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.

Vector Associative Maps: Unsupervised Real-time Error-based Learning and Control of Movement Trajectories

This article describes neural network models for adaptive control of arm movement trajectories during visually guided reaching and, more generally, a framework for unsupervised real-time error-based learning. The models clarify how a child, or untrained robot, can learn to reach for objects that it sees. Piaget has provided basic insights with his concept of a circular reaction: As an infant makes internally generated movements of its hand, the eyes automatically follow this motion. A transformation is learned between the visual representation of hand position and the motor representation of hand position. Learning of this transformation eventually enables the child to accurately reach for visually detected targets. Grossberg and Kuperstein have shown how the eye movement system can use visual error signals to correct movement parameters via cerebellar learning. Here it is shown how endogenously generated arm movements lead to adaptive tuning of arm control parameters. These movements also activate the target position representations that are used to learn the visuo-motor transformation that controls visually guided reaching. The AVITE model presented here is an adaptive neural circuit based on the Vector Integration to Endpoint (VITE) model for arm and speech trajectory generation of Bullock and Grossberg. In the VITE model, a Target Position Command (TPC) represents the location of the desired target. The Present Position Command (PPC) encodes the present hand-arm configuration. The Difference Vector (DV) population continuously.computes the difference between the PPC and the TPC. A speed-controlling GO signal multiplies DV output. The PPC integrates the (DV)·(GO) product and generates an outflow command to the arm. Integration at the PPC continues at a rate dependent on GO signal size until the DV reaches zero, at which time the PPC equals the TPC. The AVITE model explains how self-consistent TPC and PPC coordinates are autonomously generated and learned. Learning of AVITE parameters is regulated by activation of a self-regulating Endogenous Random Generator (ERG) of training vectors. Each vector is integrated at the PPC, giving rise to a movement command. The generation of each vector induces a complementary postural phase during which ERG output stops and learning occurs. Then a new vector is generated and the cycle is repeated. This cyclic, biphasic behavior is controlled by a specialized gated dipole circuit. ERG output autonomously stops in such a way that, across trials, a broad sample of workspace target positions is generated. When the ERG shuts off, a modulator gate opens, copying the PPC into the TPC. Learning of a transformation from TPC to PPC occurs using the DV as an error signal that is zeroed due to learning. This learning scheme is called a Vector Associative Map, or VAM. The VAM model is a general-purpose device for autonomous real-time error-based learning and performance of associative maps. The DV stage serves the dual function of reading out new TPCs during performance and reading in new adaptive weights during learning, without a disruption of real-time operation. YAMs thus provide an on-line unsupervised alternative to the off-line properties of supervised error-correction learning algorithms. YAMs and VAM cascades for learning motor-to-motor and spatial-to-motor maps are described. YAM models and Adaptive Resonance Theory (ART) models exhibit complementary matching, learning, and performance properties that together provide a foundation for designing a total sensory-cognitive and cognitive-motor autonomous system.

Adaptive Neural Networks for Control of Movement Trajectories Invariant under Speed and Force Rescaling

This article describes two neural network modules that form part of an emerging theory of how adaptive control of goal-directed sensory-motor skills is achieved by humans and other animals. The Vector-Integration-To-Endpoint (VITE) model suggests how synchronous multi-joint trajectories are generated and performed at variable speeds. The Factorization-of-LEngth-and-TEnsion (FLETE) model suggests how outflow movement commands from a VITE model may be performed at variable force levels without a loss of positional accuracy. The invariance of positional control under speed and force rescaling sheds new light upon a familiar strategy of motor skill development: Skill learning begins with performance at low speed and low limb compliance and proceeds to higher speeds and compliances. The VITE model helps to explain many neural and behavioral data about trajectory formation, including data about neural coding within the posterior parietal cortex, motor cortex, and globus pallidus, and behavioral properties such as Woodworth's Law, Fitts Law, peak acceleration as a function of movement amplitude and duration, isotonic arm movement properties before and after arm-deafferentation, central error correction properties of isometric contractions, motor priming without overt action, velocity amplification during target switching, velocity profile invariance across different movement distances, changes in velocity profile asymmetry across different movement durations, staggered onset times for controlling linear trajectories with synchronous offset times, changes in the ratio of maximum to average velocity during discrete versus serial movements, and shared properties of arm and speech articulator movements. The FLETE model provides new insights into how spina-muscular circuits process variable forces without a loss of positional control. These results explicate the size principle of motor neuron recruitment, descending co-contractive compliance signals, Renshaw cells, Ia interneurons, fast automatic reactive control by ascending feedback from muscle spindles, slow adaptive predictive control via cerebellar learning using muscle spindle error signals to train adaptive movement gains, fractured somatotopy in the opponent organization of cerebellar learning, adaptive compensation for variable moment-arms, and force feedback from Golgi tendon organs. More generally, the models provide a computational rationale for the use of nonspecific control signals in volitional control, or "acts of will", and of efference copies and opponent processing in both reactive and adaptive motor control tasks.

A Self-Organizing Neural Model of Motor Equivalent Reaching and Tool Use by a Multijoint Arm

This paper describes a self-organizing neural model for eye-hand coordination. Called the DIRECT model, it embodies a solution of the classical motor equivalence problem. Motor equivalence computations allow humans and other animals to flexibly employ an arm with more degrees of freedom than the space in which it moves to carry out spatially defined tasks under conditions that may require novel joint configurations. During a motor babbling phase, the model endogenously generates movement commands that activate the correlated visual, spatial, and motor information that are used to learn its internal coordinate transformations. After learning occurs, the model is capable of controlling reaching movements of the arm to prescribed spatial targets using many different combinations of joints. When allowed visual feedback, the model can automatically perform, without additional learning, reaches with tools of variable lengths, with clamped joints, with distortions of visual input by a prism, and with unexpected perturbations. These compensatory computations occur within a single accurate reaching movement. No corrective movements are needed. Blind reaches using internal feedback have also been simulated. The model achieves its competence by transforming visual information about target position and end effector position in 3-D space into a body-centered spatial representation of the direction in 3-D space that the end effector must move to contact the target. The spatial direction vector is adaptively transformed into a motor direction vector, which represents the joint rotations that move the end effector in the desired spatial direction from the present arm configuration. Properties of the model are compared with psychophysical data on human reaching movements, neurophysiological data on the tuning curves of neurons in the monkey motor cortex, and alternative models of movement control.

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.

The Construction of Arbitrary Stable Dynamics in Non-Linear Neural Networks

In this paper, two methods for constructing systems of ordinary differential equations realizing any fixed finite set of equilibria in any fixed finite dimension are introduced; no spurious equilibria are possible for either method. By using the first method, one can construct a system with the fewest number of equilibria, given a fixed set of attractors. Using a strict Lyapunov function for each of these differential equations, a large class of systems with the same set of equilibria is constructed. A method of fitting these nonlinear systems to trajectories is proposed. In addition, a general method which will produce an arbitrary number of periodic orbits of shapes of arbitrary complexity is also discussed. A more general second method is given to construct a differential equation which converges to a fixed given finite set of equilibria. This technique is much more general in that it allows this set of equilibria to have any of a large class of indices which are consistent with the Morse Inequalities. It is clear that this class is not universal, because there is a large class of additional vector fields with convergent dynamics which cannot be constructed by the above method. The easiest way to see this is to enumerate the set of Morse indices which can be obtained by the above method and compare this class with the class of Morse indices of arbitrary differential equations with convergent dynamics. The former set of indices are a proper subclass of the latter, therefore, the above construction cannot be universal. In general, it is a difficult open problem to construct a specific example of a differential equation with a given fixed set of equilibria, permissible Morse indices, and permissible connections between stable and unstable manifolds. A strict Lyapunov function is given for this second case as well. This strict Lyapunov function as above enables construction of a large class of examples consistent with these more complicated dynamics and indices. The determination of all the basins of attraction in the general case for these systems is also difficult and open.