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Biomedical subjects

G Schöner

Publications and source records attributed to G Schöner.

At least 19 recordsLinked to original sources

Effects of varying task constraints on solutions to joint coordination in a sit-to-stand task.

The question of how multijoint movement is controlled can be studied by discovering how the variance of joint trajectories is structured in relation to important task-related variables. In a previous study of the sit-to-stand task, for instance, variations of body segment postures that leave the position of the body's center of mass (CM) unchanged were significantly greater than variations of body segment posture that varied the CM position. The present experiments tested the hypothesis that such structuring of joint configuration variability is accentuated when the mechanical or perceptual task demands are made more challenging. Six subjects performed the sit-to-stand task without vision (eyes closed), either on a normal or on a narrow support surface. An additional constraint on the postural task was introduced in a third condition by requiring subjects to maintain light touch (less than 1 N) with the fingertips while coming to a standing position on the narrow base of support. The joint configurations observed at each point in normalized time were analyzed with respect to trial-to-trial variability. The task variables CM and head position were used to define goal-equivalent sets of joint configurations ("uncontrolled manifolds," UCMs) within which variation of joint configuration leaves the task variables unchanged. The variability of joint configurations across trials was decomposed into components that did not affect (within the UCM) and that did affect (orthogonal to the UCM) the values of these task variables. Our results replicate the earlier finding of much larger variability in directions of joint space that leave the CM unchanged compared with directions that affect CM position. This effect was even more pronounced here than in the previous experiment, probably because of the more difficult perceptual conditions in the current study (eyes closed). When the mechanical difficulty of the task was increased, the difference between the two types of joint variability was further accentuated, primarily through increase in goal-equivalent variance. This provides evidence for the hypothesis that under challenging task constraints increased variability is selectively directed into task-irrelevant degrees of freedom. Because differential control along different directions of joint space requires coordination among joint angles, this observation supports the view that the CNS responds to increased task difficulty through enhanced coordination among degrees of freedom. The adaptive nature of this coordination is further illustrated by the similar enhanced use of goal-equivalent joint combinations to achieve a stable CM position when subjects stood up under the additional constraint of maintaining light touch with the fingertips. This was achieved by channeling goal-equivalent variability into different directions of joint configuration space.

Adult↗

Structure of motor variability in marginally redundant multifinger force production tasks.

The framework of the uncontrolled manifold hypothesis (UCM hypothesis) was applied to the analysis of the structure of finger force variability during oscillatory force production tasks. Subjects produced cycles of force with one, two (index and middle), or three (index, middle, and ring) fingers acting in parallel against force sensors mounted inside a small frame. The frame could be placed on the top of a table (stable conditions) or on a 4-mm-wide supporting surface (unstable conditions). Subjects were less variable when they used two fingers than when using one finger; adding the third finger did not change indices of variability of the performance. Components of finger force variance that did (VUN) or did not (VCOMP) change the value of a particular functional variable were computed for two control hypotheses: (1) at each time, the subjects tried to stabilize the total value of force (force-control); and (2), at each time, the subjects tried to stabilize the total moment produced with respect to an axis parallel to the hand/forearm (moment-control). Most subjects showed selective stabilization of moment and destabilization of force throughout most of the force cycle, in both stable and unstable conditions. The shapes of VUN and VCOMP suggested a possibility of selective compensation of timing errors across fingers within force cycles. One subject showed different relations between VUN and VCOMP, suggesting that these relations did in fact reflect particular central strategies of solving the tasks. The UCM method is applicable to force production tasks. It allows the comparison of control hypotheses in a quantitative way and unveils central strategies of control of redundant motor systems. Within this approach, redundancy (rather, abundance) is not a problem but an inherent part of a solution for natural motor tasks.

Adult↗

The dynamics of embodiment: a field theory of infant perseverative reaching.

The overall goal of this target article is to demonstrate a mechanism for an embodied cognition. The particular vehicle is a much-studied, but still widely debated phenomenon seen in 7-12 month-old-infants. In Piaget's classic "A-not-B error," infants who have successfully uncovered a toy at location "A" continue to reach to that location even after they watch the toy hidden in a nearby location "B." Here, we question the traditional explanations of the error as an indicator of infants' concepts of objects or other static mental structures. Instead, we demonstrate that the A-not-B error and its previously puzzling contextual variations can be understood by the coupled dynamics of the ordinary processes of goal-directed actions: looking, planning, reaching, and remembering. We offer a formal dynamic theory and model based on cognitive embodiment that both simulates the known A-not-B effects and offers novel predictions that match new experimental results. The demonstration supports an embodied view by casting the mental events involved in perception, planning, deciding, and remembering in the same analogic dynamic language as that used to describe bodily movement, so that they may be continuously meshed. We maintain that this mesh is a pre-eminently cognitive act of "knowing" not only in infancy but also in everyday activities throughout the life span.

Child Development↗

Identifying the control structure of multijoint coordination during pistol shooting.

The question of degrees of freedom in the control of multijoint movement is posed as the problem of discovering how the motor control system constrains the many possible combinations of joint postures to stabilize task-dependent essential variables. Success at a task can be achieved, in principle, by always adopting a particular joint combination. In contrast, we propose a more selective control strategy: variations of the joint configuration that leave the values of essential task variables unchanged are predicted to be less controlled (i.e., stabilized to a lesser degree) than joint configuration changes that shift the values of the task variables. Our experimental task involved shooting with a laser pistol at a target under four conditions. The seven joint angles of the arm were obtained from the recorded positions of markers on the limb segments. The joint configurations observed at each point in normalized time were analyzed with respect to trial-to-trial variability. Different hypotheses about relevant task variables were used to define sets of joint configurations ("uncontrolled manifolds" or UCMs) that, if realized, would leave essential task variables unchanged. The variability of joint configurations was decomposed into components lying parallel to those sets and components lying in their complement. The orientation of the gun's barrel relative to a vector pointing from the gun to the target was the task variable most successful at showing a difference between the two components of joint variability. This variable determines success at the task. Throughout the movement, not only while the gun was pointing at the target, fluctuations of joint configuration that affected this variable were much reduced compared with fluctuations that did not affect this variable. The UCM principle applied to relative gun orientation thus captures the structure of the motor control system across different parts of joint configuration space as the movement evolves in time. This suggests a specific control strategy in which changes of joint configuration that are irrelevant to success at the task are selectively released from control. By contrast, constraints representing an invariant spatial position of the gun or of the arm's center of mass structured joint configuration variability in the early and mid-portion of the movement trajectory, but not at the time of shooting. This specific control strategy is not trivial, because a target can be hit successfully also by controlling irrelevant directions in joint space equally to relevant ones. The results indicate that the method can be successfully used to determine the structure of coordination in joint space that underlies the control of the essential variables for a given task.

Adult↗

The distribution of neuronal population activation (DPA) as a tool to study interaction and integration in cortical representations.

In many cortical areas, simple stimuli or task conditions activate large populations of neurons. We hypothesize that such populations support processes of interaction within parametric representations and integration of multiple sources of input and we propose to study these processes using distributions of population activation (DPAs) as a tool. Such distributions can be viewed as neuronal representations of continuous stimulus or task parameters. They are built from basis functions contributed by each neuron. These functions may be explicitly chosen based on tuning curves or receptive field profiles. Or they may be determined by minimizing the distance between chosen target distributions and the constructed DPAs. In both cases, construction of the DPA is based on a set of reference conditions in which the stimulus or task parameters are sampled experimentally. In a second step, basis functions are kept fixed, and the DPAs are used to explore time dependent processing, interaction and integration of information. For instance, stimuli which simultaneously specify multiple parameter values can be used to study interactions within the parametric representation. We review an experiment, in which the representation of retinal position is probed in this way, revealing fast excitatory interactions among neurons representing similar retinal positions and slower inhibitory interactions among neurons representing dissimilar retinal positions. Similarly, DPAs can be used to analyze different sources of input that are fused within a parametric representation. We review an experiment in which the representation of the direction of goal-directed arm movements in motor and premotor cortex is studied when prior and current information about upcoming movement tasks are integrated.

Animals↗

Parametric population representation of retinal location: neuronal interaction dynamics in cat primary visual cortex.

Neuronal interactions are an intricate part of cortical information processing generating internal representations of the environment beyond simple one-to-one mappings of the input parameter space. Here we examined functional ranges of interaction processes within ensembles of neurons in cat primary visual cortex. Seven "elementary" stimuli consisting of small squares of light were presented at contiguous horizontal positions. The population representation of these stimuli was compared to the representation of "composite" stimuli, consisting of two squares of light at varied separations. Based on receptive field measurements and by application of an Optimal Linear Estimator, the representation of retinal location was constructed as a distribution of population activation (DPA) in visual space. The spatiotemporal pattern of the DPA was investigated by obtaining the activity of each neuron for a sequence of time intervals. We found that the DPA of composite stimuli deviates from the superposition of its components because of distance-dependent (1) early excitation and (2) late inhibition. (3) The shape of the DPA of composite stimuli revealed a distance-dependent repulsion effect. We simulated these findings within the framework of dynamic neural fields. In the model, the feedforward response of neurons is modulated by spatial ranges of excitatory and inhibitory interactions within the population. A single set of model parameters was sufficient to describe the main experimental effects. Combined, our results indicate that the spatiotemporal processing of visual stimuli is characterized by a delicate, mutual interplay between stimulus-dependent and interaction-based strategies contributing to the formation of widespread cortical activation patterns.

Animals↗

The uncontrolled manifold concept: identifying control variables for a functional task.

The degrees of freedom problem is often posed by asking which of the many possible degrees of freedom does the nervous system control? By implication, other degrees of freedom are not controlled. We give an operational meaning to "controlled" and "uncontrolled" and describe a method of analysis through which hypotheses about controlled and uncontrolled degrees of freedom can be tested. In this conception, control refers to stabilization, so that lack of control implies reduced stability. The method was used to analyze an experiment on the sit-to-stand transition. By testing different hypotheses about the controlled variables, we systematically approximated the structure of control in joint space. We found that, for the task of sit-to-stand, the position of the center of mass in the sagittal plane was controlled. The horizontal head position and the position of the hand were controlled less stably, while vertical head position appears to be no more controlled than joint motions.

Biomechanical Phenomena↗

Prior information preshapes the population representation of movement direction in motor cortex.

Single neuron activity was recorded in monkey motor cortex during the execution of pointing movements in six directions. The amount of prior information was manipulated by varying the range of precued directions. A distribution of neural population activation was constructed in the space of movement directions. This population representation of movement direction was preshaped by the precue. Peak location and width reflected the precued range of movement directions. From this preshaped form, the population representation evolved continuously in time and gradually in parameter space toward a more sharply peaked distribution centered on the parameter value specified by the response signal. A theoretical model of motor programming generated a similar temporal evolution of an activation field representing movement direction.

Animals↗

Position and velocity coupling of postural sway to somatosensory drive.

Light touch contact of a fingertip to a stationary surface provides orientation information that enhances control of upright stance. Slight changes in contact force at the fingertip lead to sensory cues about the direction of body sway, allowing attenuation of sway. In the present study, the coupling of postural sway to a moving contact surface was investigated in detail. Head, center of mass, and center of pressure displacement were measured as the contact surface moved rhythmically at 0.1, 0.2, 0.4, 0.6, and 0.8 Hz. Stimulus amplitude decreased with frequency to maintain peak velocity constant across frequency. Head and body sway were highly coherent with contact surface motion at all frequencies except 0.8 Hz, where a drop-off in coherence was observed. Mean frequency of head and body sway matched the driving frequency </=0.4 Hz. At higher frequencies, non-1:1 coupling was evident. The phase of body sway relative to the touch plate averaged 20-30 degrees at 0.1-Hz drive and decreased approximately linearly to -130 degrees at 0.8-Hz drive. System gain was approximately 1 across frequency. The large phase lags observed cannot be accounted for with velocity coupling alone but indicate that body sway also was coupled to the position of the touch plate. Fitting of a linear second-order model to the data suggests that postural control parameters are not fixed but adapt to the moving frame of reference. Moreover, coupling to both position and velocity suggest that a spatial reference frame is defined by the somatosensory system.

Adaptation, Physiological↗

Coupling of fingertip somatosensory information to head and body sway.

Light touch contact of a fingertip with a stationary surface can provide orientation information that enhances control of upright stance. Slight changes in contact force at the fingertip provide sensory cues about the direction of body sway, allowing attenuation of sway. In the present study, we asked to which extent somatosensory cues are part of the postural control system, that is, which sensory signal supports this coupling? We investigated postural control not only when the contact surface was stationary, but also when it was moving rhythmically (from 0.1 to 0.5 Hz). In doing so, we brought somatosensory cues from the hand into conflict with other parts of the postural control system. Our focus was the temporal relationship between body sway and the contact surface. Postural sway was highly coherent with contact surface motion. Head and body sway assumed the frequency of the moving contact surface at all test frequencies. To account for these results, a simple model was formulated by approximating the postural control system as a second-order linear dynamical system. The influence of the touch stimulus was captured as the difference between the velocity of the contact surface and the velocity of body sway, multiplied by a coupling constant. Comparison of empirical results (relative phase, coherence, and gain) with model predictions supports the hypothesis of coupling between body sway and touch cues through the velocity of the somatosensory stimulus at the fingertip. One subject, who perceived movement of the touch surface, demonstrated weaker coupling than other subjects, suggesting that cognitive mechanisms introduce flexibility into the postural control scheme.

Adult↗

Low-frequency oscillations of visual, auditory and somatosensory cortical neurons evoked by sensory stimulation.

Low-frequency oscillations-LFOs-below 20 Hz in the activity of cortical neurons are a commonly observed property across all sensory modalities. However, the functional significance and potential role of these intrinsic oscillations are not well understood. Here, we attempt to provide a general framework for the interpretation of this phenomenon by considering its properties across several sensory modalities. In the first part, we provide a survey and a general description of low-frequency oscillations (LFOs) at a cellular level observed following adequate [Basar, and Schürmann, 1994]. Sensory stimulation of neurons recorded in three sensory modalities of neocortices in higher mammals. The second part will address some functional aspects of low-frequency oscillations (LFOs) such as stimulus selectivity and so-called 'interference' phenomena, specifically with findings related to 'resetting' and 'gating' of sensory processing streams. Finally, a hypotheses is outlined in which the low-frequency oscillations are regarded as an organizational principle by which continuity of sensory and motor states over time could be accomplished.

Acoustic Stimulation↗

The influence of adaptation and stochastic fluctuations on spontaneous perceptual changes for bistable stimuli.

Spontaneous perceptual change was studied by measuring the probabilities of the first two spontaneous pattern switches as a function of time following the onset of a bistable apparent quartet for which either horizontal or vertical motion is perceived. Contrary to the classical satiation hypothesis (Köhler & Wallach, 1944), differential time-dependent adaptation of the perceived compared with the unperceived motion directions was not necessary to account for the first spontaneous switch. In addition, adaptation of the perceived motion accompanied by recovery from adaptation of the unperceived motion was not necessary to account for the increased probability of the second spontaneous switch. It was concluded that regardless of possible adaptation effects, stochastic fluctuations are necessary for the actual reversal of activation levels that produces the spontaneous switch. When the difference in detector activation is reduced by differential adaptation of competing motion detectors (or by the occurrence of a prior spontaneous pattern change), smaller stochastic fluctuations are sufficient to reverse the relative activation of competing detectors. Thus, adaptation can increase the probability of spontaneous switches without directly causing them.

Adaptation, Ocular↗

Identification of the nonlinear state-space dynamics of the action-perception cycle for visually induced postural sway.

Human subjects standing in a sinusoidally moving visual environment display postural sway with characteristic dynamical properties. We analyzed the spatiotemporal properties of this sway in an experiment in which the frequency of the visual motion was varied. We found a constant gain near 1, which implies that the sway motion matches the spatial parameters of the visual motion for a large range of frequencies. A linear dynamical model with constant parameters was compared quantitatively with the data. Its failure to describe correctly the spatiotemporal properties of the system led us to consider adaptive and nonlinear models. To differentiate between possible alternative structures we directly fitted nonlinear differential equations to the sway and visual motion trajectories on a trial-by-trial basis. We found that the eigenfrequency of the fitted model adapts strongly to the visual motion frequency. The damping coefficient decreases with increasing frequency. This indicates that the system destabilizes its postural state in the inertial frame. This leads to a faster internal dynamics which is capable of synchronizing posture with fast-moving visual environments. Using an algorithm which allows the identification of essentially nonlinear terms of the dynamics we found small nonlinear contributions. These nonlinearities are not consistent with a limit-cycle dynamics, accounting for the robustness of the amplitude of postural sway against frequency variations. We interpret out results in terms of active generation of postural sway specified by sensory information. We derive also a number of conclusions for a behavior-oriented analysis of the postural system.

Humans↗

Perceptual stability and the selective adaptation of perceived and unperceived motion directions.

Adaptation was studied in a paradigm in which the adapting stimulus was a variably biased version of a bistable apparent motion stimulus, a motion quartet, and the post-adaptation test stimulus was a "neutral" motion quartet. Either horizontal or vertical motion was perceived, never both at the same time. When only one of these was perceived during the entire adaptation phase of a trial, and the perceived motion was highly stable, adaptation effects were greater for the perceived than the unperceived motion directions (i.e., adaptation was selective to the perceived motion). However, when the perceived motion during adaptation was relatively unstable (i.e., when the perceived motion was more likely to spontaneously change directions), similar levels of adaptation were obtained for perceived as well as unperceived, but possible motion directions. Thus, adaptation occurs prior to the determination of which of the competing motion directions will be perceived. The relationship between the stability of the adapting percept and the selectivity of adaptation is explained in terms of differences in the activation of mutually inhibitory horizontal and vertical motion detectors.

Adaptation, Ocular↗

Saccadic motor planning by integrating visual information and pre-information on neural dynamic fields.

A functional model of target selection in the saccadic system is presented, incorporating elements of visual processing, motor planning, and motor control. We address the integration of visual information with pre-information, which is provided by manipulating the probability that a target appears at a certain location. This integration is achieved within a dynamic representation of planned eye movement which is modeled through distributions of activation on a topographic field. Visual input evokes activation, which is also constrained by lateral interaction within the field and by preshaping input representing pre-information. The model describes target selection observable in paradigms in which visual goals are presented at more than one location. Specifically, we model the transition from averaging, where endpoints of first saccades fall between two visual target locations, to decision making, where endpoints of first saccades fall accurately onto one of two simultaneously presented visual targets. We make predictions about how metrical biases of first saccades are induced by pre-information about target locations acquired by learning. When coupled to a motor control stage, activation dynamics on the planning level contribute to stabilizing gaze under fixation conditions. The neurophysiological relevance of our functional model is discussed.

Computer Simulation↗

Frequency dependence of the action-perception cycle for postural control in a moving visual environment: relative phase dynamics.

When standing human subjects are exposed to a moving visual environment, the induced postural sway displays varying degrees of coherence with the visual information. In our experiment we varied the frequency of an oscillatory visual display and analysed the temporal relationship between visual motion and sway. We found that subjects maintain sizeable sway amplitudes even as temporal coherence with the display is lost. Postural sway tended to phase lead (for frequencies below 0.2 Hz) or phase lag (above 0.3 Hz). However, we also observed at a fixed frequency, highly variable phase relationships in which a preferred range of phase lags is prevalent, but phase jumps occur that return the system into the preferred range after phase has begun drifting out of the preferred regime. By comparing the results quantitatively with a dynamical model (the sine-circle map), we show that this effect can be understood as a form of relative coordination and arises through an instability of the dynamics of the action-perception cycle. Because such instabilities cannot arise in passively driven systems, we conclude that postural sway in this situation is actively generated as rhythmic movement which is coupled dynamically to the visual motion.

Humans↗

Temporal stability of the action-perception cycle for postural control in a moving visual environment.

When standing human subjects are exposed to a moving visual environment, the induced postural sway forms a stable temporal relationship with the visual information. We have investigated this relationship experimentally with a new set-up in which a computer generates video images which correspond to the motion of a 3D environment. The suggested mean distance to a sinusoidally moving wall is varied and the temporal relationship to induced sway is analysed (1) in terms of the fluctuations of relative phase between visual and sway motion and (2) in terms of the relaxation time of relative phase as determined from the rate of recovery of the stable relative phase pattern following abrupt changes in the visual motion pattern. The two measures are found to converge to a well-defined temporal stability of the action-perception cycle. Furthermore, we show that this temporal stability is a sensitive measure of the strength of the action-perception coupling. It decreases as the distance of the visual scene from the observer increases. This fact and the increase of mean relative phase are consistent with predictions of a linear second-order system driven by the visual expansion rate. However, the amplitude of visual sway decreases little as visual distance increases, in contradiction to the predictions, and is suggestive of a process that actively generates sway. The visual expansion rate on the optic array is found to decrease strongly with visual distance. This leads to the conclusion that postural control in a moving visual environment cannot be understood simply in terms of minimization of retinal slip, and that dynamic coupling of vision into the postural control system must be taken into account.

Algorithms↗

Bistability and hysteresis in the organization of apparent motion patterns.

In a paradigm for which 2 distinct patterns are perceived for the same stimulus, perceptual hysteresis (persistence of a percept despite parameter change to values favoring the alternative pattern) and temporal stability (persistence despite intrinsic propensities toward spontaneous change) are interdependent. Greater persistence during parameter change reduces temporal stability, slowing the rate of parameter change reduces hysteresis by increasing opportunity for spontaneous change, and increasing temporal stability (by enlarging the stimulus) increases hysteresis. Hysteresis results in the perception of parametrically disfavored patterns; a parameter can influence a percept without specifying it. The visual system thus exhibits time-dependent behavior analogous to dynamical behavior observed in other systems, both physical and biological, for which there is competition among alternative patterns that vary in relative stability.

Adult↗