Suppression Of Eye Movements Research Articles

12. The Role of the Human Prefrontal Cortex in the Response Suppression of Eye Movements

Increased activity in a population of prefrontal cortex neurons has been shown, in previous studies, to precede correct anti‐saccades in primates. In addition, the time courses of two competing processes in these neurons, task selection (which prepares for an eye movement) and saccade suppression (which prepares for the suppression of an eye movement), intersect at a specific time after the presentation of a coloured instruction cue. The purpose of this study is to use eye tracking behaviour to investigate this intersection point and its role in response suppression in regard to the generation of anti‐saccades in humans. Subjects were instructed before a stimulus appears, using a colour cue, to either look towards the stimulus (pro‐saccade) or away from the stimulus (anti‐saccade). Instruction period times varied from 100ms to 1400ms, in 50ms steps. Based on previous primate electrophysiological data, the ratio of direction errors (pro‐saccades on anti‐saccade trials) to correct anti‐saccades was expected to increase around 400ms to 500ms, when the processes of task selection and saccade suppression diverge. A slight ratio increase was found and full results are forthcoming.

Suppression of eye movements in visual selective attention tasks

Suppression of eye movements in visual selective attention tasks

Adaptation of a bimodal integration stage: visual input needed during neck muscle vibration to elicit a motion aftereffect

Vibratory stimulation of the neck muscles can elicit illusory drift of a visual target; after vibration stops, motion in the opposite direction is perceived. This motion aftereffect (MAE) could be due to adaptation of proprioceptive mechanisms that encode head orientation, or at a stage where visual and proprioceptive information are combined. To distinguish between these two possibilities, we applied vibratory stimulation to dorsolateral neck muscles for 15-s periods alternating with 15-s periods without vibration. Twenty-six observers used a hand-held tracker to indicate perceived motion of a stationary light-emitting diode (LED) in an otherwise dark room. In the critical condition, observers were in complete darkness during vibration, and the LED was only turned on in post-vibration periods. If adaptation was purely proprioceptive, a visual MAE should have occurred in this condition, but it did not. In a follow-up experiment (N = 9), the LED was presented intermittently to determine if there was a position aftereffect that might have been inhibited by processes signalling an absence of motion. No aftereffect occurred under these conditions either. In both experiments, a visual stimulus had to be present during the adaptation period in order to elicit an aftereffect. Results from our previous study ruled out an explanation based on suppression of eye movements. Thus, the most likely site responsible for the visual aftereffect lies with bimodal mechanisms combining proprioceptive and visual information. We conclude that the bimodal mechanisms adapted more quickly than the proprioceptive mechanisms from which they received input.

EEG activity related to preparation and suppression of eye movements in three-dimensional space.

Eight normal subjects made visually guided eye movements to four LED targets placed at two different distances (20 and 70 cm) and on either side (+/-10 degrees ) at 70 cm. Four types of eye movements were elicited: pure saccades, convergence, divergence, and combined (divergent saccades). EEG activity was recorded from 62 electrodes and was aligned to stimulus onset. A negativity peaked after 140 ms and was modulated according to the location of the stimulus in space and the type of movement prepared, mainly in central and posterior cortex. For saccade targets, we confirmed a stimulus-related negativity in the posterior and central cortical area, contralateral to target direction. For convergence and divergence targets, this negativity was bilaterally distributed; convergence targets activated a rather extended cortical network in the central and posterior area, while divergence targets activated a more confined posterior area, spreading ventrally from the occipital cortex. Cortical activity for combined targets was lateralised contralaterally to stimulus direction but its topography resembled more closely that after the divergence stimulus. When observers suppressed the relevant eye movement to the stimulus, EEG activity was enhanced on the right hemisphere, showing the more pronounced effect on the right occipital-temporal and central-parietal electrode sites.

Attention Response Functions: Characterizing Brain Areas Using fMRI Activation during Parametric Variations of Attentional Load

We derived attention response functions for different cortical areas by plotting neural activity (measured by fMRI) as a function of attentional load in a visual tracking task. In many parietal and frontal cortical areas, activation increased with load over the entire range of loads tested, suggesting that these areas are directly involved in attentional processes. However, in other areas (FEF and parietal area 7), strong activation was observed even at the lowest attentional load (compared to a passive baseline using identical stimuli), but little or no additional activation was seen with increasing load. These latter areas appear to play a different role, perhaps supporting task-relevant functions that do not vary with load, such as the suppression of eye movements.

Functional imaging of mirror and inverse reading reveals separate coactivated networks for oculomotion and spatial transformations.

Echoplanar functional magnetic resonance imaging (fMRI) was used to localize the cortical areas involved in the analysis of spatially transformed letter strings. Significant increases of the blood oxygen level-dependent (BOLD) signal for transformed vs normal reading were observed in the superior parietal lobule (SPL), along the intraparietal sulcus (IPS), in the frontal eye fields (FEF), and in the latero-occipital area LO. The respective contributions of oculomotor and spatial transformation areas to this activation pattern were separated by means of a control condition involving the execution and suppression of eye movements. Areas activated in association with the control of eye movements included the superior parietal lobule and the frontal eye fields. The cooperation of different brain areas was analysed by correlating the time course of task-dependent BOLD signal changes in these areas. This correlation analysis revealed coactivation of occipitotemporal object recognition areas and a spatial transformation area in the intraparietal sulcus during the reading of transformed letter strings. We suggest that cortical systems that are coactivated during complex cognitive tasks can be differentiated by the correlation analysis of BOLD signal time courses in spatially separate brain areas.

Acute Deprivation of the Terminal Four Hours of Sleep Does Not Increase Delta (0–3-Hz) Electroencephalograms: A Replication

This experiment evaluated further our previous finding that substitution of waking for the terminal 3-4 hr of sleep produces little or no increase in either visually scored or computer measures of delta sleep. Eleven young adults (mean age 24.5 yr) were studied on a baseline night, a night with sleep limited to an average of 188 min, and a recovery night. Visually scored sleep stages, eye movement activity and computer measures of 0-3 Hz were analyzed by nonrapid eye movement periods (NREMPs) and for all recorded sleep in each condition. In addition, we measured the heights, durations and areas under the curve manifested by the cyclic waxing and waning of 0-3-Hz integrated amplitude across sleep. Acute loss of 3.9 hr of sleep did not increase either visual or computer measures of delta electroencephalograms (EEG) on the recovery night, essentially confirming our previous findings. We hypothesize that augmentation of delta EEG above baseline levels after acute (one night's) sleep loss requires that disruption or loss of sleep from the first two NREMPs (or delta cycles). Rapid eye movement (REM) sleep durations on the recovery night were unaffected by the marked loss of REM sleep caused by partial deprivation. Although eye movements as well as stage REM were lost in the deprivation condition, eye movement density was significantly reduced rather than increased on the recovery night. This reduction is consistent with the hypothesis that REM activity varies inversely with sleep depth (or directly with central arousal level). The observations here, taken in association with previous results, suggest that a threshold for eye movement suppression by sleep deprivation in young adults lies in the range of 3-4 hr of prior sleep loss.

The role of the posterior vermis of monkey cerebellum in smooth-pursuit eye movement control. II. Target velocity-related Purkinje cell activity.

1. Purkinje cell activity was recorded from lobules VI and VII of the cerebellar vermis during the performance of visuooculomotor tasks designed to dissociate the signals related to head, smooth-pursuit eye, and retinal image movements. Task-related modulations in the simple spike discharge rates of 157 cells were observed in three alert monkeys. 2. Of 65 Purkinje cells that were completely tested for all three signals, all exhibited smooth-pursuit eye movement-related activity. An additional vestibular or visual response was observed in 17 and 11% of the cells, respectively. Eye, head, and retinal image velocity signals were all recorded in the same unit in 52% of the Purkinje cells. The responses of 5% of the fully tested cells were associated with changes in the direction of eye, head, and retinal image movement. 3. The observed sensorioculomotor responses were direction selective in 98% of the Purkinje cells. For the Purkinje cells that were fully tested, 60% of the cells exhibited peak discharge rates for ipsilateral and 40% for contralateral eye velocity. Of these Purkinje cells, 45% exhibited eye, head, and retinal image velocity signals with equivalent direction preferences. 4. Of 42 Purkinje cells tested, 88% demonstrated some kinds of interactive responses during combined eye and sensory stimulation. The interaction of eye and head velocity signals has been discussed in a companion paper (38). The modulation in discharge rate observed during tracking in the presence of a random dot background pattern could be predicted from the dissociated responses to smooth pursuit in the dark and to movements of the background pattern during suppression of eye movements. 5. The sensitivity to smooth-pursuit eye velocity averaged 1.4 times the sensitivity to head velocity. In 80% of the Purkinje cells, however, the sensitivity to eye velocity exceeded the sensitivity to head velocity by an average of only 10%. The sensitivity to smooth-pursuit eye velocity averaged 1.6 times the sensitivity to retinal image velocity. 6. An increase in Purkinje cell discharge rate was observed during the open-loop period of the initiation of smooth-pursuit eye movements. This open-loop response was consistent with the presence of a visual signal during ocular pursuit, since these cells were also shown to be responsive to a dissociated retinal image velocity signal. Furthermore, the magnitude of the open-loop response indicated an enhancement of the sensitivity to retinal image velocity when visual information became behaviorally significant.(ABSTRACT TRUNCATED AT 400 WORDS)

Activity of the striate cortex cells during saccadic eye movements of the alert cat

Neuronal activity in the striate cortex was studied during eye movements of alert cats under reinforcement of eye movements with rewards of water. Striate cells were differentiated into two groups exhibiting contrasting activities during and at intermissions of saccadic eye movements made in the presence of a visual pattern: (1) saccade-excited (SE) cells (207/271) that were excited during saccadic eye movements and were much less active at intermissions; and (2) saccade-depressed (SD) cells (55/271) that were depressed during eye movements and were strongly active at intermissions. Under suppression of eye movements by retrobulbar paralysis or by withdrawal of the rewards, most SE-cells (89/104) exhibited photic responsiveness characteristic of "complex" cells in the anesthetized cat, and almost all SD-cells (23/26) that of "simple" cells. Therefore, it is likely that the two major neuronal populations in the striate cortex provide parallel channels of visual information which are gated in an alternative way during eye movements.

Inhibition of intracranial self-stimulation in brain stem-transected cats — A proposed mechanism of aversive effects produced by brain stimulation

Effects of intracranial self-stimulation of central ‘punishment areas’ were studied on an operant conditioning of vertical eye movements in the midpontine pretrigeminal cats as well as in the ence´phale isole´cats. In 36 pretrigeminal cats, the ventromedial hypothalamus (VMH), basal amygdaloid nuclei (AMY), dorsal central gray (CG) of the midbrain and the thalamic nuclei such as the ventralis posteromedialis (VPM) and ventralis posterolateralis (VPL) were tested. No suppression of eye movements indicating a passive avoidance conditioning from stimulation of these ‘punishment areas’ was obtained in 92 electrode tip sites. In 49 ence´phale isole´cats, stimulation of the VPM associated with contraction of the facial muscles, demonstrated a marked passive avoidance effect on the eye movements. After blocking both the trigeminal (5N) and facial nerves (7N), VPM stimulation no longer produced an increase of facial EMG activity and the suppressive effect of eye movements was abolished. Extracranial blockade of 7N alone, which induced facial muscle paralysis also showed similar effects. Bilateral blockade of cranial nerves from acoustic (8N) to hypoglossal (12N) nerves had no significant effect on the avoidance conditioning. The mass neural activity recorded from the 5N showed a marked increase of discharge by VPM stimulation which was reduced significantly after 7N blockade. These results may suggest a possibility that punishing effects of brain stimulation depend on feedback from the periphery (muscles, blood vessels and visceral organs), whereas reward effects essentially depend on neural circuitry confined within the forebrain above the rostral pons.

Avoidance conditioning to tooth pulp stimulation in the cat after bulbar transection

In cats with rostral bulbar transection eliminating totally nucleus caudalis and most of nucleus interpolaris of the spinal trigeminal complex, an operant conditioning of vertical eye movements to lateral hypothalamic reward stimulation was achieved. When hypothalamic reward stimulation contingent to each eye movement was suddenly switched to tooth pulp stimulation, a passive avoidance conditioning shown as immediate suppression of eye movements was established. This suggests that a presumably strong noxious effect of tooth pulp stimulation can be conducted via the trigeminal main sensory nucleus and spinal nucleus oralis without the spinal nucleus caudalis.

Effects of stimulus-independent oculomotor activity on the occipital EEG

In order to assess the effects of oculomotor activity on the occipital alpha rhythm, 24 subjects performed each of 2 instructed tasks — random eye movement and eye movement suppression — under conditions of darkness (with eyes both open and closed) and illumination (eyes open in a Ganzfeld). Recordings of the occipital EEG indicated a general blocking of alpha during periods of oculomotor activity for 2 dependent measures, percent time alpha and mean duration of alpha bursts, by comparison with baseline intervals of relaxation. These results are interpreted as support for the activating effects of efferent, oculomotor processes on the alpha component of the EEG.