Adaptive changes in vergence eye movements induced by vergence-vestibular interaction training in monkeys

Clear vision of objects moving in three-dimensional space near an observer is attained by a combination of smooth-pursuit and vergence eye movements. The two systems must interact with the vestibular system to maintain the image of the object on the fovea. Previous studies showed that training with smooth-pursuit vestibular interactions resulted in adaptive changes in the smooth-pursuit response. Although vergence and smooth-pursuit systems are thought to have separate neural substrates, recent studies indicate that the caudal parts of the frontal eye fields that receive vestibular inputs contain neurons that discharge in response to combinations of smooth-pursuit and vergence. This combination of discharge sensitivities suggests the possibility that adaptive changes may be induced in the vergence system by vestibular inputs during vergence-pursuit training. To explore this possibility, we examined the effects of training with conflicting vestibular and vergence tracking in four head-stabilized monkeys. Animals were rewarded for tracking a laser spot that moved towards or away from them at 1 Hz in phase with sinusoidal whole-body rotation (±5°) in the pitch plane; the spot moved closer when the monkeys nose moved downward. From the monkeys point of view, the spot moved sinusoidally 10–66 cm in front of them along the mid-sagittal plane, requiring symmetrical vergence eye movements of 4.8° for each eye. Eye movements induced by equivalent spot motion at 0.3–1.0 Hz with or without chair rotation were examined before and after training for each session (0.5–1.0 h). Before training, pitch rotation alone in complete darkness did not induce vergence eye movements in any of the monkeys tested. Vergence tracking without chair rotation showed decreased gain and increased phase lag (re vergence target velocity) at frequencies above 0.5 Hz. After training, the vergence response during chair rotation with the spot showed significantly higher gains and smaller phase lags at 0.3–1.0 Hz in all monkeys. Pitch rotation alone in complete darkness induced vergence eye movements with gains (eye vergence/chair) of 0.15–0.35 after training in two monkeys. These results suggest that vestibular information can be used effectively to modify vergence tracking.     

Latency of adaptive vergence eye movements induced by vergence-vestibular interaction training in monkeys

Clear vision of objects that move in depth toward or away from an observer requires vergence eye movements. The vergence system must interact with the vestibular system to maintain the object images on the foveae of both eyes during head movement. Previous studies have shown that training with sinusoidal vergence-vestibular interaction improves the frequency response of vergence eye movements during pitch rotation: vergence eye velocity gains increase and phase-lags decrease. To further understand the changes in eye movement responses in this adaptation, we examined latencies of vergence eye movements before and after vergence-vestibular training. Two head-stabilized Japanese monkeys were rewarded for tracking a target spot moving in depth that required vergence eye movements of 10°/s. This target motion was synchronized with pitch rotation at 20°/s. Both target and chair moved in a trapezoidal waveform interspersed with random inter-trial intervals. Before training, pitch rotation in complete darkness without a target did not induce vergence eye movements. Mean latencies of convergence and divergence eye movements induced by vergence target motion alone were 182 and 169 ms, respectively. After training, mean latencies of convergence and divergence eye movements to a target synchronized with pitch rotation shortened to 65 and 53 ms, and vergence eye velocity gains (relative to vergence target velocity) at the normal latencies were 0.68 and 1.53, respectively. Pitch rotation alone without a target induced vergence eye movements with similar latencies after training. These results indicate that vestibular information can be used effectively to initiate vergence eye movements following vergence-vestibular training.     

Neural control of vergence eye movements: neurons encoding vergence velocity

Single-unit recordings were made from midbrain areas in monkeys trained to make both conjugate and disjunctive (vergence) eye movements. Previous work had identified cells with a firing rate proportional to the vergence angle, without regard to the direction of conjugate gaze. The present study describes the activity of neurons that burst for disjunctive eye movements. Convergence burst cells display a discrete burst of activity just before and during convergence eye movements. For most of these cells, the profile of the burst is correlated with instantaneous vergence velocity and the number of spikes in the burst is correlated with the size of the vergence movement. Some of these cells also have a tonic firing rate that is positively correlated with vergence angle (convergence burst-tonic cells). Divergence burst cells have similar properties, except that they fire for divergent and not convergent movements. Divergence burst cells are encountered far less often than convergence burst cells. Both convergence and divergence burst cells were found in an area of the mesencephalic reticular formation just dorsal and lateral to the oculomotor nucleus. Convergence burst cells were also recorded in another more dorsal mesencephalic region, rostral to the superior colliculus. Both of the areas also contain cells that encode vergence angle. Models of the vergence system derived from psychophysical data imply the existence of a vergence integrator, the output of which is vergence angle. Some models also suggest the presence of a parallel element that improves the frequency response of the vergence system, but has no effect on the steady-state behavior of the system. Vergence burst cells would be suitable inputs to a vergence integrator. By providing a vergence velocity signal to motoneurons, they may improve the dynamic response of the vergence system. The behavior of vergence burst cells during vergence movements is similar to that of the medium-lead burst cells during saccades. The proposed roles for vergence velocity cells are analogous to those of the saccadic burst cells. In this respect, the neural organization of the vergence system resembles that of the saccadic system, despite the distinct difference in the kinematics of these two types of eye movements.     

Latency of saccades and vergence eye movements in dyslexic children

We investigated the ability to adjust to nonlinear transformations that allow people to control external systems like machines and tools. Earlier research (Verwey and Heuer 2007) showed that in the presence of just terminal feedback participants develop an internal model of such transformations that operates at a relatively early processing level (before or at amplitude specification). In this study, we investigated the level of operation of the internal model after practicing with continuous visual feedback. Participants executed rapid aiming movements, for which a nonlinear relationship existed between the target amplitude seen on the computer screen and the required movement amplitude of the hand on a digitizing tablet. Participants adjusted to the external transformation by developing an internal model. Despite continuous feedback, explicit awareness of the transformation did not develop and the internal model still operated at the same early processing level as with terminal feedback. Thus with rapid aiming movements, the type of feedback may not matter for the locus of operation of the internal model.     

Dynamic properties of medial rectus motoneurons during vergence eye movements.

1. An early study by Keller reported that medial rectus motoneurons display a step change in firing rate during accommodative vergence movements. However, a later study by Mays and Porter reported gradual changes in firing rate during symmetrical vergence movements. Furthermore, subsequent inspection of the activity of individual medial rectus motoneurons during vergence movements indicated transient changes in their firing rate that had not been noted by Mays and Porter. For conjugate eye movements, in addition to a position signal, motoneurons display an eye velocity signal that compensates for the characteristics of the oculomotor plant. This suggested that the transient change in firing rate seen during vergence movements represented a velocity signal. Therefore the present study used single-unit recording techniques in alert rhesus monkeys to examine the dynamic behavior of medial rectus motoneurons during vergence eye movements. 2. The relationship between firing rate and eye velocity was first studied for vergence responses to step changes in binocular disparity and accommodative demand. Inspection of single trials showed that medial rectus motoneurons display transient changes in firing rate during vergence eye movements. To better visualize the dynamic signal during vergence movements, an expected firing rate (eye position multiplied by position sensitivity of the cell plus its baseline firing rate) was subtracted from the actual firing rate to yield a difference firing rate, which was displayed along with the eye velocity trace for individual trials. During all smooth symmetrical vergence movements, the profile of the difference firing rate very closely resembled the velocity profile. 3. To quantify the relationship between eye velocity and firing rate, two approaches were taken. In one, peak eye velocity was plotted against the difference firing rate. This plot yielded a measure of the velocity sensitivity of the cell (prv). In the other, a scatter plot was produced in which horizontal eye velocity throughout the vergence eye movement was plotted against the difference firing rate. This plot yielded a second measure of the velocity sensitivity of the cell (rv). 4. The behavior of 10 cells was studied during both sinusoidal vergence tracking and conjugate smooth pursuit over a range of frequencies from 0.125 to 1.0 Hz. This enabled the frequency sensitivity of the medial rectus motoneurons to be assessed for both types of movements. Both vergence velocity sensitivity and smooth pursuit velocity sensitivity decreased with increasing frequency. This is similar to a finding by Fuchs and co-workers for lateral rectus motoneurons during smooth pursuit eye movements.(ABSTRACT TRUNCATED AT 400 WORDS)     

Discharge characteristics of pursuit neurons in MST during vergence eye movements

For small objects moving smoothly in space close to the observer, smooth pursuit and vergence eye movements maintain target images near the foveae to insure high-resolution processing of visual signals about moving objects. Signals for both systems must be synthesized for pursuit-in-three-dimensions (3D). Recent studies have shown that responses of the majority of pursuit neurons in the frontal eye fields (FEF) code pursuit-in-3D. This area is known to have reciprocal connections with the medial superior temporal area (MST) where frontal pursuit neurons are found. To examine whether pursuit-in-3D signals are already present in MST and how MST neurons discharge during vergence-tracking induced by a small spot, we examined discharge of MST pursuit neurons in 2 monkeys. Of a total of 219 pursuit neurons examined during both frontal pursuit and vergence-tracking, 61% discharged only for frontal pursuit, 18% only for vergence-tracking, and 21% for both. A majority of vergence-related MST neurons exhibited sensitivity to vergence eye velocity. Their discharge was maintained during brief blanking of a vergence target. About 1/3 of vergence-related MST neurons exhibited visual responses to spot motion in depth. The preferred directions for visual motion and vergence-tracking were similar in half of our population. Some of the remaining neurons showed opposite preferred directions. A significant proportion (29%) of vergence-related neurons discharged before onset of eye movements with lead times longer than 20 ms. The results in this and previous studies indicate differences in discharge characteristics of FEF and MST pursuit neurons, suggesting different roles for the two in pursuit-in-3D.     

Speed-accuracy of saccades, vergence and combined eye movements in children with vertigo

Vergence abnormalities could lead to inappropriate vestibulo-ocular reflex (VOR), causing vertigo and imbalance (Brandt 1999). Indeed, a recent study by Anoh-Tanon et al. (2000) reported the existence of a population of children with symptoms of vertigo in the absence of vestibular dysfunction but with abnormal vergence findings in orthoptic tests. The purpose of this study was to examine in such children the accuracy, duration and mean velocity of vergence and saccades; additionally, for a few subjects, the effect of orthoptic vergence training on these parameters was also investigated. LEDs were used to stimulate saccades, pure vergence along the median plane and combined saccade-vergence movements. Movements from both eyes were recorded with a photoelectric device (Bouis). The results show that children with vertigo perform saccades as normal subjects of comparable age. In contrast, vergence, particularly convergence, shows abnormalities: poor accuracy, long duration and low speed. During combined movements, the well known reciprocal interaction between the saccade and the vergence is present only for saccades combined with divergence; for saccades combined with convergence such interaction is abnormal: the saccade is slowed down by the convergence but the convergence is not accelerated by the saccade. Orthoptic training improves significantly the accuracy of all eye movements; such improvement was significant for all types of eye movements except for divergence (pure and combined). Furthermore, convergence remains abnormal and the lack of acceleration by the saccade persists. These specific convergence deficits could be of both subcortical and cortical origin. Orthoptic training improves the accuracy presumably via visual attentional mechanisms, but cannot completely override deficits related to subcortical deficiencies.     

Short-latency vergence eye movements induced by radial optic flow in humans: dependence on ambient vergence level

Radial patterns of optic flow, such as those experienced by moving observers who look in the direction of heading, evoke vergence eye movements at short latency. We have investigated the dependence of these responses on the ambient vergence level. Human subjects faced a large tangent screen onto which two identical random-dot patterns were back-projected. A system of crossed polarizers ensured that each eye saw only one of the patterns, with mirror galvanometers to control the horizontal positions of the images and hence the vergence angle between the two eyes. After converging the subject's eyes at one of several distances ranging from 16.7 cm to infinity, both patterns were replaced with new ones (using a system of shutters and two additional projectors) so as to simulate the radial flow associated with a sudden 4% change in viewing distance with the focus of expansion/contraction imaged in or very near both foveas. Radial-flow steps induced transient vergence at latencies of 80-100 ms, expansions causing increases in convergence and contractions the converse. Based on the change in vergence 90-140 ms after the onset of the steps, responses were proportional to the preexisting vergence angle (and hence would be expected to be inversely proportional to viewing distance under normal conditions). We suggest that this property assists the observer who wants to fixate ahead while passing through a visually cluttered area (e.g., a forest) and so wants to avoid making vergence responses to the optic flow created by the nearby objects in the periphery.     

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     

Neural control of vergence eye movements: activity of abducens and oculomotor neurons

Single-unit recordings were made from neurons with horizontal eye position sensitivity in the oculomotor and abducens nuclei in alert monkeys. The animals were trained to perform a visual tracking task that resulted in conjugate eye movements or symmetrical vergence movements. Scatterplots were obtained for unit firing rate as a function of the position of the ipsilateral eye for both types of movement. The slopes of the linear regression line were computed for conjugate (kc) and vergence movements (kv). Previous recording studies implied that kv should be equal to kc for most, if not all, abducens and oculomotor neurons. Other lines of evidence suggested that kv should be zero for a substantial proportion of abducens neurons. In the abducens nucleus, we found some cells for which kv matched kc, and a few cells with a kv value of zero. However, the majority of abducens units had vergence signals that were neither equal to zero nor to their conjugate signals. Overall, kv/kc was 0.62, and the correlation between kv and kc was not significantly different from zero. Similarly, in the oculomotor nucleus, kv was significantly different from kc for a majority of the cells. A few units had kv values less than or equal to zero, whereas other cells had very high kv values. Overall, the kv/kc for oculomotor units was nearly unity (0.94), and the correlation between kv and kc was 0.31. These results confirm previous reports that most neurons in the abducens and oculomotor nuclei with a horizontal eye position sensitivity carry both conjugate and vergence eye movement signals. We do not find that the relative magnitudes of these signals are closely matched for most neurons. It is more likely that vergence and conjugate signals are matched globally, for an entire nucleus, rather than for individual motoneurons. This view is consistent with the hypothesis that conjugate and vergence signals are generated independently and combined for the first time at the motoneurons. Our results also imply that some motoneurons play a more important role than others in either vergence or conjugate movements.     

Cancelling of pursuit and saccadic eye movements in humans and monkeys

The countermanding paradigm provides a useful tool for examining the mechanisms responsible for cancelling eye movements. The key feature of this paradigm is that, on a minority of trials, a stop signal is introduced some time after the appearance of the target, indicating that the subject should cancel the incipient eye movement. If the delay in giving the stop signal is too long, subjects fail to cancel the eye movement to the target stimulus. By modeling this performance as a race between a go process triggered by the appearance of the target and a stop process triggered by the appearance of the stop signal, it is possible to estimate the processing interval associated with cancelling the movement. We have now used this paradigm to analyze the cancelling of pursuit and saccades. For pursuit, we obtained consistent estimates of the stop process regardless of our technique or assumptions--it took 50-60 ms to cancel pursuit in both humans and monkeys. For saccades, we found different values depending on our assumptions. When we assumed that saccade preparation was under inhibitory control up until movement onset, we found that saccades took longer to cancel (humans: approximately 110, monkeys: approximately 80 ms) than pursuit. However, when we assumed that saccade preparation includes a final "ballistic" interval not under inhibitory control, we found that the same rapid stop process that accounted for our pursuit results could also account for the cancelling of saccades. We favor this second interpretation because cancelling pursuit or saccades amounts to maintaining a state of fixation, and it is more parsimonious to assume that this involves a single inhibitory process associated with the fixation system, rather than two separate inhibitory processes depending on which type of eye movement will not be made. From our behavioral data, we estimate that this ballistic interval has a duration of 9-25 ms in monkeys, consistent with the known physiology of the final motor pathways for saccades, although we obtained longer values in humans (28-60 ms). Finally, we examined the effect of trial sequence during the countermanding task and found that pursuit and saccade latencies tended to be longer if the previous trial contained a stop signal than if it did not; these increases occurred regardless of whether the preceding trial was associated with the same or different type of eye movement. Together, these results suggest that a common inhibitory mechanism regulates the initiation of pursuit and saccades.     

Influence of gap and overlap paradigms on saccade latencies and vergence eye movements in seven-year-old children

The latency of eye movements is influenced by the fixation task; when the fixation stimulus is switched off before the target presentation (gap paradigm) the latency becomes short and express movements occur. In contrast, when the fixation stimulus remains on when the target appears (overlap paradigm), eye movement latency is longer. Several previous studies have shown increased rates of express saccades in children; however the presence of an express type of latency for vergence and combined movements in children has never been explored. The present study examines the effects of the gap and the overlap paradigms on horizontal saccades at far (150 cm) and at close (20 cm) viewing distances, on vergence along the median plane, and on saccades combined with convergence or divergence in 15 normal seven-year-old children. The results show that the gap paradigm produced shorter latency for all eye movements than the overlap paradigm, but the difference was only significant for saccades at close viewing distances, for divergence (pure and combined), and for saccades combined with vergence. The gap paradigm produced significantly higher rates of express latencies for saccades at close viewing distances, for divergence, and for saccades combined with divergence; in contrast, the frequencies of express latencies for saccades at far viewing distances and for convergence (pure or combined) were similar in the gap and the overlap paradigms. Interestingly, the rate of anticipatory latencies (<80 ms) was particularly high for divergence in the gap paradigm. Our collective findings suggest that the initiation of saccades at close viewing distances and of divergence is more reflexive, particularly in the gap paradigm. The finding of frequent anticipatory divergence that occurs at similar rates for seven-year-old children (this study) and for adults (Coubard et al., 2004, Exp Brain Res 154:368–381) indicates that predictive initiation of divergence is dominant.     

Discharge of frontal eye field neurons during saccadic and following eye movements in unanesthetized monkeys

Summary The cortical mechanism of eye-movement control was investigated by recording single cell activity from the frontal eye field (FEF) in unanesthetized monkeys seated in a primate chair with head restrained. Two types of cells (I and II) were found. Type I neurons fired during voluntary saccades occurring in a given direction and during the fast phase of nystagmus. Cells of this type were silent during slow pursuit movement. Type II cells showed steady discharge when the eyes were oriented in a specific direction. These cells discharged also during smooth pursuit movements and the slow phase of nystagmus, provided that the eyes were moving across positions which would have been associated with neuronal activity had the eyes come to rest there. All of Type II and a few of Type I neurons were identified by antidromic response to stimulation of the cerebral peduncle. These results indicate that cortical neurons have patterns of discharge distinctly related either to saccadic or to pursuit movements, in line with the view that these two different types of eye movement are generated by distinct neuronal mechanisms.     

Effects of voluntary blinks on saccades,vergence eye movements,and saccade-vergence interactions in humans

Blinks are known to change the kinematic properties of horizontal saccades, probably by influencing the saccadic premotor circuit. The neuronal basis of this effect could be explained by changes in the activity of omnipause neurons in the nucleus raphe interpositus or in the saccade-related burst neurons of the superior colliculus. Omnipause neurons cease discharge during both saccades and vergence movements. Because eyelid blinks can influence both sets of neurons, we hypothesized that blinks would influence the kinematic parameters of saccades in all directions, vergence, and saccade-vergence interactions. To test this hypothesis, we investigated binocular eye and lid movements in five normal healthy subjects with the magnetic search coil technique. The subjects performed conjugate horizontal and vertical saccades from gaze straight ahead to targets at 20 degrees up, down, right, or left while either attempting not to blink or voluntarily blinking. While following the same blink instruction, subjects made horizontal vergence eye movements of 7 degrees and combined saccade-vergence movements with a version amplitude of 20 degrees. The movements were performed back and forth from two targets simultaneously presented nearby (38 cm) and more distant (145 cm). Small vertical saccades accompanied most vergence movements. These results show that blinks change the kinematics (saccade duration, peak velocity, peak acceleration, peak deceleration) of not only horizontal but also of vertical saccades, of horizontal vergence eye movements, and of combined saccade-vergence eye movements. Peak velocity, acceleration, and deceleration of eye movements were decreased on the average by 30%, and their duration increased by 43% on the average when they were accompanied by blinks. The blink effect was time dependent with respect to saccade and vergence onset: the greatest effect occurred 100 ms prior to saccade onset, whereas there was no effect when the blink started after saccade onset. The effects of blinks on saccades and vergence, which are tightly coupled to latency, support the hypothesis that blinks cause profound spatiotemporal perturbations of the eye movements by interfering with the normal saccade/vergence premotor circuits. However, the measured effect may to a certain degree but not exclusively be explained by mechanical interference.     

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.     

Evidence for object permanence in the smooth-pursuit eye movements of monkeys

We recorded the smooth-pursuit eye movements of monkeys in response to targets that were extinguished (blinked) for 200 ms in mid-trajectory. Eye velocity declined considerably during the target blinks, even when the blinks were completely predictable in time and space. Eye velocity declined whether blinks were presented during steady-state pursuit of a constant-velocity target, during initiation of pursuit before target velocity was reached, or during eye accelerations induced by a change in target velocity. When a physical occluder covered the trajectory of the target during blinks, creating the impression that the target moved behind it, the decline in eye velocity was reduced or abolished. If the target was occluded once the eye had reached target velocity, pursuit was only slightly poorer than normal, uninterrupted pursuit. In contrast, if the target was occluded during the initiation of pursuit, while the eye was accelerating toward target velocity, pursuit during occlusion was very different from normal pursuit. Eye velocity remained relatively stable during target occlusion, showing much less acceleration than normal pursuit and much less of a decline than was produced by a target blink. Anticipatory or predictive eye acceleration was typically observed just prior to the reappearance of the target. Computer simulations show that these results are best understood by assuming that a mechanism of eye-velocity memory remains engaged during target occlusion but is disengaged during target blinks.     

Influence of saccadic eye movements on geniculostriate excitability in normal monkeys

Summary Using permanently implanted electrodes in squirrel monkeys and macaques, transmission through the lateral geniculate nucleus (LGN) was assayed from the amplitude of potentials evoked in optic radiation by an electrical pulse applied to optic tract. Averaging of either individually or machine selected potentials, elicited at 0.3, 1.0, 20 or 50 Hz, in all cases showed a decrease in transmission ranging from 5–60 % in the period after saccadic eye movements made ad libitum. The suppression was greater in a patterned visual environment than in diffuse illumination, which in turn was greater than that occurring following saccades in the dark. Demonstration of the effect in darkness always required data averaging and never exceeded 20%. The effect was consistently greater in the magnocellular than parvocellular component. Suppression was often abruptly terminated and replaced by a facilitation of 5–15% about 100 msec after saccade detection. Comparable effects were observed for excitability of striate cortex tested by a stimulus pulse applied to optic radiation. In addition, sharply demarcated potentials inherently arising in LGN and striate cortex were found in association with saccades made even in total darkness. Neglecting a possible but dubious contribution from eye muscle proprioceptors, the experiments establish the existence of a centrally originating modulation of visual processing at both LGN and striate cortex in relation to saccadic eye movement in primates. This modulation may partially underlie the phenomenon of saccadic suppression and hasten the acquisition of a meaningful visual sample immediately following an ocular saccade. It remains uncertain as to how it may relate to similar or greater effects accompanying changes in alertness, or to fluctuations of unknown origin occurring sometimes semirhythmically at 0.05–0.03 Hz (Fig. 7).Supported by Grant NS 03606 and Contract 70-2279 from the National Institutes of Neurological Diseases and Stroke, National Institutes of Health and by Grant GB 7522X from the National Science Foundation. B.B.L. was also aided by a travel grant from the Wellcome Trust (U.K.) and H.S. received a travel grant from the International Brain Research Organization.     

Effects of stationary textured backgrounds on the initiation of pursuit eye movements in monkeys.

1. The initial ocular pursuit of small target spots (0.25 degrees diam) that suddenly start to move at constant speed (ramps) was recorded in four rhesus monkeys with the electromagnetic search coil technique. All target motions were horizontal, and both eyes were monitored. 2. In agreement with the observations of Keller and Khan, stationary textured backgrounds substantially reduced the initial eye acceleration achieved during pursuit but did not affect its latency. Correlation techniques were used to assess the changes in the eye speed profiles and indicated that the reduction in eye acceleration due to the background was a linear function of the logarithm of target speed over the range investigated (5-40 degrees/s), averaging 60% with the fastest targets. 3. Selectively excluding the background texture from the path of the target with a horizontal strip of card (vertical width, 4 degrees) reduced the impact of the background only slightly, and, even when the vertical width of the card was increased to 60 degrees, the effect of the background was not entirely eliminated. Thus the effect involves regions of the visual field well beyond the target and is not due simply to the reduced physical salience (contrast) of the target spot. Such spatially remote interactions suggest that the neurons decoding the target's motion have very extensive visual receptive fields. 4. Textured backgrounds also caused similar reductions in the eye acceleration during initial pursuit when, before the ramps, the fixated target spots stepped forward, i.e., stepped in the direction of the subsequent ramps (step ramps). In this situation, as with no steps, initial target ramps were foveofugal. When the fixated target spots were stepped back before moving forward so that initial target ramps were foveopetal, textured backgrounds now also delayed the onset of pursuit, and the reductions in eye acceleration were not seen until some time later when tracking resulted from foveofugal target-ramp motion. Selectively excluding the texture from the path of the target with a narrow strip of card eliminated any delays in the onset of pursuit to step ramps, but the later reductions in eye acceleration were still evident. These step-ramp data indicate that the mechanisms decoding foveofugal and foveopetal target ramps differ markedly in their sensitivity to textured backgrounds. That backgrounds can influence the latency and the initial eye acceleration independently is consistent with the idea that there are independent trigger and drive mechanisms for the decoding of target motions.(ABSTRACT TRUNCATED AT 400 WORDS)