Search PubMed⌕ Search

Biomedical subjects

J A Van Gisbergen

Publications and source records attributed to J A Van Gisbergen.

At least 19 recordsLinked to original sources

The subjective vertical and the sense of self orientation during active body tilt.

Previous testing of the ability to set a luminous line to the direction of gravity in passively-tilted subjects, in darkness, has revealed a remarkable pattern of systematic errors at tilts beyond 60 degrees, as if body tilt is undercompensated or underestimated (Aubert or A-effect). We investigated whether these consistent deviations from orientation constancy can be avoided during active body tilt, where more potential cues about body tilt (e.g. proprioception and efference copy) are available. The effects of active body tilt on the subjective vertical and on the perception of self tilt were studied in six subjects. After adopting a laterally-tilted posture, while standing in a dark room, they indicated the subjective vertical by adjusting a visual line and gave their verbal estimate of head orientation, expressed on a clock scale. Head roll tilts covered the range from -150 degrees to +150 degrees. The subjective vertical results showed no sign of improvement. Actively-tilted subjects still exhibited the same pattern of systematic errors that characterised their performance during passive tilt. Random errors in this task showed a steep monotonic increase with tilt angle, as in earlier passive tilt experiments. By contrast, verbal head-tilt estimates in the active experiments showed a clear improvement and were now almost devoid of systematic errors, but the noise level remained high. Various models are discussed in an attempt to clarify how these task-related differences and the selective improvement of the self-tilt estimates in the active experiments may have come about.

Adult↗

Stimulation in the rostral pole of monkey superior colliculus: effects on vergence eye movements.

Recent studies have indicated that the superior colliculus (SC), traditionally considered to be saccade-related, may play a role in the coding of eye movements in both direction and depth. Similarly, it has been suggested that omnidirectional pause neurons are not only involved in the initiation of saccades, but can also modulate vergence eye movements. These new developments provide a challenge for current oculomotor models that attempt to describe saccade-vergence coordination and the neural mechanisms that may be involved. In this paper, we have attempted to study these aspects further by investigating the role of the rostral pole of the SC in the control of vergence eye movements. It is well-known that, by applying long-duration electrical stimulation to rostral sites in the monkey SC, saccadic responses can be prevented and interrupted. We have made use of these properties to extend this paradigm to eye movements that contain a substantial depth component. We found that electrical intervention in the rostral region also has a clear effect on vergence. For an eye movement to a near target, stimulation leads to a significant suppression and change in dynamics of the pure vergence response during the period of stimulation, but the depth component cannot be prevented entirely. When these paradigms are implemented for 3D refixations, the saccade is inactivated, as expected, while the vergence component is often suppressed more than in the case of the pure vergence. The data lead us to conclude that the rostral SC, presumably indirectly via connections with the pause neurons, can affect vergence control for both pure vergence and combined 3D responses. Suppression of the depth component is incomplete, in contrast to the directional movement, and is often different in magnitude for 3D refixations and pure vergence responses. The results are discussed in connection with current models for saccade-vergence interaction.

Animals↗

Properties of the internal representation of gravity inferred from spatial-direction and body-tilt estimates.

One of the key questions in spatial perception is whether the brain has a common representation of gravity that is generally accessible for various perceptual orientation tasks. To evaluate this idea, we compared the ability of six tilted subjects to indicate earth-centric directions in the dark with a visual and an oculomotor paradigm and to estimate their body tilt relative to gravity. Subjective earth-horizontal and -vertical data were collected, either by adjusting a visual line or by making saccades, at 37 roll-tilt angles across the entire range. These spatial perception responses and the associated body-tilt estimates were subjected to a principal-component analysis to describe their tilt dependence. This analysis allowed us to separate systematic and random errors in performance, to disentangle the effects of task (horizontal vs. vertical) and paradigm (visual vs. oculomotor) in the space-perception data, and to compare the veridicality of space perception and the sense of self-tilt. In all spatial-orientation tests, whether involving space-perception or body-tilt judgments, subjects made considerable systematic errors which mostly betrayed tilt underestimation [Aubert effect (A effect)] and peaked near 130 degrees tilt. However, the A effect was much smaller in body-tilt estimates than in spatial pointing, implying that the underlying signal processing must have been different. Pointing results obtained with the visual and the oculomotor paradigm were not identical either, but these differences, which were task-related (horizontal vs. vertical), were subtle in comparison. The tilt-dependent pattern of random errors (noisy scatter) was almost identical in visual and oculomotor pointing results, showing a steep monotonic increase with tilt angle, but was again clearly different in the body-tilt estimates. These findings are discussed in the context of a conceptual model in an attempt to explain how the different patterns of systematic and random errors in external-space and self-tilt perception may come about. The scheme proposes that basically similar computational mechanisms, working with different settings, may be responsible.

Adult↗

Kinematic strategies for upper arm-forearm coordination in three dimensions.

This study addressed the question of how the three-dimensional (3-D) control strategy for the upper arm depends on what the forearm is doing. Subjects were instructed to point a laser-attached in line with the upper arm-toward various visual targets, such that two-dimensional (2-D) pointing directions of the upper arm were held constant across different tasks. For each such task, subjects maintained one of several static upper arm-forearm configurations, i. e., each with a set elbow angle and forearm orientation. Upper arm, forearm, and eye orientations were measured with the use of 3-D search coils. The results confirmed that Donders' law (a behavioral restriction of 3-D orientation vectors to a 2-D "surface") does not hold across all pointing tasks, i.e., for a given pointing target, upper arm torsion varied widely. However, for any one static elbow configuration, torsional variance was considerably reduced and was independent of previous arm position, resulting in a thin, Donders-like surface of orientation vectors. More importantly, the shape of this surface (which describes upper arm torsion as a function of its 2-D pointing direction) depended on both elbow angle and forearm orientation. For pointing with the arm fully extended or with the elbow flexed in the horizontal plane, a Listing's-law-like strategy was observed, minimizing shoulder rotations to and from center at the cost of position-dependent tilts in the forearm. In contrast, when the arm was bent in the vertical plane, the surface of best fit showed a Fick-like twist that increased continuously as a function of static elbow flexion, thereby reducing position-dependent tilts of the forearm with respect to gravity. In each case, the torsional variance from these surfaces remained constant, suggesting that Donders' law was obeyed equally well for each task condition. Further experiments established that these kinematic rules were independent of gaze direction and eye orientation, suggesting that Donders' law of the arm does not coordinate with Listing's law for the eye. These results revive the idea that Donders' law is an important governing principle for the control of arm movements but also suggest that its various forms may only be limited manifestations of a more general set of context-dependent kinematic rules. We propose that these rules are implemented by neural velocity commands arising as a function of initial arm orientation and desired pointing direction, calculated such that the torsional orientation of the upper arm is implicitly coordinated with desired forearm posture.

Biomechanical Phenomena↗

Context compensation in the vestibuloocular reflex during active head rotations.

The vestibuloocular reflex (VOR) needs to modulate its gain depending on target distance to prevent retinal slip during head movements. We investigated gain modulation (context compensation) for binocular gaze stabilization in human subjects during voluntary yaw and pitch head rotations. Movements of each eye were recorded, both when attempting to maintain gaze on a small visual target at straight-ahead in a darkened room and after its disappearance (remembered target). In the analysis, we relied on a binocular coordinate system yielding a version and a vergence component. We examined how frequency and target distance, approached here by using vergence angle, affected the gain and phase of the version component of the VOR and compared the results to the requirements for ideal performance. Linear regression analysis on the version gain-vergence relationship yielded a slope representing the influence of target proximity and an intercept corresponding to the response at zero vergence ("default gain"). The slope of the fitted relationship, divided by the geometrically required slope, provided a measure for the quality of version context compensation ("context gain"). In both yaw and pitch experiments, we found default version gains close to one even for the remembered target condition, indicating that the active VOR for far targets is already close to ideal without visual support. In near target experiments, the presence of visual feedback yielded near unity context gains, indicating close to optimal performance (retinal slip <0.4 degrees /s). For remembered targets, the context gain deteriorated but was still superior to performance in corresponding passive studies reported in the literature. In general, context compensation in the remembered target paradigm was better for vertical than for horizontal head rotations. The phase delay of version eye velocity relative to head velocity was small (approximately 2 degrees) for both horizontal and vertical head movements. Analysis of the vergence data from the near target experiments showed that context compensation took into account that the two eyes require slightly different VORs. In the DISCUSSION, comparison of the present default VOR gains and context gains with data from earlier passive studies has led us to propose a limited role for efference copies during self-generated movements. We also discuss how our analysis can provide a framework for evaluating two different hypotheses for the generation of binocular VOR eye movements.

Adult↗

Dynamical version--vergence interactions for a binocular implementation of Donders' law.

Recent investigations of the three-dimensional (3D) binocular eye positions in near vision have shown that a full characterization of vergence requires incorporation of its torsional component. The latter has a proportional relationship with horizontal vergence and elevation, causing the eyes to have intorsion in near upgaze but extorsion in near downgaze. In this study, we focus on the dynamical implementation of the torsional vergence component in both pure vergence and combined direction-depth binocular eye movements. We report on experiments in five subjects whose eye movements were recorded binocularly with the 3D magnetic search-coil technique. In pure vergence movements at a given elevation, torsional vergence increased with almost the same time course as horizontal vergence. In addition, the dynamic relationships among torsional vergence, horizontal vergence and elevation were close to static results in all subjects. In combined direction-depth movements a similar relationship held for the complete movements, but we could not firmly establish a straight-line relationship during the saccadic portion of the movement. Possible factors determining these responses are discussed. We computed the angular velocity profiles of pure vergence movements to see how tilting of the vergence angular velocity axis relative to Listing's plane generates torsional vergence. It is widely held that both saccadic and vergence movements are controlled by dedicated pulse generators specifying velocity signals. Little thought has been given to the question of how these controllers can be coordinated to yield realistic eye movements in 3D. Our finding that this tilt was close to full-angle, suggests a model in which version and vergence velocity signals are combined before the 3D neural integrator proposed by Tweed and Vilis. The implications of this scheme for the control of binocular eye movements in three dimensions are discussed, along with possible neural correlates.

Adult↗

An alternative three-dimensional interpretation of Hering's equal-innervation law for version and vergence eye movements.

In the context of Hering's equal-innervation law, this paper discusses the problem of how the three-dimensional positions of the two eyes, each expressed by a rotation vector, can be separated into contributions of the version and vergence system. As proposed by Van Rijn and Van den Berg [(1993) Vision Research, 33, 691-708], this can be done by taking the sum and difference of the position rotation vectors of each eye. In our alternative procedure the vergence signal is defined as the rotation which transforms the left eye position into the right eye position and the version signal is the common factor in both eye positions that remains after removing the vergence signal. The version and vergence contributions, defined in this way, can be interpreted straightforwardly as rotations. When Van Rijn and Van den Berg applied their definitions to their own data, they obtained the interesting result that the reconstructed version and vergence contributions were effectively limited to two dimensions (2D). The version signal was confined to Listing's plane (no torsion) whereas the vergence signal remained within a horizontal-torsional plane (no vertical vergence). They showed theoretically that a model based on 2D version/2D vergence control will indeed produce the torsional eye positions in near fixations found in their experiments. This model cannot account for a second set of data in the literature [Mok, Ro, Cadera, Crawford & Vilis (1992) Vision Research, 32, 2055-2064]. With our definitions, we found that the simple 2D version/2D vergence control strategy cannot account for the Van Rijn and Van den Berg (1993) data but is nicely compatible with the considerably smaller amount of cyclotorsion in the data collected by Mok et al. (1992). We also show that, in such a system, having 2D vergence control is compatible with minimization of torsional disparity and provides the cyclovergence signals suitable for stabilizing the eyes in the non-Listing positions caused by a vertical saccade in near vision.

Convergence, Ocular↗

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 quantitative 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 similar cyclotorsional components were found even in convergent refixations executed in full darkness towards the location of a remembered (flashed) target. We conclude that visual feedback is unlikely to be very important in controlling cyclovergence in these various conditions.

Adult↗

Evidence for nonretinal feedback in combined version-vergence eye movements.

Recently, a quantitative model for the general of rapid eye movements in direction and depth was proposed. In this scheme, the saccadic and the vergence system share a common initiation system and are controlled by local feedback loops based on efference copy signals. We have used a remembered-target double-step paradigm to test the idea that both subsystems are guided by extraretinal signals. The subject was instructed to move the binocular point of fixation to the remembered positions indicated by a double-step movement of the target, in direction and depth. Since both binocular refixations were made in complete darkness, correct execution of this task requires information about both the stored visual coordinates of the final target and the coordinates of the first movement. Binocular eye movements from five subjects were compared with predictions from two feed-forward models and a feedback model. Analysis of the pooled direction data showed that the feedback model performed best and fitted well. Qualitatively the same result was obtained in the vergence component, but in this case the goodness of fit was considerably less. These results, confirmed in each individual subject, show that the saccadic and vergence subsystem can use nonretinal information about a prior movement in direction and depth. Further analysis showed that the gain of the direction response of the second movement was, on average, roughly correct. By contrast, the vergence component of these responses was only about 60% of the required amplitude. Since the fit procedure gave the same weighting factors to the second target and to the first movement, we propose that the low vergence gain reflects mechanisms operating after the calculation of the motor error signal, possibly at the execution stage. Finally, we discuss the possibility of a central control stage keeping track of the ongoing movement sequence, based on a comparison of desired and current eye position signals.

Adult↗

Three-dimensional analysis of strongly curved saccades elicited by double-step stimuli.

Earlier recordings of eye position in three dimensions have revealed that Listing's law is obeyed in reasonable approximation, both statically and dynamically. This implies that all eye positions are confined to a plane when using a rotation vector or quaternion representation. The orientation of the angular velocity axis is crucial in order to preserve the law. For a single-axis rotation, the eye's angular velocity axis has to tilt out of Listing's plane, otherwise the law cannot be preserved in eccentric saccades. Experimental work has confirmed that normal, visually-guided saccades resemble single-axis rotations whose angular velocity axis tilts by the right amount. We investigated how well the saccadic system implements Listing's law when the trajectory of the eyes is more complicated, as in a non-single-axis rotation where the angular velocity vector depends on instantaneous eye position. Eye position was measured in three dimensions using the magnetic scleral search coil method for five subjects. Non-single-axis rotations of the eye were evoked with a double-step paradigm. We found that Listing's law is obeyed equally well during fixations, single-axis saccades and in non-single-axis saccades. Some deviations from the law were found in both curved and single-axis eye movements, but we demonstrated that the net torsional component of eye position of these saccades is negligible compared to that expected if the angular velocity axis did not tilt at all. In addition, analysis of the angular velocity signals in the curved movements showed strong similarity to the computed signal required for implementing Listing's law. Our results show that the observed deviations from Listing's law reflect only minor failures in the mechanism underlying its dynamic implementation. We conclude that single-axis rotations are not a necessary condition for the implementation of Listing's law in saccades. Our results are compatible with the notion that the implementation of Listing's law relies upon internal feedback. Various suggestions of how models can be reconciled with recent data on the three-dimensional control of saccades are discussed.

Adult↗

Saccadic eye movements in Parkinson's disease: II. Remembered saccades--towards a unified hypothesis?

Ten patients with mild to moderate Parkinson's disease were compared with ten age-matched normal controls in a series of saccadic paradigms in order to test various hypotheses relating to the origin of the Parkinsonian saccadic defect. The paradigms comprised a reflex saccade paradigm, a standard remembered saccade paradigm, a remembered saccade paradigm with delayed centre-offset, and a remembered saccade paradigm with a second target flash immediately prior to saccade execution. Finally, subjects executed both reflex and remembered saccades in a standard remembered paradigm (the "two-saccade" paradigm). As has been reported previously, Parkinsonian subjects demonstrated hypometria on all remembered saccade paradigms, particularly the "two-saccade" paradigm. There was, however, no significant difference between the first three remembered saccade paradigms. These studies serve to refute a simple attentional capture hypothesis, and a hypothesis that suggests that the abnormality of remembered saccades is due to concurrent reflex saccade suppression. On the basis of the results, further hypotheses are advanced in an attempt to explain all published work on Parkinsonian saccades.

Aged↗

Component stretching in fast and slow oblique saccades in the human.

We have studied the dynamics of human saccades along various cardinal (horizontal and vertical) and oblique directions in two different experimental paradigms yielding fast and slow saccades, respectively. We found that the saturation of vectorial peak velocity with amplitude, which is already well known from earlier studies on fast saccades, was equally pronounced in slow saccades. In both paradigms, the saturation level had a quite similar strong dependence on saccade direction. We found that the peak-velocity/amplitude relationships (main-sequences) of fast saccades in different directions were not simply scaled versions of one another. Whereas peak velocity in small saccades showed less anisotropy, different rates of saturation with amplitude in different directions (expressed in the angular constant parameter) caused the bundle of main-sequence curves to fan out at larger amplitudes. This property is reflected in a straight-line relationship between the angular constant and the asymptotic peak velocity parameters of the main-sequence. The possible contribution of neural control signals and plant properties is discussed. We also studied the main-sequences of oblique saccade components and found clear evidence for component stretching in both paradigms which increased as the saccade vector turned away from the cardinal direction under study. We conclude that the factors determining component stretching probably reside in the final pathway common for both saccade types. These experimental findings, revealing several common features in fast and slow saccades, were compared with quantitative predictions for the dynamics of oblique saccades, made from two existing two-dimensional models, predicting that the dynamic properties of components depend upon the direction of the saccade vector.

Adult↗

An analysis of curvature in fast and slow human saccades.

In order to study the cooperation of peripheral motor subsystems, the degree of curvature of human saccades along cardinal (right, up, left, down) and oblique directions was computed from an extensive set of experimental data. Our curvature measure allows comparison of fast and slow saccade trajectories elicited in different experimental conditions, independent of the speed of execution. Although we found clear and consistent subject-specific differences, the most common pattern in oblique visually-guided (i.e., fast) saccades reflected early dominance of the horizontal velocity signal as expressed in saccade trajectories curving away from the horizontal axis. Plots of curvature against direction yielded consistent idiosyncratic patterns with periodical increases and decreases in saccade curvature which were largely independent of saccade amplitude. At the cardinal axes, mean saccade curvature was generally less, but rarely entirely absent, and fitted smoothly into the curvature pattern of neighbouring quadrants. Memory-guided saccades, which have been shown earlier to be considerably slower than visually-guided saccades and to be more variable in their dynamic properties, showed a strikingly similar dependence of curvature on saccade direction, although some small but consistent differences were noticed. This finding suggests that saccade curvature is determined by mechanisms residing in the final common pathway for both saccade types. The curvature data were compared with quantitative predictions from three different models for the generation of oblique saccades. By quantifying the curvature of human saccades and thereby revealing the shortcomings of these models, the present paper documents new constraints with which future models of the saccadic system must comply and allows certain suggestions on how these might be developed.

Adult↗

Comparison of saccades evoked by visual stimulation and collicular electrical stimulation in the alert monkey.

In the alert monkey we have compared the properties of saccades elicited by a visual stimulus (V-saccades) with those generated by electrical stimulation in the superior colliculus (E-saccades). We found that whereas there exists a graded relation between E-saccade amplitude and current strength, E-saccade direction is remarkably independent of electrical stimulation parameters. At sufficiently high current strengths (about 20 microA), E-saccades are consistently directed toward the center of the movement field of nearby cells, except when stimulation is performed at sites near the collicular borders. Further interesting differences between the amplitude and direction behaviour were observed when the variability in E-saccade vectors, obtained with fixed stimulation parameters, was analyzed. In all cases, E-saccade amplitude scatter exceeds direction scatter, suggesting the possibility of a polar coordinate organization for the coding of saccade metrics. These data are compared with V-saccade scatter data, recently obtained in the human (Van Opstal and Van Gisbergen 1989c). Finally, an analysis of saccade dynamics shows that E-saccades can reach V-saccadic velocities at higher current strengths. However, at near-threshold current strengths, where E-saccade amplitude decreases, we found at most stimulation sites (22/37) that E-saccades are consistently slower than V-saccades of the same amplitude. Possible mechanisms underlying the collicular role in saccade generation are discussed.

Animals↗

A nonlinear model for collicular spatial interactions underlying the metrical properties of electrically elicited saccades.

An earlier model for the collicular role in the generation of saccades (Van Gisbergen et al. 1987), based on ensemble coding and linear vector addition of movement contributions from independent movement cells, yields normometric saccades in all directions over a considerable range of amplitudes. The model, however, cannot account for two nonlinear phenomena which are known from collicular electrical stimulation experiments: 1) saccade amplitude has a roughly sigmoid dependence upon current strength and 2) two electrical stimuli applied simultaneously at different sites yield a response that resembles a weighted average of the individual responses. In the present paper we propose an intracollicular mechanism which, based on lateral spatial interactions in the deeper layers of the colliculus, results in nearby excitation and remote inhibition when current is applied. Both nonlinear phenomena can thus be explained. The possibility of excitatory and inhibitory collicular interactions is supported by recent evidence in the literature. The nonlinearity in the model, essential to explain the electrical stimulation findings, resides in the input-output characteristic of the deeper layer movement cells. The results, obtained by quantitative simulations with the model, are discussed together with possible alternative explanations.

Computer Simulation↗

A short-latency transition in saccade dynamics during square-wave tracking and its significance for the differentiation of visually-guided and predictive saccades.

Several recent studies indicate that saccades elicited in the absence of a visual target are slower than visually-guided movements of the same size. In addition, we have shown earlier that the slower saccades observed in two different paradigms had more asymmetrical (skewed) velocity profiles. Recently, it has been reported that predictive saccades are also slower. An interesting question, which arises if predictive and visually-guided saccades do have different velocity profiles, is whether the time when the transition occurs can be determined from their dynamic characteristics (peak velocity and skewness) and whether this transition latency can serve as a plausible criterion for distinguishing predictive and visually-guided saccades. To investigate this problem, visually-guided and predictive saccades were elicited by various experimental paradigms in six normal human subjects. Eye movements were measured using the double-magnetic induction method. We found that scatter plots of normalized peak velocity against latency showed an abrupt, small (10-20%) increase at a surprisingly short latency (about 30-70 ms). Furthermore, skewness of the saccadic velocity profile showed a significant drop at comparable latencies. There was a tight correlation between the peak velocity and skewness transition latencies of each subject. Considering the shape of the latency histograms in this and earlier studies, as well as other data, it appears unlikely that these very short transition latencies demarcate the distinction between predictive and fully visually-guided saccades. Instead, we suggest the possibility that the visual stimulus can speed up saccades at an earlier time than it can initiate and guide them. If this is the case, the very short transition latencies (mean: about 50 ms) probably represent the sum of afferent and efferent pure time delays in the system and do not include the time needed for the computation of saccade metrical properties.

Adult↗