Three issues related to Feldman and Levin's treatment of biological variability are discussed. We question the usefulness of the indirect component of δλ. We suggest that trade-offs between speed and accuracy in aimed movements support identification of δλ, rather than λ, as a control variable. We take issue with the authors' proposal for resolving redundancy in multi-joint movements, given recent data.

The experiments Feldman and Levin suggest do not definitively test their proposed solution to the problem of selecting muscle activations. Their test of the movement directions that elicit EMG activity can be interpreted without regard to the form of the central commands, and their fast elbow flexion test is based on a forward computation that obscures the insensitivity of the predicted trajectory to the details of the putative commands.

Parameters of the lambda model seem tightly linked to certain characteristics of human performance influenced by weightlessness. This commentary suggests that there is a valuable opportunity to probe the lambda model using the changed environment experienced during space flight. The likely benefits are a better model and a better understanding ofthe consequences of weightlessness for human performance.

The spring-like behaviour of a joint following a sudden change of torque is partly a result of the elastic properties of tendons. A large fall in a muscle with a long tendon may be accompanied by tendon recoil causing joint movements as large as 20°.

Philosophical controversy about the mind has flourished in the thin air of our ignorance about the brain. The humble toad, it now seems, may provide our first instance of a creature whose whole brain is within the reach of our scientific understanding. What will happen to the traditional philosophical issues as our theoretical and factual ignorance recedes? Discussion of the issues explored in the target article is, as Ewert says, "often too theoretical, sometimes philosophical and even [as if that weren't (...) bad enough?--DCD] emotion-laden." The research reported by Ewert has interesting philosophical implications, as he probably recognizes, but he wisely leaves the philosophy to the philosophers. Being one, I would like to draw some of the conclusions he eschews. (shrink)

This target article draws together two groups of experimental studies on the control of human movement through peripheral feedback and centrally generated signals of motor commands. First, during natural movement, feedback from muscle, joint, and cutaneous afferents changes; in human subjects these changes have reflex and kinesthetic consequences. Recent psychophysical and microneurographic evidence suggests that joint and even cutaneous afferents may have a proprioceptive role. Second, the role of centrally generated motor commands in the control of normal movements and movements (...) following acute and chronic deafferentation is reviewed. There is increasing evidence that subjects can perceive their motor commands under various conditions, but that this is inadequate for normal movement; deficits in motor performance arise when the reliance on proprioceptive feedback is abolished either experimentally or because of pathology. During natural movement, the CNS appears to have access to functionally useful input from a range of peripheral receptors as well as from internally generated command signals. The unanswered questions that remain suggest a number of avenues for further research. (shrink)

A theory is presented to explain how accurate, single-joint movements are controlled. The theory applies to movements across different distances, with different inertial loads, toward targets of different widths over a wide range of experimentally manipulated velocities. The theory is based on three propositions. (1) Movements are planned according to “strategies” of which there are at least two: a speed-insensitive (SI) and a speed-sensitive (SS) one. (2) These strategies can be equated with sets of rules for performing diverse movement tasks. (...) The choice between SI and SS depends on whether movement speed and/or movement time (and hence appropriate muscle forces) must be constrained to meet task requirements. (3) The electromyogram can be interpreted as a low-pass filtered version of the controlling signal to the motoneuron pools. This controlling signal can be modelled as a rectangular excitation pulse in which modulation occurs in either pulse amplitude or pulse width. Movements to different distances and with loads are controlled by the SI strategy, which modulates pulse width. Movements in which speed must be explicitly regulated are controlled by the SS strategy, which modulates pulse amplitude. The distinction between the two movement strategies reconciles many apparent conflicts in the motor control literature. (shrink)

Intermediate constructs are required as bridges between complex behaviors and realistic models of neural circuitry. For cognitive scientists in general, schemas are the appropriate functional units; brain theorists can work with neural layers as units intermediate between structures subserving schemas and small neural circuits.After an account of different levels of analysis, we describe visuomotor coordination in terms of perceptual schemas and motor schemas. The interest of schemas to cognitive science in general is illustrated with the example of perceptual schemas in (...) high-level vision and motor schemas in the control of dextrous hands.Rana computatrix, the computational frog, is introduced to show how one constructs an evolving set of model families to mediate flexible cooperation between theory and experiment. Rana computatrix may be able to do for the study of the organizational principles of neural circuitry what Aplysia has done for the study of subcellular mechanisms of learning. Approach, avoidance, and detour behavior in frogs and toads are analyzed in terms of interacting schemas. Facilitation and prey recognition are implemented as tectal-pretectal interactions, with the tectum modeled by an array of tectal columns. We show how layered neural computation enters into models of stereopsis and how depth schemas may involve the interaction of accommodation and binocular cues in anurans. (shrink)

Examination of infant spontaneous and goal-directed arm movements supports Feldman and Levin's hypothesis of a functional hierarchy. Early infant movements are dominated by biomechanical and dynamic factors without external frames of reference. Development involves not only learning to generate these frames of reference, but also protecting the higher-level goal of the movement from internal and external perturbations.

Recent results have shown that the relative activation of muscles is different for isometric contractions and for movements. These results exclude an explanation of muscle activation patterns by a combination ofreciprocal and coactivation commands. These results also indicate that joint stiffness is not uniquely determined and that it may be different for isometric contractions and movements.

The posterior parietal cortex (PPC) is the most likely site where egocentric spatial relationships are represented in the brain. PPC cells receive visual, auditory, somaesthetic, and vestibular sensory inputs; oculomotor, head, limb, and body motor signals; and strong motivational projections from the limbic system. Their discharge increases not only when an animal moves towards a sensory target, but also when it directs its attention to it. PPC lesions have the opposite effect: sensory inattention and neglect. The PPC does not seem (...) to contain a “map” of the location of objects in space but a distributed neural network for transforming one set of sensory vectors into other sensory reference frames or into various motor coordinate systems. Which set of transformation rules is used probably depends on attention, which selectively enhances the synapses needed for making a particular sensory comparison or aiming a particular movement. (shrink)

An alternative multi-joint extension to the lambda model is proposed. According to this extension, the activity of a muscle depends not only on the difference between lambda and length of that muscle, but also on the difference between lambda and length of other muscles. This 2-D extension can describe more neurophysiological experiments than the extension proposed in the target article.

Neurophysiological evidence consonant with F&L's lambda model is reviewed and results of additional experiments are presented. The evidence shows that there are neurons in the motor cortex that respond to selective band widths of passive sinusoidal movements; the additional data show how, with movement, directionally sensitive population vectors can be shown to emerge from the data.

A hypothesis about sensorimotor integration (the λ model) is described and applied to movement control and kinesthesia. The central idea is that the nervous system organizes positional frames of reference for the sensorimotor apparatus and produces active movements by shifting the frames in terms of spatial coordinates. Kinematic and electromyographic patterns are not programmed, but emerge from the dynamic interaction among the system s components, including external forces within the designated frame of reference. Motoneuronal threshold properties and proprioceptive inputs to (...) motoneurons may be cardinal components of the physiological mechanism that produces positional frames of reference. The hypothesis that intentional movements are produced by shifting the frame of reference is extended to multi-muscle and multi-degrees-of-freedom systems with a solution of the redundancy problem that allows the control of a joint alone or in combination with other joints to produce any desired limb configuration and movement trajectory. The model also implies that for each motor behavior, the nervous system uses a strategy that minimizes the number of changeable control variables and keeps the parameters of these changes invariant. Examples are provided of simulated kinematic and electromyographic signals from single- and multi-joint arm movements produced by suggested patterns of control variables. Empirical support is provided and additional tests of the model are suggested. The model is contrasted with others based on the ideas of programming of motoneuronal activity, muscle forces, stiffness, or movement kinematics. (shrink)

1. Introduction. One major concern in cognitive psychology is explaining cognitively motivated behavior. However, attempts to specify the nature of the behavior that psychology is to explain have proved to be somewhat controversial. The aim of this paper is to conduct a preliminary investigation into the kinds of behavior psychology is concerned with.

This paper is a commentary on the target article by Michael Arbib, “Levels of modeling of mechanisms of visually guided behavior”, in the same issue of the journal, pp. 407–465. -/- I focus on the importance of the inclusion of an ability of a system to entertain, at a given time, multiple instantiations of a given schema (situation template, frame, script, action plan, etc.), and complications introduced into neural/connectionist network systems by such inclusion.

Abstract“Sign stimuli” elicit specific patterns of behavior when an organism's motivation is appropriate. In the toad, visually released prey-catching involves orienting toward the prey, approaching, fixating, and snapping. For these action patterns to be selected and released, the prey must be recognized and localized in space. Toads discriminate prey from nonprey by certain spatiotemporal stimulus features. The stimulus-response relations are mediated by innate releasing mechanisms (RMs) with recognition properties partly modifiable by experience. Striato-pretecto-tectal connectivity determines the RM's recognition and localization (...) properties, whereas medialpallio-thalamo-tectal circuitry makes the system sensitive to changes in internal state and to prior history of exposure to stimuli. RMs encode the diverse stimulus conditions referring to the same prey object through different combinations of “specialized” tectal neurons, involving cells selectively tuned to prey features. The prey-selective neurons express the outcome of information processing in functional units consisting of interconnected cells. Excitatory and inhibitory interactions among feature-sensitive tectal and pretectal neurons specify the perceptual operations involved in distinguishing the prey from its background, selecting its features, and discriminating it from predators. Other connections indicate stimulus location. The results of these analyses are transmitted by specialized neurons projecting from the tectum to bulbar/spinal motor systems, providing a sensorimotor interface. Specific combinations of such projective neurons – mediating feature- and space-related messages – form “command releasing systems” that activate corresponding motor pattern generators for appropriate prey-catching action patterns. (shrink)

We argue that the concept of a control variable (CV) as described by Feldman and Levin needs to be revised because it does not account for the influence of sensory feedback from the periphery. We provide evidence from the realm of rhythmic movements that sensory feedback can permanently alter the frequency and phase of a centrally generated rhythm.

The λ model suggests that detailed kinematics arise from changes in control variables and need not be explicitly planned. However, we have shown that when moving a grasped object, grip force is precisely modulated in phase with acceleration-dependent inertial load. This suggests that the motor system can predict detailed kinematics. This prediction may be based on a forward model of the dynamics of the loaded limb.

The following questions are discussed: (1) Who determines the nature of “control variables”? (2) Is the “positional monopoly” healthy? (3) Does a descending command alter reflex threshold alone without eoncomitantly altering stiffness? (4) How does the CNS deal with history-dependent effects? (5) Should we abandon the idea that the CNS controls classical Newtonian variables such as muscle length?

The problem for the CNS in any particular movement task is to coordinate the various frames of reference appropriate to the task. Control variables are determined by this coordination. The coordination problem varies greatly from task to task, and so no single set of control variables is likely to account for a broad range of movement tasks.

The authors report that the reorganization of body configuration during weightlessness is based on an intrapersonal frame of reference such as the configuration of the support surface and the position of the body's center of gravity. These results stress the importance of “knowledge” of the state of internal geometric structures, which cannot be directly signalled by specific receptors responsible for direct dialogue with the physical external world.