Intersaccadic Interval Research Articles

Binocular quantification and characterization of microsaccades.

The significance of microsaccades in the visual process has been discussed for more than 50 years. However, only a few studies have measured microsaccades binocularly, and detailed quantification and characterization of these small movements are needed in order to further understand their nature. The amplitude, velocity, acceleration and direction of microsaccades were quantified binocularly in 10 normal test persons during a 40-s fixation task, using an infrared recording technique. All microsaccades for all test persons were performed simultaneously and individually with an almost identical amplitude in the right and left eye (a range of 0.003-0.042 deg between right and left eye mean values). The mean microsaccadic amplitude for the test persons was within a range of 0.223-1.079 deg. The directional difference between simultaneously-performed right and left eye microsaccades was less than 22.5 deg for 84.8% of the saccades, indicating that the majority of microsaccades are conjugated. Three different fixation patterns were identified and characterized: (1) a classic interplay between easily identified drifts and medium-sized microsaccades (mean amplitude range 0.328-0.413 deg); (2) long intersaccadic intervals (4-5 s) with almost absent drifts, followed by three or four large microsaccades (mean amplitude range 0.755-1.079 deg); and (3) low-amplitude drift movements interrupted by low-amplitude microsaccades (mean amplitude range 0.231-0.265 deg). Microsaccades are involuntary, predominantly conjugated, simultaneously performed, and of almost identical amplitude in the right and left eye, suggesting a central control mechanism for microsaccades at subcortical level.

Superior colliculus activity related to concurrent processing of saccade goals in a visual search task.

Saccades are typically separated by inter-saccadic fixation intervals (ISFIs) of > or =125 ms. During this time, the saccadic system selects a goal and completes the preparatory processes required prior to executing the subsequent movement. However, in tasks in which competing stimuli are presented, two sequentially executed movements to different goals can be separated by much shorter ISFIs. This suggests that the saccadic system is capable of completing many of the preparatory requirements for a second saccade concurrently with the execution of an initial movement. We recorded single neurons in the superior colliculus (SC) during rapid saccade sequences made by rhesus monkeys performing a search task. We found that during the execution of an initial saccade, activity related to the goal of a quickly following second saccade can be simultaneously maintained in the SC motor map. This activity appears to signal the selection or increased salience of the second saccade goal even before the initial saccade has ended. For movements separated by normal ISFIs (> or =125 ms), we did not observe activity related to concurrent processing, presumably because for these longer ISFI responses, the goal of the second saccade is not selected until after the end of the first saccade. These results indicate that, at the time of an initial saccade, the SC does not necessarily act as a strict winner-take-all network. Rather it appears that the salience of a second visual goal can be simultaneously maintained in the SC. This provides evidence that selection or preparatory activity related to the goal of a second saccade can overlap temporally with activity related to an initial saccade and indicates that such concurrent processing is present even in a structure which is fairly close to the motor output.

Neural discharge in the superior colliculus during target search paradigms.

Neural studies of oculomotor function in the past have been conducted with the use of very simple visual stimuli. More recently there has been a new emphasis on using more natural stimuli to extend our knowledge of oculomotor organization. Visual search paradigms are an example of the use of these more natural visual surrounds. In search a subject must locate and saccade to a target that appears simultaneously with an array of distractors. When monkeys are used in this paradigm, it is possible to record from neurons located in various central structures in the brain while the initial visual response, subsequent discrimination processes and final saccadic movement unfold. In the present study we used an array of four visual stimuli, and the target was distinguished by its odd color from three distractors of uniform color. Location of the target within the array and its color were randomly selected on each trial. Neurons located in the deeper layers of the superior colliculus (SC) were recorded by standard methods in blocks of search trials. We found several new features in the discharge of SC neurons using this search paradigm that have not previously been reported in studies using single-target visual displays. (1) In contrast to the "winner take all" behavior previously reported for the SC, we found evidence of concurrent processing of alternative movement vectors. When incorrect movements were made to distractor locations, this concurrent activity was associated with significantly shorter intersaccadic intervals. (2) The discharge profile of the visual response in many units was modified by the appearance of a second prominent peak which followed the initial phasic visual response, but which was clearly differentiated from a third burst in activity associated with a saccade into the cell's response field. In some neurons, the activity in this second peak was discriminatory for the impending saccade vector. That is, it was larger when the target appeared in the response field of the cell than when it contained a distractor. This target selection signal was thus distinct from the burst normally associated with saccades into the movement fields of SC neurons. (3) Some saccades in search had a curved trajectory bowing toward the location of a distractor. These saccades were accompanied by an elevated discharge of neurons coding that distractor.

Visual scene memory and the guidance of saccadic eye movements

An unresolved question is how much information can be remembered from visual scenes when they are inspected by saccadic eye movements. Subjects used saccadic eye movements to scan a computer-generated scene, and afterwards, recalled as many objects as they could. Scene memory was quite good: it improved with display duration, it persisted over time long after the display was removed, and it continued to accumulate with additional viewings of the same display (Melcher, D. (2001) The persistance of memory for scenes. Nature 412, 401). The occurrence of saccadic eye movements was important to ensure good recall performance, even though subjects often recalled non-fixated objects. Inter-saccadic intervals increased with display duration, showing an influence of duration on global scanning strategy. The choice of saccadic target was predicted by a Random Selection with Distance Weighting (RSDW) model, in which the target for each saccade is selected at random from all available objects, weighted according to distance from fixation, regardless of which objects had previously been fixated. The results show that the visual memory that was reflected in the recall reports was not utilized for the immediate decision about where to look in the scene. Visual memory can be excellent, but it is not always reflected in oculomotor measures, perhaps because the cost of rapid on-line memory retrieval is too great.

Opsoclonus in three dimensions: oculographic, neuropathologic and modelling correlates

Opsoclonus is a dyskinesia consisting of involuntary, arrhythmic, chaotic, multidirectional saccades, without intersaccadic intervals. We used a magnetic scleral search coil technique to study opsoclonus in two patients with paraneoplastic complications of lung carcinoma. Eye movement recordings provided evidence that opsoclonus is a three-dimensional oscillation, consisting of torsional, horizontal, and vertical components. Torsional nystagmus was also present in one patient. Antineuronal antibody study revealed the presence of anti-Ta (Ma2 onco-neuronal antigen) antibodies in one patient, which had previously been associated only with paraneoplastic limbic encephalitis and brainstem dysfunction, but not opsoclonus, and only in patients with testicular or breast cancer. Neuropathologic examination revealed mild paraneoplastic encephalitis. Normal neurons identified in the nucleus raphe interpositus (rip) do not support postulated dysfunction of omnipause cells in the pathogenesis of opsoclonus. Computer simulation of a model of the saccadic system indicated that disinhibition of the oculomotor region of the fastigial nucleus (FOR) in the cerebellum can generate opsoclonus. Histopathological examination revealed inflammation and gliosis in the fastigial nucleus. This morphological finding is consistent with, but not necessary to confirm, damage to afferent projections to the FOR, as determined by the model. Malfunction of Purkinje cells in the dorsal vermis, which inhibit the FOR, may cause opsoclonus by disinhibiting it.

Selective activation of visual cortex neurons by fixational eye movements: implications for neural coding.

During normal vision, when subjects attempt to fix their gaze on a small stimulus feature, small fixational eye movements persist. We have recorded the impulse activity of single neurons in primary visual cortex (V1) of macaque monkeys while their fixational eye movements moved the receptive-field activating region (AR) over and around a stationary stimulus. Three types of eye movement activation were found. (1) Saccade cells discharged when a fixational saccade moved the AR onto the stimulus, off the stimulus, or across the stimulus. (2) Position/drift cells discharged during the intersaccadic (drift) intervals and were not activated by saccades that swept the AR across the stimulus without remaining on it. To activate these neurons, it was essential that the AR be placed on the stimulus and many of these cells were selective for the sign of contrast. They had smaller ARs than the other cell types. (3) Mixed cells fired bursts of activity immediately following saccades and continued to fire at a lower rate during intersaccadic intervals. The tendency of each neuron to fire transient bursts or sustained trains of impulses following saccades was strongly correlated with the transiency of its response to stationary flashed stimuli. For one monkey, an extraretinal influence accompanying fixational saccades was identified. During natural viewing, the different eye movement classes probably make different contributions to visual processing. Position/drift neurons are well suited for coding spatial details of the visual scene because of their small AR size and their selectivity for sign of contrast and retinal position. However, saccade neurons transmit information that is ambiguous with respect to the spatial details of the retinal image because they are activated whether the AR lands on a stimulus contour, or the AR leaves or crosses the contour and lands in another location. Saccade neurons may be involved in constructing a stable world in spite of incessant retinal image motion, as well as in suppressing potentially confusing input associated with saccades.

Short-term priming, concurrent processing, and saccade curvature during a target selection task in the monkey

In human subjects, two mechanisms for improving the efficiency of saccades in visual search have recently been described: color priming and concurrent processing of two saccades. Since the monkey provides an important model for understanding the neural underpinnings of target selection in visual search, we sought to explore the degree to which the saccadic system of monkeys uses these same mechanisms. Therefore, we recorded the eye movements of rhesus monkeys performing a simple color-oddity pop-out search task, similar to that used previously with human subjects. The monkeys were rewarded for making a saccade to the odd-colored target, which was presented with an array of three distractors. The target and distractors were randomly chosen to be red or green in each trial. Similar to what was previously observed for humans, we found that monkeys show the influence of a cumulative, short-term priming mechanism which facilitates saccades when the color of the search target happens to repeat from trial to trial. Furthermore, we found that like humans, when monkeys make an erroneous initial saccade to a distractor, they are capable of executing a second saccade to the target after a very brief inter-saccadic interval, suggesting that the two saccades have been programmed concurrently (i.e. in parallel). These results demonstrate a close similarity between human and monkey performance. We also made a new observation: we found that when monkeys make such two-saccade responses, the trajectory of the initial saccade tends to curve toward the goal of the subsequent saccade. This provides evidence that the two saccade goals are simultaneously represented on a common motor map, supporting the idea that the movements are processed concurrently. It also indicates that concurrent processing is not limited to brain areas involved in higher-level planning; rather, such parallel programming apparently occurs at a low enough level in the saccadic system that it can affect saccade trajectory.

The ability to execute saccades on the basis of efference copy: impairments in double-step saccade performance in children with developmental co-ordination disorder.

The double-step saccade task (DSST) was used to test the hypothesis that children with developmental co-ordination disorder (DCD) who experience deficits in motor imagery have difficulty processing the visual spatial consequences of intended movements using efference copy signals. In order to ensure that the second saccade in the DSST was executed in the absence of visual cues and had to be programmed on the basis of extra-retinal information (efference copy), we analysed only those double-step ensembles where latency plus duration of first saccades was greater than 240 ms (total presentation time of the targets). No significant differences between DCD and control children were evident on measures of latency of first saccades, intersaccadic interval and first saccade error. As predicted, children with DCD who have impaired motor imagery demonstrated specific deficits on the DSST where efference copy had been used to program the saccade sequence. More specifically, these children were less accurate in terms of final eye position on second saccades. Our results raise the possibility that abnormalities in the processing of efference copy signals could underlie motor clumsiness in the majority of children with DCD. Furthermore, the origin of this deficit in efference copy probably exists at the level of the parietal lobe.

Concurrent processing of saccades in visual search

We provide evidence that the saccadic system can simultaneously program two saccades to different goals. We presented subjects with simple visual search displays in which they were required to make a saccade to an odd-colored target embedded in an array of distractors. When there was strong competition between target and distractor stimuli (due to color priming from previous trials), subjects were more likely to make a saccade to a distractor. Such error saccades were often followed, after a very short inter-saccadic interval (∼10–100 ms), by a second saccade to the target. The brevity of these inter-saccadic intervals suggests that the programming of the two saccades (one to a distractor and one to the target) overlapped in time. Using a saccade-contingent change in the search display, we show that new visual information presented during the initial saccade does not change the goal of the second saccade. This supports the idea that, by the end of the first saccade, programming of the second saccade is already well underway. We also elicited two-saccade responses (similar to those seen in search) using a double-step task, with the first saccade directed to the initial target step and the second saccade directed to the second target step. If the two saccades are programmed in parallel and programming of each saccade is triggered by one of the two target steps, the second saccade should occur at a relatively fixed time after the onset of the second target step, regardless of the timing of the initial saccade. This prediction was confirmed, supporting the idea that the two saccades are programmed in parallel. Finally, we observed that the shortest inter-saccadic intervals typically followed hypometric initial saccades, suggesting that the initial saccade may have been interrupted by the impending second saccade. Using predictions from physiological studies of interrupted saccades, we tested this hypothesis and found that the hypometric initial saccades did not appear to be interrupted in mid-flight. We discuss the significance of our findings for models of the saccadic system.

Age dependence of sensory and motor components in saccades

Humans usually perform about 3-4 saccades per second; hence, precision as well as latency and velocity of these fast eye movements are of crucial importance for analyzing complex and fast changing visual scenes, for example in traffic. Since visual performance is known to slowly decline with age, we investigated the effect of age on the most important characteristics of visually guided saccades. This investigation on age dependence of visually guided saccades included, for the first time, the gap condition, where the old fixation point disappears before the new one appears, allowing subjects to prepare for a saccade. Saccadic latencies and intersaccadic intervals increased with age, while their peak velocities and gains decreased, especially for threshold-adjusted luminance of the targets. The gap condition, however, improved reaction times (latencies) most pronounced in the older age groups, bringing performance for these groups close to the performance level of the young groups. This indicates that the slowing down of saccadic reaction times with age is not predominantly a motor problem, but, according to a common interpretation of the gap effect, a problem of disengaging attention from the old fixation spot.

Concurrent processing of saccades

We summarize several experiments indicating that the saccadic system is capable of simultaneously programming two movements toward different goals. This concurrent processing of saccades can lead to the execution of two saccades separated by an extremely short intersaccadic interval. This supports the idea of target competition proposed in Findlay & Walker's article, but suggests a greater degree of parallel processing. We provide evidence that concurrent processing of two saccades is not limited to higher-level planning subsystems; rather, it also involves both regions close enough to the motor output that it can systematically affect saccade trajectory.

Primate antisaccades. I. Behavioral characteristics.

The antisaccade task requires a subject to make a saccade to an unmarked location opposite to a flashed stimulus. This task was originally designed to study saccades made to a goal specified by instructions. Interest for this paradigm surged after the discovery that frontal lobe lesions specifically and severely affect human performance of antisaccades while prosaccades (i.e., saccades directed to the visual stimulus) are facilitated. Training monkeys to perform antisaccades was rarely attempted in the past, and this study is the first one that describes in detail the properties of such antisaccades compared with randomly intermingled prosaccades of varying amplitude in all directions. Such randomization was found essential to force the monkeys to use the instruction cue (pro- or anti-) and the location cue (peripheral stimulus) provided within a trial rather than to direct their saccades to the location of past rewards. Each trial began with the onset of a central fixation target that conveyed by its shape the instruction to make a pro- or an antisaccade to a subsequent peripheral stimulus. In one version of the task, the monkey was allowed to make an immediate saccade to the goal; in a second version, the saccade had to wait for a go signal. Analyses of the accuracy, velocity, and latency of antisaccades compared with prosaccades were performed on a sample of 7,430 pro-/antisaccades in the "immediate saccade" task (delayed saccades suffering from known distortions). Error rates fluctuated approximately 25%. Results were the same for the two monkeys with respect to accuracy and velocity, but they differed in terms of reaction time. Both monkeys generated antisaccades to stimuli in all directions, at various eccentricities, but antisaccades were significantly less accurate and slower than prosaccades elicited by the same stimuli. Interestingly, saccades to the stimulus could be followed by appropriate antisaccades with no intersaccadic interval. Such instances are here referred to as "turnaround saccades." Because they occurred sometimes with a long latency, turnaround saccades did not simply reflect the cancellation of an early foveation reflex. Consistent with human data, latencies of one monkey were longer for antisaccades than for prosaccades, but the reverse was true for the other monkey who was trained differently. In summary, this study demonstrates the feasibility of providing a subhuman primate model of antisaccade performance, but at the same time it suggests some irreducible differences between human and monkey performance.

Brain stem omnipause neurons and the control of combined eye-head gaze saccades in the alert cat.

When the head is unrestrained, rapid displacements of the visual axis-gaze shifts (eye-re-space)-are made by coordinated movements of the eyes (eye-re-head) and head (head-re-space). To address the problem of the neural control of gaze shifts, we studied and contrasted the discharges of omnipause neurons (OPNs) during a variety of combined eye-head gaze shifts and head-fixed eye saccades executed by alert cats. OPNs discharged tonically during intersaccadic intervals and at a reduced level during slow perisaccadic gaze movements sometimes accompanying saccades. Their activity ceased for the duration of the saccadic gaze shifts the animal executed, either by head-fixed eye saccades alone or by combined eye-head movements. This was true for all types of gaze shifts studied: active movements to visual targets; passive movements induced by whole-body rotation or by head rotation about stationary body; and electrically evoked movements by stimulation of the caudal part of the superior colliculus (SC), a central structure for gaze control. For combined eye-head gaze shifts, the OPN pause was therefore not correlated to the eye-in-head trajectory. For instance, in active gaze movements, the end of the pause was better correlated with the gaze end than with either the eye saccade end or the time of eye counterrotation. The hypothesis that cat OPNs participate in controlling gaze shifts is supported by these results, and also by the observation that the movements of both the eyes and the head were transiently interrupted by stimulation of OPNs during gaze shifts. However, we found that the OPN pause could be dissociated from the gaze-motor-error signal producing the gaze shift. First, OPNs resumed discharging when perturbation of head motion briefly interrupted a gaze shift before its intended amplitude was attained. Second, stimulation of caudal SC sites in head-free cat elicited large head-free gaze shifts consistent with the creation of a large gaze-motor-error signal. However, stimulation of the same sites in head-fixed cat produced small "goal-directed" eye saccades, and OPNs paused only for the duration of the latter; neither a pause nor an eye movement occurred when the same stimulation was applied with the eyes at the goal location. We conclude that OPNs can be controlled by neither a simple eye control system nor an absolute gaze control system. Our data cannot be accounted for by existing models describing the control of combined eye-head gaze shifts and therefore put new constraints on future models, which will have to incorporate all the various signals that act synergistically to control gaze shifts.

Effects of pre-cues on voluntary and reflexive saccade generation. II. Pro-cues for anti-saccades.

The reaction times of saccades (SRT) to a suddenly presented visual stimulus (pro-saccade) can be decreased and a separate mode of express saccades can occur when a gap paradigm is used (i.e. fixation-point offset precedes target onset by 200 ms). A valid peripheral cue, presented briefly (100 ms) before target onset, has been found to facilitate the generation of saccades to the target, thereby increasing the frequency of express saccades and decreasing the mean latency. This facilitation occurs only for cues that correctly indicate the direction of the subsequent target presentation (valid cues). The present study investigates the effects of valid cues on SRTs and error rate in the anti-saccade task (saccades in the direction opposite to the stimulus) by systematically varying the cue lead time (CLT) and using the gap and overlap conditions, i.e. fixation point remains on throughout the trial. For a CLT of 100 ms, both reaction times and error rates were significantly increased. With increasing CLT (200-500 ms), both the reaction times of the anti-saccades and the error rates returned to approximately control level, with CLT more than 200 ms in both the gap and the overlap condition. Additional experiments using non-informative cues in the overlap task showed that the reaction times of correct anti-saccades and the error rate were decreased when cue and stimulus appeared at the same side. Analysis of the erratic pro-saccades revealed that almost all of them were corrected, i.e. they were followed by a second saccade towards the required location. It is found that the correction times were usually very short, with intersaccadic intervals between 0 and 150 ms. We suggest that the orienting mechanism, elicited by a transient peripheral cue, relates to the command and the decision to make a pro- rather than an anti-saccade. The cue elicits pro-orienting towards its position when a pro-saccade is required, and anti-orienting when an anti-saccade is required. The orienting effect is transient and decays with CLTs of more than 200 ms; this result holds for both anti-saccades and pro-saccades. Since subjects reported that they could not prevent the erratic pro-saccades or were often not aware of them, we conclude that this orienting mechanism occurs automatically, beyond voluntary control.

Saccade Automation

The comparative analysis of basic saccade characteristics (interval, amplitude, and coefficient of asymmetry) has shown considerable similarities in adults and first-year babies. Distribution curves of intersaccadic intervals under different test conditions all have only one maximum in the range of about 0.4 s, and in all cases the most frequently met intersaccadic intervals (0.1 – 1.0) comprised comparable proportions of all (71.5% – 90.5%). The concept of saccade automation was formulated on the basis of these data (Filin and Filina, 1989 Zhurnal Vysshei Nervnoi Deyatelnosti29 603 – 607). In our opinion saccade automation is the basic law of saccadic activity, and all diversity of eye-movement activity takes place against the background of saccade automation. Microsaccades and macrosaccades, nystagmus, and rapid eye movements during sleep may be seen as special cases of saccade automation. Saccade automation is conditioned by the activity of brain structures with pacemaker function. Thus saccades are driven at one basic frequency, which is modulated by afferent influences from the retina of vestibular apparatus, proprioreceptors of eye and neck muscles, and efferent signals (forehead and occipital sections of cerebral cortex, cerebellum). At the same time, only one parameter is modulated at one particular moment, for example saccade amplitude in which case interval and orientation are given in a ready form. Saccade automation has a great functional importance. It increases the scanned area tenfold, it provides compensation of defects in the sensors and the impellent eye apparatus and for the deletion of consecutive images, maintaining the continuity of visual perception. Moving the image on the retina increases the information of the visual channel.

Saccade Mechanism Based on Processes for Depth Estimation and Incongruity Detection between Binocular Retinal Images – Analysis of Gazing Positions and Inter-saccade Intervals –

We analyzed binocular gazing positions in the existence of a rim occlusion. When a human gazes at the rim of a textured cylinder, the gazing position shifts from the fixation target, When a human gazes at the rim of a cylinder whose upper-surface is visible, the gazing points are located at the estimated surface's depth. In this paper, to investigate the mechanism changing the gazing point, we analyzed inter-saccade intervals. The result shows that an occlusion detection mechanism exists at an early stage of human visual processing, and the mechanism causes a shorter-interval saccade when incongruity regions between left and right retinal images are detected. We propose a saccade mechanism which includes processes for depth estimation and incongruity detection between binocular retinal images based on relationships between the gazing positions and the results of inter-saccade intervals.

Interaction of extraretinal eye position signals in a double-step saccade task: psychophysical estimation.

The time course of extraretinal eye position signals (EEPSs) for visually guided saccades made successively with a short intersaccadic interval was estimated on the basis of perceptual errors in localizing a visual target flashed between the two saccades. The EEPSs for the first and the second saccades were shown to interact in a specific way when the intersaccadic interval was short. The pattern of interaction depended on the direction of the second saccade. It is suggested that when the second saccade was made in the opposite direction to the first saccade, the EEPS for the first saccade was interrupted before its completion in preparation for the onset of the second saccade. When the two saccades were made in the same direction, the EEPS for the first saccade developed more quickly than in a single-saccade condition. The results are discussed in relation to the findings of recent neurophysiological studies.

The Thalamic Intralaminar Nuclei: A Role in Visual Awareness

We briefly review the anatomy and functional properties of the intralaminar nuclei (ILN) of the thalamus and the neurological disorders associated with their dysfunction. The ILN project over a wide range of cortical territories and are connected to several subcortical structures that place the ILN within the distributed networks underlying arousal, attention, working memory, and gaze control. The temporal structure of the spike discharges of ILN neurons can be controlled by levels of arousal and visuomotor behavior. Taken together, the anatomy, cellular physiology, and clinical data suggest that in the state of wakefulness, the ILN neurons promote the formation of an “event-holding” function in the cortex. In the prefrontal cortex, this function facilitates the storage of target location in working memory. In the frontal eye fields, the function produces sustained activation that anticipates the onset of intended eye movements. In the posterior parietal cortex, the sustained activation can be boosted at the start of the intersaccadic interval and can operate as an attentional gate regulating the flow of information back to the prefrontal cortex. The attentional gate is of limited capacity, as is working memory, and is best utilized during the intersaccadic interval. The ILN may help to synchronize the eye movement commands of the frontal eye fields with the episodic dynamics of the attentional gate and working memory.

Analysis of the step response of the saccadic feedback: system behavior.

We used prolonged stimulation of the monkey superior colliculus to elicit staircase eye movements. By changing the parameters of the stimulating current we were able to obtain movements with different dynamics. An increase in the current frequency resulted in the shortening of the intersaccadic interval and a decrease of the amplitudes in the staircase. In cases of high stimulation, after an initial saccade of fixed metrics, the eyes moved in an apparently smooth fashion. The movement was conjugate and in the same direction as the first saccade. By analyzing the velocity trace we found that the movement consisted of a chain of small saccades, each of which started before the previous one ended. We conducted a quantitative analysis of the staircase movements including the cases of apparently smooth movement of the eyes. We conclude that all of the movements elicited by prolonged SC stimulation were generated by the saccadic feedback circuitry. The dynamic profiles of the elicited movements changed continuously with the stimulating current parameters. On one end of the continuum we observed the classically, described staircase movements with individual movements separated in time. On the other end of the continuum we saw the apparent smooth movement as the limit case produced by high stimulation of the SC.

Monosynaptic activation of medium-lead burst neurons from the superior colliculus in the alert cat.

1. Extracellular recordings were made from medium-lead burst neurons (MLBNs) in the paramedian pontomedullary reticular formation rostral and caudal to the abducens nucleus in the alert cat. 2. Single-pulse stimulation of the contralateral superior colliculus during intersaccadic intervals evoked no response in most MLBNs. When collicular stimulation was applied at the beginning of saccades, spikes of MLBNs were consistently evoked with short latencies. The shortest latency was 0.8 ms, indicating monosynaptic activation of MLBNs from the superior colliculus. 3. Results suggest that monosynaptic excitatory effects from the colliculus are concealed by inhibitory input from omnipause neurons (OPNs) during intersaccadic intervals and that the monosynaptic collicular activation is disclosed when this inhibition is removed by a pause in OPN activity at the beginning of saccades.