve inverse dynamics problems.

Fast Synchronization of Perpetual Grouping in Laminar Visual Cortical Circuits

Perceptual grouping is well-known to be a fundamental process during visual perception, notably grouping across scenic regions that do not receive contrastive visual inputs. Illusory contours are a classical example of such groupings. Recent psychophysical and neurophysiological evidence have shown that the grouping process can facilitate rapid synchronization of the cells that are bound together by a grouping, even when the grouping must be completed across regions that receive no contrastive inputs. Synchronous grouping can hereby bind together different object parts that may have become desynchronized due to a variety of factors, and can enhance the efficiency of cortical transmission. Neural models of perceptual grouping have clarified how such fast synchronization may occur by using bipole grouping cells, whose predicted properties have been supported by psychophysical, anatomical, and neurophysiological experiments. These models have not, however, incorporated some of the realistic constraints on which groupings in the brain are conditioned, notably the measured spatial extent of long-range interactions in layer 2/3 of a grouping network, and realistic synaptic and axonal signaling delays within and across cells in different cortical layers. This work addresses the question: Can long-range interactions that obey the bipole constraint achieve fast synchronization under realistic anatomical and neurophysiological constraints that initially desynchronize grouping signals? Can the cells that synchronize retain their analog sensitivity to changing input amplitudes? Can the grouping process complete and synchronize illusory contours across gaps in bottom-up inputs? Our simulations show that the answer to these questions is Yes.

Neural Modeling and Imaging of the Cortical Interactions Underlying Syllable Production

This paper describes a neural model of speech acquisition and production that accounts for a wide range of acoustic, kinematic, and neuroimaging data concerning the control of speech movements. The model is a neural network whose components correspond to regions of the cerebral cortex and cerebellum, including premotor, motor, auditory, and somatosensory cortical areas. Computer simulations of the model verify its ability to account for compensation to lip and jaw perturbations during speech. Specific anatomical locations of the model's components are estimated, and these estimates are used to simulate fMRI experiments of simple syllable production with and without jaw perturbations.

Biologically Inspired Approaches to Automated Feature Extraction and Target Recognition

Ongoing research at Boston University has produced computational models of biological vision and learning that embody a growing corpus of scientific data and predictions. Vision models perform long-range grouping and figure/ground segmentation, and memory models create attentionally controlled recognition codes that intrinsically cornbine botton-up activation and top-down learned expectations. These two streams of research form the foundation of novel dynamically integrated systems for image understanding. Simulations using multispectral images illustrate road completion across occlusions in a cluttered scene and information fusion from incorrect labels that are simultaneously inconsistent and correct. The CNS Vision and Technology Labs (cns.bu.edulvisionlab and cns.bu.edu/techlab) are further integrating science and technology through analysis, testing, and development of cognitive and neural models for large-scale applications, complemented by software specification and code distribution.

SOVEREIGN: A Self-Organizing, Vision, Expectation, Recognition, Emotion, Intelligent, Goal-Oriented Navigation System

Both animals and mobile robots, or animats, need adaptive control systems to guide their movements through a novel environment. Such control systems need reactive mechanisms for exploration, and learned plans to efficiently reach goal objects once the environment is familiar. How reactive and planned behaviors interact together in real time, and arc released at the appropriate times, during autonomous navigation remains a major unsolved problern. This work presents an end-to-end model to address this problem, named SOVEREIGN: A Self-Organizing, Vision, Expectation, Recognition, Emotion, Intelligent, Goal-oriented Navigation system. The model comprises several interacting subsystems, governed by systems of nonlinear differential equations. As the animat explores the environment, a vision module processes visual inputs using networks that arc sensitive to visual form and motion. Targets processed within the visual form system arc categorized by real-time incremental learning. Simultaneously, visual target position is computed with respect to the animat's body. Estimates of target position activate a motor system to initiate approach movements toward the target. Motion cues from animat locomotion can elicit orienting head or camera movements to bring a never target into view. Approach and orienting movements arc alternately performed during animat navigation. Cumulative estimates of each movement, based on both visual and proprioceptive cues, arc stored within a motor working memory. Sensory cues are stored in a parallel sensory working memory. These working memories trigger learning of sensory and motor sequence chunks, which together control planned movements. Effective chunk combinations arc selectively enhanced via reinforcement learning when the animat is rewarded. The planning chunks effect a gradual transition from reactive to planned behavior. The model can read-out different motor sequences under different motivational states and learns more efficient paths to rewarded goals as exploration proceeds. Several volitional signals automatically gate the interactions between model subsystems at appropriate times. A 3-D visual simulation environment reproduces the animat's sensory experiences as it moves through a simplified spatial environment. The SOVEREIGN model exhibits robust goal-oriented learning of sequential motor behaviors. Its biomimctic structure explicates a number of brain processes which are involved in spatial navigation.

CEDI: A Neural Model of Colour Vision, with Applications to Image Processing and Classification

Air Force Office of Scientific Research (F49620-01-1-0423); National Geospatial-Intelligence Agency (NMA 201-01-1-2016); National Science Foundation (SBE-035437, DEG-0221680); Office of Naval Research (N00014-01-1-0624)

Laminar Cortical Dynamics of Visual Form and Motion Interactions During Coherent Object Motion Perception

How do visual form and motion processes cooperate to compute object motion when each process separately is insufficient? A 3D FORMOTION model specifies how 3D boundary representations, which separate figures from backgrounds within cortical area V2, capture motion signals at the appropriate depths in MT; how motion signals in MT disambiguate boundaries in V2 via MT-to-Vl-to-V2 feedback; how sparse feature tracking signals are amplified; and how a spatially anisotropic motion grouping process propagates across perceptual space via MT-MST feedback to integrate feature-tracking and ambiguous motion signals to determine a global object motion percept. Simulated data include: the degree of motion coherence of rotating shapes observed through apertures, the coherent vs. element motion percepts separated in depth during the chopsticks illusion, and the rigid vs. non-rigid appearance of rotating ellipses.

START: A Bridge between Emotion Theory and Neurobiology through Dynamic System Modeling

Lewis proposes "reconceptualization" (p. 1) of how to link the psychology and neurobiology of emotion and cognitive-emotional interactions. His main proposed themes have actually been actively and quantitatively developed in the neural modeling literature for over thirty years. This commentary summarizes some of these themes and points to areas of particularly active research in this area.

Modeling the Possible Influences of Eye Movements on the Refinement of Cortical Direction Selectivity

The second-order statistics of neural activity was examined in a model of the cat LGN and V1 during free-viewing of natural images. In the model, the specific patterns of thalamocortical activity required for a Bebbian maturation of direction-selective cells in VI were found during the periods of visual fixation, when small eye movements occurred, but not when natural images were examined in the absence of fixational eye movements. In addition, simulations of stroboscopic reming that replicated the abnormal pattern of eye movements observed in kittens chronically exposed to stroboscopic illumination produced results consistent with the reported loss of direction selectivity and preservation of orientation selectivity. These results suggest the involvement of the oculomotor activity of visual fixation in the maturation of cortical direction selectivity.

Fixational Instability and Natural Image Statistics: Implications for Early Visual Representations

Under natural viewing conditions small movements of the eye, head, and body prevent the maintenance of a steady direction of gaze. It is known that stimuli tend to fade when they a restabilized on the retina for several seconds. However; it is unclear whether the physiological motion of the retinal image serves a visual purpose during the brief periods of natural visual fixation. This study examines the impact of fixational instability on the statistics of the visua1 input to the retina and on the structure of neural activity in the early visual system. We show that fixational instability introduces a component in the retinal input signals that in the presence of natural images, lacks spatial correlations. This component strongly influences neural activity in a model of the LGN. It decorrelates cell responses even if the contrast sensitivity functions of simulated cells arc not perfectly tuned to counterbalance the power-law spectrum of natural images. A decorrelation of neural activity at the early stages of the visual system has been proposed to be beneficial for discarding statistical redundancies in the input signals. The results of this study suggest that fixational instability might contribute to establishing efficient representations of natural stimuli.

Estimating Sensor Motion from Wide-Field Optical Flow on a Log-Dipolar Sensor

Log-polar image architectures, motivated by the structure of the human visual field, have long been investigated in computer vision for use in estimating motion parameters from an optical flow vector field. Practical problems with this approach have been: (i) dependence on assumed alignment of the visual and motion axes; (ii) sensitivity to occlusion form moving and stationary objects in the central visual field, where much of the numerical sensitivity is concentrated; and (iii) inaccuracy of the log-polar architecture (which is an approximation to the central 20°) for wide-field biological vision. In the present paper, we show that an algorithm based on generalization of the log-polar architecture; termed the log-dipolar sensor, provides a large improvement in performance relative to the usual log-polar sampling. Specifically, our algorithm: (i) is tolerant of large misalignmnet of the optical and motion axes; (ii) is insensitive to significant occlusion by objects of unknown motion; and (iii) represents a more correct analogy to the wide-field structure of human vision. Using the Helmholtz-Hodge decomposition to estimate the optical flow vector field on a log-dipolar sensor, we demonstrate these advantages, using synthetic optical flow maps as well as natural image sequences.