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.

Emergence of Tri-Phasic Muscle Activation from the Non-linear Interactions of Central and Spinal Neural Network Circuits

The origin of the tri-phasic burst pattern, observed in the EMGs of opponent muscles during rapid self-terminated movements, has been controversial. Here we show by computer simulation that the pattern emerges from interactions between a central neural trajectory controller (VITE circuit) and a peripheral neuromuscularforce controller (FLETE circuit). Both neural models have been derived from simple functional constraints that have led to principled explanations of a wide variety of behavioral and neurobiological data, including, as shown here, the generation of tri-phasic bursts.

A Neural Pattern Generator that Exhibits Frequency-Dependent In-Phase and Anti-Phase Oscillations

This article describes a. neural pattern generator based on a cooperative-competitive feedback neural network. The two-channel version of the generator supports both in-phase and anti-phase oscillations. A scalar arousal level controls both the oscillation phase and frequency. As arousal increases, oscillation frequency increases and bifurcations from in-phase to anti-phase, or anti-phase to in-phase oscillations can occur. Coupled versions of the model exhibit oscillatory patterns which correspond to the gaits used in locomotion and other oscillatory movements by various animals.

3-D Vision and Figure-Ground Separation by Visual Cortex

A neural network theory of :3-D vision, called FACADE Theory, is described. The theory proposes a solution of the classical figure-ground problem for biological vision. It does so by suggesting how boundary representations and surface representations are formed within a Boundary Contour System (BCS) and a Feature Contour System (FCS). The BCS and FCS interact reciprocally to form 3-D boundary and surface representations that arc mutually consistent. Their interactions generate 3-D percepts wherein occluding and occluded object completed, and grouped. The theory clarifies how preattentive processes of 3-D perception and figure-ground separation interact reciprocally with attentive processes of spatial localization, object recognition, and visual search. A new theory of stereopsis is proposed that predicts how cells sensitive to multiple spatial frequencies, disparities, and orientations are combined by context-sensitive filtering, competition, and cooperation to form coherent BCS boundary segmentations. Several factors contribute to figure-ground pop-out, including: boundary contrast between spatially contiguous boundaries, whether due to scenic differences in luminance, color, spatial frequency, or disparity; partially ordered interactions from larger spatial scales and disparities to smaller scales and disparities; and surface filling-in restricted to regions surrounded by a connected boundary. Phenomena such as 3-D pop-out from a 2-D picture, DaVinci stereopsis, a 3-D neon color spreading, completion of partially occluded objects, and figure-ground reversals are analysed. The BCS and FCS sub-systems model aspects of how the two parvocellular cortical processing streams that join the Lateral Geniculate Nucleus to prestriate cortical area V4 interact to generate a multiplexed representation of Form-And-Color-And-Depth, or FACADE, within area V4. Area V4 is suggested to support figure-ground separation and to interact. with cortical mechanisms of spatial attention, attentive objcect learning, and visual search. Adaptive Resonance Theory (ART) mechanisms model aspects of how prestriate visual cortex interacts reciprocally with a visual object recognition system in inferotemporal cortex (IT) for purposes of attentive object learning and categorization. Object attention mechanisms of the What cortical processing stream through IT cortex are distinguished from spatial attention mechanisms of the Where cortical processing stream through parietal cortex. Parvocellular BCS and FCS signals interact with the model What stream. Parvocellular FCS and magnocellular Motion BCS signals interact with the model Where stream. Reciprocal interactions between these visual, What, and Where mechanisms arc used to discuss data about visual search and saccadic eye movements, including fast search of conjunctive targets, search of 3-D surfaces, selective search of like-colored targets, attentive tracking of multi-element groupings, and recursive search of simultaneously presented targets.

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.