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.”.

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)

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.

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.

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)

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.

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)

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.

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 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.

Extrapersonal frames of reference for aimed movements are representationally convenient. They may, however, carry associated costs when the movement is executed in terms of the complex coordination of multiple joints they require. Studies that have measured both fingertip and joint paths suggest the motor systems may seek a compromise between simplicity of extrapersonal spatial representation and computational simplicity of multi-joint execution.

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.

Kinematic properties of reaching movements reflect constraints imposed on the joint angles. Contemporary models present solutions to the redundancy problem by a pseudoinverse procedure (Whitney 1969) or without any inversion (Berkenblit et al. 1986). Feldman & Levin suggest a procedure based on a regular inversion. These procedures are considered as an outcome of a more general approach.

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.

Contemporary experimental research has emphasised the role of centrally generated signals arising from premotor areas in voluntary muscular force perception. It is therefore generally accepted that judgements of force are based on a central sense, known as the sense of effort, rather than on a sense of intra-muscular tension. Interestingly, the concept of effort is also present in the classical philosophy: to the French philosopher Maine de Biran [Maine de Biran . Mémoire sur la décomposition de la pensée , Vrin, (...) Paris ], the sense of effort is the fundamental component of self-experience, the landmark of the exercise of the will. In the present review, after a presentation of the nature and neurophysiological bases of effort sensation, we will examine its possible involvement in the neurocognitive process of agency. We will further focus on delusions of alien control in schizophrenic patients. Experimental data suggest that these patients have an abnormal awareness of effort caused by cerebral anomalies in the frontal and parietal lobes. (shrink)

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 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.

We affirm the dynamical systems approach taken by Feldman and Levin, but argue that a more mathematically rigorous and standard exposition of the model according to dynamical systems theory would greatly increase readability and testability. Such an explication would also have heuristic value, suggesting new variations of the model. We present one such variant, a new solution to the redundancy problem.

The lambda model of Feldman & Levin for intentional hand movement is compared with a hemispheric motor model (IIMM). Both models imply similar conclusions independently. This increases the validity of both models.

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)

The concept of a conservative control strategy minimizing the number of degrees of freedom used is criticised with reference to 3-D simple reaching and grasping experiments. The vector error in a redundant system would not be the prime controlled variable, but rather the posture for reaching, as exemplified by nearly straight displacements in joint space as opposed to curved ones in task space.