This paper concerns how motor actions are neurally represented and coded. Action planning and motor preparation can be studied using a specific type of representational activity, motor imagery. A close functional equivalence between motor imagery and motor preparation is suggested by the positive effects of imagining movements on motor learning, the similarity between the neural structures involved, and the similar physiological correlates observed in both imaging and preparing. The content of motor representations can be inferred from motor images at a (...) macroscopic level, based on global aspects of the action and the motor rules and constraints which predict the spatial path and kinematics of movements. A more microscopic neural account calls for a representation of object-oriented action. Object attributes are processed in different neural pathways depending on the kind of task the subject is performing. During object-oriented action, a pragmatic representation is activated in which object affordances are transformed into specific motor schemas. Animal as well as human clinical data implicate the posterior parietal and premotor cortical areas in schema instantiation. A mechanism is proposed that is able to encode the desired goal of the action and is applicable to different levels of representational organization. (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.

The model identifies a spatial coordinate frame within which the sensorimotor apparatus produces movement. Its spatial nature simplifies its coupling with spatial reference frames used concurrently by vision and proprioception. While the positional reference frame addresses the performance of spatial tasks, it seems to have little to say about movements involving energy expenditure as the principle component of the task.

Amidst the progress being made in the various (sub-)disciplines of the behavioural and brain sciences a somewhat neglected subject is the problem of how everything fits into one world and, derivatively, how the relation between different levels of discourse should be understood and to what extent different levels, domains, approaches, or disciplines are autonomous or dependent. In this paper I critically review the most recent proposals to specify the nature of interdiscourse relations, focusing on the concept of supervenience. Ideally supervenience (...) is a relation between different discourses which has all the advantages of reduction, but without its disadvantages. I apply the more abstract considerations to two concrete cases: schizophrenia and colour. Usually an interlevel or interdiscourse relation is seen as asymmetrical: the overlaying discourse depends on the underlying discourse (and not vice versa), where the out- or un-spoken assumption is that the ultimate underlying discourse is physical. Instead I argue that scientific categories referred to in interdiscourse relations are, ultimately, dependent on common sense categories and common sense normative criteria. It is the manifest categories and common sense ideas about what is reasonable and what is right that determine the relevant categorisations at the deeper, underlying levels. I suggest that the implications of this are not merely methodological or epistemological. (shrink)

Theories of the representation of specific kinetic and spatiotem-poral features of movement range from the explicit assertion that temporal aspects of movement are not represented to the idea that they are represented and that they have neurophysiological correlates. Jeannerod's thesis is that mental and visual images have common mechanisms and that there is a link between the image to move and the mechanisms involved with movement. The target article takes the position that certain parameters are coded in motor representations but (...) that the duration of an action is not one of them. This position is based on the work of Gottlieb et al. and of Decety et al.. Both these studies are worth considering in detail. In Note 1, Jeannerod suggests that: “in time-constrained tasks subjects control the amplitude parameter of force impulses, whereas in spatially constrained tasks the duration of the force impulse is affected by accuracy demands.” This is not exactly correct. Excitation pulse intensity is modulated both in tasks that require spatial and those that require temporal accuracy. Excitation pulse duration is modulated for changes in movement distance and inertial load. If subjects are required to be very accurate spatially, they will move at less than maximum speed for a given distance and this is achieved by lower levels of excitation intensity. (shrink)

Understanding of the λ model has greatly increased in recent years as evidenced by most of the commentaries. Some commentators underscored the potential of the model to integrate aspects of different sensorimotor systems in the production of movement. Other commentators focused on not-yet-fully-developed parts of the model. A few persisted in misunderstanding some of its basic concepts, and on these grounds they reject it. In responding to commentaries we continue to elaborate on some fundamental points of the model, especially control (...) variables, the idea of movement production by shifting the positional frame of reference and the hypothesis of biomechanical correspondence in motor control. We also continue to develop our ideas on the intrinsic generation of the frame of reference associated with external space and utilized for the control of arm movement and locomotion. The dynamic principles underlying the model are discussed in light of the dynamical systems approach. (shrink)

It is proposed here that the spinal network of proprioceptive feedback from length and force receptors constitutes the mechanism underlying the coordination of activation thresholds for muscles acting about the same and neighboring joints. For the most part, these circuits come between motoneurons and supraspinal signals, invalidating the idea that the activation thresholds constitute control variables for the motor system.

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.

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.

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)

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 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 lambda model predicts that the command received by each motor nucleus during locomotion is specific for the joint at which its muscle acts and is independent of external conditions. However, investigation of the commands received by motor nuclei during fictive locomotion and of the sensitivity of these commands to feedback from the limb during locomotion indicates that neither condition is satisfied.

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°.

Two aspects of the target article, (1) the extension of the equilibrium point theory to multi-joint movements, and (2) the consequence that the EMG pattern is not directly controlled by the central nervous system (CNS), are discussed in light of the experiments on upper trunk bending in humans. The principle component kinematic analysis and the analysis of the EMG data, obtained under microgravity and additional loading conditions, support the application of Feldman and Levin's for multi-joint pointing movement to equilibrium control (...) during upper trunk movement. (shrink)

A key feature of the lambda model is the hypothesis of a local spring-like muscle-reflex system defined by a central control variable that has units of position. This is intriguing, especially for a study of postural stability in large-scale systems, but it has limited direct application to skilled everyday movements. If movement is considered as a goal-directed, neuro-optimization problem, however, theavailabilityof lambda-like peripheral models (vs. conventional musculoskeletal models) deserves exploration.

The lambda model of servocontrol seems superior to the alpha model in terms of dealing with the mechanical complexities of nonlinear and multiarticular muscles. Both, however, can be trivialized by noting that the “control variable” may simply be the sum of descending influences at propriospinal interneurons in the case of the lambda model or in the muscles themselves in the case of the alpha model. The notion that the brain explicitly computes output in terms of any such control variables may (...) be an engineering metaphor, useful for conceptual understanding but not for generating predictive hypotheses about higher motor circuitry. (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.

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 search for simplifying principles in motor control motivates the target article. One method that the CNS uses to simplify the task of controlling a limb's mechanical properties is absent from the article. Evidence from multi-joint, force-field measurements and from kinematics that points to the existence of multi-joint primitives as control variables is discussed.

Jeannerod's target article describes support, through empirical and neurological findings, for the intriguing idea of motor imagery, a form of representation hypothesized to have levels of functional equivalence with motor preparation, while being consciously accessible. Jeannerod suggests that the subjectively accessible content of motor imagery allows it to be distinguished from motor preparation, which is unconscious. Motor imagery is distinguished from visual imagery in terms of content. Motor images are kinesthetic in nature; they are parametrized by variables such as force (...) and time and they are potentially governed by kinematic rules. Jeannerod acknowledges, however, that motor and visual imagery may not easily be separated, because actions take place in a spatial environment. I agree; in fact, I suggest here that visualization may generally be concomitant with, and may even subjectively dominate, motor imagery. (shrink)

It is argued that the equilibrium point model can lead to new insights regarding transition and stability processes in movement coordination. The role of movement control parameters on equifinality and phase-resetting is discussed; not only control but also external control parameters can affect the global dynamical regime.

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 equilibrium-point hypothesis (the λ-model) is superior to all other models of single-joint control and provides deep insights into the mechanisms of control of multi-joint movements. Attempts at associating control variables with neurophysiological variables look confusing rather than promising. Probabilistic mechanisms may play an important role in movement generation in redundant systems.

Feldman and Levin present a model for movement control in which the system is said to seek equilibrium points, active movement being produced by shifting frames of reference in space. It is argued that whatever merit this model might have is limited to an understanding of “the how” and not “the why” we move. In this way the authors seem to be forced into a dualistic position leaving the upper level of the proposed control hierarchy “floating.”.

Models of central control variables (CVs) that are expressed in positional reference frames and rely on proprioception as the dominant specifier of muscle activation patterns have not yet been shown to be adequate for the description of fast, voluntary movement, even of single joints. An alternative model with illustrative data is proposed.

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.

Important similarities exist between the dynamical concepts implicit in Feldman & Levin's extended λ model and those basic to a dynamical systems approach. We argue that careful application of the key concepts of control and order parameters, equilibria, and stability, can relate known facts of neuromuscular processes to the observables of functional, task-specific behavior.

We emphasize the relevance to cognitive psychology of Feldman and Levin's theoretical position. Traditional views of motor control have failed to clearly separate “production control” at the level of motor command, based on task-independent CV (control variables), from intentional “product control” based on task-dependent parameters. Because F&L's approach concentrates on the first process (trajectory formation), it can distinguish the product control stage.