Vertical gaze angle as a distance cue for programming reaching: insights from visual form agnosia II (of III)

It has been shown that a patient with visual form agnosia (DF) relies predominantly on vergence information when gauging target distance in an open-loop pointing task. This finding suggested that the programming of prehension might be severely disrupted if DF viewed target objects through ophthalmic prisms. An initial experiment showed that this prediction was not upheld; DF was able to programme reasonably accurate movements to objects located on a tabletop despite large changes in vergence angle. A second experiment, however, showed that placing the target objects at eye height whilst manipulating vergence angle caused gross disruption to prehension, with DF mis-programming the reach component in a predictable manner. Notably, the evidence for DF's reliance on vergence distance information was obtained in a task where the targets were viewed at eye height. These experiments indicate that DF uses vertical gaze angle to gauge target distance in normal prehension and suggest that this extra-retinal cue may be a useful source of distance information for the human nervous system, especially where pictorial cues are impoverished. Electronic Publication     

The role of size and binocular information in guiding reaching: insights from virtual reality and visual form agnosia III (of III)

Reaching out to grasp an object requires information about the size of the object and the distance between the object and the body. We used a virtual reality system with a control population and a patient with visual form agnosia (DF) in order to explore the use of binocular information and size cues in prehension. The experiments consisted of a perceptual matching task in addition to a prehension task. In the prehension task, control participants modified their reach distance in response to step changes in vergence in the absence of any clear reference for relative disparity. Their reach distance was unaffected by equivalent step changes in size, even though they used this information to modify grasp and showed a size bias in a distance matching task. Notably, DF showed the same pattern of results as the controls but was far more sensitive to step changes in vergence. This finding complements previous research suggesting that DF relies predominantly on vergence information when gauging target distance. The results from the perceptual matching tasks confirmed previous findings suggesting that DF is unable to make use of size information for perceptual matching, including distance comparisons. These data are discussed with regard to the properties of the pathways subserving the two visual cortical processing streams. Electronic Publication     

Getting the measure of vergence weight in nearness perception

Combining multiple sources of information allows the human nervous system to construct an approximately Euclidean representation of near (personal) space. Within this space, binocular vergence is an important source of egocentric distance information. We investigated how the nervous system determines the significance (weight) accorded to vergence information when other (retinal) distance cues are present. We found that weight decreases with (1) increasing discrepancy between vergence information and other cues and (2) reduced vergence demand. The results also provided evidence that the nervous system represents vergence related distance information in units of nearness (the reciprocal of distance).     

Vertical gaze angle: absolute height-in-scene information for the programming of prehension

One possible source of information regarding the distance of a fixated target is provided by the height of the object within the visual scene. It is accepted that this cue can provide ordinal information, but generally it has been assumed that the nervous system cannot extract "absolute" information from height-in-scene. In order to use height-in-scene, the nervous system would need to be sensitive to ocular position with respect to the head and to head orientation with respect to the shoulders (i.e. vertical gaze angle or VGA). We used a perturbation technique to establish whether the nervous system uses vertical gaze angle as a distance cue. Vertical gaze angle was perturbed using ophthalmic prisms with the base oriented either up or down. In experiment 1, participants were required to carry out an open-loop pointing task whilst wearing: (1) no prisms; (2) a base-up prism; or (3) a base-down prism. In experiment 2, the participants reached to grasp an object under closed-loop viewing conditions whilst wearing: (1) no prisms; (2) a base-up prism; or (3) a base-down prism. Experiment 1 and 2 provided clear evidence that the human nervous system uses vertical gaze angle as a distance cue. It was found that the weighting attached to VGA decreased with increasing target distance. The weighting attached to VGA was also affected by the discrepancy between the height of the target, as specified by all other distance cues, and the height indicated by the initial estimate of the position of the supporting surface. We conclude by considering the use of height-in-scene information in the perception of surface slant and highlight some of the complexities that must be involved in the computation of environmental layout.     

Reaching for virtual objects: binocular disparity and the control of prehension

Although, in principle, binocular cues provide veridical information about the three-dimensional shape of objects, our perception on the basis of these cues is distorted systematically. The consequences of these distortions may be less serious than they first appear, however, since in everyday life we rarely are required to judge the absolute shape, size or distance of objects. An important exception to this is in the control of prehension, where veridical information about an object to be grasped is required to plan the transport of the hand and to select the most appropriate grip. Here we investigate whether binocular cues provide accurate depth information for the control of prehension using disparity-defined, virtual objects and report that whilst binocular disparity can support prehensile movements, the kinematic indices, which reflect distance-reached and perceived size, show clear biases. These results suggest that accurate metric depth information for the control of prehension is not available from binocular cues in isolation. Electronic Publication     

Monocular and binocular distance cues: insights from visual form agnosia I (of III)

The human nervous system constructs a Euclidean representation of near (personal) space by combining multiple sources of information (cues). We investigated the cues used for the representation of personal space in a patient with visual form agnosia (DF). Our results indicated that DF relies predominantly on binocular vergence information when determining the distance of a target despite the presence of other (retinal) cues. Notably, DF was able to construct an Euclidean representation of personal space from vergence alone. This finding supports previous assertions that vergence provides the nervous system with veridical information for the construction of personal space. The results from the current study, together with those of others, suggest that: (i) the ventral stream is responsible for extracting depth and distance information from "monocular" retinal cues (i.e. from shading, texture, perspective) and (ii) the dorsal stream has access to binocular information (from horizontal image disparities and vergence). These results also indicate that DF was not able to use size information to gauge target distance, suggesting that intact temporal cortex is necessary for "learned size" to influence distance processing. Our findings further suggest that in neurologically intact humans, object information extracted in the ventral pathway is combined with the products of dorsal stream processing for guiding prehension. Finally, we studied the "size-distance paradox" in visual form agnosia in order to explore the cognitive use of size information. The results of this experiment were consistent with a previous suggestion that the paradox is a cognitive phenomenon. Electronic Publication     

Coordination in prehension Information-based coupling of reaching and grasping

Prehension involves the coordination of a reaching and a grasping movement, such that the hand opens and closes in tune with the transport of the hand to the object to be grasped. To investigate this coordination, we focused on the transition from hand opening to hand closing in the grasping component of prehension. Earlier research has suggested that the time taken to close the hand remains constant over varying reaching amplitudes. In the present experiment, in which subjects reached for objects at six different distances and for objects that moved away from them at three different, constant speeds, hand-closure time was found to vary as a function of experimental conditions. Moreover, initiation of hand closure did not occur at a constant value of the (perceptually available) first-order time remaining until contact with the object. However, the variations observed, occurring as a function of initial hand-object distance and object speed, could be accounted for by an abstract dynamical model of perceptually driven postural changes. Received: 25 July 1996 / Accepted: 9 October 1997     

Effects of stimulus size and eccentricity on horizontal and vertical vergence

We measured the gain and phase of horizontal and vertical vergences of five subjects as a function of stimulus area and position. Vergence eye movements were recorded by the scleral search coil method as subjects observed dichoptic displays oscillating in antiphase either from side to side or up and down with a peak-to-peak magnitude of 0.5° at either 0.1 Hz or 1.0 Hz. The stimulus was a central textured disc with diameter ranging from 0.75° to 65°, or a peripheral annulus with outer diameter 65° and inner diameter ranging from 5° to 45°. The remaining field was black. For horizontal vergence at both stimulus frequencies, gain and the phase lag were about the same for a 0.75° stimulus as for a 65° central stimulus. For vertical vergence, mean gain increased and mean phase lag decreased with increasing diameter of the central stimulus up to approximately 20°. Thus, the stimulus integration area is much smaller for horizontal vergence than for vertical vergence. The integration area for vertical vergence is similar to that for cyclovergence, as revealed in a previous study. For both types of vergence, response gains were higher and phase lags smaller at 0.1 Hz than at 1.0 Hz. Also, gain decreased and phase lag increased with increasing occlusion of the central region of the stimulus. Vergence gain was significantly higher for a 45° central disc than for a peripheral annulus with the same area. Thus, the central retina has more power to evoke horizontal or vertical vergence than the same area in the periphery. We compare the results with similar data for cyclovergence and discuss their ecological implications. Received: 30 January 1999 / Accepted: 25 June 1999     

The perception and prehension of objects oriented in the depth plane II. Dissociated orientation functions in normal subjects

 Normal human subjects were tested for their ability to discriminate the orientation of a square plaque tilted in depth, using two different tasks: a grasping task and a perceptual matching task. Both tasks were given under separate monocular and binocular conditions. Accuracy of performance was measured by use of an opto-electronic motion analysis system, which computed the hand orientation (specifically, a line joining the tips of the thumb and index finger) as the hand either approached the target during grasping or was used to match the target. In all cases there was a very strong statistical coupling between hand orientation and target orientation, irrespective of viewing conditions. However, the matching data differed from the grasping data in showing a consistent curvature in the hand-target relationship, whereby the rate of change of hand orientation as a function of object orientation was smaller for oblique orientations than for those near the horizontal or vertical. The results are interpreted as reflecting the operation of two different mechanisms for analysing orientation in depth: a visuomotor system (assumed to be located primarily in the dorsal cortical visual stream) and a perceptual system (assumed to be located in the ventral stream). It may be that the requirements of visuomotor control dictate a primary need for absolute orientation coding, whereas those of perception dictate a need for more categorical coding. Received: 13 June 1997 / Accepted: 14 October 1997     

Attention in action or obstruction of movement? A kinematic analysis of avoidance behavior in prehension

Obstacle avoidance strategies are of two basic but interrelated types: moving around an obstacle to that body parts do not come too close, and slowing down. In reaching-to-grasp, avoidance may involve the transport component, the grasp formation component, or both. There has been little research that has directly examined obstacle avoidance strategies during reaches-to-grasp. Several recent reports describe experiments in which reaches-to-grasp were made when nontarget objects were present in the workspace. The effects of these nontargets were interpreted as being due to their distracting effects rather than their obstructing effects. The results of these studies are reinterpreted as being due to the non-target’s obstructing effects. The obstacle interpretation is more parsimonious and better predicts the pattern of results than the distractor interpretation. Predictions of the obstacle interpretation were examined in an experiment in which participants were required to reach to grasp a target in the presence of another object in various locations. The results were exactly in line with the interpretation of the object as an obstacle and the data show how grasp and transport movements are subtly adjusted so as to avoid potential obstacles. It is proposed that people move so as not to bring body parts within a minimum preferred distance from nontarget objects within the workspace. What constitutes the preferred distance in a particular context appears to depend upon the speed of movement and a variety of psychological factors related to the cost that a person attaches to a collision. Received: 5 September 1997 / Accepted: 10 January 1998     

Looking at the task in hand: vergence eye movements and perceived size

A retinal afterimage of the hand changes size when the same unseen hand is moved backwards and forwards in darkness. We demonstrate that arm movements per se are not sufficient to cause a size change and that vergence eye movements are a necessary and sufficient condition for the presence of the illusory size change. We review previous literature to illustrate that changing limb position in the dark alters vergence angle and we explain the illusion via this mechanism. A discussion is provided on why altering limb position causes a change in vergence and we speculate on the underlying mechanisms. Received: 6 May 1997 / Accepted: 16 July 1997     

An analysis of spatiotemporal variability during prehension movements: effects of object size and distance

 Human prehension movements have been studied with regard to the parallel processing of motor control and sensorimotor coordination. Temporal aspects of the movement (e.g., onset time and duration) have been studied extensively, while spatial aspects have not been studied systematically. Thus, the purpose of this study was to examine spatiotemporal variability of the transport (wrist trajectory) and grasp (grip aperture between the index finger and the thumb) components. In this experiment, the extrinsic (e.g., distance) and intrinsic object properties (e.g., object size) were manipulated. Subjects were required to pick up an aluminum cylinder as quickly and accurately as possible using the index finger and the thumb. It was found that object size significantly affected both transport and grasp components. Distance mainly affected the transport component. These kinematic results were consistent with the findings of earlier studies. Furthermore, the distribution of mean within-subject variability across normalized movement time for the transport component was not the same as that of the grasp component, suggesting that the different motor control processes exist. The peak amplitudes in variability of the wrist trajectory and the grip aperture were obtained at similar points throughout movement time. Furthermore, the peak of wrist variability depended on distance not object size, while that of aperture variability depended on both distance and object size. These results strongly support the hypothesis that the grasp component is adjusted using dynamic information provided from the transport component as the wrist moves toward the object. We also found that wrist variability converged to the target point, while aperture variability was biphasic: it converged, at least, around the point of maximum aperture in the first phase and then remained constant in the second phase. This result suggests that the two components are under different control processes. We hypothesize that the transport component can be modeled as a single feedforward system, while the grasp component can be divided into two separate mechanisms. Received: 4 March 1996 / Accepted: 29 January 1997     

The stimulus integration area for horizontal vergence

Over what region of space are horizontal disparities integrated to form the stimulus for vergence? The vergence system might be expected to respond to disparities within a small area of interest to bring them into the range of precise stereoscopic processing. However, the literature suggests that disparities are integrated over a fairly large parafoveal area. We report the results of six experiments designed to explore the spatial characteristics of the stimulus for vergence. Binocular eye movements were recorded using magnetic search coils. Each dichoptic display consisted of a central target stimulus that the subject attempted to fuse, and a competing stimulus with conflicting disparity. In some conditions the target was stationary, providing a fixation stimulus. In other conditions, the disparity of the target changed to provide a vergence-tracking stimulus. The target and competing stimulus were combined in a variety of conditions including those in which (1) a transparent textured-disc target was superimposed on a competing textured background, (2) a textured-disc target filled the centre of a competing annular background, and (3) a small target was presented within the centre of a competing annular background of various inner diameters. In some conditions the target and competing stimulus were separated in stereoscopic depth. The results are consistent with a disparity integration area with a diameter of about 5°. Stimuli beyond this integration area can drive vergence in their own right, but they do not appear to be summed or averaged with a central stimulus to form a combined disparity signal. A competing stimulus had less effect on vergence when separated from the target by a disparity pedestal. As a result, we propose that it may be more useful to think in terms of an integration volume for vergence rather than a two-dimensional retinal integration area.     

Time to contact and the control of manual prehension

In the present study, a kinematic analysis was made of unconstrained, natural prehension movements directed toward an object approaching the observer on a conveyor belt at one of three constant velocities, from one of three different directions (head-on or along the fronto-parallel plane coming either from the subject′s left or right). Subjects were required to grasp the object when it reached a target located 20 cm directly in front of the hand′s start position. The kinematic analysis revealed that both the transport and grasp components of the movement changed in response to the experimental manipulations, but did so in a manner that guaranteed that, for objects approaching from a given direction, hand closure would begin at a constant time prior to object contact (regardless of the object’s approach speed). The kinematic analysis also revealed, however, that the onset of hand closure began earlier with objects approaching from the right than from other directions – an effect which would not be predicted if time to contact was the key variable controlling the onset of hand closure. These results, then, lend only partial support to the theory that temporal coordination between the transport and grasp components of prehension is ensured through their common dependence on time to contact information. Received: 20 September 1996 / Accepted: 16 June 1997     

The role of head movements in the control of manual prehension

Binocular information has been shown to be important for the programming and control of reaching and grasping. But even without binocular vision, people are still able to reach out and pick up objects accurately – albeit less efficiently. As part of a continuing investigation into the role that monocular cues play in visuomotor control, we examined whether or not subjects could use retinal motion information, derived from movements of the head, to help program and control reaching and grasping movements when binocular vision is denied. Subjects reached out in the dark to an illuminated sphere presented at eye-level, under both monocular and binocular viewing conditions with their head either free to move or restrained. When subjects viewed the display monocularly, they showed fewer on-line corrections when they were allowed to move their head. No such difference in performance was seen when subjects were allowed a full binocular view. This study, combined with previous work with neurological patients, confirms that the visuomotor system “prefers” to use binocular vision but, when this information is not available, can fall back on other monocular depth cues, such as information produced by motion of the object (and the scene) on the retina, to help program and control manual prehension. Received: 22 December 1997 / Accepted: 4 February 1998     

Influence of object position and size on human prehension movements

 Prehension movements of the right hand were recorded in normal subjects using a computerized motion analyzer. The kinematics and the spatial paths of markers placed at the wrist and at the tips of the index finger and thumb were measured. Cylindrical objects of different diameters (3, 6, 9 cm) were used as targets. They were placed at six different positions in the workspace along a circle centered on subject’s head axis. The positions were spaced by 10° starting from 10° on the left of the sagittal axis, up to 40° on the right. Both the transport and the grasp components of prehension were influenced by the distance between the resting hand position and the object position. Movement time, time to peak velocity of the wrist and time to maximum grip aperture varied as a function of distance from the object, irrespective of its size. The variability of the spatial paths of wrist and fingers sharply decreased during the phase of the movement prior to contact with the object. This indicates that the final position of the thumb and the index finger is a controlled parameter of visuomotor transformation during prehension. The orientation of the opposition axis (defined as the line connecting the tips of the thumb and the index finger at the end of the movement) was measured. Several different frames of reference were used. When an object-centered frame was used, the orientation of the opposition axis was found to change by about 10° from one object position to the next. By contrast, when a body-centered frame was used (with the head or the forearm as a reference), this orientation was found to remain relatively invariant for different object positions and sizes. The degree of wrist flexion was little affected by the position of the object. This result, together with the invariant orientation of the opposition axis, shows that prehension movements aimed at cylindrical objects are organized so as to minimize changes in posture of the lower arm. Received: 2 July 1996 / Accepted: 5 October 1996     

A three-dimensional analysis of vergence movements at various levels of elevation

Earlier studies have shown that eye positions, recorded in subjects scanning a distant visual scene with the head in a stable position, have only two degrees of freedom (Listing's law). Due to cyclovergence, this law is modified in near-vision. Two previous quantitative studies have documented that the sign of the torsional vergence component depends systematically on elevation: when fixating a nearby target, the eyes show intorsion in up gaze, extorsion in down gaze and no cyclotorsion at some intermediate elevation level (to be denoted as the null elevation). Both studies found a linear cyclovergence/elevation relation, but disagreed on the amount of cyclotorsion. A further uncertainty is how this phenomenon develops dynamically when the binocular fixation point shifts from a far to a near position. Therefore, we have investigated the dynamic coupling between the horizontal and torsional components of vergence in human subjects who were instructed to refixate a light target after it stepped in depth. The target steps were presented at various vertical and horizontal directions relative to the straight-ahead axis of the cyclopean eye. We found that the quantitative relations among horizontal vergence, torsional vergence and elevation were intermediate between those found in the two earlier near-vision studies and that they correspond reasonably to the predictions of a model by Mok and co-workers. The cyclotorsion vergence component had about the same latency and dynamics as the horizontal component. When refixations were studied at different elevations, the torsional vergence component changed from incyclotorsion in up gaze to excyclotorsion in down gaze. In agreement with expectations derived from two quantiative models, the null elevation of cyclovergence was near the binocular primary position. Furthermore, we found no consistent additional dependence on the horizontal direction of the refixation trajectory relative to the midsaggital plane. Other experiments showed that the cyclotorsional changes accompanying convergence were not critically dependent upon the visual conditions. Quantitatively similar cyclotorsional components were found even in convergent refixations executed in full darkness towards the location of a remembered (flashed) near target. We conclude that visual feedback is unlikely to be very important in controlling cyclovergence in these various conditions.     

Development of prehension movements in children: a kinematic study

 To evaluate the normal development of functional hand motor skill, the kinematics of prehension movements were analyzed in 54 healthy children (age 4–12 years). The subjects repeatedly reached out for cylindrical target objects and grasped them with a precision grip of their dominant hand. The trajectory of the reaching hand and the finger aperture were monitored by optoelectronic motion analysis. To obtain comparable conditions for the different age groups, the experimental setup was scaled according to the individual body proportions of each subject. Within the investigated age range, neither the movement duration nor the normalized (according to body proportions) peak spatial velocity of the reaching hand changed significantly. However, the hand trajectory straightened and the coordination between hand transport and grip formation improved, resulting in smooth and stereotyped kinematic profiles at the age of 12 years. The younger children opened their grip relatively wider than the older ones, thus grasping with a higher safety margin. The dependence on visual control of the movement declined during motor development. Only the oldest childen were able to scale the grip aperture adequately, according to various sizes of the target objects, when visual control of the movement was lacking. The results suggest that the development of prehensile skills during childhood lasts until the end of the first decade of life. This functional maturation is discussed in relation to the development of neuronal pathways. Received: 20 November 1997 / Accepted: 4 July 1998     

Fusional vergence ranges of the monkey: A behavioral study

Summary Fusional vergence ranges were behaviorally determined for two rhesus monkeys and two human subjects using random dot stereograms as stimuli. The subject performed a two-alternative forced choice form discrimination of discretely presented stimuli and the fusional vergence stimulus was changed during the intertrial intervals. The fusional vergence measurements for the monkey were similar to those for the human subjects.     

The removal of binocular cues disrupts the calibration of grasping in patients with visual form agnosia

 The present study tested the idea that the visuomotor systems mediating prehension do not have independent access to pictorial cues processed by perceptual mechanisms. Individuals with visual form agnosia, whose perceptual systems are compromised but who have intact visuomotor control, were examined to determine whether they could use pictorial scene cues to calibrate manual prehension when binocular information was removed. The removal of binocular cues produced considerable disruptions in size-constancy of grip aperture, which, combined with earlier observations in normal subjects, suggests that binocular cues are of primary importance in calibration of grasping. In the absence of binocular vision, normal subjects can use pictorial information, information that is severely compromised in individuals with visual form agnosia, to compute the distance (and thus the size) of the goal object. Thus, individuals with visual form agnosia must rely on a retinal image that remains uncalibrated, leading to inaccurate calibrations of grip aperture. The fact that these individuals scaled their grasp much less accurately under the monocular viewing condition, despite showing normal binocular grasping, suggests that pictorial cues to depth, which are presumably processed by mechanisms mediating our perception of objects and events in the world, can be accessed by visuomotor mechanisms only indirectly. These results, together with others, suggest that the visuomotor system ’prefers’ to use binocular information and uses pictorial cues only as a last resort. Received: 10 September 1996 / Accepted: 25 February 1997