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 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.

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.

A central issue in philosophy and neuroscience is the problem of unified visual consciousness. This problem has arisen because we now know that an object's stimulus features (e.g., its color, texture, shape, etc.) generate activity in separate areas of the visual cortex (Felleman & Van Essen, 1991). For example, recent evidence indicates that there are very few, if any, neural connections between specific visual areas, such as those that correlate with color and motion (Bartels & Zeki, 2006; Zeki, 2003). So (...) how do unified objects arise in visual consciousness? Some neuroscientists propose that neural synchrony is the mechanism that binds an object's features into a unity (e.g., see Crick, 1994; Crick & Koch, 1990; Engel, 2003; Roelfsema, 1998; Singer, 1996; von der Malsburg, 1996, 1999). I argue, on both empirical and philosophical grounds, that neural synchrony fails to explain the unity of visual consciousness. (shrink)

Many philosophical issues concern questions of objectivity and subjectivity. Of these questions, there are two kinds. The first considers whether something is objective or subjective; the second what it _means_ for something to be objective or subjective.

Many have urged that the biggest obstacles to a physicalistic understanding of consciousness are the problems raised in connection with the subjectivity of consciousness. These problems are most acutely expressed in consideration of the knowledge argument against physicalism. I develop a novel account of the subjectivity of consciousness by explicating the ways in which mental representations may be perspectival. Crucial features of my account involve analogies between the representations involved in sensory experience and the ways in which pictorial representations exhibit (...) perspectives or points of view. I argue that the resultant account of subjectivity provides a basis for the strongest response physicalists can give to the knowledge argument. (shrink)

We pose a foundational problem for those who claim that subjects are ontologically irreducible, but causally reducible (weak emergence). This problem is neuroscience’s notorious binding problem, which concerns how distributed neural areas produce unified mental objects (such as perceptions) and the unified subject that experiences them. Synchrony, synapses and other mechanisms cannot explain this. We argue that this problem seriously threatens popular claims that mental causality is reducible to neural causality. Weak emergence additionally raises evolutionary worries about how we’ve survived (...) the perils of nature. Our emergent subject hypothesis (ESH) avoids these shortcomings. Here, a singular, unified subject acts back on the neurons it emerges from and binds sensory features into unified mental objects. Serving as the mind’s controlling center, this subject is ontologically and causally irreducible (strong emergence). ESH draws on recent experimental evidence, including the motivation of a possible correlate (or “seat”) of the subject, which enhances its testability. (shrink)

Our perception of where touch occurs on our skin shapes our interactions with the world. Most accounts of cutaneous localisation emphasise spatial transformations from a skin-based reference frame into body-centred and external egocentric coordinates. We investigated another possible method of tactile localisation based on an intrinsic perception of ‘skin space’. The arrangement of cutaneous receptive fields (RFs) could allow one to track a stimulus as it moves across the skin, similarly to the way animals navigate using path integration. We applied (...) curved tactile motions to the hands of human volunteers. Participants identified the location midway between the start and end points of each motion path. Their bisection judgements were systematically biased towards the integrated motion path, consistent with the characteristic inward error that occurs in navigation by path integration. We thus showed that integration of continuous sensory inputs across several tactile RFs provides an intrinsic mechanism for spatial perception. (shrink)

Vision is so closely identified with visual phenomenology that we sometimes forget that the visual system does more than deliver our experience of the world. Vision also plays a critical role in the control of our movements, from picking up our coffee cups to playing tennis. But the visual control of movement has, until recently, been relatively neglected. Indeed, traditional accounts of vision, while acknowledging the role of vision in motor control, have simply regarded such control as part of a (...) larger function – that of constructing an internal model of the external world. Even though such accounts might postulate separate ‘modules’ for the processing of different visual features, such as motion, colour, texture, and form, in most of these accounts there is an implicit assumption that, in the end, vision delivers a single representation of the external world – a kind of simulacrum of the real thing that serves as the perceptual foundation for all visually driven thought and action. (shrink)

The question of how the brain can construct a stable representation of the external world despite eye movements is a very old one. If there have been some wrong statements of problems (such as the inverted retinal image), other statements are less naive and have led to analytic solutions possibly adopted by the brain to counteract the spurious effects of eye movements. Following the MacKay (1973) objections to the analytic view of perceptual stability, Bridgeman et al. claim that the idea (...) that signals canceling the effects of saccadic eye movements are needed is also a misconception, as is the claim that stability and position encoding are two distinct problems. It must be remembered, however, that what made the theory of “cancellation” formulated by von Holst and Mittelstaedt (1950) so appealing was the clinical observation of perceptual instability following ocular paralysis. Following the concept of corollary discharge, the theory of efference copy had the advantage of simultaneously solving three problems: the stability of the visual world during the saccade, the same visual stability across saccades, and the visual constancy problem of allowing the subject to know where an object in space is. (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.

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.

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.

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.

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.

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 lambda model provides a physiologically grounded terminology for describing muscle function and emphasizes the important influence of environmental and reflex-mediated effects on final states. However, lambda itself is only a convenient point on the length-tension curve; its importance should not be overemphasized. Ascribing movement to changes in a lambda-based frame of reference is generally valid, but it leaves unanswered a number of questions concerning mechanisms.

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 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 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)

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?

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.

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)

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.

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.

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.”.

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)

There are now various approaches to understand where and how in the brain consciousness arises from neural activity, none of which is universally accepted. Difficulties among these approaches are reviewed, and a missing ingredient is proposed here to help adjudicate between them, that of ''perspectivalness.'' In addition to a suitable temporal duration and information content of the relevant bound brain activity, this extra component is posited as being a further important ingredient for the creation of consciousness from neural activity. It (...) guides the development of what is termed the ''Central Representation,'' which is supposed to be present in all mammals and extended in humans to support self-consciousness as well as phenomenal consciousness. Experimental evidence and a theoretical framework for the existence of the central representation are presented, which relates the extra component to specific buffer working memory sites in the inferior parietal lobes, acting as attentional coordinators on the spatial maps making up the central representation. The article closes with a discussion of various open questions. (shrink)

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.

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.

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)