We identify two aspects of the problem of maintaining perceptual stability despite an observer's eye movements. The first, visual direction constancy, is the (egocentric) stability of apparent positions of objects in the visual world relative to the perceiver. The second, visual position constancy, is the (exocentric) stability of positions of objects relative to each other. We analyze the constancy of visual direction despite saccadic eye movements.Three information sources have been proposed to enable the visual system to achieve stability: the structure (...) of the visual field, proprioceptive inflow, and a copy of neural efference or outflow to the extraocular muscles. None of these sources by itself provides adequate information to achieve visual direction constancy; present evidence indicates that all three are used.Our final question concerns how information processing operations result in a stable world. The three traditionally suggested means have been elimination, translation, or evaluation. All are rejected. From a review of the physiological and psychological evidence we conclude that no subtraction, compensation, or evaluation need take place. The problem for which these solutions were developed turns out to be a false one. We propose a “calibration” solution: correct spatiotopic positions are calculated anew for each fixation. Inflow, outflow, and retinal sources are used in this calculation: saccadic suppression of displacement bridges the errors between these sources and the actual extent of movement. (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.

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.

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.

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 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°.

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.

The conservative strategy proposed by the authors suggests a solution of the degrees-of-freedom problem of the controller. However, several simple motor control tasks cannot be explained by this strategy. A nonconservative strategy, in which more parameters of the control signal vary, can account for these simple motor tasks. However, the simplicity that distinguishes the proposed model from many others is lost.

The historical debate on representation in cognitive science and neuroscience construes representations as theoretical posits and discusses the degree to which we have reason to posit them. We reject the premise of that debate. We argue that experimental neuroscientists routinely observe and manipulate neural representations in their laboratory. Therefore, neural representations are as real as neurons, action potentials, or any other well-established entities in our ontology.

The calibration solution to the stability of the world despite eye movements depends, according to Bridgeman et al., upon a combination of three factors which presumably all need to operate to achieve the goal of stability. Although the authors admit (sect. 4.3, para. 5) that the relative contributions of retinal and extraretinal factors will depend on the particular viewing situation, Figure 5 (sect. 4.3) makes it clear in its representation that the role of perceptual factors is relatively minor compared to (...) extraretinal ones. It is with this representation that this commentary wishes to take issue, believing that it occurs as a result of some assumptions about terminology that may be ambiguous, as well as some misconceptions about the circumstances in which there is a need for stability. (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 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 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)

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.

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.

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.

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.

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.

We describe a solution to the redundancy problem related to that proposed in Feldman & Levin's target article. We suggest that the system may use a fixed mapping between commands organized at the level of degrees of freedom and commands to individual muscles. This proposal eliminates the need to maintain an explicit representation of musculoskeletalgeometry in planning movements.

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.

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.”.

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 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)

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.

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.

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.

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)

Whereas the λ model provides a useful technique to describe complex movements, the focus on control variables in this model limits its potential for interpreting the activity and function of many cells in motor areas of the CNS.

Generation of swing phase limb trajectory over obstacles during locomotion should be a reasonable test for the λ model proposed by Feldman and Levin. The observed features such as lack of simple amplitude scaling of endpoint (toe) trajectories for different obstacle heights, complex shaped toe velocity profiles, and exploitation of passive intersegmental dynamics to control limb elevation cannot be adequately explained by the λ model.

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.

Generalizing the notion that muscles are positional frames of reference, a high-dimensional muscle space is defined for multi-muscle systems with an embedded low-dimensional motor manifold of functional articulators. A central representation of such a manifold is proposed as computational body schema. The example of the jaw-tongue system is presented, discussing the relation of functional articulators with kinematic invariances and control problems.