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)

Engineers use neural networks to control systems too complex for conventional engineering solutions. To examine the behavior of individual hidden units would defeat the purpose of this approach because it would be largely uninterpretable. Yet neurophysiologists spend their careers doing just that! Hidden units contain bits and scraps of signals that yield only arcane hints about network function and no information about how its individual units process signals. Most literature on single-unit recordings attests to this grim fact. On the other (...) hand, knowing a system's function and describing it with elegant mathematics tell one very little about what to expect of interneuronal behavior. Examples of simple networks based on neurophysiology are taken from the oculomotor literature to suggest how single-unit interpretability might decrease with increasing task complexity. It is argued that trying to explain how any real neural network works on a cell-by-cell, reductionist basis is futile and we may have to be content with trying to understand the brain at higher levels of organization. (shrink)

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)

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 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.

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.”.

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)

Time underlies many interesting human behaviors. Thus, the question of how to represent time in connectionist models is very important. One approach is to represent time implicitly by its effects on processing rather than explicitly (as in a spatial representation). The current report develops a proposal along these lines first described by Jordan (1986) which involves the use of recurrent links in order to provide networks with a dynamic memory. In this approach, hidden unit patterns are fed back to themselves: (...) the internal representations which develop thus reflect task demands in the context of prior internal states. A set of simulations is reported which range from relatively simple problems (temporal version of XOR) to discovering syntactic/semantic features for words. The networks are able to learn interesting internal representations which incorporate task demands with memory demands: indeed, in this approach the notion of memory is inextricably bound up with task processing. These representations reveal a rich structure, which allows them to be highly context‐dependent, while also expressing generalizations across classes of items. These representations suggest a method for representing lexical categories and the type/token distinction. (shrink)

1. The Situation in Cognition 2. Situated Cognition: A Potted Recent History 3. Extensions in Biology, Computation, and Cognition 4. Articulating the Idea of Cognitive Extension 5. Are Some Resources Intrinsically Non-Cognitive? 6. Is Cognition Extended or Only Embedded? 7. Letting Nature Take Its Course.

The uncertainty associated with paradigmatic families has been shown to correlate with their phonetic characteristics in speech, suggesting that representations of complex sublexical relations between words are part of speaker knowledge. To better understand this, recent studies have used two-layer neural network models to examine the way paradigmatic uncertainty emerges in learning. However, to date this work has largely ignored the way choices about the representation of inflectional and grammatical functions in models strongly influence what they subsequently learn. To explore (...) the consequences of this, we investigate how representations of IFS in the input-output structures of learning models affect the capacity of uncertainty estimates derived from them to account for phonetic variability in speech. Specifically, we examine whether IFS are best represented as outputs to neural networks or as inputs by building models that embody both choices and examining their capacity to account for uncertainty effects in the formant trajectories of word final [ɐ], which in German discriminates around sixty different IFS. Overall, we find that formants are enhanced as the uncertainty associated with IFS decreases. This result dovetails with a growing number of studies of morphological and inflectional families that have shown that enhancement is associated with lower uncertainty in context. Importantly, we also find that in models where IFS serve as inputs—as our theoretical analysis suggests they ought to—its uncertainty measures provide better fits to the empirical variance observed in [ɐ] formants than models where IFS serve as outputs. This supports our suggestion that IFS serve as cognitive cues during speech production, and should be treated as such in modeling. It is also consistent with the idea that when IFS serve as inputs to a learning network. This maintains the distinction between those parts of the network that represent message and those that represent signal. We conclude by describing how maintaining a “signal-message-uncertainty distinction” can allow us to reconcile a range of apparently contradictory findings about the relationship between articulation and uncertainty in context. (shrink)

Musical improvisation is a sophisticated activity in which a performer realizes, real-time, melodic, and rhythmic sequences in harmony with those from other musicians. The study of musical improvisation helps one to understand not only the cognition of creativity, but also the complex neuronal basis of executive functions, the relation between conscious and unconscious action, and even more. So far, the prevailing models, founded on the brain imaging method, have focused on the connection between the cortical areas and their cognitive processes. (...) Little attention, on the other hand, has been given to the huge variety of subcortical activities, especially in the basal ganglia. This fundamental subcortical component, through its implicit procedures and the role it plays in memory, is responsible to produce new information all the time, allowing the prefrontal cortex to transform a huge and disordered amount of data in explicit creative acts. The basal ganglia are strongly related to the activation of chemical signals generated by dissonance or lack of symmetry between perception and expectation, participating even in the responses to environmental demands according to the circumstances. Thus, they interact with the frontal cortex and with the limbic system, playing a key role in planning and selecting appropriate actions and in decision making. In this text, we try to explain in which sense improvisation is connected to the processes of executive functions, to creativity and to the integrated activity of cortical–subcortical areas controlling the free flow of ideas and to expressive spontaneity. Eventually, we purpose a model according to which structure and process take part at the centrencephalic space of functional integration, which, through both competing and cooperating dynamics, gives way to spontaneous composition. (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.

Brain damage may doubly dissociate cognitive modules, but the practice of revealing dissociations is predicated on modularity being true (T. Shallice, 1988). This article questions the utility of assuming modularity, as it examines a paradigmatic double dissociation of reading modules. Reading modules illustrate two general problems. First, modularity fails to converge on a fixed set of exclusionary criteria that define pure cases. As a consequence, competing modular theories force perennial quests for purer cases, which simply perpetuates growth in the list (...) of exclusionary criteria. The first problem leads, in part, to the second problem. Modularity fails to converge on a fixed set of pure cases. The second failure perpetuates unending fractionation into more modules. (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)

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 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)