Biomedical subjects

J Randall Flanagan

Publications and source records attributed to J Randall Flanagan.

17 recordsLinked to original sources

PubMed2006

Neural correlates of internal-model loading.

Skilled object manipulation requires knowledge, or internal models, of object dynamics relating applied force to motion , and our ability to handle myriad objects indicates that the brain maintains multiple models . Recent behavioral studies have shown that once learned, an internal model of an object with novel dynamics can be rapidly recruited and derecruited as the object is grasped and released . We used event-related fMRI to investigate neural activity linked to grasping an object with recently learned dynamics in preparation for moving it after a delay. Subjects also performed two control tasks in which they either moved without the object in hand or applied isometric forces to the object. In all trials, subjects received a cue indicating which task to perform in response to a go signal delivered 5-10 s later. We examined BOLD responses during the interval between the cue and go and assessed the conjunction of the two contrasts formed by comparing the primary task to each control. The analysis revealed significant activity in the ipsilateral cerebellum and the contralateral and supplementary motor areas. We propose that these regions are involved in internal-model recruitment in preparation for movement execution.

Adult↗

PubMed2006

Control strategies in object manipulation tasks.

The remarkable manipulative skill of the human hand is not the result of rapid sensorimotor processes, nor of fast or powerful effector mechanisms. Rather, the secret lies in the way manual tasks are organized and controlled by the nervous system. At the heart of this organization is prediction. Successful manipulation requires the ability both to predict the motor commands required to grasp, lift, and move objects and to predict the sensory events that arise as a consequence of these commands.

Animals↗

PubMed2006

Eye movements when observing predictable and unpredictable actions.

We previously showed that, when observers watch an actor performing a predictable block-stacking task, the coordination between the observer's gaze and the actor's hand is similar to the coordination between the actor's gaze and hand. Both the observer and the actor direct gaze to forthcoming grasp and block landing sites and shift their gaze to the next grasp or landing site at around the time the hand contacts the block or the block contacts the landing site. Here we compare observers' gaze behavior in a block manipulation task when the observers did and when they did not know, in advance, which of two blocks the actor would pick up first. In both cases, observers managed to fixate the target ahead of the actor's hand and showed proactive gaze behavior. However, these target fixations occurred later, relative to the actor's movement, when observers did not know the target block in advance. In perceptual tests, in which observers watched animations of the actor reaching partway to the target and had to guess which block was the target, we found that the time at which observers were able to correctly do so was very similar to the time at which they would make saccades to the target block. Overall, our results indicate that observers use gaze in a fashion that is appropriate for hand movement planning and control. This in turn suggests that they implement representations of the manual actions required in the task and representations that direct task-specific eye movements.

Adult↗

PubMed2006

Attenuation of self-generated tactile sensations is predictive, not postdictive.

When one finger touches the other, the resulting tactile sensation is perceived as weaker than the same stimulus externally imposed. This attenuation of sensation could result from a predictive process that subtracts the expected sensory consequences of the action, or from a postdictive process that alters the perception of sensations that are judged after the event to be self-generated. In this study we observe attenuation even when the fingers unexpectedly fail to make contact, supporting a predictive process. This predictive attenuation of self-generated sensation may have evolved to enhance the perception of sensations with an external cause.

Adolescent↗

PubMed2005

Eye-hand coordination during learning of a novel visuomotor task.

We investigated how gaze behavior and eye-hand coordination change when subjects learned a challenging visuomotor task that required acquisition of a novel mapping between bimanual actions and their visual sensory consequences. By applying isometric forces and torques to a rigid tool held freely between the two hands, subjects learned to control a cursor on a computer screen to hit successively displayed targets as quickly as possible. The learning occurred in stages that could be distinguished by changes in performance (target-hit rate) as well as by gaze behavior and eye-hand coordination. In a first exploratory stage, the hit rate was consistently low, the cursor position varied widely, and gaze typically pursued the cursor. In a second skill acquisition stage, the hit rate improved rapidly, and gaze fixations began to mark predictively desired cursor positions, indicating that subjects started to program spatially congruent eye and hand motor commands. In a third skill refinement stage, performance continued to improve gradually, and gaze shifted directly toward the target. We suggest that during the exploratory stage, the learner attempts to establish basic mapping rules between manual actions and eye-movement commands. In this process, subjects may establish correlations between hand motor commands and their visual sensory consequences, primarily in fovea-anchored, gaze-centered coordinates, and correlations between recent hand motor commands and eye motor commands. The established mapping rules are then implemented and refined in the skill acquisition and refinement stages.

Adult↗

PubMed2005

Perception of the consequences of self-action is temporally tuned and event driven.

It has been proposed that in order to increase the salience of sensations with an external cause, sensations that are predictable based on one's own actions are attenuated [1 and 2]. This may explain why self-imposed tickle [3 and 4] or constant forces [5] are perceived as less intense than the same stimuli externally imposed. Here, subjects used their right index finger to tap a force sensor mounted above their left index finger. When a motor generated a tap on the left finger synchronously with the right tap, simulating contact between the fingers, the perception of force in the left finger was attenuated compared to the same tap experienced during rest. Attenuation gradually reduced as the left tap was either delayed or advanced relative to the active right tap. However, no attenuation was seen to left taps triggered by right-finger movements that stopped above or passed wide of the sensor. We conclude that there is a window of sensory attenuation that is broadly temporally tuned and centered on the time at which the fingers would normally make contact. That is, predictive tactile sensory attenuation is linked to specific external events arising from movement rather than to the movement per se.

Adolescent↗

PubMed2005

Learning and recall of incremental kinematic and dynamic sensorimotor transformations.

Numerous studies have shown that when people encounter a sudden and novel sensorimotor transformation that alters perceived or actual movement, they gradually adapt and can later recall what they have learned if they encounter the transformation again. In this study, we tested whether retention and recall of learning is also observed when kinematic and dynamic transformations are introduced incrementally such that participants never experience large movement errors. Participants adapted their reaching movements to either a visuomotor rotation of hand position (kinematic transformation) or a rotary viscous force-field applied to the hand (dynamic transformation). These perturbations were introduced either incrementally or instantaneously. Thus, four groups of participants were tested with an incremental and an instantaneous group for both the kinematic and dynamic perturbations. To evaluate retention of learning, participants from all four groups were tested a day later on the same kinematic or dynamic perturbation presented instantaneously (at full strength). As expected, we found that subjects in the instantaneous group retained learning across days. We also found that, for both kinematic and dynamic perturbations, retention was equally good or better when the transformation was introduced incrementally. Because large and clearly detectable movement errors were not observed during adaptation to incremental perturbations, we conclude that such errors are not required for the learning and retention of internal models of kinematic and dynamic sensorimotor transformations.

Adolescent↗

PubMed2005

Common encoding of novel dynamic loads applied to the hand and arm.

In manual action, the relationship between a given motor command and the ensuing movement depends on the dynamics of both the arm and hand-held objects. Skilled performance relies on the brain learning both these dynamics, and previous studies have examined how people adapt to novel loads applied to either the hand or the arm. In this study, we ask whether these different kinds of load are represented independently as a result of changes in cutaneous feedback and hand-arm coordination. We used a robotic apparatus that could either apply forces to an object held in the subject's hand or directly to the segments of the arm. We tested whether subjects could retain learning of a force field applied to the hand after subsequently experiencing the opposing field applied to the arm (or vice versa), or whether retrograde interference would be observed. In separate experiments, we used force fields and torque fields that were linearly related to either hand or joint velocities, respectively. Our finding of complete interference between opposing fields suggests that loads applied to the arm and hand are not represented independently by the sensorimotor system. This interference occurred despite markedly different cutaneous inputs that were directly related to the movement task. This result suggests that the brain represents dynamics independently of these sensory inputs. In addition, we found that the rate at which subjects adapted to a given force field, specified either in hand or joint coordinates, was independent of whether the forces were applied to the hand or arm segments.

Adult↗

PubMed2005

Interference between velocity-dependent and position-dependent force-fields indicates that tasks depending on different kinematic parameters compete for motor working memory.

Humans demonstrate motor learning when exposed to changes in the dynamics of movement or changes in the visuomotor map. However, when two opposing dynamic transformations are learned in succession, the memory of the first is overwritten by learning of the second; the same is true for two opposing visuomotor rotations. This retrograde interference is not seen for all combinations of transformations, however. When a dynamic transformation is learned subsequent to a visuomotor rotation, the presence or absence of interference appears to depend crucially on the structure of the dynamic task: a force-field dependent on the position of the hand produces interference, whereas an inertial load applied lateral to the hand does not. To explain these results, it has been hypothesized that two transformations can be learned without interference if they depend on two different kinematic parameters of movement (such as position and velocity of the hand). Here we demonstrate, contrary to this hypothesis, interference between a dynamic transformation that depends on the position of the hand and one that depends on its velocity. However, the interference was found to be incomplete, supporting the view that the ability to retain motor memories for different tasks depends on the degree to which their representations conflict in working memory.

Adaptation, Physiological↗

PubMed2004

Failure to consolidate the consolidation theory of learning for sensorimotor adaptation tasks.

An influential idea in human motor learning is that there is a consolidation period during which motor memories are transformed from a fragile to a permanent state, no longer susceptible to interference from new learning. The evidence supporting this idea comes from studies showing that the motor memory of a task (A) is lost when an opposing task (B) is experienced soon after, but not if sufficient time is allowed to pass (approximately 6 hr). We report results from three laboratories challenging this consolidation idea. We used an ABA paradigm in the context of a reaching task to assess the influence of experiencing B after A on the retention of A. In two experiments using visuomotor rotations, we found that B fully interferes with the retention of A even when B is experienced 24 hr after A. Contrary to previous reports, in four experiments on learning force fields, we also observed full interference between A and B when they are separated by 24 hr or even 1 week. This latter result holds for both position-dependent and velocity-dependent force fields. For both the visuomotor and force-field tasks, complete interference is still observed when the possible affects of anterograde interference are controlled through the use of washout trials. Our results fail to support the idea that motor memories become consolidated into a protected state. Rather, they are consistent with recent ideas of memory formation, which propose that memories can shift between active and inactive states.

Adaptation, Physiological↗

PubMed2003

Action plans used in action observation.

How do we understand the actions of others? According to the direct matching hypothesis, action understanding results from a mechanism that maps an observed action onto motor representations of that action. Although supported by neurophysiological and brain-imaging studies, direct evidence for this hypothesis is sparse. In visually guided actions, task-specific proactive eye movements are crucial for planning and control. Because the eyes are free to move when observing such actions, the direct matching hypothesis predicts that subjects should produce eye movements similar to those produced when they perform the tasks. If an observer analyses action through purely visual means, however, eye movements will be linked reactively to the observed action. Here we show that when subjects observe a block stacking task, the coordination between their gaze and the actor's hand is predictive, rather than reactive, and is highly similar to the gaze-hand coordination when they perform the task themselves. These results indicate that during action observation subjects implement eye motor programs directed by motor representations of manual actions and thus provide strong evidence for the direct matching hypothesis.

Adult↗

PubMed2003

Prediction precedes control in motor learning.

Skilled motor behavior relies on the brain learning both to control the body and predict the consequences of this control. Prediction turns motor commands into expected sensory consequences, whereas control turns desired consequences into motor commands. To capture this symmetry, the neural processes underlying prediction and control are termed the forward and inverse internal models, respectively. Here, we investigate how these two fundamental processes are related during motor learning. We used an object manipulation task in which subjects learned to move a hand-held object with novel dynamic properties along a prescribed path. We independently and simultaneously measured subjects' ability to control their actions and to predict their consequences. We found different time courses for predictor and controller learning, with prediction being learned far more rapidly than control. In early stages of manipulating the object, subjects could predict the consequences of their actions, as measured by the grip force they used to grasp the object, but could not generate appropriate actions for control, as measured by their hand trajectory. As predicted by several recent theoretical models of sensorimotor control, our results indicate that people can learn to predict the consequences of their actions before they can learn to control their actions.

Biophysical Phenomena↗

PubMed2003

Task-specific internal models for kinematic transformations.

Numerous studies of motor learning have focused on how people adapt their reaching movements to novel dynamic and visuomotor perturbations that alter the actual or visually perceived motion of the hand. An important finding from this work is that learning of novel dynamics generalizes across different movement tasks. Thus adaptation to an unusual force field generalizes from center-out reaching movements to circular movements (Conditt et al. 1997). This suggests that subjects acquired an internal model of the dynamic environment that could be used to determine the motor commands needed for untrained movements. Using a task interference paradigm, we investigated whether transfer across tasks is also observed when learning visuomotor transformations. On day 1, all subjects adapted to a +30 degrees rotation while making center-out-and-back reaching movements. After a delay of 5 min, different groups of subjects then adapted to a -30 degrees rotation while performing either a continuous tracking task, a figure-eight drawing task, or the center-out-and-back reaching task. All subjects were then retested the next day on the +30 degrees rotation in the reaching task. As expected, subjects who experienced the opposing rotations while performing the same reaching tasks showed no retention of learning for the first rotation when tested on day 2 (Krakauer et al. 1999). In contrast, such retrograde interference was not observed in the two groups of subjects who experienced the opposing rotations while performing different tasks. In fact, their performance on day 2 was similar to that of control subjects who never experienced the opposite rotation. This lack of interference suggests that memory resources for visuomotor rotations are task specific.

Adult↗

PubMed2002

Kinematics and dynamics are not represented independently in motor working memory: evidence from an interference study.

Our capacity to learn multiple dynamic and visuomotor tasks is limited by the time between the presentations of the tasks. When subjects are required to adapt to equal and opposite position-dependent visuomotor rotations (Krakauer et al., 1999) or velocity-dependent force fields (Brashers-Krug et al., 1996) in quick succession, interference occurs that prevents the first task from being consolidated in memory. In contrast, such interference is not observed between learning a position-dependent visuomotor rotation and an acceleration-dependent force field. On the basis of this finding, it has been argued that internal models of kinematic and dynamic sensorimotor transformations are learned independently (Krakauer et al., 1999). However, these findings are also consistent with the perturbations interfering only if they depend on the same kinematic variable. We evaluated this hypothesis using kinematic and dynamic transformations matched in terms of the kinematic variable on which they depend. Subjects adapted to a position-dependent visuomotor rotation followed 5 min later by a position-dependent rotary force field either with or without visual feedback of arm position. The force field tended to rotate the hand in the direction opposite to the visuomotor rotation. To assess learning, all subjects were retested 24 hr later on the visuomotor rotation, and their performance was compared with a control group exposed only to the visuomotor rotation on both days. Adapting to the position-dependent force field, both with and without visual feedback, impaired learning of the visuomotor rotation. Thus, interference between our kinematic and dynamic transformations was observed, suggesting that the key determinant of interference is the kinematic variable on which the transformation depends.

Adolescent↗

PubMed2002

Visuomotor rotations of varying size and direction compete for a single internal model in motor working memory.

When participants adapt to equal and opposite visuomotor rotations in close temporal proximity, memory of the 1st is not consolidated. The authors investigated whether this retrograde interference depends on the use of equal and opposite rotations. On Day 1, different groups of participants adapted to a -30 degrees rotation followed 5 min later by rotations of +30 degrees, +60 degrees, or -60 degrees. On Day 2, all groups were retested on the -30 degrees rotation. Either retrograde interference (in groups who adapted to rotations of opposite sign on Day 1) or retrograde facilitation (in the remaining group) was observed. In all groups, learning of the 2nd rotation resulted in unlearning of the first, indicating that all visuomotor rotations compete for common working memory resources.

Arm↗

PubMed2002

Engagement of gaze in capturing targets for future sequential manual actions.

We investigated the role of saccadic gaze fixations in encoding target locations for planning a future manual task consisting of a sequence of discrete target-oriented actions. We hypothesized that fixations of the individual targets are necessary for accurate encoding of target locations and that there is a transfer of sequence information from visual encoding to manual recall. Subjects viewed four targets presented at random positions on a screen. After various delays following target extinction, the subjects marked the remembered target locations on the screen with the tip of a hand-held stick. When the targets were presented simultaneously among distracting elements, the overall accuracy of marking increased with presentation time and total number of targets fixated because the subjects had to serially fixate the individual targets to locate them. Without distractors, the marking accuracy was similarly high regardless of duration of target presentation (0.25-8 s) and number of targets fixated; it was comparable to that with distractors when all four targets had been fixated. This indicates parallel encoding of target locations largely based on peripheral vision. Location memory was stable in these tasks over the delay periods investigated (0.5-8 s). With parallel encoding there was a "shrinkage" in the visuomotor transformation, i.e., the distances between the markings were systematically smaller than the corresponding inter-target distances. When the targets were presented sequentially without distractors, marking accuracy improved with the total number of targets fixated and shrinkage in the visuomotor transformation occurred only with parallel encoding, i.e., when subjects did not fixate the targets. In all experimental conditions for trials in which targets were fixated during encoding, there was little correspondence between the marking sequence and the sequence in which the targets were fixated. We conclude that subjects benefit from fixating targets for subsequent target-oriented manual actions when the targets are presented among distractors and when presented sequentially; when distinct targets are presented simultaneously against a blank background, they are efficiently encoded in parallel largely by peripheral vision.

Adult↗

PubMed2001

Perspectives and problems in motor learning.

Movement provides the only means we have to interact with both the world and other people. Such interactions can be hard-wired or learned through experience with the environment. Learning allows us to adapt to a changing physical environment as well as to novel conventions developed by society. Here we review motor learning from a computational perspective, exploring the need for motor learning, what is learned and how it is represented, and the mechanisms of learning. We relate these computational issues to empirical studies on motor learning in humans.

Journal Article↗