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Contribution of the forebrain archistriatal gaze fields to auditory orienting behavior in the barn owl.

A region in the barn owl forebrain, referred to as the archistriatal gaze fields (AGF), is shown to be involved in auditory orienting behavior. In a previous study, electrical microstimulation of the AGF was shown to produce saccadic movements of the eyes and head, and anatomical data revealed that neurons in the AGF region of the archistriatum project directly to brainstem tegmental nuclei that mediate gaze changes. In this study, we investigated the effects of AGF inactivation on the auditory orienting responses of trained barn owls. The AGF and/or the optic tectum (OT) were inactivated pharmacologically using the GABAA agonist muscimol. Inactivation of the AGF alone had no effect on the probability or accuracy of orienting responses to contralateral acoustic stimuli. Inactivation of the OT alone decreased the probability of responses to contralateral stimuli, but the animals were still capable of orienting accurately toward stimuli on about 60% of the trials. Inactivation of both the AGF and the OT drastically decreased the probability of responses to 16-21% and, on the few trials that the animals did respond, there was no relationship between the final direction of gaze and the location of the stimulus. Thus, with the AGF and OT both inactivated, the animals were no longer capable of orienting accurately toward acoustic stimuli located on the contralateral side. These data confirm that the AGF is involved in gaze control and that the AGF and the OT have parallel access to gaze control circuitry in the brainstem tegmentum. In these respects, the AGF in barn owls is functionally equivalent to the frontal eye fields in primates.

Acoustic Stimulation↗

Quantitative determination of orientational and directional components in the response of visual cortical cells to moving stimuli.

The response characteristic of visual cortical cells to moving oriented stimuli consists mainly of directional (D) and orientational (O) components superimposed to a spontaneous activity (S). Commonly used polar plot diagrams reflect the maximal responses for different orientations and directions of stimulus movement with a periodicity of 360 degrees in the visual field. Fast Fourier analysis (FFT) is applied to polar plot data in order to determine the intermingled S, D, and O components. The zero order gain component of the spectrum corresponds to a (virtual) spontaneous activity. The first order component is interpreted as the strength of the direction selectivity and the second order component as the strength of the orientation specificity. The axes of the preferred direction and optimal orientation are represented by the respective phase values. Experimental data are well described with these parameters and relative changes of the shape of a polar plot can be detected with an accuracy better than 1%. The results are compatible with a model of converging excitatory and inhibitory inputs weighted according to the zero to second order components of the Fourier analysis. The easily performed quantitative determination of the S, D, and O components allows the study of pharmacologically induced changes in the dynamic response characteristics of single visual cortical cells.

Animals↗

Orientation behaviour of toads (Bufo bufo) displaced from the breeding site.

The sensory basis and spatial range of orientation to the breeding site were studied in the toad Bufo bufo, during two breeding seasons. Toads were displaced passively from their breeding pond and fitted with a tracking device to record the path of migration in individuals. The directional choice and the straightness of trails after release were used to quantify the effect of experimental treatments. In both years, control (untreated) toads headed to the breeding site with the same precision at all release sites. The initial orientation of toads blinded by opaque tape over their eyes did not differ from controls, but the return paths were not as direct. The directional choice of anosmic toads was apparently random, however, individuals followed a straight path in a chosen direction. Anosmic toads also blinded were completely disoriented, moving in cycloid trails. Bar magnets glued to the head caused an increase in dispersion of toads. However, in some individual releases a directional bias without increased dispersion was observed. Sky conditions (clear or overcast) did not influence the initial orientation or the dispersion of toads. Nevertheless, the breeding site component was significantly correlated with wind direction in relation to the breeding site. Wind blowing from the breeding site improved the initial orientation, whereas wind from the opposite direction reduced the breeding site component. The spatial range for the ability to relocate the breeding pond after displacement exceeded 3 km, but the time taken to select the correct direction increased with the displacement distance. The results indicate that after displacement the initial orientation of B. bufo is based mainly on olfactory and magnetic cues, with visual control of straightness.

Animals↗

Magnetic compass orientation in the Eastern red-spotted newt (Notophthalmus viridescens).

Laboratory tests were carried out to examine the orientation behavior of adult Eastern red-spotted newts (Notophthalmus viridescens) to earth-strength magnetic fields. Groups of 30 to 40 newts were housed in water-filled, all-glass aquaria with an artificial shoreline at one end. The aquaria were located in a greenhouse or outdoors adjacent to the laboratory building, and aligned on either the magnetic north-south or east-west axis. Tests were carried out in an enclosed indoor arena. Newts were tested in four horizontal alignments of the magnetic field: the ambient magnetic field (magnetic north at North) and three altered fields (magnetic north rotated to East, South or West). Data were analyzed after pooling the magnetic bearings from all four conditions in such a way as to retain the component of the newts' orientation that was a consistent response to the magnetic field. Elevation of training tank water temperature was used to increase the newts' motivation to orient in the direction of shore. Newts exposed to a training tank water temperature of 33-34 degrees C just prior to testing exhibited consistent unimodal magnetic compass orientation. The direction of orientation was altered predictably by changing training tank alignment and location relative to the laboratory building. The results provide the first evidence of a strong, replicable magnetic compass response in a terrestrial vertebrate under controlled laboratory conditions. Further, the present study demonstrates that the Eastern newt is able to learn a directional response relative to the earth's magnetic field.

Animals↗

The effect of amygdalectomy on orienting and classical conditioning in monkeys.

The basic findings of these two studies are as follows: a failure of orientating and conditioning of viscero-autonomic responses, but essentially normal orienting and conditioning of a temporal muscle response in amygdalectomized animals. Small procedural differences exist between the two studies,and a possible order effect exists as a result of the use of the same subjects consecutively. However, we fell that the difference between the responses of the viscero-autonomic system and the skeletal system would still be found were all of the measures gathered simultaneously in an optimal conditioning situation. The definitive study--simultaneous viscero-autonomic and skeletal (behavioral) recording--remains to be done. Trial-by-trial analysis of the correlations or lack of correlations between these responses in such a study should be highly informative. These two studies offer substantial information relevant to the original questions. When behavior is modified by simple repetition of experience the effect of amygdalectomy is restricted to the viscero-autonomic components of orienting and classical conditioning, not to the entire spectrum of responses. The issue raised, therefore, is the significance of the viscero-autonomic components of orienting and classical conditioning. The suggestion has been proposed (Pribram, 1969) that these components serve as mechanisms of internal rehearsal necessary to the registration (as novel or familiar) of the orienting and conditioning experience. This proposal requires further testing.

Amygdala↗

The coding of head orientations in neurons of bilateral vestibular nuclei of cats after unilateral labyrinthectomy: response to off-vertical axis rotation.

In decerebrate cats that had been acutely hemilabyrinthectomized (HL), the extracellular activities of vestibular nuclear neurons on the lesioned and labyrinth-intact sides were studied during constant-velocity off-vertical axis rotations (OVAR) in the clockwise (CW) and counterclockwise (CCW) directions (at 10 degrees tilt). Over the range of 1.75-15 degrees/s, two types of neuronal responses were identified on both sides. Some neurons showed symmetric and velocity-stable bidirectional response sensitivity (delta defined as the CW gain over the CCW gain) while other neurons exhibited asymmetric and velocity-variable delta. The mathematically derived gain tuning ratios of these two groups of neurons were within the range of one-dimensional and two-dimensional neurons respectively. The best response orientations in one-dimensional neurons and the orientations of the maximum response vector, S(max), in two-dimensional neurons were found to point in all directions on the horizontal plane. On the labyrinth-intact side, both the one-dimensional and two-dimensional neurons showed asymmetry in the neuron numbers and/or the response gains between the two roll quadrants as well as between the two pitch quadrants. In addition, both the neuron number and gain were significantly higher for neurons in the head-down/ipsilateral-side-down half-circle than those in the head-up/contralateral-side-down half-circle. None of the aforementioned asymmetries was observed on the lesioned side. That a comparable pattern of distribution was observed in the one-dimensional and two-dimensional neurons suggests that these neurons maintain a common spatial reference frame in encoding head orientational signals arising from the ipsilateral and contralateral otoliths. Furthermore, a predominance of two-dimensional neurons that exhibited a greater gain with CW rotations was observed on both sides of HL cats. Of the response dynamics observed amongst neurons on the two sides of HL cats, no difference was found with regard to the response gain and the pattern of response lead. However, a difference in response lag was observed between neurons on the two sides of HL cats. These suggest that there is a segregation of otolithic signals to reach the ipsilateral and contralateral vestibular nuclei. Taken together, the present study demonstrates that one-dimensional and two-dimensional neuronal responses could be elicited with inputs arising solely from the ipsilateral or contralateral otoliths. The observed orientational tuning and the CW-CCW asymmetry to bidirectional rotation may provide the essential directional coding of head orientations. Further, the imbalance of spatial/dynamic response patterns between the bilateral vestibular nuclei following the restriction of otolith inputs by HL implies that converging otolithic inputs from the bilateral labyrinths are essential for producing the neuronal responses in control animals. The results are also discussed in terms of the possible contribution of the various neural asymmetries between neuronal subpopulations in the bilateral vestibular nuclei to the behavioral symptoms accompanying acute HL.

Animals↗

Orienting and alerting: effect of 24 h of prolonged wakefulness.

It is well known that a decrease in vigilance can easily occur during 24 h of continuous wakefulness, but no study has assessed whether and in what way extended wakefulness might affect spatial orienting. In other words, it is not clear what happens when a subject has to orient his attention during a state of poor vigilance, resulting from sleep loss or sustained wakefulness. The aim of this study was to investigate this issue. Twelve right-handed male subjects participated in the experiment, which took place on two consecutive days. On the first day, in order to evaluate baseline orienting attention, the subjects performed a covert orienting task (in which the cue stimuli generated endogenous shifts of attention), lasting 20 min; on the second day, during 24 h of prolonged wakefulness, the same task was performed 12 times, about every 120 min, beginning at 10.00 a.m. Results showed an overall slowing of reaction time across the sessions, indicating a linear decrease of vigilance. However, this vigilance decrease did not seem to affect attention-orienting mechanisms, suggesting that the two systems are independent of each other.

Adult↗

Adaptation of postural orientation to changes in surface inclination.

We previously showed that standing on an inclined surface resulted in an after-effect of leaning in many healthy, blindfolded subjects when they returned to standing on a horizontal surface (Kluzik et al. in Exp Brain Res 162:474-489, 2005). The direction of leaning depended on the direction of prior surface inclination, always in a direction that preserved the relative alignment between the body and the support surface. For example, subjects leaned forward after they stood on a toes-up-inclined surface. In the present study, we investigated how the amplitude of surface inclination affected postural muscle activity, joint position, body segment orientation, and body center of mass (CoM) and foot center of pressure (CoP) locations before, during, and after subjects stood on an inclined surface. We asked whether the mechanism that underlies the lean after-effect involves regulation of local postural variables, such as the position of the ankle joint or the level of muscle activity, or whether instead, the mechanism involves regulation of global, whole-body postural variables that can only be determined by multisensory processing, such as orientation of the trunk or the body's CoM. In one experiment, we found that varying the amplitude of a toes-up surface inclination between 2.5 degrees and 10 degrees had a systematic, linear, effect on the post-incline orientation of the trunk and head, but did not systematically affect the post-incline orientation of the legs, position of the ankle joint, the level of EMG activity, or the location of the CoP. In a second experiment, we found that preventing the legs from leaning in the post-incline period did not abolish leaning of the upper body. These findings suggest that (1) the body-to-support-surface relationship is an important reference for the CNS internal representation of postural orientation which is subject to adaptive modification and (2) the adaptive mechanism underlying the post-incline after-effect of leaning acts at the level of global, whole-body postural variables.

Adaptation, Physiological↗

Spatial orientation and balance control changes induced by altered gravitoinertial force vectors.

To better understand the mechanisms of human adaptation to rotating environments, we exposed 19 healthy subjects and 8 vestibular-deficient subjects ("abnormal"; four bilateral and four unilateral lesions) to an interaural centripetal acceleration of 1 g (resultant 45 degrees roll-tilt of 1.4 g) on a 0.8-m-radius centrifuge for periods of 90 min. The subjects sat upright (body z-axis parallel to centrifuge rotation axis) in the dark with head stationary, except during 4 min of every 10 min, when they performed head saccades toward visual targets switched on at 3- to 5-s intervals at random locations (within +/- 30 degrees) in the earth-horizontal plane. Eight of the normal subjects also performed the head saccade protocol in a stationary chair adjusted to a static roll-tilt angle of 45 degrees for 90 min (reproducing the change in orientation but not the magnitude of the gravitoinertial force on the centrifuge). Eye movements, including voluntary saccades directed along perceived earth- and head-referenced planes, were recorded before, during, and immediately after centrifugation. Postural center of pressure (COP) and multisegment body kinematics were also gathered before and within 10 min after centrifugation. Normal subjects overestimated roll-tilt during centrifugation and revealed errors in perception of head-vertical provided by directed saccades. Errors in this perceptual response tended to increase with time and became significant after approximately 30 min. Motion-sickness symptoms caused approximately 25% of normal subjects to limit their head movements during centrifugation and led three normal subjects to stop the test early. Immediately after centrifugation, subjects reported feeling tilted 10 degrees in the opposite direction, which was in agreement with the direction of their earth-referenced directed saccades. Postural COP, segmental body motion amplitude, and hip-sway frequency increased significantly after centrifugation. These postural effects were short-lived, however, with a recovery time of several postural test trials (minutes). There were also asymmetries in the direction of postcentrifugation COP and head tilt which depended on the subject's orientation during the centrifugation adaptation period (left ear or right ear out). The amount of total head movements during centrifugation correlated poorly or inversely with postcentrifugation postural stability, and the most unstable subject made no head movements. There was no decrease in postural stability after static tilt, although these subjects also reported a perceived tilt briefly after return to upright, and they also had COP asymmetries. Abnormal subjects underestimated roll-tilt during centrifugation, and their directed saccades revealed permanent spatial distortions. Bilateral abnormal subjects started out with poor postural control, but showed no postural decrements after centrifugation, while unilateral abnormal subjects had varying degrees of postural decrement, both in their everyday function and as a result of experiencing the centrifugation. In addition, three unilateral, abnormal subjects, who rode twice in opposite orientations, revealed a consistent orthogonal pattern of COP offsets after centrifugation. These results suggest that both orientation and magnitude of the gravitoinertial vector are used by the central nervous system for calibration of multiple orientation systems. A change in the background gravitoinertial force (otolith input) can rapidly initiate postural and perceptual adaptation in several sensorimotor systems, independent of a structured visual surround.

Acceleration↗

Orienting of visual attention in dyslexia: evidence for asymmetric hemispheric control of attention.

The control of attentional orienting was studied in children with specific reading disorder (SRD) or dyslexia, and it was compared with that of normal readers. We used the covert orienting paradigm to measure subjects' reaction times for target detection both in valid and invalid cue conditions, either in the left or in the right visual fields. In experiment 1, we investigated exogenous orienting. The cue consisted of a peripheral abrupt onset and the cue-target delay was 350 ms. As compared with normal readers, in dyslexics the cue effect was absent in the right visual field, whereas in the left visual field a greater cue effect was observed. No visual field asymmetry was found in normal readers. In experiment 2, we investigated endogenous orienting. The cue was shown centrally and the cue-target delay was 750 ms. In dyslexics and normal readers, orienting of attention was present in both visual fields. However, in the invalid condition, dyslexic children showed significantly slower reaction times in the left visual field than in the right visual field. These results were interpreted as being due to an asymmetric control of visual spatial attention, possibly related with a posterior attention mechanism deficit in the right parietal cortex and/or an interhemispheric dysfunction and/or an impairment of cerebellar functions.

Adolescent↗

Effects of stimulus orientation on spatial frequency function of the visual evoked potential.

Visual performance is better in response to vertical and horizontal stimuli than oblique ones in many visual tasks; this is called the orientation effect. In order to elucidate the electrophysiological basis of this psychophysical effect, we studied the effects of stimulus orientation on the amplitudes and latencies of visual evoked potentials (VEPs) over different spatial frequencies of the visual stimulation. VEPs to sinusoidal gratings at four orientations (vertical, horizontal, and oblique at 45 degrees and 135 degrees) with eight spatial frequencies (0.5-10.7 cycles/deg) at reversal rates of 1 Hz and 4 Hz were recorded in nine subjects. At 1-Hz stimulation, the amplitude and latency of P100 were measured. At 4-Hz stimulation, VEPs were Fourier-analyzed to obtain phase and amplitude of the second harmonic response (2F). At 1-Hz stimulation, P100 latencies were decreased for oblique stimuli compared with those for horizontal and vertical stimuli at lower spatial frequencies. Conversely, those for oblique stimuli were increased compared with those for horizontal and vertical stimuli at higher spatial frequencies. At 4-Hz stimulation, spatial tuning observed in 2F amplitude of the oblique gratings shifted to lower spatial frequencies when compared with those of vertical stimulation. The alteration of the VEP spatial frequency function caused by the oblique stimuli was in good agreement with the orientation effect observed in psychophysical studies. Our study may have a clinical implication in that VEP testing with stimuli in more than one orientation at slow and fast temporal modulations can be useful in evaluating neurological disease affecting the visual system.

Adult↗

Visual cues used by ball-rolling dung beetles for orientation.

Ball rolling by dung beetles is considered to be a derived behaviour that evolved under pressure for space, and from competitors at the dung pat. Straight-line orientation away from the pat using a celestial cue should be the most successful rolling strategy to move dung to an unknown burial site. We tested this hypothesis in the field and the laboratory by presenting five species of ball-rolling beetles with different orientation tasks, involving reaction to obstacles as well as to reflected sunlight and artificial light sources. Beetles were found to consistently orientate along a chosen route, usually in the direction of the sun. Beetles rolling dung balls successfully negotiated barriers and returned to the original path as did beetles falling from ramps, or rotated about a fixed point while rolling a ball. The sun was found to be the main orientation cue, which could be substituted by reflected or artificial light. However, beetles reoriented themselves less accurately in response to lights in the laboratory, than they did to the reflected sun in the field. It is probable that phototactic orientation using the sun, which is widespread amongst arthropods, has been incorporated in the straight-line foraging behaviour that has evolved in ball-rolling dung beetles.

Animals↗

An explanation of contextual modulation by short-range isotropic connections and orientation map geometry in the primary visual cortex.

Recent experimental studies on the primary visual cortex have revealed complicated nonclassical neuronal activities. Contextual modulation on orientation-contrast is one typical example of nonclassical neuronal behavior. This modulation by surrounding stimuli in a nonclassical receptive field is mainly thought to be mediated by short- and long-range horizontal connections within the primary visual cortex. Short-range connections are circularly symmetrical and relatively independent of orientation preferences, while long-range connections are patchy, asymmetrical, and orientation specific. Although this modulation can be explained by long-range specific connections qualitatively, recent studies suggest that long-range connections alone may be insufficient with respect to the balance between two types of connections. Here, in order to clarify the role of short-range connections in the process of contextual modulation, we propose a model of the primary visual cortex with isotropic short-range connections and a geometric orientation map. Computational simulations using the model have demonstrated that contextual modulation can be explained by short-range connections alone. This is due to the interaction between the spatial periodicity of orientation domains and the excitatory-inhibitory regions arising from the propagation of activities.

Action Potentials↗

Effect of imbalance in activities between ON- and OFF-center LGN cells on orientation map formation.

It has been reported that the OFF responses of cells in the visual pathway are stronger, on average, than the ON responses early in the life of cats and ferrets. In this study, we theoretically investigate the effects of this imbalance in activity on the orientation map formation. We carry out computer simulations based on our previously proposed self-organization model, in which the correlated activities between ON- and OFF-center cells in the lateral geniculate nucleus regulate the formation of orientation maps in the visual cortex. When imbalance between the activities of these ON- and OFF-center cells is assumed, we obtain orientation maps with spatial periodicity, as observed in the experiments. On the other hand, when balanced activities are assumed, orientation maps do not show periodicity. This suggests that the imbalance in activities between ON- and OFF-center cells contributes to the elaboration of orientation maps during the critical period.

Algorithms↗

Orientational behaviors and EAG responses of male codling moth after exposure to synthetic sex pheromone from various dispensers.

The effect of brief pheromone exposures on responses of codling moth (Cydia pomonella L.) males was tested by flight-tunnel and electroantennogram (EAG) studies. Males were preexposed to pheromone for up to 3 min as they sat in release cages or for shorter times (a few seconds to several min) upon initiating flights or orienting in plumes. Brief exposures to Isomate-C Plus dispensers nearly eliminated moth orientations to 0.1 mg codlemone [(E,E)-8,10-dodecadien-1-ol] and 0.1 mg three-component (codlemone/12OH/14OH, 100:20:5 ratio) lures 15 min later. However, there was no associated change in EAG responses between preexposed and control moths. Behavioral responses of Isomate-C Plus-exposed males were normal 24 hr following exposure. The reduced sexual responsiveness observed following exposure to Isomate dispensers appeared to be associated with an elevation of response threshold. Brief preexposure to 0.1 mg codlemone and three-component lures also reduced orientational behavior of males 15 min later, but to a lesser degree than when preexposed to Isomate-C Plus dispensers. Male behavior following preexposure to a 0.1 mg codlemone/pear ester [(2E,4Z)-2,4-decadienoate] lure (1:1 ratio) was no different from exposure to codlemone only. Orientational disruption in plots treated with 10 dispensers of Isomate-C Plus per tree was 88.3 and 95.9% for 1.0 and 0.1 mg codlemone lures, respectively. Some males did orient to 0.1 mg codlemone lures so we caution that flight-tunnel experiments on preexposure may overestimate the actual pheromone exposure dosage received by feral moths in treated orchards. Importantly, this work documents that a portion of feral males within a population has the capacity to overcome communicational disruption by high densities of Isomate-C Plus dispensers.

Animals↗

Distribution in the visual field of the costs of voluntarily allocated attention and of the inhibitory after-effects of covert orienting.

By using a simple reaction time (RT) paradigm we have investigated the spatial distribution of the benefits and costs of voluntarily directed attention and of the inhibitory after-effects of covert orienting. In the first experiment subjects deliberately allocated attention to each one of five stimulus positions disposed along the horizontal meridian, while at the same time fixing their eyes on the central position. The separation in visual angle between the central position and the two nearest positions, one on the left and the other on the right, was 10 degrees; that between the central position and the two most eccentric positions was 30 degrees. By comparing RT to brief flashes of light presented at each position during directed attention with RT to identical flashes at the same position during diffuse attention (i.e. in a condition in which subjects paid equal attention to all five positions), it was possible to determine that benefits, that is RT decreases relative to the diffuse-attention condition, were strictly limited to the attended position. Costs, i.e. RT increases relative to the diffuse-attention condition, showed a more diffuse and complex spatial pattern. When attention was directed to one of the noncentral positions, costs were apparent at the two contralateral positions and at the central position, but not at the ipsilateral position. When attention was directed to the central position, costs occurred at all other positions. This suggests a special role for the vertical meridian in delimiting the area of costs when one covertly orients towards the opposite right or left visual half field. Work of others and our preliminary evidence indicate that the area of costs is similarly limited by the horizontal meridian when one orients toward the opposite upper or lower visual field. In the second experiment we studied the inhibitory after-effect of covert orienting. Orienting to a light stimulus without moving the eyes to it may induce a short-lived facilitation of the speed of response to a second stimulus presented at the same position, but this facilitation is followed by a profound and prolonged RT retardation. By using a two-flashes paradigm we observed this RT retardation not only when the two stimuli appeared at the same position, but also when they occurred at different locations in the same altitudinal or lateral visual hemifield. There were no inhibitory after-effects when the two stimuli appeared on opposite sides of the vertical or horizontal meridian.(ABSTRACT TRUNCATED AT 400 WORDS)

Adult↗

Focal right temporo-occipital blood flow changes associated with judgment of line orientation.

Focal cerebral blood flow changes for judgment of line orientation were determined using the 133Xe inhalation technique with 18 normal right-handed male subjects. Measurements were made during three conditions in the same session: a line orientation task, a sensorimotor control task, and normal rest. Blood flow changes attributable to the judgment of line orientation were found for the measure f1 at a detector centered over the right temporo-occipital region. For the measure IS this activation was significant in the temporo-occipital region in both hemispheres but significantly greater in the right hemisphere. For f1 the change in blood flow and for IS the percent change in blood flow in the right temporo-occipital region attributable to judgment of line orientation decreased as performance on the line orientation task improved.

Adolescent↗

Visuomotor sensitivity for shape and orientation in a patient with visual form agnosia.

We have previously demonstrated that a patient with visual form agnosia (DF), who is unable to report the orientation or size of visual targets, can nevertheless use these same visual attributes to control motor acts. In the first of three new experiments, we found that DF is able to grasp everyday tools and utensils proficiently (i.e. with a well-formed hand posture) but has difficulty in visually selecting the correct part of the object to grasp (e.g. the handle) for subsequent use of that object. A second experiment revealed that DF's visuomotor system is able to adjust concurrently to variations in both the size and orientation of target objects; when these visual attributes were both varied, she adjusted both her grip aperture and the orientation of her hand well in advance of target contact. These spared visuomotor abilities do not seem to extend to shape processing per se, however. In the final experiment we found that DF was insensitive to changes in the orientation of a cross-shaped object, where no single principal axis could be extracted to control orientation of the grasp. These observations extend our knowledge of DF's residual visuomotor abilities, and suggest limitations on the visual processing capacities of the human dorsal stream.

Adult↗