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Biomedical subjects

Melvyn A Goodale

Publications and source records attributed to Melvyn A Goodale.

At least 19 recordsLinked to original sources

Naming and grasping common objects: a priming study.

This study examined the effects of priming on response latency when participants named and/or grasped common objects. A repetition-priming paradigm was used. An object was presented during a study phase and then was presented again during testing along with other objects that had not been seen before. In experiment 1, the studied objects were either named twice or grasped twice, named first and then grasped, or grasped first and then named. We found a strong priming effect (i.e., decreased latency) when naming was preceded by naming, as well as by grasping, but no priming effect when grasping was preceded by either naming or grasping. In experiment 2, we investigated the effects of priming in naming-naming and grasping-grasping paradigms, with and without a change in object orientation from study to test. As expected, we found significant priming of naming by naming, and the effect was not reduced by orientation change. Again, we found no evidence of priming in grasping. Experiment 3 was designed to examine how different kinds of perceptual and visuomotor processing (naming, orientation matching, orientation discrimination, simple observation, and grasping) during the study phase affect naming at a later test phase. We found significant priming of naming following all study conditions. Notably, the effect differed depending on how much "perceptual" processing was involved in the study phase. The results clearly indicate that perceptual/semantic processing is more dependent on memory than visuomotor processing, which instead relies more on moment-to-moment computations.

Adolescent↗

When two eyes are better than one in prehension: monocular viewing and end-point variance.

Previous research has suggested that binocular vision plays an important role in prehension. It has been shown that removing binocular vision affects (negatively) both the planning and on-line control of prehension. It has been suggested that the adverse impact of removing binocular vision is because monocular viewing results in an underestimation of target distance in visuomotor tasks. This suggestion is based on the observation that the kinematics of prehension are altered when viewing monocularly. We argue that it is not possible to draw unambiguous conclusions regarding the accuracy of distance perception from these data. In experiment 1, we found data that contradict the idea that a consistent visuomotor underestimation of target distance is an inevitable consequence of monocular viewing. Our data did show, however, that positional variance increases under monocular viewing. We provide an alternative explanation for the kinematic changes found when binocular vision is removed. Our account is based on the changes in movement kinematics that occur when end-point variance is altered following the removal of binocular vision. We suggest that the removal of binocular vision leads to greater perceptual uncertainty (e.g. less precise stimulus cues), resulting in changes in the kinematics of the movement (longer duration movements). Our alternative account reconciles some differences within the research literature. We conducted a series of experiments to explore further the issue of when binocular information is advantageous in prehension. Three subsequent experiments were employed which varied binocular/monocular viewing in selectively lit conditions. Experiment 2 explored the differences in prehension measured between monocular and binocular viewing in a full cue environment with a continuous view of the target object. Experiment 3 required participants to reach, under a monocular or binocular view, for a continuously visible self-illuminated target object in an otherwise dark room. In Experiment 3, the participant could neither see the target object nor the reaching hand following initiation of the prehension movement. Our results suggest that binocular vision contributes to prehension by providing additional information (cues) to the nervous system. These cues appear to be weighted differentially according to the particular constellation of stimulus cues available to the participants when reaching to grasp. One constant advantage of a binocular view appears to be the provision of on-line information regarding the position of the hand relative to the target. In reduced cue conditions (i.e. where a view of the target object is lost following initiation of the movement), binocular information regarding target location appears to be particularly useful in the initial programming of reach distance. Our results are a step towards establishing the specific contributions that binocular vision makes to the control of prehension.

Biomechanical Phenomena↗

An evolving view of duplex vision: separate but interacting cortical pathways for perception and action.

In 1992, Goodale and Milner proposed a division of labour in the visual pathways of the primate cerebral cortex between a dorsal stream specialised for the visual control of action and a ventral stream dedicated to the perception of the visual world. In the years since this original proposal, support for the perception-action hypothesis has come from neuroimaging experiments, human neuropsychology, monkey neurophysiology, and human psychophysical experiments. Indeed, some of the strongest support for this hypothesis has come from behavioural experiments showing that visually guided actions are largely refractory to perceptual illusions. Although controversial, the findings from this literature both support the original hypothesis and suggest important modifications. The ongoing challenge for neurobiologists is to map these behavioural findings onto their corresponding neural substrates.

Animals↗

Spared somatomotor and cognitive functions in a patient with a large porencephalic cyst revealed by fMRI.

To date functional magnetic resonance imaging (fMRI) has not been extensively used in presurgical evaluation of patients with intractable epilepsy. Patient S.P. presented with left frontal originating seizures, secondary to a large porencephalic cyst that encompassed much of his occipital and temporal cortex and a substantial portion of ipsilateral parietal cortex. Nevertheless, S.P. did not demonstrate any gross impairments of praxis or speech. Scalp electroencephalogram (EEG) telemetry revealed reduced background activity in the left hemisphere, an absence of identifiable normal sleep states, and confirmed the left frontal origin of his seizures with a prolonged postictal state, suggesting that the remaining cortex in S.P.'s left hemisphere did not function normally despite his apparently normal appearance. Dichotic listening results also suggested that S.P. had an atypical language representation suggestive of either bilateral or right hemisphere speech representation. Surgical intervention to remove the remaining left hemisphere cortex was a serious consideration for treatment of S.P.'s seizures. We used fMRI to evaluate whether or not the remaining cortex in S.P.'s left hemisphere supported any cognitive or motor functions. Even though the volume of cerebral cortex was severely reduced and displaced in the left hemisphere, fMRI revealed significant activation in this remaining tissue in response to motor, somatosensory, and word generation tasks. In other words, we were able to demonstrate using fMRI that the remaining tissue in S.P.'s left hemisphere continued to support some motor and cognitive functions. The possible implications of these findings in terms of functional reorganisation are discussed briefly.

Brain Diseases↗

Two distinct modes of control for object-directed action.

There are multiple routes from vision to action that play a role in the production of visually guided reaching and grasping. What remain to be resolved, however, are the conditions under which these various routes are recruited in the generation of actions and the nature of the information they convey. We argue in this chapter that the production of real-time actions to visible targets depends on pathways that are separate from those mediating memory-driven actions. Furthermore, the transition from real-time to memory-driven control occurs as soon as the intended target is no longer visible. Real-time movements depend on pathways from the early visual areas through to relatively encapsulated visuomotor mechanisms in the dorsal stream. These dedicated visuomotor mechanisms, together with motor centers in the premotor cortex and brainstem, compute the absolute metrics of the target object and its position in the egocentric coordinates of the effector used to perform the action. Such real-time programming is essential for the production of accurate and efficient movements in a world where the location and disposition of a goal object with respect to the observer can change quickly and often unpredictably. In contrast, we argue that memory-driven actions make use of a perceptual representation of the target object generated by the ventral stream. Unlike the real-time visuomotor mechanisms, perception-based movement planning makes use of relational metrics and scene-based coordinates. Such computations make it possible, however, to plan and execute actions upon objects long after they have vanished from view.

Humans↗

Visual control of action but not perception requires analytical processing of object shape.

The visual perception of object shape depends on 'holistic' processing in which a given dimension cannot be perceptually isolated from the other dimensions of the object. The visual control of action (such as grasping an object), however, which is mediated by cortical areas that are largely independent of those mediating conscious perception, must take into account only the most action-relevant dimension of an object without being misled by other non-relevant object features. Here we report the results of two experiments showing that vision for perception and vision for action deal with objects in a fundamentally different manner. We tested participants' ability to make perceptual judgements of the width of different rectangular objects or to grasp them across their width, while in both cases ignoring length. Participants could not ignore length when making perceptual judgements of width but they could completely ignore length when grasping the same objects. These results suggest that in situations in which the elementary dimensions of an object's shape are perceived in a holistic manner, the same dimensions are treated analytically when a visually guided action is directed at that same object.

Form Perception↗

Ventral occipital lesions impair object recognition but not object-directed grasping: an fMRI study.

D.F., a patient with severe visual form agnosia, has been the subject of extensive research during the past decade. The fact that she could process visual input accurately for the purposes of guiding action despite being unable to perform visual discriminations on the same visual input inspired a novel interpretation of the functions of the two main cortical visual pathways or 'streams'. Within this theoretical context, the authors proposed that D.F. had suffered severe bilateral damage to her occipitotemporal visual system (the 'ventral stream'), while retaining the use of her occipitoparietal visual system (the 'dorsal stream'). The present paper reports a direct test of this idea, which was initially derived from purely behavioural data, before the advent of modern functional neuroimaging. We used functional MRI to examine activation in her ventral and dorsal streams during object recognition and object-directed grasping tasks. We found that D.F. showed no difference in activation when presented with line drawings of common objects compared with scrambled line drawings in the lateral occipital cortex (LO) of the ventral stream, an area that responded differentially to these stimuli in healthy individuals. Moreover, high-resolution anatomical MRI showed that her lesion corresponded bilaterally with the location of LO in healthy participants. The lack of activation with line drawings in D.F. mirrors her poor performance in identifying the objects depicted in the drawings. With coloured and greyscale pictures, stimuli that she can identify more often, D.F. did show some ventral-stream activation. These activations were, however, more widely distributed than those seen in control participants and did not include LO. In contrast to the absent or abnormal activation observed during these perceptual tasks, D.F. showed robust activation in the expected dorsal stream regions during object grasping, despite considerable atrophy in some regions of the parietal lobes. In particular, an area in the anterior intraparietal sulcus was activated more for grasping an object than for just reaching to that object, for both D.F. and controls. In conclusion, we have been able to confirm directly that D.F.'s visual form agnosia is associated with extensive damage to the ventral stream, and that her spared visuomotor skills are associated with visual processing in the dorsal stream.

Agnosia↗

Flexible retinotopy: motion-dependent position coding in the visual cortex.

Although the visual cortex is organized retinotopically, it is not clear whether the cortical representation of position necessarily reflects perceived position. Using functional magnetic resonance imaging (fMRI), we show that the retinotopic representation of a stationary object in the cortex was systematically shifted when visual motion was present in the scene. Whereas the object could appear shifted in the direction of the visual motion, the representation of the object in the visual cortex was always shifted in the opposite direction. The results show that the representation of position in the primary visual cortex, as revealed by fMRI, can be dissociated from perceived location.

Attention↗

Visually guided grasping produces fMRI activation in dorsal but not ventral stream brain areas.

Although both reaching and grasping require transporting the hand to the object location, only grasping also requires processing of object shape, size and orientation to preshape the hand. Behavioural and neuropsychological evidence suggests that the object processing required for grasping relies on different neural substrates from those mediating object recognition. Specifically, whereas object recognition is believed to rely on structures in the ventral (occipitotemporal) stream, object grasping appears to rely on structures in the dorsal (occipitoparietal) stream. We used functional magnetic resonance imaging (fMRI) to determine whether grasping (compared to reaching) produced activation in dorsal areas, ventral areas, or both. We found greater activity for grasping than reaching in several regions, including anterior intraparietal (AIP) cortex. We also performed a standard object perception localizer (comparing intact vs. scrambled 2D object images) in the same subjects to identify the lateral occipital complex (LOC), a ventral stream area believed to play a critical role in object recognition. Although LOC was activated by the objects presented on both grasping and reaching trials, there was no greater activity for grasping compared to reaching. These results suggest that dorsal areas, including AIP, but not ventral areas such as LOC, play a fundamental role in computing object properties during grasping.

Adult↗

FMRI evidence for a 'parietal reach region' in the human brain.

Event-related functional magnetic resonance imaging was used to examine activation in the posterior parietal cortex when subjects made pointing movements or saccades to the same spatial location. One region, well positioned to be homologous to the monkey parietal reach region (PRR), responded preferentially during memory-delay trials in which the subject planned to point to a specific location as compared to trials in which the subject planned to make a saccade to that same location. We therefore conclude that activation in this region is related to specific motor intent; i.e. it encodes information related to the subject's intention to make a specific movement to a particular spatial location.

Arm↗

A haptic size-contrast illusion affects size perception but not grasping.

Object features (e.g. size, shape and orientation) are relevant for recognition and identification, but also for the control of manual actions. Converging evidence suggests a dissociation between the visual systems that mediate object perception and object-directed action. Here we present evidence suggesting that a similar dissociation might exist in the haptic domain. We demonstrate that a haptic variation of a size-contrast illusion influences the perceived size of a target object, but not the degree to which the hand is opened when that object is the target of a grasping movement. This finding is consistent with the view that object perception is "scene-based" and takes into consideration not only the size of the target object but also the sizes of other nearby objects. In contrast, the control of object-directed action is primarily driven by the absolute size of the target object independent of the relative sizes of other objects in the environment, suggesting a "actor-based" frame of reference. The present findings suggest that dissociations between action and perception are not unique to the visual system, but might instead reflect a general organizational principle of sensory processing.

Adult↗

The influence of visual motion on fast reaching movements to a stationary object.

One of the most important functions of vision is to direct actions to objects. However, every time that vision is used to guide an action, retinal motion signals are produced by the movement of the eye and head as the person looks at the object or by the motion of other objects in the scene. To reach for the object accurately, the visuomotor system must separate information about the position of the stationary target from background retinal motion signals-a long-standing problem that is poorly understood. Here we show that the visuomotor system does not distinguish between these two information sources: when observers made fast reaching movements to a briefly presented stationary target, their hand shifted in a direction consistent with the motion of a distant and unrelated stimulus, a result contrary to most other findings. This can be seen early in the hand's trajectory (approximately 120 ms) and occurs continuously from programming of the movement through to its execution. The visuomotor system might make use of the motion signals arising from eye and head movements to update the positions of targets rapidly and redirect the hand to compensate for body movements.

Eye Movements↗

Measuring unconscious actions in action-blindsight: exploring the kinematics of pointing movements to targets in the blind field of two patients with cortical hemianopia.

We tested two patients with posterior cerebral lesions on two pointing tasks. In the first task, the patients pointed to targets presented on a touch screen monitor and pointing accuracy was recorded. One patient (JR) demonstrated good localisation of targets presented to her blind field while the other patient (YP) did not. Movement kinematics were measured in the second task to compare the kinematics of movements made to sighted field targets with those made to blind field targets. For this version of the task both patients demonstrated above chance localisation of blind field targets although the slope of the relationship between the end of pointing movements and the target locations was significantly steeper for JR than for YP. Furthermore, JR showed a kinematic profile for movements made to blind field targets that mirrored the profile of kinematics to sighted field targets. That is, both peak velocity and time to peak velocity increased with increasing target eccentricity for movements made to blind and sighted field targets alike. Although patient YP now showed more reliable spatial localisation on this pointing task when compared with the touch screen task, his kinematics for movements made to targets in his blind field were quite different from those made to targets in his sighted field. Based on the patients' CT scans, we suggest that the superior performance of patient JR is a consequence of greater sparing of her parietal cortex in the damaged hemisphere.

Adult↗

The effects of different aperture-viewing conditions on the recognition of novel objects.

The process of learning the structure of novel objects involves the selective use of information available in the distal stimulus. By allowing participants to explore the object within a limited field of view, we were able to examine more rigorously what regions of the object are actually selected in the learning process. Participants explored objects either by moving a circular aperture over a stationary novel object (the aperture-movement condition), or by moving the object behind a stationary aperture (the object-movement condition). Given the differences in how the spatial layout of object parts is revealed in the two study conditions, we expected that exploration would be more systematic in the aperture-movement condition than it would be in the object-movement condition, and would lead to better object recognition. We show evidence that in the aperture-movement condition exploration patterns were more related to the structure of the object and, as a consequence, the aperture-movement condition resulted in more accurate recognition in a later old--new discrimination test.

Adult↗

Perceptual illusion and the real-time control of action.

Participants were cued by an auditory tone to grasp a target object from within a size-contrast display. The peak grip aperture was unaffected by the perceptual size illusion when the target array was visible between the response cue and movement onset (vision trials). The grasp was sensitive to the illusion, however, when the target array was occluded from view when the response was cued (occlusion trials). This was true when the occlusion occurred 2.5 s before the response cue (delay), but also when the occlusion coincided with the response cue (no-delay). Unlike previous experiments, vision and occlusion trials were presented in random sequence. The results suggest that dedicated, real-time visuomotor mechanisms are engaged for the control of action only after the response is cued, and only if the target is visible. These visuomotor mechanisms compute the absolute metrics of the target object and therefore resist size-contrast illusions. In other situations (e.g. prior to the response cue, or if the target is no longer visible), a perceptual representation of the target object can be used for action planning. Unlike the real-time visuomotor mechanisms, perception-based movement planning makes use of relational metrics, and is therefore sensitive to size-contrast illusions.

Cues↗

Target selection for reaching and saccades share a similar behavioral reference frame in the macaque.

The selection of one of two visual stimuli as a target for a motor action may depend on external as well as internal variables. We examined whether the preference to select a leftward or rightward target depends on the action that is performed (eye or arm movement) and to what extent the choice is influenced by the target location. Two targets were presented at the same distance to the left and right of a fixation position and the stimulus onset asynchrony (SOA) was adjusted until both targets were selected equally often. This balanced SOA time is then a quantitative measure of selection preference. In two macaque monkeys tested, we found the balanced SOA shifted to the left side for left-arm movements and to the right side for right-arm movements. Target selection strongly depended on the horizontal target location. By varying eye, head, and trunk position, we found this dependency embedded in a head-centered behavioral reference frame for saccade targets and, somewhat counter-intuitively, for reach targets as well. Target selection for reach movements was influenced by the eye position, while saccade target selection was unaffected by the arm position. These findings suggest that the neural processes underlying target selection for a reaching movement are to a large extent independent of the coordinate frame ultimately used to make the limb movement, but are instead closely linked to the coordinate frame used to plan a saccade to that target. This similarity may be indicative of a common spatial framework for hand-eye coordination.

Animals↗

Learned perceptual associations influence visuomotor programming under limited conditions: cues as surface patterns.

The present set of three experiments was designed to extend the findings that visuomotor programming can make use of learned size information under some, but not all, conditions. An association was established between the size of square wooden blocks and a perceptual cue in all experiments. In Experiment 1 the perceptual cue to size was a small two-dimensional drawing of a shape affixed to the top of the blocks (e.g. triangle = large; circle = small, or vice versa). In Experiment 2 size and shape were again associated but this time a pattern of two-dimensional shapes covered the visible surface of the blocks. In Experiment 3 block size was associated with the colour of a small circle affixed to the top of the blocks (e.g. red = large; yellow = small, or vice versa). All of the subjects grasped the blocks, and on other trials estimated the size of the blocks by opening their thumb and finger a matching amount. Consistent with previous reports, in all experiments, the learned associations changed the perceived size of two test blocks halfway in size between the large and small blocks: estimations of the test block matched by shape or colour to the group of large objects were smaller than estimations of the test block matched to the group of small objects. The effect appears to result from relative-size comparisons being made between the medium-sized test blocks and the size category (large or small) associated with the matching shape or colour cue. Despite the significant effect of the learned perceptual associations on manual estimations, no effect on grip scaling was seen when the cues associated with size were single small elements centred on the top of the block (Experiment 1 and Experiment 3). Changes in grip scaling corresponded to the change in perceived size only when the cue to size covered the entire block (Experiment 2), forming a surface pattern. These findings suggest that visuomotor programming is more likely to use learned size information when the cue providing the size association covers the visible surface of the target objects, perhaps by acting as a texture that provides reliable information about the target's material and identity.

Association Learning↗