Transfer of adaptation between ocular saccades and arm movements

Previous studies found little or no transfer of adaptation from reactive saccades to arm pointing movements, which suggests that the two motor systems rely on distinct adaptive mechanisms. However, this conclusion is based on experiments about the adaptation of response amplitudes, which is known to follow somewhat different principles than the adaptation of response directions. In the present study, we therefore investigate whether adapting the direction of reactive saccades will transfer to arm movements. We also test transfer in the opposite direction, from the arm to the eyes. Participants executed aimed saccades or arm movements from a central starting point towards visual targets in the participants' frontal plane. Targets were presented in eight possible locations along a circle of 20 cm radius about the starting point; each remained for 200 ms in one position, and was then displaced along the circle by -15 degrees . Participants from group E adapted to these double-stepped targets while executing eye movements, and were then tested for transfer while executing arm movements. The reciprocal design was used in participants from group A. Adaptive change in group A was about 14 degrees , while in group E it was only about 7 degrees . Transfer of adaptation was substantial, and was more pronounced when using the arm (i.e., eye-to-arm transfer in group E) rather than the eyes (i.e., arm-to-eye transfer in group A). Strong aftereffects were yielded in both groups. This pattern of findings implies that the adaptive change observed in our study was mainly based on recalibration rather than on cognitive strategies (strong aftereffects), that eyes and arm had access to a common adaptive mechanism (substantial transfer), and that the arm had better access than the eyes (larger adaptation and transfer when using the arm). When considering this outcome along with the available literature, it appears that arm and eyes may rely sometimes on a common and sometimes on distinct adaptive mechanisms, depending on the adapted parameter and on the nature of the motor task.     

Reduced felt arm sensation effects on visual adaptation

Proprioception is often considered to be critically involved in producing adaptation to a prism-induced visual displacement. The present study focused on reduction of proprioceptive feedback during prism exposure by means of hypnotically induced anesthesia in the adapting arm. In addition, intermanual transfer was considered. Results showed adaptation occurring in situations where S could feel arm sensations while viewing arm movement during a prism exposure. However, if the adapting arm was hypnotically anesthetized while still remaining mobile, adaptation did not occur. No intermanual transfer was found between the adapted arm and the unadapted arm.     

Adaptation in the constancy of visual direction measured by a one-trial method

Adaptation to field displacement during head movements in the direction with the head rotation and in the direction against it was produced under otherwise identical conditions and compared; the field displacement rate was also varied. A rapid training procedure was used, and a novel one-trial test was employed that could measure the adaptation well enough to compare the effects of various training conditions. The one-trial test measured the magnitude of one of the manifestations of adaptation, the apparent displacement of a stationary target during head movements. This apparent horizontal target displacement was transformed into an oblique one by having the head movements that brought forth the apparent target displacement simultaneously cause an objective vertical target displacement. The slant of the resultant apparent motion path varied with the magnitude of the apparent horizontal target displacement. It was measured by having S reproduce its slant angle. It was found that adaptation to field displacement in the direction with the head rotation was consistently greater than adaptation to the opposite displacement conditions. An explanation for this result is offered.     

Additivity of components of prismatic adaptation

Ss pointed with each hand at a light or at the unseen toe and looked in the direction of the unseen toe before, during, and after training one arm to point to a visual target which was progressively displaced to one side by a prism. Results show that a proprioceptive change in the trained arm is a universal component of the adaptation. When a change in the eye-head system occurs, it and the proprioceptive change in the arm sum to the total adaptation and it is accompanied by a predictable degree of intermanual transfer of the adaptation, as a felt-position theory of adaptation would predict. However, when there is no change in the eye-head system, the proprioceptive shift is not always sufficient to account for the total adaptive shift.     

Properties of intermodal transfer after dual visuo- and auditory-motor adaptation

Previous work documented that sensorimotor adaptation transfers between sensory modalities: When subjects adapt with one arm to a visuomotor distortion while responding to visual targets, they also appear to be adapted when they are subsequently tested with auditory targets. Vice versa, when they adapt to an auditory-motor distortion while pointing to auditory targets, they appear to be adapted when they are subsequently tested with visual targets. Therefore, it was concluded that visuomotor as well as auditory-motor adaptation use the same adaptation mechanism. Furthermore, it has been proposed that sensory information from the trained modality is weighted larger than sensory information from an untrained one, because transfer between sensory modalities is incomplete. The present study tested these hypotheses for dual arm adaptation. One arm adapted to an auditory-motor distortion and the other either to an opposite directed auditory-motor or visuomotor distortion. We found that both arms adapted significantly. However, compared to reference data on single arm adaptation, adaptation in the dominant arm was reduced indicating interference from the non-dominant to the dominant arm. We further found that arm-specific aftereffects of adaptation, which reflect recalibration of sensorimotor transformation rules, were stronger or equally strong when targets were presented in the previously adapted compared to the non-adapted sensory modality, even when one arm adapted visually and the other auditorily. The findings are discussed with respect to a recently published schematic model on sensorimotor adaptation.     

Adaptation to field displacement during head movement unrelated to the constancy of visual direction

Adaptation to vertical field displacements dependent on head turning about a vertical axis was demonstrated under two conditions, rapid training with 100 head movements and 1-h-long training with continuous head turning. The effect of rapid training was measured with the slant estimation method. Adaptation after the longer training was ascertained by comparing the uncertainty ranges for apparent target immobility before and after the adaptation period. Adaptation to field displacements in directions parallel to the plane of the head rotation obtained under corresponding conditions was also measured and found to be somewhat greater than adaptation to vertical field displacements. The result of work by Wallach and Frey that adaptation to field displacement in the direction with the head rotation is greater than to displacement against it was corroborated. While the previous result had, been obtained with rapid adaptation and with the slant estimation method, we confirmed it with 1-h training and by measuring the uncertainty ranges before and after the adaptation period.     

Target distance and adaptation in distance perception in the constancy of visual direction

Hay and Sawyer recently demonstrated that the constancy of visual direction (CVD) also operates for near targets. A luminous spot in the dark, 40 cm from the eyes, was perceived as stationary when S nodded his head. This implies that CVD takes target distance, as well as head rotation, into account as a stationary environment is perceived during head movements. Distance is a variable in CVD because, during a turning or nodding of the head, the eyes become displaced relative to the main target direction, the line between the target and the rotation axis of the head. This displacement of the eyes during head rotation causes an additional change in the target direction, i.e., a total angular change greater than the angle of the head rotation. The extent of this additional angular displacement is greater the nearer the target. We demonstrated that the natural combination of accommodation and convergence can supply the information needed by the nervous system to compensate for this additional target displacement. We also found that wearing glasses that alter the relation between these oculomotor adjustments and target distance produces an adaptation in CVD. An adaptation period of 1.5 h produced a large adaptation effect. This effect was not entirely accounted for by an adaptation in distance perception. Measurements of the alteration between oculomotor cues and registered distance with two kinds of tests for distance perception yielded effects significantly smaller than the effect measured with the CVD test. We concluded that the wearing of the glasses had also produced an adaptation within CVD.     

Transfer to intermediate forms following concept discrimination by pigeons: chimeras and morphs

Two experiments examined pigeons' generalization to intermediate forms following training of concept discriminations. In Experiment 1, the training stimuli were sets of images of dogs and cats, and the transfer stimuli were head/body chimeras, which humans tend to categorize more readily in terms of the head part rather than the body part. In Experiment 2, the training stimuli were sets of images of heads of dogs and cats, and the intermediate stimuli were computer-generated morphs. In both experiments, pigeons learned the concept discrimination quickly and generalized with some decrement to novel instances of the categories. In both experiments, transfer tests were carried out with intermediate forms generated from both familiar and novel exemplars of the training sets. In Experiment 1, the pigeons' transfer performance, unlike that of human infants exposed to similar stimuli, was best predicted by the body part of the stimulus when the chimeras were formed from familiar exemplars. Spatial frequency analysis of the stimuli showed that the body parts were richer in high spatial frequencies than the head parts, so these data are consistent with the hypothesis that categorization is more dependent on local stimulus features in pigeons than in humans. There was no corresponding trend when the chimeras were formed from novel exemplars. In Experiment 2, when morphs of training stimuli were used, response rates declined smoothly as the proportion of the morph contributed by the positive stimulus fell, although results with morphs of novel stimuli were again less orderly.     

Does hand dominance affect the use of motor abundance when reaching to uncertain targets?

This study investigated hemispheric differences in utilizing motor abundance to achieve flexible patterns of joint coordination when reaching to uncertain target locations. Right-handed participants reached with each arm to the same central target when its final location was certain or when there was a 66% probability that its location could change after movement initiation. Use of greater motor abundance was observed when participants reached to the central target under target location uncertainty regardless of the arm used to reach. Joint variance associated with variability of movement direction was larger when reaching with the left, non-dominant arm. This arm also exhibited higher hand path variability compared to the dominant arm. These arm differences were not found when the final (central) target location was known in advance. The results provide preliminary evidence for a greater ability of the dominant (right) arm/left hemisphere to decouple directions in joint space. That is, to increase the use of motor abundance without simultaneously inducing unwanted hand path variability requires that joint variations be restricted to a limited subspace of joint space. Hemispheric differences in motor planning did not appear to account for arm differences related to the use of motor abundance.     

The constancy of the orientation of the visual field

Evidence is presented that the perceived immobility of the environment during tilting of the head from side to side results from a compensating process. This compensating process operates well only when peripheral vision is present. An objectively stationary environment was, for instance, not perceived as immobile during head tilting when vision was confined to the macular region of the retina. The compensating process could be rapidly altered by exposure to environmental tilting during and dependent on head tilting. Such adaptation had the result that some environmental tilting that normally is perceived led to apparent immobility.     

Counteradaptation after exposure to displaced visual direction

Counteradaptation, previously demonstrated in connection with adaptation in distance perception, was obtained after exposure to displaced visual direction. When S adapted to a laterally displacing wedge prism by walking during the exposure period, there was not only a change in the perceived visual direction, but also a change m the proprioceptively perceived walking direction. When S adapts to lateral displacement of the visual direction by looking at his stationary or his moving arm, visual adaptation is obtained in the latter, but not in the former, case (Held & Hein, 1958). We obtained a change in the proprioceptively perceived position of the arm when it was stationary during the exposure period, a condition which had not yielded visual adaptation, and a much smaller, not significant, change in the felt position in the case of the actively moved arm. In the present experiments, changes in proprioceptively perceived direction or position amounted to counteradaptation.     

Analogical transfer from a simulated physical system

Previous research has consistently found that spontaneous analogical transfer is strongly tied to concrete and contextual similarities between the cases. However, that work has largely failed to acknowledge that the relevant factor in transfer is the similarity between individuals' mental representations of the situations rather than the overt similarities between the cases themselves. Across several studies, we found that participants were able to transfer strategies learned from a perceptually concrete simulation of a physical system to a task with very dissimilar content and appearance. This transfer was reflected in better performance on the transfer task when its underlying dynamics were consistent rather than inconsistent with the preceding training task. Our data indicate that transfer in these tasks relies on the perceptual and spatial nature of the training task but does not depend on direct interaction with the system, with participants performing equally well after simply observing the concrete simulation. We argue that participants generated a spatial, dynamic, and force-based mental model while interacting with the training simulation and tended to spontaneously interpret the transfer task according to this primed model. Unexpectedly, our data consistently show that transfer was independent of reported recognition of the analogy between tasks: Although such recognition was associated with better overall performance, it was not associated with better transfer (in terms of applying an appropriate strategy). Together, these findings suggest that analogical transfer between overtly dissimilar cases may be much more common--and much more relevant to our cognitive processing--than is generally assumed.     

Dual adaptation of apparent concomitant motion contingent on head rotation frequency

Perceived movement of a stationary visual stimulus during head motion was measured before and after adaptation intervals during which participants performed voluntary head oscillations while viewing a moving spot. During these intervals, participants viewed the spot stimulus moving alternately in the same direction as the head was moving during either .25- or 2.0-Hz oscillations, and then in the opposite direction as the head at the other of the two frequencies. Postadaptation measures indicated that the visual stimuli were perceived as stationary only if traveling in the same direction as that viewed during adaptation at the same frequency of head motion. Thus, opposite directions of spot motion were perceived as stationary following adaptation depending on head movement frequency. The results provide an example of the ability to establish dual (or “context-specific”) adaptations to altered visual—vestibular feedback.     

Direct-effects and after-effects of dynamic adaptation on intralimb and interlimb transfer

After-effects following sensorimotor adaptation are generally considered as evidence for the formation of an internal model, although evidence lacks on whether the absence of after-effects necessarily indicates that the adaptation did not result in the formation of an internal model. Here, we examined direct- and after-effects of dynamic adaptation with one arm at one workspace on subsequent performance with the other arm, as well as the same arm at another workspace. During training, subjects performed reaching movements under a novel dynamic condition with the right arm; during testing, they performed reaching movements with the left or right arm at a new workspace, under either the same dynamic condition (direct-effects) or a normal condition (after-effects). Results showed significant transfer within the same arm in terms of both direct- and after-effects, but significant transfer across the arms only in terms of direct-effects. These findings suggest that the formation of an internal model does not always result in after-effects. They also support the idea that the neural representation developed after sensorimotor adaptation comprise some aspects that are effector independent and other aspects that are effector dependent; and that direct- and after-effects following sensorimotor adaptation mainly reflect the effector-independent and the effector-dependent aspects, respectively.     

Shifts in kinesthesis through time and after active and passive movement.

The position sense of a stationary arm was investigated subsequent to an horizontally adductive movement with axis the shoulder joint. The right arm was the treated arm: it reached a test position actively, using minimal voluntary effort, or passively from each of 10 starting positons. The blind-folded S localized the index finger of the treated arm by attempting to touch it with the index finger of his left hand. The results indicate that subsequent to active movement the final position of a limb is more accurately known than a position resulting from passive movement. A second finding is that concomitant with both forms of limb placement there is a unidirectional drift of perceived limb position over trials.     

Visuomotor coordination and intermanual transfer for a proprioceptive reaching task

Two experiments were conducted to determine the role of head constraint, whether present or absent and arm exposure type (terminal or continuous) on the production of intermanual transfer to two types of visual distortion. Experiment 1 investigated intermanual transfer to binocular, lateral prism displacement where the prism base orientation for both eyes was in the same direction. Experiment 2 determined whether intermanual transfer could be produced to squint prism viewing where the prism base orientation for each eye was in an opposite direction (base-out prisms). In both experiments transfer was produced when either head movement during prism exposure was unconstrained or when a terminal arm exposure was employed. Maximal transfer was produced when both of these conditions were employed.     

Learning motor synergies makes use of information on muscular load

Prism adaptation, a form of procedural learning, requires the integration of visual and motor information for its proper acquisition. Although the role of the visual feedback has begun to be understood, the nature of the motor information necessary for the development of the adaptation remains unknown. In this work we have tested the idea that modifying the arm load at different stages of the adaptation process, and the ensuing change of motor information perceived by the subjects, would modify the final properties of the adaptation. We trained a set of subjects to throw balls to a target while wearing prism glasses and varied the weight of their arms at different time points during the task. We observed that the acquisition of the adaptation was not affected by the change in load. However, its persistence (i.e., the aftereffect) was reduced when tested under a weight condition different from the training trials. Furthermore, when the training weight conditions were restored later during testing, a second, late aftereffect was unmasked, suggesting that the missing aftereffect did not disappear but had remained latent. Our results show that the internal representation of a motor memory incorporates information about load conditions and that the memory stored under a specific weight condition can be fully retrieved only when the original training condition is restored.     

The effect of training and handedness on the performance of two simple motor tasks

In the first experimental series nine right-handed and nine left-handed subjects were tested on each side on two tasks. The first task measured the accuracy of reproduction of pressure in attempted extension of the elbow joint, and the second the speed of oscillation of attempted flexion and extension of the elbow joint. The results showed no significant difference in performance on the two sides in the accuracy task but a significant difference in performance on the speed task.

In the second experimental series the effect of 15-minutes daily training on each of two tasks over a four-week period was investigated on six subjects. The two tasks were the same as those used in the first experimental series, and attempted movement at the metacarpo-phalangeal joint as well as the elbow joint was studied. The mean results of all subjects showed a significant reduction in the mean error of the “accuracy” task over the training period for both the finger and arm. However, further analysis suggests that this may have been due to a clarifying of the subjects' concept of the target value rather than an improvement in his ability to grade the appropriate muscle contractions. A significant increase in the mean speed of oscillation over the training period was also recorded for both finger and arm. This improvement was accompanied in most instances by a corresponding decrease in variation of cycle length of the individual oscillations, which is interpreted as an improvement in the subjects' “timing” of the appropriate muscle contractions.     

Discriminative conditioning of prism adaptation

Two experiments were used to demonstrate that adaptation to ll-deg prism displacement can be conditioned to the stimuli associated with the goggles in which the prisms are housed. In Experiment 1 it was found that repeated alternation between a series of target-pointing responses while wearing prism goggles and a series of responses without prism goggles led to larger adaptive shift when S was tested with nondisplacing goggles than when tested without goggles. The results of Experiment 2 indicated that the adaptation revealed in the first experiment was primarily proprioceptive, rather than visual. Surprisingly, most Ss reported greater difficulty during the exposure period in overcoming the negative aftereffect than they did the prism-induced error.     

Learning and transfer of a relative phase pattern and a joint amplitude ratio in a rhythmic multijoint arm movement

According to the coordination dynamics perspective, one can characterize the learning of novel relative phase patterns as the formation of a stable attractor in the coordination landscape of the order parameter relative phase. The authors examined 18 participants' learning and transfer of a 90 degrees relative phase pattern and a 0.6-joint-amplitude ratio between the elbow and wrist. Variability in the relative phasing and the joint amplitude ratio between the elbow and wrist decreased with practice. Positive transfer of the 90 degrees relative phase pattern was not dependent on the learning arm (dominant or nondominant). Positive transfer of the joint amplitude ratio was dependent on the learning arm and the direction of transfer. The results demonstrated that relative phase is an order parameter that characterizes the coordination dynamics of learning and transferring multijoint arm movements, and they provide preliminary evidence that joint amplitude ratios act as order parameters in the learning and transfer of multijoint arm movements.