It is uncertain how vision and proprioception contribute to adaptation of voluntary arm movements. In normal participants, adaptation to imposed forces is possible with or without vision, suggesting that proprioception is sufficient; in participants with proprioceptive loss (PL), adaptation is possible with visual feedback, suggesting that proprioception is unnecessary. In experiment 1 adaptation to, and retention of, perturbing forces were evaluated in three chronically deafferented participants. They made rapid reaching movements to move a cursor toward a visual target, and a planar robot arm applied orthogonal velocity-dependent forces. Trial-by-trial error correction was observed in all participants. Such adaptation has been characterized with a dual-rate model: a fast process that learns quickly, but retains poorly and a slow process that learns slowly and retains well. Experiment 2 showed that the PL participants had large individual differences in learning and retention rates compared to normal controls. Experiment 3 tested participants’ perception of applied forces. With visual feedback, the PL participants could report the perturbation’s direction as well as controls; without visual feedback, thresholds were elevated. Experiment 4 showed, in healthy participants, that force direction could be estimated from head motion, at levels close to the no-vision threshold for the PL participants. Our results show that proprioceptive loss influences perception, motor control and adaptation but that proprioception from the moving limb is not essential for adaptation to, or detection of, force fields. The differences in learning and retention seen between the three deafferented participants suggest that they achieve these tasks in idiosyncratic ways after proprioceptive loss, possibly integrating visual and vestibular information with individual cognitive strategies.
The intention to execute a movement can modulate our perception of sensory events, and this modulation is observed ahead of both ocular and upper limb movements. However, theoretical accounts of these effects, and also the empirical data, are often contradictory. Accounts of "active touch", and the premotor theory of attention, have emphasized how movement intention leads to enhanced perceptual processing at the target of a movement, or on the to-be-moved effector. By contrast, recent theories of motor control emphasize how internal "forward" model (FM) estimates may be used to cancel or attenuate sensory signals that arise as a result of self-generated movements. We used behavioural and functional brain imaging (functional magnetic resonance imaging, fMRI) to investigate how perception of a somatosensory stimulus differed according to whether it was delivered to a hand that was about to execute a reaching movement or the alternative, nonmoving, hand. The results of our study demonstrate that a somatosensory stimulus delivered to a hand that is being prepared for movement is perceived to have occurred later than when that same stimulus is delivered to a nonmoving hand. This result indicates that it takes longer for a tactile stimulus to be detected when it is delivered to a moving limb and may correspond to a change in perceptual threshold. Our behavioural results are paralleled by the results of our fMRI study that demonstrated that there were significantly reduced blood-oxygen-level-dependent (BOLD) responses within the parietal operculum and insula following somatosensory stimulation of the hand being prepared for movement, compared to when an identical stimulus was delivered to a nonmoving hand. These findings are consistent with the prediction of FM accounts of motor control that postulate that central sensory suppression of somatosensation accompanies self-generated limb movements, and with previous reports indicating that effects of sensory suppression are observed in higher order somatosensory regions.
Alternating the point of gaze between an original (model or sitter, object, or scene) and a picture (paper, canvas, or digital touch screen) is the most common observational drawing strategy. However, a number of investigations into eye-hand interactions in drawing have revealed the existence of some "blind" drawing taking place (drawing the picture while the eye remains on the original or during gaze shifts between the original and the drawing). These observations of a direct visual-to-motor transformation challenge the commonly held assumption that the gaze-shifting strategy reflects a memory process in which the gaze on the original is used to encode a visual detail to short or long term memory, subsequently retrieved during the gaze on the picture. To study the blind drawing strategy in more depth during naturalistic drawing, we compared 3 basic drawing tasks-copying, contouring, and drawing of graded zones as lines, where original and picture were placed side by side on a vertical plane. We found that subjects drew almost continuously, thus exhibiting periods of blind drawing while the eye was on the original. The amount of blind drawing increased progressively between the copying task, the contouring task, and the graded zone task. When gaze shifted to the picture, it was generally to a fixation point located in advance of the hand on the part of the line not yet drawn. For individual tests, gaze ratios (gaze duration on original divided by gaze duration on picture) were approximately equal to drawing ratios (drawing duration during original gaze divided by drawing duration during picture gaze). We propose a general gaze-shift strategy that takes into account these observations. Gaze shifting is probably the most common form of eye move ment during observational drawing or drawing from life. It consists of the eyes alternating between an original-the model or sitter,
To copy a natural visual image as a line drawing, visual identification and extraction of features in the image must be guided by top-down decisions, and is usually influenced by prior knowledge. In parallel with other behavioral studies testing the relationship between eye and hand movements when drawing, we report here a functional brain imaging study in which we compared drawing of faces and abstract objects: the former can be strongly guided by prior knowledge, the latter less so. To manipulate the difficulty in extracting features to be drawn, each original image was presented in four formats including high contrast line drawings and silhouettes, and as high and low contrast photographic images. We confirmed the detailed eye–hand interaction measures reported in our other behavioral studies by using in-scanner eye-tracking and recording of pen movements with a touch screen. We also show that the brain activation pattern reflects the changes in presentation formats. In particular, by identifying the ventral and lateral occipital areas that were more highly activated during drawing of faces than abstract objects, we found a systematic increase in differential activation for the face-drawing condition, as the presentation format made the decisions more challenging. This study therefore supports theoretical models of how prior knowledge may influence perception in untrained participants, and lead to experience-driven perceptual modulation by trained artists.
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