Impairment of extraretinal eye position signals after central thalamic lesions in humans

Accuracy of four different types of memory-guided saccades was studied in two patients with a small central thalamic lesion, probably involving the region of the internal medullary lamina (IML), and in a control group. In the first paradigm, the eyes and head remained immobile between the time of the presentation of the visual target to be remembered and the memory-guided saccade. In the other three paradigms, the eyes were displaced during the same period (before the memory-guided saccade) by either visually-guided saccades, a smooth pursuit eye movement or a body movement (with vestibulo-ocular reflex suppression). Therefore, in these three paradigms, the initial eye displacement required the use of extraretinal eye position to produce accurate memory-guided saccades. Compared with the control group, the two patients had normal accuracy in the first memory-guided saccade paradigm, in which there was no initial eye displacement, but markedly impaired saccade accuracy in the other three paradigms. These results suggest that the cortical areas triggering saccades did not receive correct extraretinal eye position signals. They are consistent with an impairment of the efference copy, which could be distributed to the cortical ocular motor areas by the IML.     

Memory-guided saccades: what is memorized?

Summary In order to find out whether extraretinal (oculomotor, internal) input suffices to provide the oculomotor system with the information necessary for saccadic control, two subjects were asked to make memoryguided saccades in complete darkness, after three different location acquisition conditions. These conditions were visually-guided saccades (SA), providing retinal (external) and extraretinal input, visual peripheral target presentation during central target fixation (FI) (external input only), and smooth pursuit (PU) (internal input only). Either 2 or 12 s (delay) after locating the target, the subjects had to make a memory-guided saccade toward it in complete darkness. The results show that whereas these memory-guided saccades were quite accurate for trials with preceding external input, this was not the case with acquisition through internal input alone. Moreover, the accuracy of memory-guided saccades decreased when the delay increased from 2 to 12s for both conditions with retinal input, whereas the accuracy increased for the one condition without retinal input, i.e., the smooth pursuit location acquisition. Furthermore, when both retinal and oculomotor inputs were provided, better accuracy of the memory-guided saccades was observed than with single input.     

Effects of switching between leftward and rightward pro- and antisaccades

Previous studies suggested that random switching between pro- and antisaccades increases errors in both tasks. However, little is known about the effects of switching between leftward and rightward saccades (response switching). The present study investigated task and response switching using an alternating runs procedure. Tasks (i.e., prosaccades versus antisaccades) were switched every second trial. Response switches (i.e., leftward saccades versus rightward saccades) were counterbalanced across tasks and task-switching conditions. Task switching increased errors in both tasks. Response switching increased errors when antisaccades were preceded by antisaccades but not when antisaccades were preceded by prosaccades or for prosaccades regardless of the preceding saccade type. The task-switch effects suggest that both pro- and antisaccade trials activate specific production rules that can persist in a subsequent trial. The differential response-switch effects may reflect different modes of response activation in pro- and antisaccades (sensorimotor transformation of visual information versus selection of motor programs).     

Role of the different frontal lobe areas in the control of the horizontal component of memory-guided saccades in man

Two paradigms of memory-guided saccades were studied in 14 patients with focal vascular lesions affecting either the frontal eye field (FEF), or the supplementary eye field (SEF) or Brodmann's area 46 in the prefrontal cortex (PFC), and in 13 age-matched control subjects. In the first paradigm, the subject had to remember the position of a visual target with the body immobile and, in the second, the position towards which gaze was directed before a body rotation, i.e. with a vestibular input. In control subjects, the percentage of error in saccade accuracy (horizontal component) was greater in the second than in the first paradigm (37% and 14% on average, respectively). Compared with controls, amplitude error was significantly increased in the FEF group for the first paradigm only, in the SEF group for the second paradigm only, and in the PFC group for both paradigms. These results are consistent with (1) the PFC providing an improvement in the utilization by the saccade system of the visual and vestibular signals used in the two paradigms, and (2) the FEF and SEF providing an improvement in the utilization of visual signals in the first paradigm and vestibular signals in the second paradigm, respectively. Furthermore, from these findings and experimental data, it may be hypothesized (1) that the PFC is a part of the network contributing to short-term memorization of both visual and vestibular signals, and (2) that the FEF and SEF control two different types of memory-guided saccades, with separate calculation modes to determine their amplitude.     

Muscimol-induced inactivation of monkey frontal eye field: effects on visually and memory-guided saccades.

Muscimol-induced inactivation of the monkey frontal eye field: effects on visually and memory-guided saccades. Although neurophysiological, anatomic, and imaging evidence suggest that the frontal eye field (FEF) participates in the generation of eye movements, chronic lesions of the FEF in both humans and monkeys appear to cause only minor deficits in visually guided saccade generation. Stronger effects are observed when subjects are tested in tasks with more cognitive requirements. We tested oculomotor function after acutely inactivating regions of the FEF to minimize the effects of plasticity and reallocation of function after the loss of the FEF and gain more insight into the FEF contribution to the guidance of eye movements in the intact brain. Inactivation was induced by microinjecting muscimol directly into physiologically defined sites in the FEF of three monkeys. FEF inactivation severely impaired the monkeys' performance of both visually guided and memory-guided saccades. The monkeys initiated fewer saccades to the retinotopic representation of the inactivated FEF site than to any other location in the visual field. The saccades that were initiated had longer latencies, slower velocities, and larger targeting errors than controls. These effects were present both for visually guided and for memory-guided saccades, although the memory-guided saccades were more disrupted. Initially, the effects were restricted spatially, concentrating around the retinotopic representation at the center of the inactivated site, but, during the course of several hours, these effects spread to flanking representations. Predictability of target location and motivation of the monkey also affected saccadic performance. For memory-guided saccades, increases in the time during which the monkey had to remember the spatial location of a target resulted in further decreases in the accuracy of the saccades and in smaller peak velocities, suggesting a progressive loss of the capacity to maintain a representation of target location in relation to the fovea after FEF inactivation. In addition, the monkeys frequently made premature saccades to targets in the hemifield ipsilateral to the injection site when performing the memory task, indicating a deficit in the control of fixation that could be a consequence of an imbalance between ipsilateral and contralateral FEF activity after the injection. There was also a progressive loss of fixation accuracy, and the monkeys tended to restrict spontaneous visual scanning to the ipsilateral hemifield. These results emphasize the strong role of the FEF in the intact monkey in the generation of all voluntary saccadic eye movements, as well as in the control of fixation.     

The frontal eye field is involved in spatial short-term memory but not in reflexive saccade inhibition

Physiological studies in monkeys have shown that the frontal eye field (FEF) is involved in the preparation and triggering of purposive saccades. However, several questions of FEF function remain unclear: the role of the FEF in visual short-term memory, its ability to update its spatial map and its role in reflexive saccade inhibition. We have addressed these issues in a patient with a small acute ischemic lesion whose location corresponded very accurately to the region of the left FEF according to the most recent cerebral blood flow studies. An initial study was conducted on days 7 and 8 after the stroke, i.e., before substantial recovery. A first group of paradigms (smooth pursuit, simple saccade tasks) was performed to assess FEF dysfunction. In a second group of paradigms, (1) visual short-term memory was tested by means of memory-guided saccade paradigms with short and long delays (1 and 7 s), (2) spatial updating abilities were tested by a double-step saccade task and two memory-guided saccade tasks in which the central fixation point was displaced during the memorization delay, and (3) reflexive saccade inhibition was tested by the antisaccade task. Results show that the FEF is involved in short-term memorization of the parameters of the forthcoming memory-guided saccade encoded in oculocentric coordinates. Normal results in the antisaccade task suggest that the FEF is not involved in reflexive saccade inhibition. Received: 26 January 1999 / Accepted: 3 June 1999     

Processing of retinal and extraretinal signals for memory-guided saccades during smooth pursuit

It is an essential feature for the visual system to keep track of self-motion to maintain space constancy. Therefore the saccadic system uses extraretinal information about previous saccades to update the internal representation of memorized targets, an ability that has been identified in behavioral and electrophysiological studies. However, a smooth eye movement induced in the latency period of a memory-guided saccade yielded contradictory results. Indeed some studies described spatially accurate saccades, whereas others reported retinal coding of saccades. Today, it is still unclear how the saccadic system keeps track of smooth eye movements in the absence of vision. Here, we developed an original two-dimensional behavioral paradigm to further investigate how smooth eye displacements could be compensated to ensure space constancy. Human subjects were required to pursue a moving target and to orient their eyes toward the memorized position of a briefly presented second target (flash) once it appeared. The analysis of the first orientation saccade revealed a bimodal latency distribution related to two different saccade programming strategies. Short-latency (<175 ms) saccades were coded using the only available retinal information, i.e., position error. In addition to position error, longer-latency (>175 ms) saccades used extraretinal information about the smooth eye displacement during the latency period to program spatially more accurate saccades. Sensory parameters at the moment of the flash (retinal position error and eye velocity) influenced the choice between both strategies. We hypothesize that this tradeoff between speed and accuracy of the saccadic response reveals the presence of two coupled neural pathways for saccadic programming. A fast striatal-collicular pathway might only use retinal information about the flash location to program the first saccade. The slower pathway could involve the posterior parietal cortex to update the internal representation of the flash once extraretinal smooth eye displacement information becomes available to the system.     

Interaction between smooth anticipation and saccades during ocular orientation in darkness

A saccade triggered during sustained smooth pursuit is programmed using retinal information about the relative position and velocity of the target with respect to the eye. Thus the smooth pursuit and saccadic systems are coordinated by using common retinal inputs. Yet, in the absence of retinal information about the relative motion of the eye with respect to the target, the question arises whether the smooth and saccadic systems are still able to be coordinated possibly by using extraretinal information to account for the saccadic and smooth eye movements. To address this question, we flashed a target during smooth anticipatory eye movements in darkness, and the subjects were asked to orient their visual axis to the remembered location of the flash. We observed multiple orientation saccades (typically 2-3) toward the memorized location of the flash. The first orienting saccade was programmed using only the position error at the moment of the flash, and the smooth eye movement was ignored. However, subsequent saccades executed in darkness compensated gradually for the smooth eye displacement (mean compensation congruent with 70%). This behavior revealed a 400-ms delay in the time course of orientation for the compensation of the ongoing smooth eye displacement. We conclude that extraretinal information about the smooth motor command is available to the saccadic system in the absence of visual input. There is a 400-ms delay for smooth movement integration, saccade programming and execution.     

Predictive smooth eye pursuit in a population of young men: I. Effects of age, IQ, oculomotor and cognitive tasks

Smooth eye pursuit is believed to involve the integration of an extraretinal signal formed by an internal representation of the moving target and a retinal signal using the visual feedback to evaluate performance. A variation of the smooth eye pursuit paradigm (in which the moving target is occluded for a short period of time and subjects are asked to continue tracking) designed to isolate the predictive processes that drive the extraretinal signal was performed by 1,187 young men. The latency to the onset of change in pursuit speed, the time of decelerating eye-movement speed and the steady state residual gain were measured for each subject and correlated with measures of other oculomotor (closed-loop smooth eye pursuit, saccade, antisaccade, active fixation) and cognitive tasks (measuring sustained attention and working memory). Deceleration time increased with increasing age, while education, general IQ and cognitive variables had no effect on predictive pursuit performance. Predictive pursuit indices were correlated to those of closed-loop pursuit and antisaccade performance, but these correlations were very weak except for a positive correlation of residual gain to saccade frequency in the fixation task with distracters. This correlation suggested that the maintenance of active fixation is negatively correlated with the ability to maintain predictive pursuit speed. In conclusion, this study presents predictive pursuit performance in a large sample of apparently healthy individuals. Surprisingly, predictive pursuit was weakly if at all related to closed-loop pursuit or other oculomotor and cognitive tasks, supporting the usefulness of this phenotype in the study of frontal lobe integrity in normal and patient populations.     

Identifying sites of saccade amplitude plasticity in humans: transfer of adaptation between different types of saccade

To view different objects of interest, primates use fast, accurate eye movements called saccades. If saccades become inaccurate, the brain adjusts their amplitudes so they again land on target, a process known as saccade adaptation. The different types of saccades elicited in different behavioral circumstances appear to utilize different parts of the oculomotor circuitry. To gain insight into where adaptation occurs in different saccade pathways, we adapted saccades of one type and examined how that adaptation affected or transferred to saccades of a different type. If adaptation of one type of saccade causes a substantial change in the amplitude of another, that adaptation may occur at a site used in the generation of both types of saccade. Alternatively, if adaptation of one type of saccade transfers only partially, or not at all, to another, adaptation occurs at least in part at a location that is not common to the generation of both types of saccade. We produced significant amplitude reductions in memory-guided, delayed, targeting and express saccades by moving the target backward during the saccade. After memory-guided saccades were adapted, the amplitude of express, targeting and delayed saccades exhibited only a partial reduction. In contrast, when express, targeting, or delayed saccades were adapted, amplitude transfer to memory-guided saccades was more substantial. These results, combined with previously published data, suggest that there are at least two sites of adaptation within the saccadic system. One is used communally in the generation of express, targeting, delayed and memory-guided saccades, whereas the other is specific for the generation of memory-guided saccades.     

Effects of direction on saccadic performance in relation to lateral preferences

A sample of 676 healthy young males performed visually guided saccades and antisaccades and completed the Porac-Coren questionnaire measuring lateral preferences. There was no difference in mean latency between rightward versus leftward saccades or for saccades executed in the left versus right hemispace. There was also no right/left asymmetry for individuals with left or right dominance as assessed by the lateral preferences questionnaire. The same results were observed for the latency of antisaccades and for the error rate in the antisaccade task. Finally, we did not confirm any substantial subpopulation of individuals with idiosyncratic left/right latency asymmetries that persisted both in the saccade and antisaccade task. These results suggest that neither latency nor antisaccade error rate are good indicators of lateral preferences in these tasks. Other oculomotor tasks might be more sensitive to hemifield differences, or cerebral hemispheric asymmetry is not present at the level of cortical organization of saccades and antisaccades.     

Cortical control of memory-guided saccades in man

Summary Memory-guided saccades were electro-oculographically recorded in 30 patients with limited unilateral cerebral infarction, documented by computerized tomographic scan and/or magnetic resonance imaging. The lesions affected either (1) the posterior parietal cortex (PPC), (2) the dorsolateral frontal cortex (DLFC), involving the frontal eye field (FEF) and/or the prefrontal cortex (PFC) (area 46 of Brodmann), or (3) the supplementary motor area in the dorsomedial frontal cortex (DMFC). Patients were divided into 6 groups according to the location (PPC, DLFC, DMFC) and side of the lesions. Both latency and accuracy (expressed as a percentage of error in amplitude) of memory-guided saccades were compared in each group of patients to values obtained from 20 age-matched normal subjects. Latency was significantly increased, for both directions of saccades in the two DLFC groups and in the right PPC group, and for leftward saccades in the left PPC group. The percentage of error in amplitude was also significantly increased for both directions of saccades in the right PPC group and the left DLFC group, and for leftward saccades in the right DLFC group. Results were near the normal values in patients with lesions affecting the DMFC. Thus, both the PPC (essentially on the right side) and the DLFC appear to play a role in the control of memory-guided saccades. It is suggested that the cortical pathway involved in these saccades includes the PPC, the PFC and the FEF, successively. The PPC could have a dual role: visuospatial integration, and early selection and preparation of certain collicular cells by pre-excitation. Both functions could be ensured by two different types of cells, corresponding, in the monkey, to area 7a and to the lateral intraparietal area, respectively. The DLFC could also have a dual role: memorization of visuospatial information by the PFC, and triggering of memory-guided saccades by the FEF.     

An assessment of optokinetic nystagmus (OKN) in persons with down syndrome

Down syndrome (DS), the most common genetically defined cause of intellectual disability, is the phenotypic consequence of a supernumerary chromosome 21. Persons with DS commonly display deficits in visuomotor integration, motor coordination, and balance. Despite the key roles of the optokinetic and vestibular systems in these submodalities of motor function, a systematic investigation of the optokinetic nystagmus (OKN) and vestibulo-ocular reflex (VOR) in persons with DS had lacked in the literature. Accordingly, this study generated quantitative data on oculomotor function in persons with DS under optokinetic and sinusoidal smooth pursuit stimulation. Thirty-two participants with DS (14–36 years old, equally divided by gender) and 32 chronological age- and gender-matched typically developing controls were recruited from the community. Eye movements were recorded by binocular video oculography and an LCD projector produced visual stimulation. Assessments of the gain and frequency of slow phase of OKN beats and number and mean amplitude of intruding saccades during smooth pursuit were performed. Individuals with DS displayed angular velocity-dependent reduction in OKN gain and number of produced nystagmus beats compared to controls. The gain of the smooth pursuit was not significantly different between participants with DS and control participants. However, the number and mean amplitude of intruding saccades during smooth pursuit were increased in participants with DS compared to control participants. These findings may have implications to the understanding of the neurological basis of the motor dysfunction that affects performance in many practical tasks persons with DS encounter in their everyday lives.     

Adaptation of catch-up saccades during the initiation of smooth pursuit eye movements

Reduction of retinal speed and alignment of the line of sight are believed to be the respective primary functions of smooth pursuit and saccadic eye movements. As the eye muscles strength can change in the short-term, continuous adjustments of motor signals are required to achieve constant accuracy. While adaptation of saccade amplitude to systematic position errors has been extensively studied, we know less about the adaptive response to position errors during smooth pursuit initiation, when target motion has to be taken into account to program saccades, and when position errors at the saccade endpoint could also be corrected by increasing pursuit velocity. To study short-term adaptation (250 adaptation trials) of tracking eye movements, we introduced a position error during the first catch-up saccade made during the initiation of smooth pursuit—in a ramp-step-ramp paradigm. The target position was either shifted in the direction of the horizontally moving target (forward step), against it (backward step) or orthogonally to it (vertical step). Results indicate adaptation of catch-up saccade amplitude to back and forward steps. With vertical steps, saccades became oblique, by an inflexion of the early or late saccade trajectory. With a similar time course, post-saccadic pursuit velocity was increased in the step direction, adding further evidence that under some conditions pursuit and saccades can act synergistically to reduce position errors.     

Visual versus motor vector inversions in the antisaccade task: a behavioral investigation with saccadic adaptation

In the antisaccade task, subjects must execute an eye movement away from a visual target. Correctly executing an antisaccade requires inhibiting a prosaccade toward the visual target and programming a movement to the opposite side. This movement could be based on the inversion of the visual vector, corresponding to the distance between the fixation point and the visual target, or the motor vector of the unwanted prosaccade. We dissociated the two vectors by means of saccadic adaptation. Adaptation can be observed when systematic targeting errors are caused by the displacement of the visual target during the saccade. Adaptation progressively modifies saccade amplitude (defined by the motor vector) such that it becomes appropriate to the postsaccadic stimulus position and thus different from the visual vector of the target. If antisaccade preparation depended on visual vector inversion, rightward prosaccade adaptation should not transfer to leftward antisaccades (which are based on the same visual vector) but should transfer to rightward antisaccades (which are based on a visual vector inside the adaptation field). If antisaccade preparation depended on motor vector inversion, rightward prosaccade adaptation should transfer to leftward antisaccades (which are based on the same, adapted motor vector) but should not transfer to rightward antisaccades (which are based on a nonadapted motor vector). The results are in line with the first hypothesis, showing that vector inversion precedes saccadic adaptation and suggesting that antisaccade preparation depends on the inversion of the visual target vector.     

Spatial memory following shifts of gaze. I. Saccades to memorized world-fixed and gaze-fixed targets

During a shift of gaze, an object can move along with gaze or stay fixed in the world. To examine the effect of an object's reference frame on spatial working memory, we trained monkeys to memorize locations of visual stimuli as either fixed in the world or fixed to gaze. Each trial consisted of an initial reference frame instruction, followed by a peripheral visual flash, a memory-period gaze shift, and finally a memory-guided saccade to the location consistent with the instructed reference frame. The memory-period gaze shift was either rapid (a saccade) or slow (smooth pursuit or whole body rotation). This design allowed a comparison of memory-guided saccade performance under various conditions. Our data indicate that after a rotation or smooth-pursuit eye movement, saccades to memorized world-fixed targets are more variable than saccades to memorized gaze-fixed targets. In contrast, memory-guided saccades to world- and gaze-fixed targets are equally variable following a visually guided saccade. Across all conditions, accuracy, latency, and main sequence characteristics of memory-guided saccades are not influenced by the target's reference frame. Memory-guided saccades are, however, more accurate after fast compared with slow gaze shifts. These results are most consistent with an eye-centered representational system for storing the spatial locations of memorized objects but suggest that the visual system may engage different mechanisms to update the stored signal depending on how gaze is shifted.     

Event-related fMRI responses in the human frontal eye fields in a randomized pro- and antisaccade task

We examined whether the frontal eye fields (FEF) are involved in the suppression of reflexive saccades. Simultaneous recording of horizontal eye movements and functional magnetic resonance imaging enabled us to perform a randomized pro- and antisaccade task and to sort blood oxygenation level dependent (BOLD) time series on the basis of task performance. Saccadic reaction time distributions were comparable across tasks indicating a similar effort in preprocessing of the saccades. Furthermore, we found similar BOLD activation in FEF during both correctly performed pro- and antisaccades. Frontal eye field activation started prior to target presentation and saccade generation. While we observed only few erroneous antisaccades, these were associated with a decrease in BOLD activity prior to target presentation, and increased BOLD activity after target presentation relative to correctly performed antisaccades. These findings are consistent with a role of the FEF in the suppression of reflexive saccades. The increase in activity after target presentation for antisaccade errors can only be indirectly linked to such a role but may also reflect activity related to the generation of a correction saccade. Frontal eye field BOLD activity may further represent general arousal, preparatory set, short-term memory, or salience-map related activity.     

Quantitative analysis of catch-up saccades during sustained pursuit

During visual tracking of a moving stimulus, primates orient their visual axis by combining two very different types of eye movements, smooth pursuit and saccades. The purpose of this paper was to investigate quantitatively the catch-up saccades occurring during sustained pursuit. We used a ramp-step-ramp paradigm to evoke catch-up saccades during sustained pursuit. In general, catch-up saccades followed the unexpected steps in position and velocity of the target. We observed catch-up saccades in the same direction as the smooth eye movement (forward saccades) as well as in the opposite direction (reverse saccades). We made a comparison of the main sequences of forward saccades, reverse saccades, and control saccades made to stationary targets. They were all three significantly different from each other and were fully compatible with the hypothesis that the smooth pursuit component is added to the saccadic component during catch-up saccades. A multiple linear regression analysis was performed on the saccadic component to find the parameters determining the amplitude of catch-up saccades. We found that both position error and retinal slip are taken into account in catch-up saccade programming to predict the future trajectory of the moving target. We also demonstrated that the saccadic system needs a minimum period of approximately 90 ms for taking into account changes in target trajectory. Finally, we reported a saturation (above 15 degrees /s) in the contribution of retinal slip to the amplitude of catch-up saccades.     

Effects of occipital lobectomy upon eye movements in primate

1. Eye movements were recorded before and after bilateral occipital lobectomy in six rhesus monkeys trained to fixate and to follow small targets. Striate cortex was completely removed in two animals; small islands islands remained in the others. In all animals portions of extrastriate cortex were also removed but the medial superior temporal area in the superior temporal sulcus was largely spared. Optokinetic nystagmus (OKN) was markedly altered but not abolished in all animals. The immediate pursuit component of OKN was eliminated leading to a poor response to stimuli comprised of high frequencies. The velocity-storage component of OKN was present, but the maximum value of OKN that could be achieved was decreased to 6 and 16 degrees/s in the two most severely affected animals (preop, 65-116 degrees/s). The residual OKN was similar to that of afoveate animals with a diminished response to high velocities of retinal-image motion and a temporal to nasal predominance during monocular viewing. 2. In the initial postoperative period all animals appeared completely blind. Within 1-6 mo, however, they regained an ability to make visually guided saccades to, and smooth pursuit of, small targets. Saccades were nearly as accurate as preoperatively, but saccade amplitudes were more variable and saccade latencies increased. In the two animals with a complete removal of striate cortex, gains (eye velocity/target velocity) of smooth pursuit during sinusoidal tracking (60 degrees/s, 0.5 Hz) were 0.9 and 0.95. During tracking of step-ramp (Rashbass) stimuli with 60 degrees/s ramps, the average acceleration of the eyes during the first 120 ms of smooth pursuit was 189-278 degrees.s-1.s-1 (preop range, 154-418 degrees.s-1.s-1). In other respects, though, smooth pursuit was not normal. Latencies were increased two- to threefold, and tracking was more variable. 3. Paradoxically, as visually guided saccades and pursuit recovered, some other ocular motor functions deteriorated. Spontaneous and gaze-evoked nystagmus developed 3-6 mo after occipital lobectomy; the time constant of the neural eye-position integrator dropped to values as low as 2.6-4.8 s. The maximum slow-phase velocity of OKN also decreased. 4. The findings immediately after occipital lobectomy indicate that in normal primates occipital cortex is necessary for visually guided saccades and smooth pursuit as well as for the immediate component of OKN. Occipital cortex also makes the predominant contribution toward the generation of the velocity-storage component of OKN.(ABSTRACT TRUNCATED AT 400 WORDS)     

The “ways” we look at dreams: evidence from unilateral spatial neglect (with an evolutionary account of dream bizarreness)

Despite decades of research, the question of whether the rapid eye movements (REMs) of paradoxical sleep (PS) are equivalent to waking saccades and whether their direction is congruent with visual spatial events in the dream scene is still very controversial. We gained an insight into these questions through the study of a right brain damaged patient suffering attentional neglect for the left side of space and drop of the optokinetic nystagmus (OKN) with alternating rightward slow/leftward fast phases evoked by rightward optic flow. During PS the patient had frequent Nystagmoid REMs with alternating leftward slow/rightward fast phases and reported dreams with visual events evoking corresponding OKN such as a train running leftward. By contrast, just as in waking OKN, Nystagmoid REMs with alternating rightward slow/leftward fast phases were virtually absent. REMs followed by staring eye position or by consecutive REMs were also observed: these showed no asymmetry comparable to that of Nystagmoid ones. The selective disappearance of Nystagmoid REMs in one horizontal direction proves, for the first time, that in humans different types of REMs exists and that these are driven by different premotor mechanisms. Concomitant drop of OKN and Nystagmoid REMs toward the same horizontal direction demonstrates that phylogenetically ancient oculomotor mechanisms, such as the OKN, are shared by waking and PS. On this evidence and converging findings from animal, neuropsychological and brain imaging studies, a new evolutionary account of dream bizarreness is proposed. Classification and labelling of the different types of REMs are also provided.