Neuronal responses in macaque area PEc to saccades and eye position

Neurons in area PEc in the superior parietal cortex encode signals from different modalities, such as visual, extraretinal and somatosensory, probably combining them to encode spatial parameter of extrapersonal space to prepare body movements. This study reports the characterization of the functional properties of PEc non-visual neurons that showed saccade-related activity. We analyzed the pre- and post-saccadic firing activity in 189 neurons recorded in five hemispheres of three behaving monkeys. Spiking activity of PEc single neurons was recorded while the monkeys performed visually-guided saccades in a reaction time task. We found that 84% of neurons recorded from area PEc showed pre-saccadic activity with directional tuning. In 26% of neurons, we found inhibition of activity in the pre-saccadic period. The onset of this "pause" always started before the saccade and, in 51% of neurons, it was invariant among different gaze directions. The post-saccadic activity in these cells was either a phasic response with directional tuning (77%) and/or an eye position tuning (75%). The analysis of the preferred direction did not show hemispheric preference, however, for the majority of neurons, the angular difference in the preferred direction, in the pre- and post-saccadic period, was more than 60 degrees . By confirming, therefore, that PEc neurons carry information about eye position, these novel findings open new horizons on PEc function that, to date, is not well documented. The pre-saccadic activity may reflect an involvement in saccade control, whereas post-saccadic activity may indicate a role in informing on the new eye position. These novel results about saccade and eye position processing may imply a role of area PEc in gaze direction mechanisms and, possibly, in remapping visual space after eye movements.     

The parietal reach region codes the next planned movement in a sequential reach task

Distinct subregions of the posterior parietal cortex contribute to planning different movements. The parietal reach region (PRR) is active during the delay period of a memory-guided reach task but generally not active during a memory-guided saccade task. We explored whether the reach planning activity in PRR is related to remembering targets for reaches or if it is related to specifying the reach that the monkey is about to perform. Monkeys were required to remember two target locations and then reach to them in sequence. Before the movements were executed, PRR neurons predominantly represented the reach about to be performed and only rarely represented the remembered target for the second reach. This indicates the area plays a role in specifying the target for the impending reach and may not contribute to storing the memory of subsequent reach targets.     

Auditory signals evolve from hybrid- to eye-centered coordinates in the primate superior colliculus

Visual and auditory spatial signals initially arise in different reference frames. It has been postulated that auditory signals are translated from a head-centered to an eye-centered frame of reference compatible with the visual spatial maps, but, to date, only various forms of hybrid reference frames for sound have been identified. Here, we show that the auditory representation of space in the superior colliculus involves a hybrid reference frame immediately after the sound onset but evolves to become predominantly eye centered, and more similar to the visual representation, by the time of a saccade to that sound. Specifically, during the first 500 ms after the sound onset, auditory response patterns (N = 103) were usually neither head nor eye centered: 64% of neurons showed such a hybrid pattern, whereas 29% were more eye centered and 8% were more head centered. This differed from the pattern observed for visual targets (N = 156): 86% were eye centered, <1% were head centered, and only 13% exhibited a hybrid of both reference frames. For auditory-evoked activity observed within 20 ms of the saccade (N = 154), the proportion of eye-centered response patterns increased to 69%, whereas the hybrid and head-centered response patterns dropped to 30% and <1%, respectively. This pattern approached, although did not quite reach, that observed for saccade-related activity for visual targets: 89% were eye centered, 11% were hybrid, and <1% were head centered (N = 162). The plainly eye-centered visual response patterns and predominantly eye-centered auditory motor response patterns lie in marked contrast to our previous study of the intraparietal cortex, where both visual and auditory sensory and motor-related activity used a predominantly hybrid reference frame (Mullette-Gillman et al. 2005, 2009). Our present findings indicate that auditory signals are ultimately translated into a reference frame roughly similar to that used for vision, but suggest that such signals might emerge only in motor areas responsible for directing gaze to visual and auditory stimuli.     

A quantitative analysis of the correlations between eye movements and neural activity in the pretectum

The study of the saccadic system has focused mainly on neurons active before the beginning of saccades, in order to determine their contribution in movement planning and execution. However, most oculomotor structures contain also neurons whose activity starts only after the onset of saccades, the maximum of their activity sometimes occurring near saccade end. Their characteristics are still largely unknown. We investigated pretectal neurons with saccade-related activity in the alert cat during eye movements towards a moving target. They emitted a high-frequency burst of action potentials after the onset of saccades, irrespective of their direction, and will be referred to as "pretectal saccade-related neurons". The delay between saccade onset and cell activity varied from 17 to 66 ms on average. We found that burst parameters were correlated with the parameters of saccades; the peak eye velocity was correlated with the peak of the spike density function, the saccade amplitude with the number of spikes in the burst, and burst duration increased with saccade duration. The activity of six pretectal saccade-related neurons was studied during smooth pursuit at different velocities. A correlation was found between smooth pursuit velocity and mean firing rate. A minority of these neurons (2/6) were also visually responsive. Their visual activity was proportional to the difference between eye and target velocity during smooth pursuit (retinal slip). These results indicate that the activity of pretectal saccade-related neurons is correlated with the characteristics of eye movements. This finding is in agreement with the known anatomical projections from premotor regions of the saccadic system to the pretectum.     

Distribution of eye- and arm-movement-related neuronal activity in the SEF and in the SMA and Pre-SMA of monkeys

We analyzed neuronal activity in the supplementary eye field (SEF), supplementary motor area (SMA), and presupplementary motor area (pre-SMA) during the performance of three motor tasks: capturing a visual target with a saccade, reaching one arm to a target while gazing at a visual fixation point, or capturing a target with a saccade and arm-reach together. Our data demonstrated that each area was involved in controlling the arm and eye movements in a different manner. Saccade-related neurons were found mainly in the SEF. In contrast, arm-movement-related neurons were found primarily in the SMA and pre-SMA. In addition, we found that the activity of both arm-movement- and saccade-related neurons differed depending on the presence or absence of an accompanying saccade or arm movement. Such context dependency was found in all three areas. We also discovered that activity preceding eye or arm movement alone, and eye and arm movement combined, appeared more often in the pre-SMA and SEF, suggesting their involvement in effector-independent aspects of motor behavior. Subsequent analysis revealed that the laterality of arm representation differed in the three areas: it was predominantly contralateral in the SMA but largely bilateral in the pre-SMA and SEF.     

Properties of spike train spectra in two parietal reach areas

In the lateral intraparietal area (LIP), a saccade-related region of the posterior parietal cortex (PPC), spiking activity recorded during the memory period of an instructed-delay task exhibits temporal structure that is spatially tuned. These results provide evidence for the existence of 'dynamic memory fields' which can be read-out by other brain areas, along with information contained in the mean firing rate, to give the direction of a planned movement. We looked for evidence of dynamic memory fields in spiking activity in two parietal reach areas, the parietal reach region (PRR) and area 5. Monkeys made center-out reaches to eight target locations in an instructed-delay task with a memory component. Neurons in both areas exhibited sustained activity during the delay period that was spatially tuned. Many single cell PRR spectra exhibited spatially tuned temporal structure, as evidenced by a significant and spatially tuned peak in the 20–50 Hz band. The PRR population spectrum of spike trains was also tuned, with the peak power centered on approximately 25 Hz. In contrast, area 5 spiking activity did not exhibit any significant temporal structure. These results suggest that different mechanisms underlie sustained delay period activity in these two areas and that dynamic memory fields, as revealed by our techniques, are more prominent in PRR than in area 5. Temporal structure in the spike train and local field potential (LFP) are related in at least one other brain area (LIP). The present results suggest then that LFP activity obtained from PRR may be better suited than area 5 LFP activity for use in neural prosthetic systems that incorporate analysis of temporal structure as part of a decode mechanism for extracting intended movement goals.     

The coincidence between late non-phase-locked gamma synchronization response and saccadic eye movements.

The event-related response in the gamma (30-45 Hz) frequency band was studied in healthy subjects (n=45) viewing sequentially presented pictures from the International Affective Picture System. The distinct non-phase-locked gamma response was obtained in characteristic time window (200-400 ms) with clear-cut centro-parietal location. The strong coincidence between induced gamma oscillations and saccadic eye movements was revealed. We suggest that saccade-related gamma increase is another manifestation of the phenomenon known as presaccadic spike potential, which is commonly registered over parietal scalp leads at 10-20 ms prior to saccade onset. It is hypothesized that late non-phase-locked gamma synchronization mainly reflects activity of a system responsible for attentional tuning and motor planning/execution of saccadic eye movements.     

Eye-hand coordination: saccades are faster when accompanied by a coordinated arm movement

When primates reach for an object, they very often direct an eye movement toward the object as well. This pattern of directing both eye and limb movements to the same object appears to be fundamental to eye-hand coordination. We investigated interactions between saccades and reaching movements in a rhesus monkey model system. The amplitude and peak velocity of isolated eye movements are positively correlated with one another. This relationship is called the main sequence. We now report that the main sequence relationship for saccades is changed during coordinated eye and arm movements. In particular, peak eye velocity is approximately 4% faster for the same size saccade when the saccade is accompanied by a coordinated arm movement. Saccade duration is reduced by an equivalent amount. The main sequence relationship is unperturbed when the arm moves simultaneously but in the opposite direction as the eyes, suggesting that eye and arm movements must be tightly coordinated to produce the effect. Candidate areas mediating this interaction include the posterior parietal cortex and the superior colliculus.     

Saccade-related Purkinje cells in the cerebellar hemispheres of the monkey

Summary Extracellular single unit discharges of cerebellar Purkinje cells (P-cells) were recorded from the cerebellar hemispheres of two Japanese monkeys (Macaca fuscata) during spontaneous and visually guided eye movements. We found that saccade-related P-cells, whose simple-spike (SS) discharge rates were modulated in close correlation with saccadic eye movements, were localized in fairly restricted areas in the hemisphere, mostly in Crus IIa with some in the deep folia of Crus I. P-cells located in simple lobules, superficial folia of Crus I or in Crus IIp did not change their discharge rate during voluntary eye movements. Fifty-five saccade-related P-cells recorded from Crus I and II showed modulation of SS discharge rate related to both spontaneous and visually triggered saccades, with the modulation closely time-locked to the saccades. Two thirds (37/55) of saccade-related P-cells began to change their SS discharge rate 20–100 ms prior to the onset of saccades. The remaining one third (18/55) changed their activity approximately at the same time as the saccade onset. These saccade-related P-cells did not show changes in activity during smooth pursuit eye movements, and we did not find any P-cells in the cerebellar hemisphere which showed changes of activity preferentially during smooth pursuit eye movements. In about half (26/55) of the saccade-related P-cells, the pattern of modulation prior to and during saccades was biphasic: increase-decrease or decrease-increase. The other half (29/55) showed monophasic increases or decreases. For a given P-cell, the discharge pattern during saccades was similar for saccades of all directions, though there was a preferred direction in the amount of discharge rate modulation. The present findings suggest that the cerebellar hemisphere (Crus I and IIa) plays an important role in the control of voluntary saccadic eye movements, in addition to other cerebellar cortical areas (flocculus and posterior vermis) which are known to participate in the control of saccades.     

Activity of neurons in the lateral intraparietal area of the monkey during an antisaccade task.

The close relationship between saccadic eye movements and vision complicates the identification of neural responses associated with each function. Visual and saccade-related responses are especially closely intertwined in a subdivision of posterior parietal cortex, the lateral parietal area (LIP). We analyzed LIP neurons using an antisaccade task in which monkeys made saccades away from a salient visual cue. The vast majority of neurons reliably signaled the location of the visual cue. In contrast, most neurons had only weak, if any, saccade-related activity independent of visual stimulation. Thus, whereas the great majority of LIP neurons reliably encoded cue location, only a small minority encoded the direction of the upcoming saccade.     

Premotor inhibitory neurons carry signals related to saccade adaptation in the monkey

Cerebellar output changes during motor learning. How these changes cause alterations of motoneuron activity and movement remains an unresolved question for voluntary movements. To answer this question, we examined premotor neurons for saccadic eye movement. Previous studies indicate that cells in the fastigial oculomotor region (FOR) within the cerebellar nuclei on one side exhibit a gradual increase in their saccade-related discharge as the amplitude of ipsiversive saccades adaptively decreases. This change in FOR activity could cause the adaptive change in saccade amplitude because neurons in the FOR project directly to the brain stem region containing premotor burst neurons (BNs). To test this possibility, we recorded the activity of saccade-related burst neurons in the area that houses premotor inhibitory burst neurons (IBNs) and examined their discharge during amplitude-reducing adaptation elicited by intrasaccadic target steps. We specifically analyzed their activity for off-direction (contraversive) saccades, in which the IBN activity would increase to reduce saccade size. Before adaptation, 29 of 42 BNs examined discharged, at least occasionally, for contraversive saccades. As the amplitude of contraversive saccades decreased adaptively, half of BNs with off-direction spike activity showed an increase in the number of spikes (14/29) or an earlier occurrence of spikes (7/14). BNs that were silent during off-direction saccades before adaptation remained silent after adaptation. These results indicate that the changes in the off-direction activity of BNs are closely related to adaptive changes in saccade size and are appropriate to cause these changes.     

Saccade-related neurons in the primate fastigial nucleus: what do they encode?

The cerebellar fastigial oculomotor region (FOR) and the overlying oculomotor vermis (OV) are involved in the control of saccadic eye movements, but nature and function of their saccade-related neuronal signals are not fully understood. There is controversy in at least two major aspects: first, lesion studies in OV/FOR reported eye-position-dependent dysmetria-with FOR lesions, centripetal saccades became more hypermetric than centrifugal saccades-suggesting that the cerebellum may compensate for orbital mechanics. However, single-unit studies failed to reveal corresponding eye-position dependencies in FOR saccade-related discharge patterns. Second, some single-unit studies reported precise correlation between burst and saccade duration in the FOR. However, others stated that FOR bursts were only weakly related to saccade properties. In an attempt to resolve these discrepancies, we recorded single FOR units in monkeys that made horizontal saccades (16 degrees ) from different starting positions. Sampling saccades of one fixed amplitude and application of an objective, computer-based burst-detection-routine allowed us to correlate burst parameters (onset latency, peak latency, peak amplitude, number of spikes, duration) and kinematic properties of individual saccades. FOR bursts were found to start and peak earlier and exhibit higher peak burst amplitudes for faster than for slower saccades of the same amplitude. While these correlations between FOR bursts and saccade properties were statistically significant for a minority of approximately 20-25% of individual units, the same effects were also predominant in the remainder of the neuronal sample and statistically significant on the population level. Neuronal activity was not significantly modulated by eye position itself. However, reflecting differences in saccade velocities but not an actual influence of eye position per se, FOR bursts for centripetal and centrifugal saccades exhibited subtle but systematic differences, which closely paralleled, and hence probably explain, the eye-position dependency of deficits observed after FOR inactivation. Our findings indicate that FOR signals reflect much of the kinematic properties of the saccade. Moreover, they are consistent with the idea that the FOR output is purposefully modified according to these kinematic properties to maintain saccadic accuracy.     

Early- and late-responding cells to saccadic eye movements in the cortical area V6A of macaque monkey

The cortical area V6A, located in the dorsal part of the anterior bank of the parieto-occipital sulcus, contains retino- and craniocentric visual neurones together with neurones sensitive to gaze direction and/or saccadic eye movements, somatosensory stimulation and arm movements. The aim of this work was to study the dynamic characteristics of V6A saccade-related activity. Extracellular recordings were carried out in six macaque monkeys performing a visually guided saccade task with the head restrained. The task was performed in the dark, in both the dark and light, and sometimes in the light only. The discharge of certain neurones during saccades is due to their responsiveness to visual stimuli. We used a statistical method to distinguish responses due to visual stimulation from those responsible for saccadic control. Out of 597 V6A neurones tested, 66 (11%) showed responses correlated with saccades; 26 of 66 responded also to visual stimulation and 31 of 66 did not; the remaining 9 were not visually tested. We calculated the response latency to saccade onset and its inter-trial variance in 24 of 66 neurones. Saccade neurones could respond before, during or after the saccade. Neurones responding before saccade-onset or during saccades had much higher latency variance than neurones responding after saccades. The early-responding cells had a mean latency (±SD) of –64±62 ms, while the late-responding cells a mean latency of +89±20 ms. The responses to saccadic eye movements were directionally sensitive and varied with the amplitude of the saccade. Responses of late-responding cells disappeared in complete darkness. We suggest that the activity of early-responding cells represents the intended saccadic eye movement or the shift of attention towards another part of the visual space, whereas that of late-responding cells is a visual response due to retinal stimulation during saccades. Electronic Publication     

Different learned coordinate frames for planning trajectories and final positions in reaching

We previously reported that the kinematics of reaching movements reflect the superimposition of two separate control mechanisms specifying the hand's spatial trajectory and its final equilibrium position. We now asked whether the brain maintains separate representations of the spatial goals for planning hand trajectory and final position. One group of subjects learned a 30 degrees visuomotor rotation about the hand's starting point while performing a movement reversal task ("slicing") in which they reversed direction at one target and terminated movement at another. This task required accuracy in acquiring a target mid-movement. A second group adapted while moving to -- and stabilizing at -- a single target ("reaching"). This task required accuracy in specifying an intended final position. We examined how learning in the two tasks generalized both to movements made from untrained initial positions and to movements directed toward untrained targets. Shifting initial hand position had differential effects on the location of reversals and final positions: Trajectory directions remained unchanged and reversal locations were displaced in slicing whereas final positions of both reaches and slices were relatively unchanged. Generalization across directions in slicing was consistent with a hand-centered representation of desired reversal point as demonstrated previously for this task whereas the distributions of final positions were consistent with an eye-centered representation as found previously in studies of pointing in three-dimensional space. Our findings indicate that the intended trajectory and final position are represented in different coordinate frames, reconciling previous conflicting claims of hand-centered (vectorial) and eye-centered representations in reach planning.     

Target selection for reaching and saccades share a similar behavioral reference frame in the macaque

The selection of one of two visual stimuli as a target for a motor action may depend on external as well as internal variables. We examined whether the preference to select a leftward or rightward target depends on the action that is performed (eye or arm movement) and to what extent the choice is influenced by the target location. Two targets were presented at the same distance to the left and right of a fixation position and the stimulus onset asynchrony (SOA) was adjusted until both targets were selected equally often. This balanced SOA time is then a quantitative measure of selection preference. In two macaque monkeys tested, we found the balanced SOA shifted to the left side for left-arm movements and to the right side for right-arm movements. Target selection strongly depended on the horizontal target location. By varying eye, head, and trunk position, we found this dependency embedded in a head-centered behavioral reference frame for saccade targets and, somewhat counter-intuitively, for reach targets as well. Target selection for reach movements was influenced by the eye position, while saccade target selection was unaffected by the arm position. These findings suggest that the neural processes underlying target selection for a reaching movement are to a large extent independent of the coordinate frame ultimately used to make the limb movement, but are instead closely linked to the coordinate frame used to plan a saccade to that target. This similarity may be indicative of a common spatial framework for hand-eye coordination.     

Non-spatial,motor-specific activation in posterior parietal cortex

A localized cluster of neurons in macaque posterior parietal cortex, termed the parietal reach region (PRR), is activated when a reach is planned to a visible or remembered target. To explore the role of PRR in sensorimotor transformations, we tested whether cells would be activated when a reach is planned to an as-yet unspecified goal. Over one-third of PRR cells increased their firing after an instruction to prepare a reach, but not after an instruction to prepare a saccade, when the target of the movement remained unknown. A partially overlapping population (two-thirds of cells) was activated when the monkey was informed of the target location but not the type of movement to be made. Thus a subset of PRR neurons separately code spatial and effector-specific information, consistent with a role in specifying potential motor responses to particular targets.     

Relationship between saccadic eye movements and cortical activity as measured by fMRI: quantitative and qualitative aspects

We investigated the quantitative relationship between saccadic activity (as reflected in frequency of occurrence and amplitude of saccades) and blood oxygenation level dependent (BOLD) changes in the cerebral cortex using functional magnetic resonance imaging (fMRI). Furthermore, we investigated quantitative changes in cortical activity associated with qualitative changes in the saccade task for comparable levels of saccadic activity. All experiments required the simultaneous acquisition of eye movement and fMRI data. For this purpose we used a new high-resolution limbus-tracking technique for recording eye movements in the magnetic resonance tomograph. In the first two experimental series we varied both frequency and amplitude of saccade stimuli (target jumps). In the third series we varied task difficulty; subjects performed either pro-saccades or anti-saccades. The brain volume investigated comprised the frontal and supplementary eye fields, parietal as well as striate cortex, and the motion sensitive area of the parieto-occipital cortex. All these regions showed saccade-related BOLD responses. The responses in these regions were highly correlated with saccade frequency, indicating that repeated processing of saccades is integrated over time in the BOLD response. In contrast, there was no comparable BOLD change with variation of saccade amplitude. This finding speaks for a topological rather than activity-dependent coding of saccade amplitudes in most cortical regions. In the experiments comparing pro- vs anti-saccades we found higher BOLD activation in the "anti" task than in the "pro" task. A comparison of saccade parameters revealed that saccade frequency and cumulative amplitude were comparable between the two tasks, whereas reaction times were longer in the "anti" task than the pro task. The latter finding is taken to indicate a more demanding cortical processing in the "anti" task than the "pro" task, which could explain the observed difference in BOLD activation. We hold that a quantitative analysis of saccade parameters (especially saccade frequency and latency) is important for the interpretation of the BOLD changes observed with visual stimuli in fMRI.     

Electrophysiological recordings in humans reveal reduced location-specific attentional-shift activity prior to recentering saccades

Being able to effectively explore the visual world is of fundamental importance, and it has been suggested that the straight-ahead gaze position within the egocentric reference frame ("primary position") might play a special role in this context. In the present study we employed human electroencephalography (EEG) to examine neural activity related to the spatial guidance of saccadic eye movements. Moreover, we sought to investigate whether such activity would be modulated by the spatial relation of saccade direction to the primary gaze position (recentering saccades). Participants executed endogenously cued saccades between five equidistant locations along the horizontal meridian. This design allowed for the comparison of isoamplitude saccades from the same starting position that were oriented either toward the primary position (centripetal) or further away from it (centrifugal). By back-averaging time-locked to the saccade onset on each trial, we identified a parietally distributed, negative-polarity EEG deflection contralateral to the direction of the upcoming saccade. Importantly, this contralateral presaccadic negativity, which appeared to reflect the location-specific attentional guidance of the eye movement, was attenuated for recentering saccades relative to isoamplitude centrifugal saccades. This differential electrophysiological signature was paralleled by faster saccadic reaction times and was substantially more apparent when time-locking the data to the onset of the saccade rather than to the onset of the cue, suggesting a tight temporal association with saccade initiation. The diminished level of this presaccadic component for recentering saccades may reflect the preferential coding of the straight-ahead gaze position, in which both the eye-centered and head-centered reference frames are perfectly aligned and from which the visual world can be effectively explored.     

Visuomotor interactions in responses of neurons in the middle and lateral suprasylvian cortices of the behaving cat

Summary We studied visuomotor processing in the middle (MS) and lateral suprasylvian (LS) cortices of the alert cat by making single cell recordings while the cat was working in a behavioral task requiring visual fixation and visually guided eye movements. We found responses with three different components: visual sensory, saccaderelated motor, and fixation. Some cells exhibited purely visual responses and all of their activity during visuomotor tasks could be attributed to the sensory aspects of the task. Other cells showed no sensory response properties, but discharged in relation to the saccadic eye movements that the cat made to visual targets. A smaller number of fixation cells displayed increased discharge when the cat fixated a target light and usually only when that target was in a particular region of the visual field.These response components could be present in a variety of combinations in different cells, of which the largest proportion combined visuomotor responses and could take five general forms: simple visuomotor, saccadic enhanced, visually triggered movement (VTM), enhanced VTM, and disenhanced. Simple visuomotor responses had both a visual and saccade-related component. Saccadic enhanced responses had a visual response to the appearance of a spot in the cell's receptive field that became enhanced when the cat subsequently made a saccade to that spot. The VTM responses were synchronized better to the visual stimulus than to the saccade, but they also exhibited properties expected of motor responses. The last two classes of visuomotor responses were rare: one we termed enhanced VTM and the other disenhanced. Cells could combine different visuomotor response components or even sensory, saccade-related and fixation responses in different combinations for different directions of eye movements. Generally, the timing of the saccade-related responses occurred too late to play a role in the initiation of saccades: most (83%) saccade-related responses occurred between 40 ms before to 80 ms after the onset of the eye movement. Cells of all different types could be found in both the MS and LS areas, though in general the responses in LS were more sensory in nature while those in MS were more closely related to the eye movement. About a quarter of the cells were unresponsive during any aspect of our tasks.     

Similarity of superior colliculus involvement in microsaccade and saccade generation

The characteristics of microsaccades, or small fixational saccades, and their influence on visual function have been studied extensively. However, the detailed mechanisms for generating these movements are less understood. We recently found that the superior colliculus (SC), a midbrain structure involved in saccade generation, also plays a role in microsaccade generation. Here we compared the dynamics of neuronal activity in the SC associated with microsaccades to those observed in this structure in association with larger voluntary saccades. We found that microsaccade-related activity in the SC is characterized by a gradual increase in firing rate starting ～100 ms prior to microsaccade onset, a peak of neuronal discharge just after movement onset, and a subsequent gradual decrease in firing rate until ～100 ms after movement onset. These properties were shared with saccade-related SC neurons, recorded from the same monkeys but preferring larger eye movements, suggesting that at the level of the SC the neuronal control of microsaccades is similar to that for larger voluntary saccades. We also found that neurons exhibiting microsaccade-related activity often also exhibited saccade-related activity for slightly larger movements of similar direction, suggesting a continuity of the spatial representation in the SC, in both amplitude and direction, down to the smallest movements. Our results indicate that the mechanisms controlling microsaccades may be fundamentally the same as those for larger saccades, and thus shed new light on the functional role of these eye movements and their possible influence on sensory and sensory-motor processes.