Luminance neurons in the pretectal olivary nucleus mediate the pupillary light reflex in the rhesus monkey

In humans and other primates, an increase in luminance in either eye elicits bilateral pupilloconstriction that is essentially equal in both eyes. Current models of the neural substrate for this clinically important light reflex propose that a retinorecipient pretectal nucleus projects bilaterally to the Edinger-Westphal nucleus (EW), which contains the parasympathetic, preganglionic neurons controlling pupilloconstriction. Based on single-unit recording studies in anesthetized cats and rats, it has been further suggested that luminance neurons in only one pretectal nucleus, the pretectal olivary nucleus, mediate this reflex. However, to our knowledge, there have been no comparable electrophysiological studies in primates of the pupillary light reflex or the pretectal luminance neurons that mediate this reflex. To address this issue, single-unit recording and electrical microstimulation studies were carried out in the pretectum of alert, trained, rhesus monkeys. These studies demonstrated that the primate pretectum contains luminance neurons with the characteristics appropriate for mediating the pupillary light reflex and that these neurons are located in one retinorecipient pretectal nucleus, the pretectal olivary nucleus. Electrical microstimulation at the site of these neurons often elicited pupilloconstriction. Our results provide clear evidence for the involvement of the pretectum, and more specifically the pretectal olivary nucleus, in mediating the pupillary light reflex in primates.     

Role of the pretectal nucleus of the optic tract in short-latency ocular following responses in monkeys

When a large-field image is suddenly moved in front of an observer, an ocular following response (OFR) with short latency (<60 ms in monkey and <85 ms in human) is observed. Previous studies have shown that neurons in the pretectal nucleus of the optic tract (NOT) of the monkey respond to movements of large-field visual stimuli. To understand the potential role of the NOT in the OFR, we first recorded single-unit activity in the NOT of four monkeys (Macaca fuscata). Sixty-six NOT neurons preferred large-field ipsiversive visual motion. In 86% (49/57) of the neurons, optimal directions were distributed over +/-30 degrees from ipsilateral. NOT units were sensitive to the speed of the visual motion; 54% (27/50) preferred slow (< or =20 degrees/s), 22% (11/50) preferred fast (> or =80 degrees/s) and the remainder intermediate speeds. Their response latencies to the moving visual scene were very short (approximately 51 ms), and 44% of them led the onset of the OFR by 10 ms or more. To characterize the response properties of these neurons, we reconstructed the temporal firing patterns of 17 NOT neurons, using the acceleration, velocity, position and bias components of retinal image slip or eye movements during the OFR by a least squares error method. For each stimulus speed fitting condition, using either retinal slip or eye movements, their firing patterns were matched to some extent although the goodness of fit was better using retinal slip than when eye movements were used. Neither of these models could be applied independently of stimulus speed, suggesting that the firing pattern of the NOT neurons represented information associated with retinal slip or eye movements during the OFR, over a limited range. To provide further evidence that the NOT is involved in generating the OFR, we placed unilateral microinjections of muscimol into the NOT. Following the muscimol injection, we observed a approximately 50% decrease in eye velocity of the OFR toward the side of injection regardless of stimulus speed, while only a weak effect was observed in the OFR during contraversive or vertical image motion. These results suggest that the NOT may play a role in the initiation and support of the short-latency ocular following response.     

Primate pupillary light reflex: receptive field characteristics of pretectal luminance neurons

This study examined the response properties of luminance neurons found within the pretectal olivary nucleus (PON), which is the pretectal nucleus that mediates the primate pupillary light reflex. We recorded the activity of 121 single units in alert, behaving rhesus monkeys trained to fixate a back-projected laser spot while a luminance stimulus was presented. The change in the firing rate of luminance neurons was measured as a function of changes in the size, retinal illuminance, and position of the stimulus. We found that these neurons possessed large receptive fields, which were sufficiently distinct that they could be placed into three classes. Approximately 40% of the PON luminance neurons responded well to stimuli presented in either the contralateral or ipsilateral hemifield. These neurons were classified as "bilateral" neurons. In the primate, retinal projections to the pretectum and other retinorecipient nuclei are organized such that direct retinal input can only account for the contralateral hemifield responses of these neurons. Thus the representation of the ipsilateral hemifield in "bilateral" PON cells must result from input from a nonretinal source. Approximately 30% of PON neurons responded only to stimuli presented in the contralateral hemifield. These neurons were classified as "contralateral" neurons. Finally, approximately 30% of PON neurons responded to stimuli presented at or near the animal's fixation point. These neurons were classified as "macular" neurons. The mean firing rates of all classes of neurons increased with increases in stimulus size and luminance within their receptive fields. The thresholds and magnitude of these responses closely matched those that would be appropriate for mediating the pupillary light reflex. In summary, these results suggest that all three classes of PON neurons contribute to the behaviorally observed pupillomotor field characteristics in which stimuli at the macular produce substantially larger pupillary responses than more peripheral stimuli. The contributions of "bilateral" and "contralateral" cells account for pupillary responses evoked by peripheral changes in luminance, whereas the contributions of all three cell classes account for the larger pupillary responses evoked by stimuli in the central visual field.     

The ultrastructure of the olivary pretectal nucleus in rats. A tracing and GABA immunohistochemical study

 The olivary pretectal nucleus is a primary visual centre, involved in the pupillary light reflex. In the present study an ultrastructural analysis was made of the olivary pretectal nucleus by means of separate, anterograde and retrograde tracing techniques and immunohistochemistry of gamma-aminobutyric acid. Large-projection neurons and two types of gamma-aminobutyric acid-immunoreactive (GABA-ir) neurons are observed in the olivary pretectal nucleus. The primary dendrites of the projection neurons have a dichotomous appearance, the secondary dendrites a multipolar appearance. At the ultrastructural level the projection neurons have well-developed Golgi fields, abundant rough endoplasmic reticulum and the nucleus is always heavily indented. Numerous small GABA-ir neurons and a few medium-sized GABA-ir neurons are found. The small GABA-ir neurons contain a few stacks of rough endoplasmic reticulum and the nucleus is oval-shaped. The medium-sized GABA-ir neurons have well-developed Golgi fields, a moderate number of rough endoplasmic reticulum stacks and an indented nucleus. GABA-positive dendritic profiles containing vesicles also are observed. In the neuropil of the olivary pretectal nucleus, retinal terminals are found that contain round clear vesicles and electron-lucent mitochondria. They make asymmetric synaptic contacts (Gray type I) with dendritic profiles and with profiles containing vesicles. Terminals originating from the contralateral olivary pretectal nucleus exhibit small, round clear vesicles, electron-dense mitochondria and make asymmetric synaptic contacts (Gray type I) mainly with dendritic profiles. Two types of GABA-ir terminals were found. One type is incorporated in glomerulus-like arrangements, whereas the other type is not. GABA-ir terminals contain pleomorphic vesicles, electron-dense mitochondria and make symmetric synaptic contacts (Gray type II). Retinal terminals, terminals originating from the contralateral olivary pretectal nucleus and GABA-ir terminals are organized in glomerulus-like structures, in which dendrites of the large projection neurons form the central elements. Triadic arrangements are observed in these structures; a retinal terminal contacts a dendrite and a GABA-ir terminal and the GABA-ir terminal also contacts the dendrite. The complexity of the synaptic organization and the abundancy of inhibitory elements in the olivary pretectal nucleus suggest that the olivary pretectal nucleus is strongly involved in processing visual information in the pupillary light reflex arc. Received: 17 July 1996 / Accepted: 24 September 1996     

Synaptic organization in the olivary pretectal nucleus of the adult rat

The fine structure of the glomerular neuropil of the olivary pretectal nucleus has been studied in adult rats. Four presynaptic components and two postsynaptic components of the neuropil are described and the synaptic relationships established between these components, including complex serial synapses, are defined. A conspicuous feature of this neuropil is the presence of triplet (triadic) synapses involving retinal afferents, appendages of the presynaptic dendrites of intrinsic neurones, and projection cell dendrites.     

Differential effects of aging on the distribution of calcium-binding proteins in a pretectal nucleus of the chicken brain

The nucleus pretectalis (PT) of birds is an ovoid-shaped visuomotor cell group of the pretectum that receives tectal input and projects back to the optic tectum. We performed immunohistochemical single- and double-labeling to determine the distribution and abundance of neurons containing three calcium-binding proteins, parvalbumin (PV), calretinin (CR), and calbindin (CB), in the PT in chickens at three ages. We found that PV-positive and CR-positive cells co-localize and are largely found in the outer part of PT at all ages. The GluR4 subunit of the AMPA-type glutamate receptor was selectively localized to these neurons. CB-positive neurons, however, were largely absent from the PT in young and adult chickens. The abundance of PV-positive and CR-positive neurons in PT in old birds was indistinguishable from that in the younger birds, but CB-positive perikarya were 10-20-fold more common than in young birds, and were again mainly found in the outer part of PT. The overall abundance of neurons in PT was reduced to about 50% of its former abundance in the old birds, with this loss restricted to the central part of the nucleus. These data indicate that a cell loss process develops in PT as birds age, that parvalbuminergic and calretinergic neurons resist this process, and that this process is associated with increased expression of CB.     

Antidromic identification of midbrain near response cells projecting to the oculomotor nucleus

Summary Medial rectus motoneurons carry both conjugate and vergence eye position signals. Abducens internuclear neurons, whose axons travel in the medial longitudinal fasciculus, provide these motoneurons with the major signal for conjugate eye movements but not for vergence eye movements. A vergence signal appropriate for these motoneurons is seen on the near response cells that are found in the mesencephalic reticular formation within 2 mm of the oculomotor nucleus. The goal of the present study was to determine if midbrain near response cells project to the medial rectus subdivision of the oculomotor nucleus. Near response cells were recorded in two trained rhesus monkeys with ocular search coils. A stimulating electrode was positioned within the medial rectus subdivision of the oculomotor nucleus. Twenty-eight near response cells were found that could be driven by single pulse microstimulation of the ipsilateral medial rectus subdivision. In all cases, antidromic activation was confirmed by collision testing. Attempts to antidromically activate midbrain near response cells from the contralateral medial rectus subdivision were unsuccessful. Most antidromically activated cells had a steady state firing rate proportional to vergence angle. One cell also showed burst activity during the vergence eye movements. Divergence cells were not antidromically activated.     

Electrophysiological study of inferior olivary neurons in staggerer mutant mice

Summary Extracellular recordings were obtained from neurons of the dorsal and medial accessory olives (DAO and MAO) in staggerer and normal mice. The anti- and orthodromic responses of these neurons to stimulation of the contralateral cerebellum were very similar in control and staggerer mice. On the other hand, the mean value of the spontaneous discharge of inferior olivary (I.O.) cells was significantly lower and the modal interval of the discharge was significantly shorter in staggerer than in control mice. These results demonstrate that I.O. neurons and climbing fibers are functional in staggerer mice, although their spontaneous discharge is somewhat different from that of I.O. neurons in control mice.Attaché de Recherche INSERM. This work was performed while the author was at the Laboratoire de Physiologie Comparée, Université Pierre et Marie Curie, Paris, France     

Luminance detectors in the olivary pretectal nucleus and their relationship to the pupillary light reflex in the rat. II. Studies using sinusoidal light

Summary The luminance detectors in the olivary pretectal nucleus, which are likely candidates mediating the pupillary light reflex, responded to all frequencies of sinusoidally modulated light up to 12–25 Hz. At low frequencies (0.05–4.0 Hz) the luminance detectors responded with modulated firing to different stimulation rates. The modulation depth of the cell response increased with the increase in stimulation frequency up to 20 Hz, then rapidly fell. There was a delay between the peak intensity of the stimulus and the peak firing of cell response of about 30–40 ms. The amplitudes of the consensual pupil responses to the same sinusoidal stimulus, on the other hand, decreased with an increase in frequency and no discernible response was recorded above 2 Hz. The pupil responses were little affected by sympathectomy. The differences in the frequency response characteristics of luminance detectors and the pupil were attributed to the sluggish dynamic properties of the pupil muscles. This was demonstrated using an electronic model of the iris muscle which modified the responses of the luminance detectors giving output waveforms broadly resembling pupil responses to square and sinusoidally modulated lights.The work was supported by a grant from the Special Trustees of St. Thomas' Hospital to H.I.; R.J.C. was a holder of a studentship of the Science Research Council of Great Britain (SERC)     

Behavior of preoculomotor burst neurons during eye-head coordination

Summary Single-unit recordings from the pontine reticular formation in four monkeys have shown the presence of two classes of short-lead burst neurons firing during coordinated eye-head movements. The activity of one class showed a correlation with the size of saccadic movements performed during head movements; the other showed a correlation between firing pattern and the combined eye-head movement. Anatomical reconstructions of the recording sites point to an intermingling of the two cell types.This research was supported by National Institutes of Health Research Grant NS09343, awarded by National Institute of Neurological Diseases and Stroke, by National Aeronautics and Space Administration Grant NGR 22-099-798, and by National Eye Institute Grant EY02621     

Axon terminals of the projection from the superior colliculus to the olivary pretectal nucleus in the rat

The terminals of axons projecting to the olivary pretectal nucleus have been identified by electron microscopy following injections of horseradish peroxidase into the superior colliculus of adult albino rats. The labelled terminals were equated with RD-terminals described in previous studies of this nucleus. They were 0.3-1.3 micron in diameter and contained round synaptic vesicles. Most also contained small dark mitochondria. They established Gray type 1 synaptic contacts with the dendrites of presumptive projection cells. Most terminated within non-glomerular neuropil, chiefly in the peripheral 'shell' of the nucleus; a few terminated in regions of glomerular neuropil.     

Characteristics of the pupillary light reflex in the macaque monkey: discharge patterns of pretectal neurons

Anatomical and physiological data have implicated the pretectal olivary nucleus (PON) as the midbrain relay for the pupillary light reflex in a variety of species. To determine the nature of the discharge of pretectal light reflex relay neurons, we recorded their activity in monkeys that were fixating a stationary spot while a full-field random-dot stimulus was flashed on for 1 s. Based on their discharge patterns, neurons in or near the PON came in two varieties. The most prevalent neuron discharged a burst of spikes 56 ms (on average) after the light came on followed by a sustained rate for the duration of the stimulus (burst-sustained neurons). When the light went off, nearly all neurons (33/34) ceased firing, and then all the neurons with a resting response in the dark (n = 15) resumed firing. Both the firing rate within the burst and the sustained discharge rate increased with log light intensity and the latency of the burst decreased. The burst and cessation of firing were better aligned with the stimulus occurrence than with the onset of pupillary constriction or dilation. Taken together, these data suggest that burst-sustained neurons respond to the visual stimulus eliciting the pupillary change rather than dictating the metrics of the subsequent pupillary response. Electrical stimulation at the site of four of five burst-sustained neurons elicited pupillary constriction at low stimulus strengths after a latency of approximately 100 ms. When the electrode was moved 250 microm away from the burst-sustained neuron, the elicited response disappeared. Reconstructions of the locations of burst-sustained luminance neurons place them in the PON or its immediate vicinity. We suggest that PON burst-sustained neurons constitute the pretectal relay for the pupillary light reflex. A minority of our recorded pretectal neurons discharged a burst of spikes at both light onset and light offset. For most of these transient neurons, neither the burst rate nor the interburst rate was significantly related to light intensity. We conclude that these neurons are not involved in the light reflex but subserve some other pretectal function.     

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.     

Discharge of primate magnocellular red nucleus neurons during reaching to grasp in different spatial locations

Reaching to grasp is of fundamental importance to primate motor behavior. One descending motor pathway that contributes to the control of this behavior is the rubrospinal tract. An important source of origin of the rubrospinal tract is the magnocellular red nucleus (RNm). Forelimb RNm neurons discharge vigorously during reach-to-grasp movements. RNm discharge is important for hand use, as coordinated whole-limb movements without hand use are not associated with strong discharge. Because RNm is functionally linked to muscles of the entire forelimb, RNm discharge may also contribute to use of the proximal limb that accompanies hand use. If RNm contributes to proximal limb use, we predict discharge to differ for reaches that differ in proximal limb involvement but require the same grasp. We tested this prediction by measuring discharge of individual RNm neurons while monkeys reached to grasp objects in four spatial locations in front of them. The animals reached from the waist to locations to the left, right, above, and below the shoulder of the "reaching" limb. RNm neurons of our sample were activated strongly during reach-to-grasp, and discharge of a third of the neurons tested depended on the spatial location of the object grasped. Discharge of RNm neurons and EMG activity of many of the distal and proximal forelimb muscles we tested were larger for reaching to grasp in the upper and/or right than lower and left target locations. Based on comparisons of each individual neuron's discharge patterns during reaches with and without preshaping the hand, we conclude that target location-dependent modulations in discharge rate of the majority of RNm neurons whose discharge differed for reaching to grasp in the four target locations contributed to aspects of hand preshaping that covaried with reach direction.     

Morphology of neurons in the lateral geniculate nucleus of the monkey

Summary Rapid Golgi preparations of the lateral geniculate nucleus of old- and new-world monkeys were analysed in an attempt to classify the neuronal types. Four main types are described. The commonest, the multipolar neurons, are found in all laminae. Their somata can be large, medium or small and bear dendrites with sparse spines. Some have a radiate dendritic arbor and others have dendrites grouped in tufts. The next most frequent class is of bipolar neurons with two thick dendrites arising from opposite poles of the soma, which is usually large. Otherwise the dendrites are similar to those of multipolar neurons. Relatively rare is a class of medium-sized neurons with beaded dendrites, found here only in magnocellular laminae. There is a fourth class of small neurons distinguished by fine axonlike dendritic processes. They are in all laminae and form two subgroups, one with very long, cylindrical dendrites and few axon-like processes, the other with shorter dendritic arbors and many axon-like processes. In addition, a class of capsular neurons is found in the circumgeniculate capsule between layer 6 and the pregeniculate nucleus. They are large neurons with umbrella-like dendritic arbors sending ramifications into layer 6. The interlaminar zones contain scattered somata of all types except beaded and capsular neurons.The work described in this paper forms part of a study for a doctoral dissertation in the University of Lausanne by K.D. Saini     

Visual and vergence eye movement-related responses of pursuit neurons in the caudal frontal eye fields to motion-in-depth stimuli

The caudal parts of the frontal eye fields (FEF) contain smooth-pursuit related neurons. Previous studies show that most FEF pursuit neurons carry visual signals in relation to frontal spot motion and discharge before the initiation of smooth-pursuit. It has also been demonstrated that most FEF pursuit neurons discharge during vergence tracking. Accurate vergence tracking requires information about target motion-in-depth. To further understand the role of the FEF in vergence tracking and to determine whether FEF pursuit neurons carry visual information about target motion-in-depth, we examined visual and vergence eye movement-related responses of FEF pursuit neurons to sinusoidal spot motion-in-depth. During vergence tracking, most FEF pursuit neurons exhibited both vergence eye position and velocity sensitivity. Phase shifts (re target velocity) of most neurons remained virtually constant up to 1.5 Hz. About half of FEF pursuit neurons exhibited visual responses to spot motion-in-depth. The preferred directions for visual responses of most neurons were similar to those during vergence tracking. Visual responses of most of these neurons exhibited sensitivity to the velocity of spot motion-in-depth. Phase shifts of most of the responding neurons remained virtually constant up to 2.0 Hz. Neurons that exhibited visual responses in-depth were mostly separate from neurons that showed visual responses in the frontal plane. To further examine whether FEF pursuit neurons could participate in initiation of vergence tracking, we examined latencies of neuronal responses with respect to vergence eye movements induced by step target motion-in-depth. About half of FEF pursuit neurons discharged before the onset of vergence eye movements with lead times longer than 20 ms. These results together with previous observations suggest that the caudal FEF carries visual signals appropriate to be converted into motor commands for pursuit in depth and frontal plane.     

Luminance and darkness detectors in the olivary and posterior pretectal nuclei and their relationship to the pupillary light reflex in the rat

Summary In order to identify the pretectal nucleus which contains pupillomotor cells in the rat, cells were sought which were sensitive to changes in luminance level at the eye. Two types were found: Luminance detectors which showed a graded increase in firing with increase in luminance, and darkness detectors which showed a graded increase in firing rate with graded dimming of luminance intensity. All luminance detectors were located in the olivary pretectal nucleus, whereas darkness detectors were located in the posterior pretectal nucleus. Consensual pupil responses were recorded in conscious normal and sympathectomised rats using an infra-red sensitive T.V. pupillometer. Pupil diameter varied 2mm in an approximately linear fashion over six log units range in luminance intensity. Sympathectomy produced a general constriction of the pupil, but the overall response to light was unaffected. The changes in pupil size occurred over the same range of luminance that the firing rates of both luminance and darkness detectors changed. The olivary pretectal nucleus may therefore be involved in pupilloconstruction in the light, and the posterior pretectal nucleus, with pupillodilation in the dark.This work was supported by a grant from the Special Trustees of St. Thomas' Hospital to H.I.; R.J.C. was a holder of a studentship of the Science Research Council of Great Britain during 1976–1979     

In vitro properties of neurons in the rat pretectal nucleus of the optic tract

The nucleus of the optic tract (NOT) has been implicated in the initiation of the optokinetic reflex (OKR) and in the modulation of visual activity during saccades. The present experiments demonstrate that these two functions are served by separate cell populations that can be distinguished by differences in both their cellular physiology and their efferent projections. We compared the response properties of NOT cells in rats using target-directed whole cell patch-clamp recording in vitro. To identify the cells at the time of the recording experiments, they were prelabeled by retrograde axonal transport of WGA-apo-HRP-gold (15 nm), which was injected into their primary projection targets, either the ipsilateral superior colliculus (iSC), or the contralateral NOT (cNOT), or the ipsilateral inferior olive (iIO). Retrograde labeling after injections in single animals of either WGA-apo-HRP-gold with different particle sizes (10 and 20 nm) or two different fluorescent dyes distinguished two NOT cell populations. One projects to both the iSC and cNOT. These cells are spontaneously active in vitro and respond to intracellular depolarizations with temporally regular tonic firing. The other population projects to the iIO and consists of cells that show no spontaneous activity, respond phasically to intracellular depolarization, and show irregular firing patterns. We propose that the spontaneously active pathway to iSC and cNOT is involved in modulating the level of visual activity during saccades and that the phasically active pathway to iIO provides a short-latency relay from the retina to premotor mechanisms involved in reducing retinal slip.     

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.     

Development of the rabbit visual cortex: A quantitative Golgi analysis

Summary The horizontal and vertical components of the positions of both eyes of rhesus monkeys were measured during periods of binocularly stable eye positions (eye pauses) while the animals fixated a small target. Differences between monocular and binocular viewing, as well as effects of target size and background illumination, were assessed and found to be comparable to similar measures for humans. The scatter of eye position for either eye during binocular viewing had a standard deviation of 6–8 min arc in the horizontal and 7–13 min arc in the vertical meridia. Measurements of vergence and vertical misalignment, taken from binocular positional disparity, showed that for nearly 60% of eye pause time the eyes were misaligned on the fixation target by more than 7 min arc along both horizontal and vertical axes. In addition, the line of gaze during the trial was found to follow certain idiosyncratic tendencies for each monkey, although the positional variability remained relatively constant throughout the fixation trial. These observations suggest that during binocular fusion and stereopsis a mechanism exists that dynamically compensates for the relatively large shifts in retinal image position during fixation.Supported by NIH Grant EY02966