Centrifugal effects in the avian retina.

Electrical stimulation of the centrifugal fibers to the avian retina can disturb the balance between the excitatory and inhibitory system within the receptive fields of individual retinal ganglion cells. Although the mechanisms may vary from one unit to another, the effect is always to make them fire more readily and to a wider range of visual inputs.

Ocular responses to linear motion are inversely proportional to viewing distance.

Eye movements exist to improve vision, in part by preventing excessive retinal image slip. A major threat to the stability of the retinal image comes from the observer's own movement, and there are visual and vestibular reflexes that operate to meet this challenge by generating compensatory eye movements. The ocular responses to translational disturbances of the observer and of the scene were recorded from monkeys. The associated vestibular and visual responses were both linearly dependent on the inverse of the viewing distance. Such dependence on proximity is appropriate for the vestibular reflex, which must transform signals from Cartesian to polar coordinates, but not for the visual reflex, which operates entirely in polar coordinates. However, such shared proximity effects in the visual reflex could compensate for known intrinsic limitations that would otherwise compromise performance at near viewing.

Visual tracking and the primate flocculus.

Purkinje cells in the primate flocculus discharge specifically in relation to visual tracking, effectively generating a velocity profile of the target during pursuit. It is suggested that these neurons supply oculomotor centers with the velocity command signals needed to support pursuit eye movements.

The initial torsional Ocular Following Response (tOFR) in humans: a response to the total motion energy in the stimulus?

We recorded the initial torsional Ocular Following Responses (tOFRs) elicited at short latency by visual images that occupied the frontal plane and rotated about the lines of sight. Using 1-D radial gratings, the local spatio-temporal characteristics of these tOFRs closely resembled those we previously reported for the hOFRs to horizontal motion with 1-D vertical gratings. When the 1-D radial grating was subdivided into a number of concentric annuli, each with the same radial thickness, tOFRs were less than predicted from the sum of the responses to the individual annuli: spatial normalization. However, the normalization was much weaker than that which we previously reported for the hOFRs. Further, when the number, thickness and contrast of these concentric annuli were varied systematically, the latency and magnitude of the tOFRs were well described by single monotonic functions when plotted against the product of the total area of the annuli and the square of their Michelson contrast ("A*C(2)"), consistent with the hypothesis that the onset and magnitude of the initial tOFR are determined by the total motion energy in the stimulus. When our previously published hOFR data were plotted against A*C(2), a single monotonic function sufficed to describe the latency but not the magnitude.

Deficits in short-latency tracking eye movements after chemical lesions in monkey cortical areas MT and MST.

Past work has suggested that the medial superior temporal area (MST) is involved in the initiation of three kinds of eye movements at short latency by large-field visual stimuli. These eye movements consist of (1) version elicited by linear motion (the ocular following response), (2) vergence elicited by binocular parallax (the disparity vergence response), and (3) vergence elicited by global motion toward or away from the fovea (the radial-flow vergence response). We investigated this hypothesis by recording the effects of ibotenic acid injections in the superior temporal sulcus (STS) of both hemispheres in five monkeys. After the injections, all three kinds of eye movements were significantly impaired, with the magnitude of the impairments often showing a strong correlation with the extent of the morphological damage in the three subregions of the STS: dorsal MST on the anterior bank, lateral MST and middle temporal area on the posterior bank. However, the extent of the lesions in the three subregions often covaried, rendering it difficult to assess their relative contributions to the various deficits. The effects of the lesions on other aspects of oculomotor behavior that are known to be important for the normal functioning of the three tracking mechanisms (e.g., ocular stability, fixation disparity) were judged to be generally minor and to contribute little to the impairments. We conclude that, insofar as MST sustained significant damage in all injected hemispheres, our findings are consistent with the hypothesis that MST is a primary site for initiating all three visual tracking eye movements at ultra-short latencies.

Ocular following responses of monkeys to the competing motions of two sinusoidal gratings.

Ocular following responses (OFRs) were elicited in monkeys at short latencies ( approximately 50ms) by applying motion in the form of successive 1/4-wavelength steps to each of two overlapping vertical sine-wave gratings that had different spatial frequencies. In the first experiment, the two sine waves had spatial frequencies in the ratio 3:5 and moved in opposite directions. The initial OFRs showed a highly nonlinear dependence on the relative contrasts of the competing sine waves. On average, when the contrast of one was less than a third of that of the other then the one with the lower contrast became ineffective - as though suppressed - and the OFR was entirely determined by the sine wave of higher contrast: winner-take-all. In a second experiment, the two sine waves had spatial frequencies in the ratio 3:7 and moved in the same direction (though at different speeds). The initial OFRs again showed a highly nonlinear dependence on the relative contrasts of the competing sine waves, with a winner-take-all outcome when the contrasts of the two sine waves were sufficiently different. In both experiments, the nonlinear dependence on the relative contrasts of the competing sine waves was well described by a contrast-weighted-average model with just two free parameters. These findings were very similar to those of [Sheliga, B.M., Kodaka, Y., FitzGibbon, E.J., Miles, F.A., 2006c. Human ocular following initiated by competing image motions: evidence for a winner-take-all mechanism. Vision Res. 46, 2041-2060] on the human OFR, indicating that the monkey is a good animal model for studying the nonlinear interactions that emerge when competing motions are used.

Human vergence eye movements to oblique disparity stimuli: evidence for an anisotropy favoring horizontal disparities.

Binocular disparities applied to large-field patterns elicit vergence eye movements at ultra-short latencies. We used the electromagnetic search coil technique to record the horizontal and vertical positions of both eyes while subjects briefly viewed (150 ms) large patterns that were identical at the two eyes except for a difference in position (binocular disparity) that was varied in direction from trial to trial. For accurate alignment with the stimuli, the horizontal and vertical disparity vergence responses (HDVRs, VDVRs) should vary as the sine and cosine, respectively, of the direction of the disparity stimulus vector. In a first experiment, using random-dots patterns (RDs) with a binocular disparity of 0.2 degrees , this was indeed the case. In a second experiment, using 1-D sine-wave gratings with a binocular phase difference (disparity) of 1/4-wavelength, it was not the case: HDVRs were maximal when the grating was vertical and showed little decrement until the grating was oriented more than approximately 65 degrees away from vertical, whereas VDVRs were maximal when the grating was horizontal and began to decrement roughly linearly when the grating was oriented away from the horizontal. We attribute these complex directional dependencies with gratings to the aperture problem, and the HDVR data strongly resemble the stereothresholds for 1-D gratings, which are minimal when the gratings are vertical and remain constant for orientations up to approximately 80 degrees away from the vertical when expressed as spatial phase disparities [Morgan, M. J., & Castet, E. (1997). The aperture problem in stereopsis. Vision Research, 37, 2737-2744.]. To explain this constancy of stereothresholds, Morgan and Castet (1997) postulated detectors sensitive to the phase disparity of the gratings seen by the two eyes (rather than their linear separation along some fixed axis, such as the horizontal). However, because (1) our VDVR data with gratings did not show this constancy and (2) the available evidence strongly suggests that there are no major differences in the disparity detectors mediating the initial HDVR and VDVR, we sought an alternative explanation for our data. We show that the dependence of the initial HDVR and VDVR on grating orientation can be successfully modeled by a bias in the number and/or efficacy of the detectors that favors horizontal disparities.

Spatial summation properties of the human ocular following response (OFR): evidence for nonlinearities due to local and global inhibitory interactions.

Ocular following responses (OFRs) are the initial tracking eye movements that can be elicited at ultra-short latency by sudden motion of a textured pattern. A recent study used motion stimuli consisting of two large coextensive sine-wave gratings with the same orientation but different spatial frequency and moving in (1/4)-wavelength steps in the same or opposite directions: when the two gratings differed in contrast by more than about an octave then the one with the higher contrast completely dominated the OFR and the one with lower contrast lost its influence as though suppressed [Sheliga, B. M., Kodaka, Y., FitzGibbon, E. J., & Miles, F. A. (2006). Human ocular following initiated by competing image motions: Evidence for a winner-take-all mechanism. Vision Research, 46, 2041-2060]. This winner-take-all (WTA) outcome was attributed to nonlinear interactions in the form of mutual inhibition between the mechanisms sensing the competing motions. In the present study, we recorded the initial horizontal OFRs to the horizontal motion of two vertical sine-wave gratings that differed in spatial frequency and were each confined to horizontal strips that extended the full width of our display (45 degrees ) but were only 1-2 degrees high. The two gratings could be coextensive or separated by a vertical gap of up to 8 degrees , and each underwent motion consisting of successive (1/4)-wavelength steps. Initial OFRs showed strong dependence on the relative contrasts of the competing gratings and when these were coextensive this dependence was always highly nonlinear (WTA), regardless of whether the two gratings moved in the same or opposite direction. When the two gratings moved in opposite directions the nonlinear interactions were purely local: with a vertical gap of 1 degrees or more between the gratings OFRs approximated the linear sum of the responses to each grating alone. On the other hand, when the two gratings moved in the same direction the nonlinear interactions were more global: even with a gap of 8 degrees -the largest separation tried-OFRs were still substantially less than predicted by the linear sum. When the motions were in the same direction, we postulate two nonlinear interactions: local mutual inhibition (resulting in WTA) and global divisive inhibition (resulting in normalization). Motion stimuli whose responses were totally suppressed by coextensive opponent motion of higher contrast were rendered invisible to normalization, suggesting that the local interactions responsible for the WTA behavior here occur at an earlier stage of neural processing than the global interactions responsible for normalization.

Human vergence eye movements initiated by competing disparities: evidence for a winner-take-all mechanism.

Vergence eye movements were elicited in human subjects at short latencies (approximately 70 ms) by applying binocular disparities briefly (200 ms) to large grating patterns (46 degrees wide, 35 degrees high). The positions of both eyes were recorded with the electromagnetic search coil technique. Using a dichoptic viewing arrangement (Wheatstone stereoscope), each eye viewed two overlapping 1-D sine waves that had the same orientation but different spatial frequencies. These two sine waves each had a binocular disparity that was 1/4 of its wavelength and the effect of varying their relative contrasts was examined (15 contrast ratios ranging from 0.125 to 8). The first experiment used horizontal gratings and recorded the vertical vergence responses when the two sine waves had spatial frequencies in the ratio 3:5 and vertical disparities of opposite sign. Initial vergence responses showed a highly nonlinear dependence on the contrast ratio. On average, when the contrast of one sine wave exceeded that of the other by a factor of >2.2, the sine wave with the higher contrast dominated responses and the sine wave with the lower contrast had almost no influence: winner-take-all. A second experiment, which used vertical gratings and recorded the horizontal vergence responses when the two sine waves had spatial frequencies in the ratio 3:5 and horizontal disparities of opposite sign, also uncovered nonlinear interactions but these were much more variable from one subject to another and, on average, one sine wave did not achieve complete dominance until its contrast exceeded that of the other by a factor of >4.5. When these two experiments were repeated with grating patterns in which the two sine waves had spatial frequencies in the ratio 3:7 and disparities of the same sign, similar nonlinear interactions were apparent. We attribute the nonlinear dependence on relative contrast to mutual inhibition between the neural elements processing the disparities of the two sine waves. We further suggest that this interaction will help to maintain binocular alignment on the objects in the plane of regard because the retinal images of those objects will tend to be better focused-and hence tend to have higher contrasts-than the images of objects in other depth planes.

The vergence eye movements induced by radial optic flow: some fundamental properties of the underlying local-motion detectors.

Radial optic flow applied to large random dot patterns is known to elicit horizontal vergence eye movements at short latency, expansion causing convergence and contraction causing divergence: the Radial Flow Vergence Response (RFVR). We elicited RFVRs in human subjects by applying radial motion to concentric circular patterns whose radial luminance modulation was that of a square wave lacking the fundamental: the missing fundamental (mf) stimulus. The radial motion consisted of successive 1/4-wavelength steps, so that the overall pattern and the 4n+1 harmonics (where n=integer) underwent radial expansion (or contraction), whereas the 4n-1 harmonics--including the strongest Fourier component (the 3rd harmonic)--underwent the opposite radial motion. Radial motion commenced only after the subject had fixated the center of the pattern. The initial RFVRs were always in the direction of the 3rd harmonic, e.g., expansion of the mf pattern causing divergence. Thus, the earliest RFVRs were strongly dependent on the motion of the major Fourier component, consistent with early spatio-temporal filtering prior to motion detection, as in the well-known energy model of motion analysis. If the radial mf stimulus was reduced to just two competing harmonics--the 3rd and 5th--the initial RFVRs showed a nonlinear dependence on their relative contrasts: when the two harmonics differed in contrast by more than about an octave then the one with the higher contrast completely dominated the RFVRs and the one with lower contrast lost its influence: winner-take-all. We suggest that these nonlinear interactions result from mutual inhibition between the mechanisms sensing the motion of the different competing harmonics. If single radial-flow steps were used, a brief inter-stimulus interval resulted in reversed RFVRs, consistent with the idea that the motion detectors mediating these responses receive a visual input whose temporal impulse response function is strongly biphasic. Lastly, all of these characteristics of the RFVR, which we attribute to the early cortical processing of visual motion, are known to be shared by the Ocular Following Response (OFR)--a conjugate tracking (version) response elicited at short-latency by linear motion-and even the quantitative details are generally very similar. Thus, although the RFVR and OFR respond to very different patterns of global motion-radial vs. linear-they have very similar local spatiotemporal properties as though mediated by the same low-level, local-motion detectors, which we suggest are in the striate cortex.

Visual stabilization of the eyes in primates.

Observers moving through a textured three-dimensional world experience potentially confusing patterns of optic flow. Recent experiments on monkeys and humans have revealed the existence of rapid, yet cortically mediated, reflex eye movements that automatically single out images in the plane of fixation for stabilization and ignore images that are nearer or further.

Short-latency disparity vergence eye movements: a response to disparity energy.

Vergence eye movements were elicited in human subjects by applying disparities to square-wave gratings lacking the fundamental ("missing fundamental", mf). Using a dichoptic arrangement, subjects viewed gratings that were identical at the two eyes except for a phase difference of 1/4 wavelength so that, based on the nearest-neighbor matches, the features and the 4n+1 harmonics (5th, 9th, etc.) all had binocular disparities of one sign, whereas the 4n-1 harmonics (3rd, 7th, etc.) all had disparities of the opposite sign. Further, the amplitude of the ith harmonic was proportional to 1/i. Using the electromagnetic search coil technique to record the positions of both eyes indicated that the earliest vergence eye movements elicited by these disparity stimuli had ultra-short latencies (minimum, <65 ms) and were always in the direction of the most prominent harmonic, the 3rd, but their magnitudes fell short of those elicited when the same disparities were applied to pure sinusoids whose spatial frequency and contrast matched those of the 3rd harmonic. This shortfall was evident in both the horizontal vergence responses recorded with vertical grating stimuli and the vertical vergence responses recorded with horizontal grating stimuli. When the next most prominent harmonic, the 5th, was removed from the mf stimulus (creating the "mf-5" stimulus) the vertical vergence responses showed almost no shortfall-indicating that it had been almost entirely due to that 5th harmonic-but the horizontal vergence responses still showed a small shortfall, at least with higher contrast stimuli. This small shortfall might represent a very minor contribution from higher harmonics and/or distortion products and/or a feature-based mechanism. We conclude that the earliest disparity vergence responses-especially vertical-were strongly dependent on the major Fourier components of the binocular images, consistent with early spatial filtering of the monocular visual inputs prior to their binocular combination as in the disparity-energy model of complex cells in striate cortex [Ohzawa, I., DeAngelis, G. C., & Freeman, R. D. (1990). Stereoscopic depth discrimination in the visual cortex: neurons ideally suited as disparity detectors. Science, 249, 1037-1041].

The initial ocular following responses elicited by apparent-motion stimuli: reversal by inter-stimulus intervals.

Transient apparent-motion stimuli, consisting of single 1/4-wavelength steps applied to square-wave gratings lacking the fundamental ("missing fundamental stimulus") and to sinusoidal gratings, were used to elicit ocular following responses (OFRs) in humans. As previously reported [Sheliga, B. M., Chen, K. J., FitzGibbon, E. J., & Miles, F. A. (2005). Initial ocular following in humans: a response to first-order motion energy. Vision Research, in press], the earliest OFRs were strongly dependent on the motion of the major Fourier component, consistent with early spatio-temporal filtering prior to motion detection, as in the well-known energy model of motion analysis. Introducing inter-stimulus intervals (ISIs) of 10-200 ms, during which the screen was gray with the same mean luminance, reversed the initial direction of the OFR, the peak reversed responses (with ISIs of 20-40 ms) being substantially greater than the non-reversed responses (with an ISI of 0 ms). When the mean luminance was reduced to scotopic levels, reversals now occurred only with ISIs > or=60 ms and the peak reversed responses (with ISIs of 60-100 ms) were substantially smaller than the non-reversed responses (with an ISI of 0 ms). These findings are consistent with the idea that initial OFRs are mediated by first-order motion-energy-sensing mechanisms that receive a visual input whose temporal impulse response function is strongly biphasic in photopic conditions and almost monophasic in scotopic conditions.

Human ocular following initiated by competing image motions: evidence for a winner-take-all mechanism.

The initial ocular following responses (OFRs) elicited by 1/4-wavelength steps applied to the missing fundamental (mf) stimulus are in the backward direction and largely determined by the principal Fourier component, the 3rd harmonic [Sheliga, B. M., Chen, K. J., FitzGibbon, E. J., & Miles, F. A. (2005). Initial ocular following in humans: A response to first-order motion energy. Vision Research, 45, 3307-3321]. When the contrast of the 3rd harmonic was selectively reduced below that of the next most prominent harmonic-the 5th, which moves in the opposite (forward) direction-then the OFR reversed direction and the 3rd harmonic effectively lost all of its influence as the OFR was now largely determined by the 5th harmonic. Restricting the stimulus to just two sine waves (of equal efficacy when of equal contrast and presented singly) with the spatial frequencies of the 3rd and 5th harmonics of the mf stimulus indicated that the critical factor was the ratio of their two contrasts: when of similar contrast both were effective (vector sum/averaging), but when the contrast of one was <1/2 that of the other then the one with the lower contrast became ineffective (winner-take-all). This nonlinear dependence on the contrast ratio was attributed to mutual inhibition and was well described by a weighted-average model with just two free parameters. Further experiments with broadband and dual-grating stimuli indicated that nonlinear interactions occur not only in the neural processing of stimuli moving in opposite directions but also of stimuli that share the same direction and differ only in their spatial frequency and speed. Clearly, broad-band and dual-grating stimuli can uncover significant nonlinearities in visual information processing that are not evident with single sine-wave stimuli.

Short-latency disparity vergence in humans: evidence for early spatial filtering.

Our study was concerned with the disparity detectors underlying the initial disparity vergence responses (DVRs) that are elicited at ultrashort latencies by binocular disparities applied to large images. DVRs were elicited in humans by applying horizontal disparity to vertical square-wave gratings lacking the fundamental (termed here, the "missing fundamental"). In the frequency domain, a pure square wave is composed of odd harmonics--first, third, fifth, seventh, etc.--such that the third, fifth, seventh, etc., have amplitudes that are one-third, one-fifth, one-seventh, etc., that of the first, and the missing fundamental lacks the first harmonic. The patterns seen by the two eyes have a phase difference of one-quarter wavelength, so the disparity of the features and 4n + 1 harmonics (where n = integer) has one sign (crossed or uncrossed), whereas the 4n - 1 harmonics--including the strongest Fourier component (the third harmonic)--has the opposite sign (uncrossed or crossed): spatial aliasing. The earliest DVRs, recorded with the search-coil technique, had minimum latencies of 70 to 80 ms and were generally in the direction of the third harmonic, that is, uncrossed disparities resulted in convergent eye movements. In other experiments on the DVRs, one eye saw a missing fundamental and the other saw a pure sine wave with the contrast and wavelength of the third harmonic but differing in phase by one-quarter wavelength. This resulted in short-latency vergence in accordance with matching of the third harmonic. These data all indicate the importance of the Fourier components, consistent with early spatial filtering prior to binocular matching.

Initial ocular following in humans depends critically on the fourier components of the motion stimulus.

Visual motion is sensed by low-level (energy-based) and high-level (feature-based) mechanisms. Our interest is in the motion detectors underlying the initial ocular following responses (OFR) that are elicited at ultrashort latencies by sudden motions of large images. OFR were elicited in humans by applying horizontal motion to vertical square-wave gratings lacking the fundamental. In the frequency domain, a pure square wave is composed of the odd harmonics--first, third, fifth, seventh, etc.--such that the third, fifth, seventh, etc., have amplitudes that are one-third, one-fifth, one-seventh, etc., that of the first, and the missing fundamental stimulus lacks the first harmonic. Motion consisted of successive quarter-wavelength steps, so the features and 4n+1 harmonics (where n = integer) shifted forward, whereas the 4n-1 harmonics--including the strongest Fourier component (the third harmonic)--shifted backward (spatial aliasing). Thus, the net Fourier energy and the non-Fourier features moved in opposite directions. Initial OFR, recorded with the search coil technique, had minimum latencies of 60 to 70 ms and were always in the direction of the third harmonic, for example, leftward steps resulted in rightward OFR. Thus, the earliest OFR were strongly dependent on the motion of the major Fourier component, consistent with mediation by oriented spatiotemporal visual filters as in the well-known energy model of motion detection. Introducing interstimulus intervals of 10 to 100 ms (during which the screen was uniform gray) reversed the initial direction of tracking, consistent with extensive neurophysiological and psychophysical data suggesting that the visual input to the motion detectors has a biphasic temporal impulse response.

Initial ocular following in humans: a response to first-order motion energy.

Visual motion is sensed by low-level (energy-based) and high-level (feature-based) mechanisms. Ocular following responses (OFR) were elicited in humans by applying horizontal motion to vertical square-wave gratings lacking the fundamental ("missing fundamental stimulus"). Motion consisted of successive 1/4-wavelength steps, so the features and 4n+1 harmonics (where n=integer) shifted forwards, whereas the 4n-1 harmonics--including the strongest Fourier component (the 3rd harmonic)--shifted backwards (spatial aliasing). Initial OFR, recorded with the electromagnetic search coil technique, were always in the direction of the 3rd harmonic, e.g., leftward steps resulted in rightward OFR. Thus, the earliest OFR were strongly dependent on the motion of the major Fourier component, consistent with early spatio-temporal filtering prior to motion detection, as in the well-known energy model of motion analysis.

Short-latency visual stabilization mechanisms that help to compensate for translational disturbances of gaze.

Recent studies in primates have revealed short-latency visual tracking mechanisms that help to stabilize the eyes during translational disturbances of the observer, and so operate as backups to otolith-mediated vestibulo-ocular reflexes. One such mechanism generates version eye movements to help stabilize gaze when the moving observer looks off to one side, utilizing binocular disparity to help single out the images in the plane of fixation (ocular following). Two others generate vergence eye movements to help maintain binocular alignment on objects that lie ahead: one responds to the radial patterns of optic flow (radial-flow vergence) and the other to the changes in binocular parallax (disparity vergence). Accumulating evidence suggests that, despite their short latency, all are mediated by the medial superior temporal area of cortex.

Ocular compensation for self-motion. Visual mechanisms.

In monkeys, there are several reflexes that generate eye movements to compensate for the observer's own movements. Two vestibuloocular reflexes compensate selectively for rotational (RVOR) and translational (TVOR) disturbances of the head, receiving their inputs from the semicircular canals and otolith organs, respectively. Two independent visual tracking systems that deal with residual disturbances of gaze are manifest in the two components of the optokinetic response: the indirect or delayed component (OKNd) and the direct or early component (OKNe). We hypothesize that OKNd--like the RVOR--is phylogenetically old, being found in all animals with mobile eyes, and that it evolved as a backup to the RVOR to compensate for rotational disturbances of gaze. Indeed, optically induced changes in the gain of the RVOR result in parallel changes in the gain of OKNd, consistent with the idea of shared pathways as well as shared functions. In contrast, OKNe--like the TVOR--seems to have evolved much more recently in frontal-eyed animals and, we suggest, acts as a backup to the TVOR to deal primarily with translational disturbances of gaze. Frontal-eyed animals with good binocular vision must be able to keep both eyes directed at the object of regard irrespective of proximity, and in order to achieve this during translational disturbances, the output of the TVOR is modulated inversely with the viewing distance. OKNe shares this sensitivity to absolute depth, consistent with the idea that it is synergistic with the TVOR and shares some of its central pathways. There is evidence that OKNe is also sensitive to relative depth cues such as motion parallax, which we suggest helps the system to segregate the object of regard from other elements in the scene. However, there are occasions when the global optic flow cannot be resolved into a single vector useful to the oculomotor system (e.g., when the moving observer looks towards the direction of heading). We suggest that on such occasions a third independent tracking mechanism, the smooth pursuit system, is deployed to stabilize gaze on the local feature of interest. In this scheme, the pursuit system has an attentional focusing mechanism that spatially filters the visual motion inputs driving the oculomotor system. The major distinguishing features of the 3 visual tracking mechanisms are summarized in Table 1.

Population coding in cortical area MST.

Disparity steps applied to large patterns elicit vergence eye movements at ultrashort latencies. Disparity tuning curves, describing the dependence of the amplitude of the initial vergence responses on the amplitude of the disparity steps, resemble the derivative of a gaussian and indicate that appropriate servo-like behavior occurs only with small disparity steps (<1 degree). Lesion data from monkeys suggest that these vergence responses are mediated, at least in part, by neurons in the medial superior temporal area of the cerebral cortex, and we here review a recent study of the associated single unit activity in that area. Few medial superior temporal neurons have disparity tuning curves whose shapes resemble the tuning curve for vergence. Yet, when the disparity tuning curves for all of the disparity-sensitive cells recorded from a given monkey are summed together, they match the tuning curves for the vergence responses of that monkey very closely, even reproducing that animal's idiosyncracies. When all of the spike trains elicited by a given disparity step are summed together to give an average discharge profile for the whole population of recorded cells, many are noisy, but others that are less so match the temporal profile of the motor response, vergence velocity, quite well. We conclude that the discharges of the disparity-sensitive cells in the medial superior temporal area each represent only a very limited aspect of the sensory stimulus (and/or associated motor response?), but when pooled together, they provide a complete description of the vergence velocity motor response: population coding.