Microstimulation in visual area MT: effects of varying pulse amplitude and frequency.

We have previously shown that perceptual judgements of motion direction are based in part on the activity of direction selective neurons in extrastriate visual area MT (Salzman et al., 1990, 1992). In those experiments, we applied low-amplitude microstimulation pulses (10 microA, 200 Hz) to clusters of MT neurons whose preferred directions were similar. The effect of microstimulation was to bias the monkeys' choices on a direction discrimination task toward the preferred direction of neurons at the stimulation site. The results suggest that microstimulation generated a directionally specific cortical signal by activating selectively neurons near the electrode tip. To test this notion more directly, we have now examined the behavioral effects of varying current amplitude, current frequency, and electrode position. In the majority of experiments, the directional bias in the monkeys' choices was reduced or eliminated as current amplitude increased to 80 microA. In addition, 80 microA stimulating pulses frequently impaired overall performance as measured by the percentage of correct responses. This decrement in performance indicated that 80 microA pulses introduced "noise" into the neural circuitry encoding motion direction, presumably by increasing current spread to activate a larger population of neurons representing all directions of motion. In contrast, increasing current frequency to 500 Hz (10 microA pulses) preserved the directional specificity of microstimulation effects. The precise position of the stimulating electrode also influenced the magnitude of microstimulation effects; in some cases, differences in position on the order of 100 microns determined whether an experiment yielded a very large effect or no effect at all. Thus, directionally specific activation of cortical circuitry within MT can be disrupted by increases in current spread or by small changes in electrode position. These observations suggest that the effects of low-amplitude microstimulation depend upon direct activation of a well-localized population of neurons.

Microstimulation in visual area MT: effects on direction discrimination performance.

Physiological and behavioral evidence suggests that the activity of direction selective neurons in visual cortex underlies the perception of moving visual stimuli. We tested this hypothesis by measuring the effects of cortical microstimulation on perceptual judgements of motion direction. To accomplish this, rhesus monkeys were trained to discriminate the direction of motion in a near-threshold, stochastic motion display. For each experiment, we positioned a microelectrode in the middle of a cluster of neurons that shared a common preferred direction of motion. The psychophysical task was then adjusted so that the visual display was presented directly over the neurons' receptive field. The monkeys were required to discriminate between motion shown either in the direction preferred by the neurons or in the opposite direction. On half the trials of an experiment, we applied electrical microstimulation while monkeys viewed the motion display. We hypothesized that enhancing the neurons' discharge rate would introduce a directionally specific signal into the cortex and thereby influence the monkeys' choices on the discrimination task. We compared the monkeys' performance on "stimulated" and "nonstimulated" trials in 139 experiments; all trials within an experiment were presented in random order. Statistically significant effects of microstimulation were obtained in 89 experiments. In 86 of the 89 experiments with significant effects (97%), the monkeys indicated that motion was in the neurons' preferred direction more frequently on stimulated trials than on nonstimulated trials. The data demonstrate a functional link between the activity of direction selective neurons and perceptual judgements of motion direction.

Human optokinetic nystagmus: competition between stationary and moving displays.

We reported earlier that occlusion of the central retina and stationary edges have highly interactive effects on the gain of optokinetic nystagmus (OKN; Murasugi, Howard, & Ohmi, 1986). In this study, we explored this effect in more detail. A central occluding band of variable height, flanked by vertical bars, was superimposed onto an array of dots moving at 30 degrees per second. The height of the occluding band required to abolish OKN increased with the separation of the vertical bars. For bars 3.5 degrees apart, OKN was abolished in most subjects when a band only 6' high ran between them. For bars 75 degrees apart, a band at least 20 degrees in height was required to abolish the response. The effects of the stationary figure depended to some extent on the subject's attention, but only at intermediate values of bar separation. Both low- and high-level mechanisms are proposed to account for the results.

Human horizontal optokinetic nystagmus elicited by the upper versus the lower visual fields.

A 30-deg-high horizontally rotating random-dot display was presented to the central field, and with its more central edge at vertical eccentricities of 0, 2.5, 5, and 10 deg above or below the horizon. Stimulus velocities of 25-100 deg/s and two directions of motion were presented. The mean gain of the slow phases of optokinetic nystagmus (OKN) for five subjects was significantly higher when the stimulus was presented to the lower visual field than when the stimulus was presented to the upper field. This difference was most pronounced when the display was displaced 5 deg from the fovea and moving below 100 deg/s. Our results are consistent with existing psychophysical and physiological evidence for the superiority of the upper retina. In addition, four of the five observors showed significant directional asymmetries.

Suppression of OKN and VOR by afterimages and imaginary objects.

Optokinetic nystagmus (OKN) is suppressed if attention is directed to a centrally placed afterimage superimposed on a moving display. Imagining a stationary object has little or no effect. An afterimage does not provide the retinal slip and misfoveation error signals provided by a stationary object and we have shown that an effective error signal does not arise from occlusion or masking of the display by the afterimage. Although a lack of relative motion between afterimage and moving display could indicate when OKN gain is one, there is no unique relative motion signal associated with a gain of zero. Subjects could partially inhibit the vestibulo-ocular reflex (VOR) in the dark when they imagined a head-fixed object. They could suppress the response more effectively by attending to an afterimage, but the suppression was still only partial. When OKN and VOR were evoked simultaneously, pursuit movements of the eyes could not be suppressed until the vestibular inputs had subsided. We conclude that signals associated with OKN, are fully available to the mechanism that assesses the headcentric motion of objects but that signals associated with VOR are only partially available to that mechanism.

Up-down asymmetry in human vertical optokinetic nystagmus and afternystagmus: contributions of the central and peripheral retinae.

The vertical optokinetic nystagmus (OKN) of 10 normal subjects and the optokinetic afternystagmus (OKAN) of 3 subjects were measured with the magnetic search coil technique. In order to assess the relative contributions of various retinal areas to the up-down asymmetry in OKN the central and peripheral visual fields were selectively stimulated in four OKN conditions. In the full-field OKN condition the stimulus was a 61 degrees x 64 degrees display of moving random-dots. Overall, full-field OKN gains elicited by upward motion were significantly higher than those elicited by downward motion at stimulus velocities between 30 and 70 degrees/s. In the periphery-only OKN condition a 3 degrees or 6 degrees-wide vertical band occluded the center of the full-field display. Nine of the 10 subjects displayed OKN in this condition. For 6 subjects, the addition of the 6 degrees band to the full field resulted in an increase in the up-down asymmetry at stimulus velocities above 30 degrees/s. For the other three subjects there was a decline in the gains of both upward and downward OKN when the 3 degrees or 6 degrees band was present; the result was directionally symmetric OKN gains. In the central-strip OKN condition only a 6 degrees-wide central vertical strip of moving dots was visible. The gains of central-strip OKN were not significantly different from the full-field responses. A servo controlled centrally-located 10 degrees x 6 degrees moving display was used in the center-only OKN condition. In this condition both upward and downward gains were attenuated and there was no up-down asymmetry. OKAN was measured following a 50-s exposure to either the full-field or center-only OKN display. The stimulus velocity was 30 degrees/s. After viewing the full-field display the 3 subjects displayed OKAN with slow phases upward following upward OKN but there was no downward OKAN following downward OKN. In contrast, there was no consistent directional asymmetry following exposure to the center-only display. The disappearance of the upward preponderance in OKN and OKAN with occlusion of the peripheral retina suggests that the directional asymmetry in vertical OKN exists in the slow OKN system.

Anisotropy in the chromatic channel: a horizontal-vertical effect.

We compared the chromatic contrast thresholds of drifting (2Hz) red-green sine-wave gratings of horizontal, vertical, and two oblique orientations at three spatial frequencies (2, 4, 8 cpd). Luminance contrast thresholds for yellow-black gratings were also obtained. The classic oblique effect was found for high spatial frequency luminance and chromatic stimuli. For chromatic thresholds, a significant difference was found between the horizontal and vertical thresholds of all observers. One observer was retested with her head tilted 45 deg and demonstrated that the anisotropy was specific to retinal coordinates. These results give evidence for orientation selectivity in the chromatic channel which is at least partially independent of that in the luminance channel. We estimated the degree of lateral chromatic aberration in our observers' eyes and discuss the possible contribution of this aberration to the horizontal-vertical difference in the chromatic channel.

Optokinetic nystagmus: the effects of stationary edges, alone and in combination with central occlusion.

In Experiment 1 we investigated the independent and combined effects of horizontal OKN of stationary edges and occlusion of the central retina. For a display 60 degrees wide moving at 30 degrees/sec a symmetrically placed pair of vertical nonoccluding bars suppressed OKN when near the center of the display but had no effect when 30 degrees apart. A 7 degrees-high 60 degrees-wide central occluder reduced OKN gain by 37%. However, a central occluder with edges only 30 degrees wide abolished OKN. In Experiment 2 this interaction between central occlusion and stationary edges was confirmed with a wider display over a range of stimulus velocities and configurations. A functional explanation of this interaction is presented.