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Schnier, Fabian, i Markus Lappe. "Differences in intersaccadic adaptation transfer between inward and outward adaptation". Journal of Neurophysiology 106, nr 3 (wrzesień 2011): 1399–410. http://dx.doi.org/10.1152/jn.00236.2011.
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Saccadic adaptation is a mechanism to increase or decrease the amplitude gain of subsequent saccades, if a saccade is not on target. Recent research has shown that the mechanism of gain increasing, or outward adaptation, and the mechanism of gain decreasing, or inward adaptation, rely on partly different processes. We investigate how outward and inward adaptation of reactive saccades transfer to other types of saccades, namely scanning, overlap, memory-guided, and gap saccades. Previous research has shown that inward adaptation of reactive saccades transfers only partially to these other saccade types, suggesting differences in the control mechanisms between these saccade categories. We show that outward adaptation transfers stronger to scanning and overlap saccades than inward adaptation, and that the strength of transfer depends on the duration for which the saccade target is visible before saccade onset. Furthermore, we show that this transfer is mainly driven by an increase in saccade duration, which is apparent for all saccade categories. Inward adaptation, in contrast, is accompanied by a decrease in duration and in peak velocity, but only the peak velocity decrease transfers from reactive saccades to other saccade categories, i.e., saccadic duration remains constant or even increases for test saccades of the other categories. Our results, therefore, show that duration and peak velocity are independent parameters of saccadic adaptation and that they are differently involved in the transfer of adaptation between saccade categories. Furthermore, our results add evidence that inward and outward adaptation are different processes.
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Dombrowe, Isabel. "Saccadic inhibition in a guided saccade task". PeerJ 6 (14.03.2018): e4493. http://dx.doi.org/10.7717/peerj.4493.
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The eye movement system reacts very systematically to visual transients that are presented during the planning phase of a saccade. About 50 to 70 ms after the onset of a transient, the number of saccades that are started decreases, a phenomenon that has been termed saccadic inhibition. Saccades started just before this time window are hypometric compared to regular saccades, presumably because the presentation of the transient stops them in mid-flight. Recent research investigating the properties of repeated saccades to fixed locations found that these early saccades were additionally faster than expected from the main sequence relation, suggesting that a saccadic dead time during which saccades can no longer be modified does not exist. The present study investigated the properties of saccades to random locations in a guided saccade task. As expected, early saccades starting just before the saccadic inhibition dip in frequency were hypometric. Their velocity profiles implied that these saccades were actively stopped after reaching peak velocity. However, the peak velocities of these saccades did not generally deviate from the main sequence relation. The question whether an active stop of early saccades is incompatible with the idea of a saccadic dead time is open to debate.
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Awater, Holger, i Markus Lappe. "Perception of Visual Space at the Time of Pro- and Anti-Saccades". Journal of Neurophysiology 91, nr 6 (czerwiec 2004): 2457–64. http://dx.doi.org/10.1152/jn.00821.2003.
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The localization of peri-saccadically flashed objects shows two types of errors: first, a uniform shift in saccade direction, and second, a compression of visual space around the saccade target. Whereas the uniform shift occurs when the experiment is performed in complete darkness compression occurs when additional visual references are available. Thus peri-saccadic mislocalization contains motor and visual components. To distinguish between both factors we compared peri-saccadic localization errors during pro- and anti-saccades. In the case of anti-saccades, the visual cue that elicits the saccade and the actual eye movement are in opposite directions. We asked whether peri-saccadic compression can be observed with anti-saccades, and if so, whether the compression is directed toward the visual cue or follows the actual eye movement. In blocked trials, subjects performed saccades either toward a visual cue (pro-saccade) or to the mirrored position opposite to a visual cue (anti-saccade). Peri-saccadically, we flashed a thin vertical bar at one of four possible locations. Subjects had to indicate the perceived position of the bar with a mouse pointer about 500 ms after the saccade. Experiments were performed in complete darkness and with visual references. Peri-saccadic mislocalizations occurred during anti-saccades. The mislocalizations were very similar for pro- and anti-saccades in magnitude and direction. For both, pro- and anti-saccades, mislocalizations were directed toward the actual eye movement and not the visual cue.
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Handel, Ari, i Paul W. Glimcher. "Contextual Modulation of Substantia Nigra Pars Reticulata Neurons". Journal of Neurophysiology 83, nr 5 (1.05.2000): 3042–48. http://dx.doi.org/10.1152/jn.2000.83.5.3042.
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Neurons in the substantia nigra pars reticulata (SNr) are known to encode saccadic eye movements within some, but not all, behavioral contexts. However, the precise contextual factors that effect the modulations of nigral activity are still uncertain. To further examine the effect of behavioral context on the SNr, we recorded the activity of 72 neurons while monkeys made saccades during a delayed saccade task and during periods of free viewing. We quantified and compared the movement fields of each neuron for saccades made under three different conditions: 1) spontaneous saccades, which shifted gaze during periods of free viewing when no stimuli were presented and no reinforcements were delivered; 2) fixational saccades, which brought gaze into alignment with a fixation target at the start of a delayed saccade trial, were necessary for trial completion, but were not directly followed by reinforcement; and 3) terminal saccades, which brought gaze into alignment with a visual target at the end of a delayed saccade trial and were directly followed by reinforcement. For three of the four SNr neuron classes, saccade-related modulations were only present before terminal saccades. For the fourth class, discrete pausers, saccade-related modulations were substantially larger for terminal saccades than for fixational saccades, and modulations were absent for spontaneous saccades. These results and other recent work on the basal ganglia suggest that some saccade-related signals in the SNr may be influenced by the reinforcement associated with a particular saccadic eye movement.
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Henik, Avishai, Robert Rafal i Dell Rhodes. "Endogenously Generated and Visually Guided Saccades after Lesions of the Human Frontal Eye Fields". Journal of Cognitive Neuroscience 6, nr 4 (lipiec 1994): 400–411. http://dx.doi.org/10.1162/jocn.1994.6.4.400.
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Nine patients with chronic, unilateral lesions of the dorso-lateral prefrontal cortex including the frontal eye fields (FEF) made saccades toward contralesional and ipsilesional fields. The saccades were either voluntarily directed in response to arrows in the center of a visual display, or were reflexively summoned by a peripheral visual signal. Saccade latencies were compared to those made by seven neurologic control patients with chronic, unilateral lesions of dorsolateral prefrontal cortex sparing the FEF, and by 13 normal control subjects. In both the normal and neurologic control subjects, reflexive saccades had shorter Latencies than voluntary saccades. In the FEF lesion patients, voluntary saccades had longer latencies toward the contralesional field than toward the ipsilesional field. The opposite pattern was found for reflexive saccades: latencies of saccades to targets in the contralesional field were shorter than saccades summoned to ipsilesional targets. Reflexive saccades toward the ipsilesional field had abnormally prolonged latencies; they were comparable to the latencies observed for voluntary Saccades. The effect of FEF lesions on saccacles contrasted with those observed in a second experiment requiring a key press response: FEF lesion patients were slower in making key press responses to signals detected in the contralesional field. To assess covert attention and preparatory set the effects of precues providing advance information were measured in both saccade and key press experiments. Neither patient group showed any deficiency in using precues to shift attention or to prepare saccades. The FEF facilitates the generation of voluntary saccatles and also inhibits reflexive saccades to exogenous signals. FEF lesions may disinhibit the ipsilesional midbrain which in turn may inhibit the opposite colliculus to slow reflexive saccades toward the ipsilesional field.
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Havermann, Katharina, Eckart Zimmermann i Markus Lappe. "Eye position effects in saccadic adaptation". Journal of Neurophysiology 106, nr 5 (listopad 2011): 2536–45. http://dx.doi.org/10.1152/jn.00023.2011.
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Saccades are used by the visual system to explore visual space with the high accuracy of the fovea. The visual error after the saccade is used to adapt the control of subsequent eye movements of the same amplitude and direction in order to keep saccades accurate. Saccadic adaptation is thus specific to saccade amplitude and direction. In the present study we show that saccadic adaptation is also specific to the initial position of the eye in the orbit. This is useful, because saccades are normally accompanied by head movements and the control of combined head and eye movements depends on eye position. Many parts of the saccadic system contain eye position information. Using the intrasaccadic target step paradigm, we adaptively reduced the amplitude of reactive saccades to a suddenly appearing target at a selective position of the eyes in the orbitae and tested the resulting amplitude changes for the same saccade vector at other starting positions. For central adaptation positions the saccade amplitude reduction transferred completely to eccentric starting positions. However, for adaptation at eccentric starting positions, there was a reduced transfer to saccades from central starting positions or from eccentric starting positions in the opposite hemifield. Thus eye position information modifies the transfer of saccadic amplitude changes in the adaptation of reactive saccades. A gain field mechanism may explain the eye position dependence found.
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Ohtsuka, K., i H. Noda. "Saccadic burst neurons in the oculomotor region of the fastigial nucleus of macaque monkeys". Journal of Neurophysiology 65, nr 6 (1.06.1991): 1422–34. http://dx.doi.org/10.1152/jn.1991.65.6.1422.
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1. The discharge of 255 neurons in the fastigial nuclei of three trained macaque monkeys was investigated during visually guided saccades. Responses of these neurons were examined also during horizontal head rotation and during microstimulation of the oculomotor vermis (lobules VIc and VII). 2. One hundred and two units were characterized by bursts of firing in response to visually guided saccades. Ninety-eight of these (96.1%) were located within the anatomic confines of the fastigial oculomotor region (FOR), on the basis of reconstruction of recording sites. During contralateral saccades, these neurons showed bursts that preceded the onset of saccades (presaccadic burst), whereas, during ipsilateral saccades, they showed bursts associated with the end of saccades (late saccadic burst). They were hence named saccadic burst neurons. Sixty-one saccadic burst neurons (62.2%) were inhibited during microstimulation of the oculomotor vermis with currents less than 10 microA. 3. All saccadic burst neurons were spontaneously active, and the resting firing rate varied considerably among units, ranging from 10 to 50 imp/s. The tonic levels of activity did not correlate significantly with eye position. 4. The presaccadic burst started 18.5 +/- 4.7 (SD) ms (n = 45) before the onset of saccades in the optimal direction (the direction associated with the maximum values of burst lead time, number of spikes per burst, and burst duration). Optimal directions covered the entire contralateral hemifield, although there was a slightly higher incidence in both horizontal and upper-oblique directions in the present sample. The duration of the presaccadic burst was highly correlated with the duration of saccade (0.85 less than or equal to r less than or equal to 0.97). 5. The late saccadic burst was most robust in the direction opposite to the optimal in each unit (the nonoptimal direction). Its onset preceded the completion of ipsilateral saccade by 30.4 +/- 5.9 ms. The lead time to the end of saccade was consistent among different units and was constant also for saccades of various sizes. Thus the late saccadic burst started even before the saccade onset when the saccade duration was less than 30 ms. Unlike the presaccadic burst, its duration was not related to the duration of saccade. 6. Discharge rates of saccadic burst neurons were correlated neither to eye positions during fixation nor to the initial eye positions before saccades. 7. Eye-position units and horizontal head-velocity units were located rostral to the FOR.(ABSTRACT TRUNCATED AT 400 WORDS)
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Ohtsuka, K., i H. Noda. "Discharge properties of Purkinje cells in the oculomotor vermis during visually guided saccades in the macaque monkey". Journal of Neurophysiology 74, nr 5 (1.11.1995): 1828–40. http://dx.doi.org/10.1152/jn.1995.74.5.1828.
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1. We previously described discharge properties of cerebellar output cells in the fastigial nucleus during ipsilateral and contralateral saccades. Fastigial cells exhibited unique responses depending on the direction of saccades and were involved in execution of accurate targeting saccades. Purkinje cells in the oculomotor vermis (lobules VIc and VII) are thought to modulate these discharges of fastigial cells. In this study we reexamine discharge properties of Purkinje cells on the basis of this hypothesis. 2. Initially we physiologically identified the right and left sides of the oculomotor vermis. Saccade-related discharges of 79 Purkinje cells were recorded from both sides of the vermis during visually guided saccades toward the sides ipsilateral and contralateral to the recording side in two trained macaque monkeys. To clarify the correlation of Purkinje cell discharge with burst activities in the fastigial nucleus during saccadic eye movements, we analyzed our data by employing methods used in the study of fastigial neurons. 3. Among the 79 cells, 56 (71%) showed burst discharges during saccades (saccadic burst cells). Of the 56 cells, 29 exhibited a peak of burst discharges in both the contralateral and ipsilateral directions (bidirectional cells). The remaining 27 saccadic burst cells showed a peak of burst discharges during either contralateral or ipsilateral saccades (unidirectional cells). Among the 79 cells, 14 (18%) exhibited a pause of discharges during contralateral saccades (pause cells). Among the 79 cells, 9 (11%) showed burst discharge during contralateral saccades followed by tonic discharge that was correlated with eye position (burst tonic cells). 4. The timing of bursts in bidirectional cells with respect to saccade onset was dependent on the direction of saccade. During ipsilateral saccades, Purkinje cells exhibited a long lead burst that built up gradually, peaked near the onset of the saccade, and terminated sharply near midsaccade. The mean lead time relative to saccade onset was 29.3 +/- 24.5 (SD) ms. During contralateral saccades, Purkinje cells exhibited a short lead/late burst that built up sharply, peaked near midsaccade, and terminated gradually after the end of the saccade. The mean lead time relative to saccade onset was 10.7 +/- 20.8 ms. The burst onset time during contralateral saccades and the burst offset time during ipsilateral saccades preceded the saccade offset time by about the same interval regardless of the saccade amplitude. 5. In pause cells the pause preceded saccade onset by 17.5 +/- 10.6 ms. The duration of the pause was not correlated with the duration of saccades. There was little trial-to-trial variability in the onset time of the pause with respect to the onset of saccades, whereas there was large trial-to-trial variability in the offset time of the pause with respect to the offset of saccades. In addition, the mean onset time of the pause for each cell had a relatively narrow distribution. 6. The burst lead time of burst tonic cells relative to saccade onset was 9.5 +/- 3.9 ms. The tonic discharge rate of burst tonic cells was a nonlinear function of eye position. The regression of each cell was fit to two lines. The regression coefficient ranged from 0.95 to 0.99 (mean = 0.97). 7. Axons of Purkinje cells in the oculomotor vermis are thought to project exclusively to saccadic burst cells in the fastigial oculomotor region (FOR), which is located in the caudal portion of the fastigial nucleus. Our previous studies indicated that FOR cells provide temporal signals for controlling targeting saccades. The present results suggest that Purkinje cells in the oculomotor vermis modify the temporal signals of FOR cells for saccades in different directions and amplitudes. The modification of FOR cell activity by Purkinje cells is thought to be essential for the function of the cerebellum in the control of saccadic eye movements.
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Crowder, Nathan A., Nicholas S. C. Price, Michael J. Mustari i Michael R. Ibbotson. "Direction and Contrast Tuning of Macaque MSTd Neurons During Saccades". Journal of Neurophysiology 101, nr 6 (czerwiec 2009): 3100–3107. http://dx.doi.org/10.1152/jn.91254.2008.
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Saccades are rapid eye movements that change the direction of gaze, although the full-field image motion associated with these movements is rarely perceived. The attenuation of visual perception during saccades is referred to as saccadic suppression. The mechanisms that produce saccadic suppression are not well understood. We recorded from neurons in the dorsal medial superior temporal area (MSTd) of alert macaque monkeys and compared the neural responses produced by the retinal slip associated with saccades (active motion) to responses evoked by identical motion presented during fixation (passive motion). We provide evidence for a neural correlate of saccadic suppression and expand on two contentious results from previous studies. First, we confirm the finding that some neurons in MSTd reverse their preferred direction during saccades. We quantify this effect by calculating changes in direction tuning index for a large cell population. Second, it has been noted that neural activity associated with saccades can arrive in the parietal cortex ≤30 ms earlier than activity produced by similar visual stimulation during fixation. This led to the question of whether the saccade-related responses were visual in origin or were motor signals arising from saccade-planning areas of the brain. By comparing the responses to saccades made over textured backgrounds of different contrasts, we provide strong evidence that saccade-related responses were visual in origin. Refinements of the possible models of saccadic suppression are discussed.
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Schnier, Fabian, i Markus Lappe. "Mislocalization of stationary and flashed bars after saccadic inward and outward adaptation of reactive saccades". Journal of Neurophysiology 107, nr 11 (1.06.2012): 3062–70. http://dx.doi.org/10.1152/jn.00877.2011.
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Recent studies have shown that saccadic inward adaptation (i.e., the shortening of saccade amplitude) and saccadic outward adaptation (i.e., the lengthening of saccade amplitude) rely on partially different neuronal mechanisms. There is increasing evidence that these differences are based on differences at the target registration or planning stages since outward but not inward adaptation transfers to hand-pointing and perceptual localization of flashed targets. Furthermore, the transfer of reactive saccade adaptation to long-duration overlap and scanning saccades is stronger after saccadic outward adaptation than that after saccadic inward adaptation, suggesting that modulated target registration stages during outward adaptation are increasingly used in the execution of saccades when the saccade target is visually available for a longer time. The difference in target presentation duration between reactive and scanning saccades is also linked to a difference in perceptual localization of different targets. Flashed targets are mislocalized after inward adaptation of reactive and scanning saccades but targets that are presented for a longer time (stationary targets) are mislocalized stronger after scanning than after reactive saccades. This link between perceptual localization and adaptation specificity suggests that mislocalization of stationary bars should be higher after outward than that after inward adaptation of reactive saccades. In the present study we test this prediction. We show that the relative amount of mislocalization of stationary versus flashed bars is higher after outward than that after inward adaptation of reactive saccades. Furthermore, during fixation stationary and flashed bars were mislocalized after outward but not after inward adaptation. Thus, our results give further evidence for different adaptation mechanisms between inward and outward adaptation and harmonize some recent research.
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Lueck, C. J., S. Tanyeri, T. J. Crawford, L. Henderson i C. Kennard. "Saccadic Eye Movements in Parkinson's Disease: I. Delayed Saccades". Quarterly Journal of Experimental Psychology Section A 45, nr 2 (sierpień 1992): 193–210. http://dx.doi.org/10.1080/14640749208401324.
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The saccadic eye movements of nine patients with Parkinson's disease were compared to those of nine age-matched controls in two paradigms generating volitional saccades. In both paradigms, subjects had to make delayed saccades to peripheral LED targets: a peripheral target appeared 700 msec before a buzzer sounded, the buzzer being the signal to make a saccade to the target. In the first paradigm (“centre-off”), the fixation target was extinguished simultaneously with buzzer onset. In the second (“centre-remain”) it was not extinguished until 1000 msec later. The results showed that for outward saccades in both paradigms, there was no difference between Parkinsonian patients and controls, but saccadic latencies were significantly shorter in the “centre-remain” paradigm. The initial outward saccades were indistinguishable from the normal, reflex saccades of the same subjects. However, saccades returning to the centre (a type of remembered target saccade) were hypometric and showed multistepping. Both effects were more pronounced in patients with Parkinson's disease. The significance of these findings in terms of current hypotheses about the nature of the Parkinsonian saccadic deficit is discussed.
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Bibi, Raquel, i Jay A. Edelman. "The Influence of Motor Training on Human Express Saccade Production". Journal of Neurophysiology 102, nr 6 (grudzień 2009): 3101–10. http://dx.doi.org/10.1152/jn.90710.2008.
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Express saccadic eye movements are saccades of extremely short latency. In monkey, express saccades have been shown to occur much more frequently when the monkey has been trained to make saccades in a particular direction to targets that appear in predictable locations. Such results suggest that express saccades occur in large number only under highly specific conditions, leading to the view that vector-specific training and motor preparatory processes are required to make an express saccade of a particular magnitude and direction. To evaluate this hypothesis in humans, we trained subjects to make saccades quickly to particular locations and then examined whether the frequency of express saccades depended on training and the number of possible target locations. Training significantly decreased saccade latency and increased express saccade production to both trained and untrained locations. Increasing the number of possible target locations (two vs. eight possible targets) led to only a modest increase of saccade latency. For most subjects, the probability of express saccade occurrence was much higher than that expected if vector-specific movement preparation were necessary for their production. These results suggest that vector-specific motor preparation and vector-specific saccade training are not necessary for express saccade production in humans and that increases in express saccade production are due in part to a facilitation in fixation disengagement or else a general enhancement in the ability of the saccadic system to respond to suddenly appearing visual stimuli.
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Aizawa, Hiroshi, i Robert H. Wurtz. "Reversible Inactivation of Monkey Superior Colliculus. I. Curvature of Saccadic Trajectory". Journal of Neurophysiology 79, nr 4 (1.04.1998): 2082–96. http://dx.doi.org/10.1152/jn.1998.79.4.2082.
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Aizawa, Hiroshi and Robert H. Wurtz. Reversible inactivation of monkey superior colliculus. I. Curvature of saccadic trajectory. J. Neurophysiol. 79: 2082–2096, 1998. The neurons in the intermediate layers of the monkey superior colliculus (SC) that discharge before saccadic eye movements can be divided into at least two types, burst and buildup neurons, and the differences in their characteristics are compatible with different functional contributions of the two cell types. It has been suggested that a spread of activity across the population of the buildup neurons during saccade generation may contribute to the control of saccadic eye movements. The influence of any such spread should be on both the horizontal and vertical components of the saccade because the map of the movement fields on the SC is a two-dimensional one; it should affect the trajectory of saccade. The present experiments used muscimol injections to inactivate areas within the SC to determine the functional contribution of such a spread of activity on the trajectory of the saccades. The analysis concentrated on saccades made to areas of the visual field that should be affected primarily by alteration of buildup neuron activity. Muscimol injections produced saccades with altered trajectories; they became consistently curved after the injection, and successive saccades to the same targets had similar curvatures. The curved saccades showed changes in their direction and speed at the very beginning of the saccade, and for those saccades that reached the target, the direction of the saccade was altered near the end to compensate for the initially incorrect direction. Postinjection saccades had lower peak speeds, longer durations, and longer latencies for initiation. The changes in saccadic trajectories resulting from muscimol injections, along with the previous observations on changes in speed of saccades with such injections, indicate that the SC is involved in influencing the eye position during the saccade as well as at the end of the saccade. The changes in trajectory when injections were made more rostral in the SC than the most active burst neurons also are consistent with a contribution of the buildup neurons to the control of the eye trajectory. The results do not, however, support the hypothesis that the buildup neurons in the SC act as a spatial integrator.
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Burman, Douglas D., i Charles J. Bruce. "Suppression of Task-Related Saccades by Electrical Stimulation in the Primate's Frontal Eye Field". Journal of Neurophysiology 77, nr 5 (1.05.1997): 2252–67. http://dx.doi.org/10.1152/jn.1997.77.5.2252.
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Burman, Douglas D. and Charles J. Bruce. Suppression of task-related saccades by electrical stimulation in the primate's frontal eye field. J. Neurophysiol. 77: 2252–2267, 1997. Patients with frontal lobe damage have difficulty suppressing reflexive saccades to salient visual stimuli, indicating that frontal lobe neocortex helps to suppress saccades as well as to produce them. In the present study, a role for the frontal eye field (FEF) in suppressing saccades was demonstrated in macaque monkeys by application of intracortical microstimulation during the performance of a visually guided saccade task, a memory prosaccade task, and a memory antisaccade task. A train of low-intensity (20–50 μA) electrical pulses was applied simultaneously with the disappearance of a central fixation target, which was always the cue to initiate a saccade. Trials with and without stimulation were compared, and significantly longer saccade latencies on stimulation trials were considered evidence of suppression. Low-intensity stimulation suppressed task-related saccades at 30 of 77 sites tested. In many cases saccades were suppressed throughout the microstimulation period (usually 450 ms) and then executed shortly after the train ended. Memory-guided saccades were most dramatically suppressed and were often rendered hypometric, whereas visually guided saccades were less severely suppressed by stimulation. At 18 FEF sites, the suppression of saccades was the only observable effect of electrical stimulation. Contraversive saccades were usually more strongly suppressed than ipsiversive ones, and cells recorded at such purely suppressive sites commonly had either foveal receptive fields or postsaccadic responses. At 12 other FEF sites at which saccadic eye movements were elicited at low thresholds, task-related saccades whose vectors differed from that of the electrically elicited saccade were suppressed by electrical stimulation. Such suppression at saccade sites was observed even with currents below the threshold for eliciting saccades. Pure suppression sites tended to be located near or in the fundus, deeper in the anterior bank of the arcuate than elicited saccade sites. Stimulation in the prefrontal association cortex anterior to FEF did not suppress saccades, nor did stimulation in premotor cortex posterior to FEF. These findings indicate that the primate FEF can help orchestrate saccadic eye movements by suppressing inappropriate saccade vectors as well as by selecting, specifying, and triggering appropriate saccades. We hypothesize that saccades could be suppressed both through local FEF interactions and through FEF projections to subcortical regions involved in maintaining fixation.
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Lévy-Bencheton, Delphine, Denis Pélisson, Muriel Panouillères, Christian Urquizar, Caroline Tilikete i Laure Pisella. "Adaptation of scanning saccades co-occurs in different coordinate systems". Journal of Neurophysiology 111, nr 12 (15.06.2014): 2505–15. http://dx.doi.org/10.1152/jn.00733.2013.
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Plastic changes of saccades (i.e., following saccadic adaptation) do not transfer between oppositely directed saccades, except when multiple directions are trained simultaneously, suggesting a saccadic planning in retinotopic coordinates. Interestingly, a recent study in human healthy subjects revealed that after an adaptive increase of rightward-scanning saccades, both leftward and rightward double-step, memory-guided saccades, triggered toward the adapted endpoint, were modified, revealing that target location was coded in spatial coordinates ( Zimmermann et al. 2011 ). However, as the computer screen provided a visual frame, one alternative hypothesis could be a coding in allocentric coordinates. Here, we questioned whether adaptive modifications of saccadic planning occur in multiple coordinate systems. We reproduced the paradigm of Zimmermann et al. (2011) using target light-emitting diodes in the dark, with and without a visual frame, and tested different saccades before and after adaptation. With double-step, memory-guided saccades, we reproduced the transfer of adaptation to leftward saccades with the visual frame but not without, suggesting that the coordinate system used for saccade planning, when the frame is visible, is allocentric rather than spatiotopic. With single-step, memory-guided saccades, adaptation transferred to leftward saccades, both with and without the visual frame, revealing a target localization in a coordinate system that is neither retinotopic nor allocentric. Finally, with single-step, visually guided saccades, the classical, unidirectional pattern of amplitude change was reproduced, revealing retinotopic coordinate coding. These experiments indicate that the same procedure of adaptation modifies saccadic planning in multiple coordinate systems in parallel—each of them revealed by the use of different saccade tasks in postadaptation.
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Wallman, Josh, i Albert F. Fuchs. "Saccadic Gain Modification: Visual Error Drives Motor Adaptation". Journal of Neurophysiology 80, nr 5 (1.11.1998): 2405–16. http://dx.doi.org/10.1152/jn.1998.80.5.2405.
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Wallman, Josh and Albert F. Fuchs. Saccadic gain modification: visual error drives motor adaptation. J. Neurophysiol. 80: 2405–2416, 1998. The brain maintains the accuracy of saccadic eye movements by adjusting saccadic amplitude relative to the target distance (i.e., saccade gain) on the basis of the performance of recent saccades. If an experimenter surreptitiously moves the target backward during each saccade, thereby causing the eyes to land beyond their targets, saccades undergo a gradual gain reduction. The error signal driving this conventional saccadic gain adaptation could be either visual (the postsaccadic distance of the target from the fovea) or motoric (the direction and size of the corrective saccade that brings the eye onto the back-stepped target). Similarly, the adaptation itself might be a motor adjustment (change in the size of saccade for a given perceived target distance) or a visual remapping (change in the perceived target distance). We studied these possibilities in experiments both with rhesus macaques and with humans. To test whether the error signal is motoric, we used a paradigm devised by Heiner Deubel. The Deubel paradigm differed from the conventional adaptation paradigm in that the backward step that occurred during the saccade was brief, and the target then returned to its original displaced location. This ploy replaced most of the usual backward corrective saccades with forward ones. Nevertheless, saccadic gain gradually decreased over hundreds of trials. Therefore, we conclude that the direction of saccadic gain adaptation is not determined by the direction of corrective saccades. To test whether gain adaptation is a manifestation of a static visual remapping, we decreased the gain of 10° horizontal saccades by conventional adaptation and then tested the gain to targets appearing at retinal locations unused during adaptation. To make the target appear in such “virgin territory,” we had it jump first vertically and then 10° horizontally; both jumps were completed and the target spot extinguished before saccades were made sequentially to the remembered target locations. Conventional adaptation decreased the gain of the second, horizontal saccade even though the target was in a nonadapted retinal location. In contrast, the horizontal component of oblique saccades made directly to the same virgin location showed much less gain decrease, suggesting that the adaptation is specific to saccade direction rather than to target location. Thus visual remapping cannot account for the entire reduction of saccadic gain. We conclude that saccadic gain adaptation involves an error signal that is primarily visual, not motor, but that the adaptation itself is primarily motor, not visual.
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King, W. M., S. G. Lisberger i A. F. Fuchs. "Oblique saccadic eye movements of primates". Journal of Neurophysiology 56, nr 3 (1.09.1986): 769–84. http://dx.doi.org/10.1152/jn.1986.56.3.769.
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The objective of these experiments was to determine whether the trajectories of the horizontal and vertical components of oblique saccades in primates were coupled. Human and monkey eye movements were recorded during a visual tracking task that jumped a small visible target spot to different locations on a tangent screen. For oblique saccades larger than ca. 3 deg, there was coupling between the horizontal and vertical components so that the duration of the smaller component was longer ("stretched") than would have been expected from its amplitude-duration relationship. The duration of a stretched component of an oblique saccade was linearly related to the vector amplitude of the eye movement but not to the amplitude of the stretched component. Stretched components of oblique saccades had lower peak and average velocities than would have occurred with pure horizontal or vertical saccades of the same size. Decreased component velocity was not caused by low-velocity eye movement components inserted at the beginning or end of the saccade, but was a function of the saccade's direction and component amplitude. For any saccade, there was a linear relationship between peak and average component velocity. We compared the discharge of monkey abducens neurons with the characteristics of the on-direction horizontal components of oblique saccades. The burst duration of an abducens neuron was lengthened when the horizontal component of an oblique saccade was stretched. Intraburst firing frequency was also decreased in correspondence with a decrease in horizontal component velocity. For an oblique saccade, the duration of the neuron's burst was correlated with the duration of the horizontal component and with the vector amplitude of the saccade, but was not correlated with the amplitude of the horizontal component itself. The duration of the smaller component of an oblique saccade was proportional but not always equal to the duration of the larger component. Usually, the smaller component began later and ended earlier than the larger component. These results show that the horizontal and vertical components of oblique saccades are coupled centrally so that the velocity of the smaller component is decreased and its duration is increased. For oblique saccades, larger than ca. 3 deg, amplitude-duration and amplitude-velocity relationships based on pure horizontal or vertical saccade data are not applicable. These findings are discussed in relation to three recently proposed models of coupled saccadic burst generators.
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Fuchs, A. F., D. Reiner i M. Pong. "Transfer of gain changes from targeting to other types of saccade in the monkey: constraints on possible sites of saccadic gain adaptation". Journal of Neurophysiology 76, nr 4 (1.10.1996): 2522–35. http://dx.doi.org/10.1152/jn.1996.76.4.2522.
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1. Our goal was to use behavioral experiments to delimit where in the simian oculomotor system the gain of horizontal saccadic eye movements might be controlled. Our strategy was to change the gain of saccades to visual target steps (called targeting saccades) and to examine whether these changes transferred to other types of saccades. We reduced the gain of targeting saccades by jumping the target backward as a saccade was made so that the saccade appeared to overshoot. After 1,000-1,500 saccades to such backstepping targets, the average overshoot, and therefore the saccadic gain, had decreased substantially. 2. After the gain of targeting saccades had been reduced by 15-22%, several kinds of saccades were tested. Most were elicited by various visual targets. Some were made to jumping targets, which were timed to elicit saccades with longer (delayed saccades) or shorter (express saccades) latencies than normal or to targets that disappeared after a brief exposure (memory-guided saccades). Others were elicited to stationary targets (self-paced saccades) or in pursuit of a smoothly moving target (catchup saccades). Finally, we tested the saccadic fast phases of vestibular and optokinetic nystagmus. 3. Gain reduction of targeting saccades transferred at least partially to all the other types of saccades made to target jumps. The percentage gain transfer was calculated as (gain reduction of test saccades)/(gain reduction of adapted targeting saccades). The average percent transfer to delayed, memory-guided, and express saccades was 96, 88, and 91%, respectively. 4. Monkeys also showed substantial gain transfer to self-paced saccades, which scanned stationary targets. The average percentage gain transfer was 69% in the four animals tested. When two humans performed the same task, there was no transfer at all. These data suggest that saccadic gain adjustment involves different processes in monkeys and humans. 5. The transfer of gain to the catchup saccades of smooth pursuit varied from 41 to 100% across the four monkeys tested. Nevertheless, the average percentage gain transfer for all the animals was 75%. 6. As judged by the amplitude distribution of fast phases before and after adaptation, there was little, if any, saccadic gain transfer to the fast phases of vestibular or optokinetic nystagmus. In 12 of 13 experiments, there was no significant decrease in fast phase amplitude after a gain reduction of targeting saccades (P > 0.1). 7. This study shows that the average percentage gain transfer from targeting to delayed, express, memory-guided, self-paced, and catchup saccades was never < 69%. Although there was substantial transfer to saccades elicited by jumping, stationary, remembered, or slowly moving visual targets, there was relatively little to the saccadelike fast phases of nystagmus. The transfer of saccadic gain to the very short-latency express saccades suggests that adaptation modifies a subcortical locus. Moreover, the major locus must lie only in the premotor pathway for visual saccades, because saccadic gain adaptation is only poorly transferred to the fast phases of vestibular and optokinetic nystagmus.
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Goossens, H. H. L. M., i A. J. Van Opstal. "Blink-Perturbed Saccades in Monkey. I. Behavioral Analysis". Journal of Neurophysiology 83, nr 6 (1.06.2000): 3411–29. http://dx.doi.org/10.1152/jn.2000.83.6.3411.
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Saccadic eye movements are thought to be influenced by blinking through premotor interactions, but it is still unclear how. The present paper describes the properties of blink-associated eye movements and quantifies the effect of reflex blinks on the latencies, metrics, and kinematics of saccades in the monkey. In particular, it is examined to what extent the saccadic system accounts for blink-related perturbations of the saccade trajectory. Trigeminal reflex blinks were elicited near the onset of visually evoked saccades by means of air puffs directed on the eye. Reflex blinks were also evoked during a straight-ahead fixation task. Eye and eyelid movements were measured with the magnetic-induction technique. The data show that saccade latencies were reduced substantially when reflex blinks were evoked prior to the impending visual saccades as if these saccades were triggered by the blink. The evoked blinks also caused profound spatial-temporal perturbations of the saccades. Deflections of the saccade trajectory, usually upward, extended up to ∼15°. Saccade peak velocities were reduced, and a two- to threefold increase in saccade duration was typically observed. In general, these perturbations were largely compensated in saccade mid-flight, despite the absence of visual feedback, yielding near-normal endpoint accuracies. Further analysis revealed that blink-perturbed saccades could not be described as a linear superposition of a pure blink-associated eye movement and an unperturbed saccade. When evoked during straight-ahead fixation, blinks were accompanied by initially upward and slightly abducting eye rotations of ∼2–15°. Back and forth wiggles of the eye were frequently seen; but in many cases the return movement was incomplete. Rather than drifting back to its starting position, the eye then maintained its eccentric orbital position until a downward corrective saccade toward the fixation spot followed. Blink-associated eye movements were quite rapid, albeit slower than saccades, and the velocity-amplitude-duration characteristics of the initial excursions as well as the return movements were approximately linear. These data strongly support the idea that blinks interfere with the saccade premotor circuit, presumably upstream from the neural eye-position integrator. They also indicated that a neural mechanism, rather than passive elastic restoring forces within the oculomotor plant, underlies the compensatory behavior. The tight latency coupling between saccades and blinks is consistent with an inhibition of omnipause neurons by the blink system, suggesting that the observed changes in saccade kinematics arise elsewhere in the saccadic premotor system.
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Asscheman, Susanne J., Katharine N. Thakkar i Sebastiaan F. W. Neggers. "Changes in Effective Connectivity of the Superior Parietal Lobe during Inhibition and Redirection of Eye Movements". Journal of Experimental Neuroscience 9s1 (styczeń 2015): JEN.S32736. http://dx.doi.org/10.4137/jen.s32736.
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Executive control is the ability to flexibly control behavior and is frequently studied with saccadic eye movements. Contrary to frontal oculomotor areas, the role of the superior parietal lobe (SPL) in the executive control of saccades remains unknown. To explore the role of SPL networks in saccade control, we performed a saccadic search-step task while acquiring functional magnetic resonance imaging data for 41 participants. Psychophysiological interaction analyses assessed task-related differences in the effective connectivity of SPL with other brain regions during the inhibition and redirection of saccades. Results indicate an increased coupling of SPL with frontal, posterior, and striatal oculomotor areas for redirected saccades versus visually guided saccades. Saccade inhibition versus unsuccessful inhibition revealed an increased coupling of SPL with dorsolateral prefrontal cortex and anterior cingulate cortex. We discuss how these findings relate to ongoing debates about the implementation of executive control and conclude that early attentional control and rapid updating of saccade goals are important signals for executive control.
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Lueck, C. J., T. J. Crawford, L. Henderson, J. A. M. Van Gisbergen, J. Duysens i C. Kennard. "Saccadic Eye Movements in Parkinson's Disease: II. Remembered Saccades— towards a Unified Hypothesis?" Quarterly Journal of Experimental Psychology Section A 45, nr 2 (sierpień 1992): 211–33. http://dx.doi.org/10.1080/14640749208401325.
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Ten patients with mild to moderate Parkinson's disease were compared with ten age-matched normal controls in a series of saccadic paradigms in order to test various hypotheses relating to the origin of the Parkinsonian saccadic defect. The paradigms comprised a reflex saccade paradigm, a standard remembered saccade paradigm, a remembered saccade paradigm with delayed centre-offset, and a remembered saccade paradigm with a second target flash immediately prior to saccade execution. Finally, subjects executed both reflex and remembered saccades in a standard remembered paradigm (the “two-saccade” paradigm). As has been reported previously, Parkinsonian subjects demonstrated hypometria on all remembered saccade paradigms, particularly the “two-saccade” paradigm. There was, however, no significant difference between the first three remembered saccade paradigms. These studies serve to refute a simple attentional capture hypothesis, and a hypothesis that suggests that the abnormality of remembered saccades is due to concurrent reflex saccade suppression. On the basis of the results, further hypotheses are advanced in an attempt to explain all published work on Parkinsonian saccades.
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Chen, Jing, Matteo Valsecchi i Karl R. Gegenfurtner. "Saccadic suppression measured by steady-state visual evoked potentials". Journal of Neurophysiology 122, nr 1 (1.07.2019): 251–58. http://dx.doi.org/10.1152/jn.00712.2018.
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Visual sensitivity is severely impaired during the execution of saccadic eye movements. This phenomenon has been extensively characterized in human psychophysics and nonhuman primate single-neuron studies, but a physiological characterization in humans is less established. Here, we used a method based on steady-state visually evoked potential (SSVEP), an oscillatory brain response to periodic visual stimulation, to examine how saccades affect visual sensitivity. Observers made horizontal saccades back and forth, while horizontal black-and-white gratings flickered at 5–30 Hz in the background. We analyzed EEG epochs with a length of 0.3 s either centered at saccade onset (saccade epochs) or centered at fixations half a second before the saccade (fixation epochs). Compared with fixation epochs, saccade epochs showed a broadband power increase, which most likely resulted from saccade-related EEG activity. The execution of saccades, however, led to an average reduction of 57% in the SSVEP amplitude at the stimulation frequency. This result provides additional evidence for an active saccadic suppression in the early visual cortex in humans. Compared with previous functional MRI and EEG studies, an advantage of this approach lies in its capability to trace the temporal dynamics of neural activity throughout the time course of a saccade. In contrast to previous electrophysiological studies in nonhuman primates, we did not find any evidence for postsaccadic enhancement, even though simulation results show that our method would have been able to detect it. We conclude that SSVEP is a useful technique to investigate the neural correlates of visual perception during saccadic eye movements in humans. NEW & NOTEWORTHY We make fast ballistic saccadic eye movements a few times every second. At the time of saccades, visual sensitivity is severely impaired. The present study uses steady-state visually evoked potentials to reveal a neural correlate of the fine temporal dynamics of these modulations at the time of saccades in humans. We observed a strong reduction (57%) of visually driven neural activity associated with saccades but did not find any evidence for postsaccadic enhancement.
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Soetedjo, Robijanto, Chris R. S. Kaneko i Albert F. Fuchs. "Evidence That the Superior Colliculus Participates in the Feedback Control of Saccadic Eye Movements". Journal of Neurophysiology 87, nr 2 (1.02.2002): 679–95. http://dx.doi.org/10.1152/jn.00886.2000.
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There is general agreement that saccades are guided to their targets by means of a motor error signal, which is produced by a local feedback circuit that calculates the difference between desired saccadic amplitude and an internal copy of actual saccadic amplitude. Although the superior colliculus (SC) is thought to provide the desired saccadic amplitude signal, it is unclear whether the SC resides in the feedback loop. To test this possibility, we injected muscimol into the brain stem region containing omnipause neurons (OPNs) to slow saccades and then determined whether the firing of neurons at different sites in the SC was altered. In 14 experiments, we produced saccadic slowing while simultaneously recording the activity of a single SC neuron. Eleven of the 14 neurons were saccade-related burst neurons (SRBNs), which discharged their most vigorous burst for saccades with an optimal amplitude and direction (optimal vector). The optimal directions for the 11 SRBNs ranged from nearly horizontal to nearly vertical, with optimal amplitudes between 4 and 17°. Although muscimol injections into the OPN region produced little change in the optimal vector, they did increase mean saccade duration by 25 to 192.8% and decrease mean saccade peak velocity by 20.5 to 69.8%. For optimal vector saccades, both the acceleration and deceleration phases increased in duration. However, during 10 of 14 experiments, the duration of deceleration increased as fast as or faster than that of acceleration as saccade duration increased, indicating that most of the increase in duration occurred during the deceleration phase. SRBNs in the SC changed their burst duration and firing rate concomitantly with changes in saccadic duration and velocity, respectively. All SRBNs showed a robust increase in burst duration as saccadic duration increased. Five of 11 SRBNs also exhibited a decrease in burst peak firing rate as saccadic velocity decreased. On average across the neurons, the number of spikes in the burst was constant. There was no consistent change in the discharge of the three SC neurons that did not exhibit bursts with saccades. Our data show that the SC receives feedback from downstream saccade-related neurons about the ongoing saccades. However, the changes in SC firing produced in our study do not suggest that the feedback is involved with producing motor error. Instead, the feedback seems to be involved with regulating the duration of the discharge of SRBNs so that the desired saccadic amplitude signal remains present throughout the saccade.
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Blohm, Gunnar, Marcus Missal i Philippe Lefèvre. "Processing of Retinal and Extraretinal Signals for Memory-Guided Saccades During Smooth Pursuit". Journal of Neurophysiology 93, nr 3 (marzec 2005): 1510–22. http://dx.doi.org/10.1152/jn.00543.2004.
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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.
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Irwin, David E., i Laura A. Carlson-Radvansky. "Cognitive Suppression During Saccadic Eye Movements". Psychological Science 7, nr 2 (marzec 1996): 83–88. http://dx.doi.org/10.1111/j.1467-9280.1996.tb00334.x.
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Saccadic eye movements are made at least 100,000 times each day It is well known that sensitivity to visual input is suppressed during saccades, we examined whether cognitive activity (specifically, mental rotation) is suppressed as well If cognitive processing occurs during saccades, a prime viewed in one fixation should exert a larger influence on a target viewed in a second fixation when a long rather than a short saccade separates their viewing No such effect was found, even though the time difference between long and short saccades was effective in a no-saccade control These results indicate that at least some cognitive operations are suppressed during saccades
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Bruce, C. J., M. E. Goldberg, M. C. Bushnell i G. B. Stanton. "Primate frontal eye fields. II. Physiological and anatomical correlates of electrically evoked eye movements". Journal of Neurophysiology 54, nr 3 (1.09.1985): 714–34. http://dx.doi.org/10.1152/jn.1985.54.3.714.
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We studied single neurons in the frontal eye fields of awake macaque monkeys and compared their activity with the saccadic eye movements elicited by microstimulation at the sites of these neurons. Saccades could be elicited from electrical stimulation in the cortical gray matter of the frontal eye fields with currents as small as 10 microA. Low thresholds for eliciting saccades were found at the sites of cells with presaccadic activity. Presaccadic neurons classified as visuomovement or movement were most associated with low (less than 50 microA) thresholds. High thresholds (greater than 100 microA) or no elicited saccades were associated with other classes of frontal eye field neurons, including neurons responding only after saccades and presaccadic neurons, classified as purely visual. Throughout the frontal eye fields, the optimal saccade for eliciting presaccadic neural activity at a given recording site predicted both the direction and amplitude of the saccades that were evoked by microstimulation at that site. In contrast, the movement fields of postsaccadic cells were usually different from the saccades evoked by stimulation at the sites of such cells. We defined the low-threshold frontal eye fields as cortex yielding saccades with stimulation currents less than or equal to 50 microA. It lies along the posterior portion of the arcuate sulcus and is largely contained in the anterior bank of that sulcus. It is smaller than Brodmann's area 8 but corresponds with the union of Walker's cytoarchitectonic areas 8A and 45. Saccade amplitude was topographically organized across the frontal eye fields. Amplitudes of elicited saccades ranged from less than 1 degree to greater than 30 degrees. Smaller saccades were evoked from the ventrolateral portion, and larger saccades were evoked from the dorsomedial portion. Within the arcuate sulcus, evoked saccades were usually larger near the lip and smaller near the fundus. Saccade direction had no global organization across the frontal eye fields; however, saccade direction changed in systematic progressions with small advances of the microelectrode, and all contralateral saccadic directions were often represented in a single electrode penetration down the bank of the arcuate sulcus. Furthermore, the direction of change in these progressions periodically reversed, allowing particular saccade directions to be multiply represented in nearby regions of cortex.(ABSTRACT TRUNCATED AT 400 WORDS)
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Van Horn, Marion R., i Kathleen E. Cullen. "Dynamic Coding of Vertical Facilitated Vergence by Premotor Saccadic Burst Neurons". Journal of Neurophysiology 100, nr 4 (październik 2008): 1967–82. http://dx.doi.org/10.1152/jn.90580.2008.
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To redirect our gaze in three-dimensional space we frequently combine saccades and vergence. These eye movements, known as disconjugate saccades, are characterized by eyes rotating by different amounts, with markedly different dynamics, and occur whenever gaze is shifted between near and far objects. How the brain ensures the precise control of binocular positioning remains controversial. It has been proposed that the traditionally assumed “conjugate” saccadic premotor pathway does not encode conjugate commands but rather encodes monocular commands for the right or left eye during saccades. Here, we directly test this proposal by recording from the premotor neurons of the horizontal saccade generator during a dissociation task that required a vergence but no horizontal conjugate saccadic command. Specifically, saccadic burst neurons (SBNs) in the paramedian pontine reticular formation were recorded while rhesus monkeys made vertical saccades made between near and far targets. During this task, we first show that peak vergence velocities were enhanced to saccade-like speeds (e.g., >150 vs. <100°/s during saccade-free movements for comparable changes in vergence angle). We then quantified the discharge dynamics of SBNs during these movements and found that the majority of the neurons preferentially encode the velocity of the ipsilateral eye. Notably, a given neuron typically encoded the movement of the same eye during horizontal saccades that were made in depth. Taken together, our findings demonstrate that the brain stem saccadic burst generator encodes integrated conjugate and vergence commands, thus providing strong evidence for the proposal that the classic saccadic premotor pathway controls gaze in three-dimensional space.
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Munoz, D. P., D. M. Waitzman i R. H. Wurtz. "Activity of neurons in monkey superior colliculus during interrupted saccades". Journal of Neurophysiology 75, nr 6 (1.06.1996): 2562–80. http://dx.doi.org/10.1152/jn.1996.75.6.2562.
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1. Recent studies of the monkey superior colliculus (SC) have identified several types of cells in the intermediate layers (including burst, buildup, and fixation neurons) and the sequence of changes in their activity during the generation of saccadic eye movements. On the basis of these observations, several hypotheses about the organization of the SC leading to saccade generation have placed the SC in a feedback loop controlling the amplitude and direction of the impending saccade. We tested these hypotheses about the organization of the SC by perturbing the system while recording the activity of neurons within the SC. 2. We applied a brief high-frequency train of electrical stimulation among the fixation cells in the rostral pole of the SC. This momentarily interrupted the saccade in midflight: after the initial eye acceleration, the eye velocity decreased (frequently to 0) and then again accelerated. Despite the break in the saccade, these interrupted saccades were of about the same amplitude as normal saccades. The postinterruption saccades were usually initiated immediately after the termination of stimulation and occurred regardless of whether the saccade target was visible or not. The velocity-amplitude relationship of the preinterruption component of the saccade fell slightly above the main sequence for control saccades of that amplitude, whereas postinterruption saccades fell near the main sequence. 3. Collicular burst neurons are silent during fixation and discharge a robust burst of action potentials for saccades to a restricted region of the visual field that define a closed movement field. During the stimulation-induced saccadic interruption, these burst neurons all showed a pause in their high-frequency discharge. During an interrupted saccade to a visual target, the typical saccade-related burst was broken into two parts: the first part of the burst began before the initial preinterruption saccade; the second burst began before the postinterruption saccade. 4. We quantified three aspects of the resumption of activity of burst neurons following saccade interruption: 1) the total number of spikes in the pre- and postinterruption bursts, was very similar to the total number of spikes in the control saccade burst; 2) the increase in total duration of the burst (preinterruption period + interruption + postinterruption period) was highly correlated with the increase in total saccade duration (preinterruption saccade + interruption + postinterruption saccade); and 3) the time course of the postinterruption saccade and the resumed cell discharge both followed the same monotonic trajectory as the control saccade in most cells. 5. The same population of burst neurons was active for both the preinterruption and the postinterruption saccades, provided that the stimulation was brief enough to allow the postinterruption saccade to occur immediately. If the postinterruption saccade was delayed by > 100 ms, then burst neurons at a new and more rostral locus related to such smaller saccades became active in association with the smaller remaining saccade. We interpret this shift in active locations within the SC as a termination of the initial saccadic error command and the triggering of a new one. 6. Buildup neurons usually had two aspects to their discharge: a high-frequency burst for saccades of the optimal amplitude and direction (similar to burst neurons), and a low-frequency discharge for saccades of optimal direction whose amplitudes were equal to or greater than the optimal (different from burst neurons). The stimulation-induced interruption in saccade trajectory differentially affected these two components of buildup neuron discharge. The high-frequency burst component was affected in a manner very similar to the burst neurons.(ABSTRACT TRUNCATED)
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Pallus, Adam C., Mark M. G. Walton i Michael J. Mustari. "Response of supraoculomotor area neurons during combined saccade-vergence movements". Journal of Neurophysiology 119, nr 2 (1.02.2018): 585–96. http://dx.doi.org/10.1152/jn.00193.2017.
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Combined saccade-vergence movements allow humans and other primates to align their eyes with objects of interest in three-dimensions. In the absence of saccades, vergence movements are typically slow, symmetrical movements of the two eyes in opposite directions. However, combined saccade-vergence movements produce vergence velocities that exceed values observed during vergence alone. This phenomenon is often called “vergence enhancement”, or “saccade-facilitated vergence,” though it is important to consider that rapid vergence changes, known as “vergence transients,” are also observed during conjugate saccades. We developed a visual target array that allows monkeys to make saccades in all directions between targets spaced at distances that correspond to ~1° intervals of vergence angle relative to the monkey. We recorded the activity of vergence-sensitive neurons in the supra-oculomotor area (SOA), located dorsal and lateral to the oculomotor nucleus while monkeys made saccades with vergence amplitudes ranging from 0 to 10°. The primary focus of this study was to test the hypothesis that neurons in the SOA fire a high frequency burst of spikes during saccades that could generate the enhanced vergence. We found that individual neurons encode vergence velocity during both saccadic and non-saccadic vergence, yet firing rates were insufficient to produce the observed enhancement of vergence velocity. Our results are consistent with the hypothesis that slow vergence changes are encoded by the SOA while fast vergence movements require an additional contribution from the saccadic system. NEW & NOTEWORTHY Research into combined saccade-vergence movements has so far focused on exploring the saccadic neural circuitry, leading to diverging hypotheses regarding the role of the vergence system in this behavior. In this study, we report the first quantitative analysis of the discharge of individual neurons that encode vergence velocity in the monkey brain stem during combined saccade-vergence movements.
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Noto, Christopher T., Shoji Watanabe i Albert F. Fuchs. "Characteristics of Simian Adaptation Fields Produced by Behavioral Changes in Saccade Size and Direction". Journal of Neurophysiology 81, nr 6 (1.06.1999): 2798–813. http://dx.doi.org/10.1152/jn.1999.81.6.2798.
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Characteristics of adaptation fields produced by behavioral changes in saccade size and direction. The gain of saccadic eye movements can be altered gradually by moving targets either forward or backward during targeting saccades. If the gain of saccades to targets of only one size is adapted, the gain change generalizes or transfers only to saccades with similar vectors. In this study, we examined the spatial extent of such saccadic size adaptation, i.e., the gain adaptation field. We also attempted to adapt saccade direction by moving the target orthogonally during the targeting saccade to document the extent of a direction or cross-axis adaptation field. After adaptive gain decreases of horizontal saccades to 15° target steps, >82% of the gain reduction transferred to saccades to 25° horizontal target steps but only ∼30% transferred to saccades to 5° steps. For the horizontal component of oblique saccades to target steps with 15° horizontal components and 10° upward or downward vertical components, the transfer was similar at 51 and 60%, respectively. Thus the gain decrease adaptation field was quite asymmetric in the horizontal dimension but symmetric in the vertical dimension. Although gain increase adaptation produced a smaller gain change (13% increase for a 30% forward adapting target step) than did gain decrease adaptation (20% decrease for a 30% backward adapting target step), the spatial extent of gain transfer was quite similar. In particular, the gain increase adaptation field displayed asymmetry in the horizontal dimension (58% transfer to 25° saccades but only 32% transfer to 5° saccades) and symmetry in the vertical direction (50% transfer to the horizontal component of 10° upward and 40% transfer to 10° downward oblique saccades). When a 5° vertical target movement was made to occur during a saccade to a horizontal 10° target step, a vertical component gradually appeared in saccades to horizontal targets. More than 88% of the cross-axis change in the vertical component produced in 10° saccades transferred to 20° saccades but only 12% transferred to 4° saccades. The transfer was similar to the vertical component of oblique saccades to target steps with either 10° upward (46%) or 10° downward (46%) vertical components. Therefore both gain and cross-axis adaptation fields have similar spatial profiles. These profiles resemble those of movement fields of neurons in the frontal eye fields and superior colliculus. How those structures might participate in the adaptation process is considered in the discussion.
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Pare, M., i D. P. Munoz. "Saccadic reaction time in the monkey: advanced preparation of oculomotor programs is primarily responsible for express saccade occurrence". Journal of Neurophysiology 76, nr 6 (1.12.1996): 3666–81. http://dx.doi.org/10.1152/jn.1996.76.6.3666.
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1. The introduction of a period of darkness between the disappearance of an initial fixation target and the appearance of a peripheral saccade target produces a general reduction in saccadic reaction time (SRT)-known as the gap effect- and often very short latency express saccades. To account for these phenomena, premotor processes may be facilitated by release of visual fixation and advanced preparation of saccadic programs. The experiments described in this paper were designed to test the relevance of the ocular fixation disengagement and oculomotor preparation hypotheses by identifying the influence of different factors on SRTs and the occurrence of express saccades in the monkey. 2. The SRTs of two monkeys were measured in two behavioral paradigms. A peripheral saccade target appeared at the time of disappearance of a central fixation target in the no-gap task, whereas a 200-ms period of no stimuli was interposed between the fixation target disappearance and the saccade target appearance in the gap task. The distribution of SRTs in these tasks was generally bimodal; the first and second mode was composed of express and regular saccades, respectively. We measured the mean SRT, mean regular saccade latency, mean express saccade latency, and percentage of express saccades in both tasks. We also estimated the gap effect, i.e., the difference between the SRTs in no-gap trial and the SRTs in gap trials. 3. Once the animals were trained to make saccades to a single target location and produce express saccades, SRTs in both no-gap and gap trials displayed a broad tuning with respect to the spatial location of the trained target when the target location was varied randomly in a block of trials. Express saccades were made only to a restricted region of the visual field surrounding the trained target location. A gap effect was present for nearly all target locations tested, irrespective of express saccade occurrence. Finally, the probability of generating an express saccade at the trained target location decreased with the introduction of uncertainty about target location. 4. The occurrence of express saccades increased with the duration of the visual and nonvisual (gap) fixation that the animal was required to maintain before the onset of a saccade target. The gap duration was effective in reducing the mean SRT for gaps < or = 300 ms, and it was more influential than comparable variation in the visual fixation duration. 5. The occurrence of express saccades made to targets of identical eccentricity increased when the initial eye fixation position was shifted eccentric in a direction opposite to the saccade direction. Concomitantly, mean SRT decreased by approximately 2 ms for each 1-deg change in initial eye fixation position. 6. The occurrence of express saccades depended upon contextual factors, i.e., on both the behavioral task (no-gap or gap) and the latency of the saccade that the monkey executed to the same target in the preceding trial. The highest percentage of express saccades was observed after an express saccade in a no-gap trial, whereas the lowest percentage was obtained after a regular saccade in a gap trial. 7. These findings indicate that training-dependent express saccades are restricted to a specific spatial location dictated by the training target, and their incidence is facilitated by high predictability of target presentation, long-duration foreperiod, absence of visual fixation, eccentric initial eye position opposite to the saccade direction, and express saccade occurrence in the previous trial. The release of fixation afforded by the gap accounts for the general gap effect, but has only a modulatory influence on express saccade generation. We conclude that advanced motor preparation of saccadic programs generally reduces SRT and is primarily responsible for the occurrence of express saccades, which therefore may be caused mainly by neuronal changes restricted to a specific locus-coding for the trained movemen
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Steinmetz, Nicholas A., i Tirin Moore. "Changes in the Response Rate and Response Variability of Area V4 Neurons During the Preparation of Saccadic Eye Movements". Journal of Neurophysiology 103, nr 3 (marzec 2010): 1171–78. http://dx.doi.org/10.1152/jn.00689.2009.
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The visually driven responses of macaque area V4 neurons are modulated during the preparation of saccadic eye movements, but the relationship between presaccadic modulation in area V4 and saccade preparation is poorly understood. Recent neurophysiological studies suggest that the variability across trials of spiking responses provides a more reliable signature of motor preparation than mean firing rate across trials. We compared the dynamics of the response rate and the variability in the rate across trials for area V4 neurons during the preparation of visually guided saccades. As in previous reports, we found that the mean firing rate of V4 neurons was enhanced when saccades were prepared to stimuli within a neuron's receptive field (RF) in comparison with saccades to a non-RF location. Further, we found robust decreases in response variability prior to saccades and found that these decreases predicted saccadic reaction times for saccades both to RF and non-RF stimuli. Importantly, response variability predicted reaction time whether or not there were any accompanying changes in mean firing rate. In addition to predicting saccade direction, the mean firing rate could also predict reaction time, but only for saccades directed to the RF stimuli. These results demonstrate that response variability of area V4 neurons, like mean response rate, provides a signature of saccade preparation. However, the two signatures reflect complementary aspects of that preparation.
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Harris, C. M., J. Wallman i C. A. Scudder. "Fourier analysis of saccades in monkeys and humans". Journal of Neurophysiology 63, nr 4 (1.04.1990): 877–86. http://dx.doi.org/10.1152/jn.1990.63.4.877.
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1. By recording eye movements with the search coil technique and subjecting them to an accurate infinite Fourier transform algorithm, we describe the Fourier spectra of human and monkey saccades. In both species we find heretofore undescribed features consisting of a regular pattern of local minima in the power plot, which cannot be attributed to noise. The frequency of these minima is well correlated with saccade duration. 2. Computer simulation shows that if the pulse component of the saccade is considered to be rectangular, then the first of these minima (called M1) occurs at a frequency that is the reciprocal of the duration of the pulse. 3. Comparing the position of this component during individual monkey saccades with electrophysiological recordings of motoneurons during the same saccades leads to the conclusion that these minima are related to the burst components in ocular motoneuron discharges. Specifically, the reciprocals of the frequencies of these minima are correlated with the duration of the burst component in the motoneuron discharge. 4. In the Fourier spectra of human saccades, the relationship of the frequency of M1 to saccadic duration is a function similar to that in the monkey. This adds to the evidence that the human saccade also is driven by a pulse-step signal. 5. In both monkeys and humans, T1, the reciprocal of the frequency of M1, is shorter than both the saccade duration and the burst duration of individual motoneurons, even though neurophysiological studies in monkeys generally report the saccadic burst duration to be equal to the saccade duration. This probably arises because the saccadic pulse is not rectangular, with the extremes contributing very little energy to the Fourier spectrum. By further computer modeling we show these shape effects explicitly: as the rise- and falltime increase, making the pulse less rectangular, T1 becomes shorter; in addition, as the asymmetry of rise and fall increases, the depth of the minima is reduced. We conclude that T1 measures the "effective pulse" duration of the motoneuron. 6. There is a difference in the relationship of effective pulse duration to the saccade duration between short and long saccades. For saccades shorter than approximately 40 ms in the human and 50 ms in the monkey, the pulse width as measured by this technique varies little with saccade duration. For longer saccades, effective pulse width increases linearly with duration. We agree with others that for short saccades the pulse is both height- and width-modulated; but for longer saccades, height modulation saturates and only width modulation remains.(ABSTRACT TRUNCATED AT 400 WORDS)
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Dandekar, Sangita, Claudio Privitera, Thom Carney i Stanley A. Klein. "Neural saccadic response estimation during natural viewing". Journal of Neurophysiology 107, nr 6 (15.03.2012): 1776–90. http://dx.doi.org/10.1152/jn.00237.2011.
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Studying neural activity during natural viewing conditions is not often attempted. Isolating the neural response of a single saccade is necessary to study neural activity during natural viewing; however, the close temporal spacing of saccades that occurs during natural viewing makes it difficult to determine the response to a single saccade. Herein, a general linear model (GLM) approach is applied to estimate the EEG neural saccadic response for different segments of the saccadic main sequence separately. It is determined that, in visual search conditions, neural responses estimated by conventional event-related averaging are significantly and systematically distorted relative to GLM estimates due to the close temporal spacing of saccades during visual search. Before the GLM is applied, analyses are applied that demonstrate that saccades during visual search with intersaccadic spacings as low as 100–150 ms do not exhibit significant refractory effects. Therefore, saccades displaying different intersaccadic spacings during visual search can be modeled using the same regressor in a GLM. With the use of the GLM approach, neural responses were separately estimated for five different ranges of saccade amplitudes during visual search. Occipital responses time locked to the onsets of saccades during visual search were found to account for, on average, 79 percent of the variance of EEG activity in a window 90–200 ms after the onsets of saccades for all five saccade amplitude ranges that spanned a range of 0.2–6.0 degrees. A GLM approach was also used to examine the lateralized ocular artifacts associated with saccades. Possible extensions of the methods presented here to account for the superposition of microsaccades in event-related EEG studies conducted in nominal fixation conditions are discussed.
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Kleine, J. F., Y. Guan i U. Büttner. "Saccade-Related Neurons in the Primate Fastigial Nucleus: What Do They Encode?" Journal of Neurophysiology 90, nr 5 (listopad 2003): 3137–54. http://dx.doi.org/10.1152/jn.00021.2003.
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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°) 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 ∼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.
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Cotti, Julien, Alain Guillaume, Nadia Alahyane, Denis Pelisson i Jean-Louis Vercher. "Adaptation of Voluntary Saccades, But Not of Reactive Saccades, Transfers to Hand Pointing Movements". Journal of Neurophysiology 98, nr 2 (sierpień 2007): 602–12. http://dx.doi.org/10.1152/jn.00293.2007.
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Studying the transfer of visuomotor adaptation from a given effector (e.g., the eye) to another (e.g., the hand) allows us to question whether sensorimotor processes influenced by adaptation are common to both effector control systems and thus to address the level where adaptation takes place. Previous studies have shown only very weak transfer of the amplitude adaptation of reactive saccades—i.e., produced automatically in response to the sudden appearance of visual targets—to hand pointing movements. Here we compared the amplitude of hand pointing movements recorded before and after adaptation of either reactive or voluntary saccades, produced either in a saccade sequence task or in a single saccade task. No transfer to hand pointing movements was found after adaptation of reactive saccades. In contrast, a substantial transfer to the hand was obtained following adaptation of voluntary saccades produced in sequence. Large amounts of transfer between the two saccade types were also found. These results demonstrate that the visuomotor processes influenced by saccadic adaptation depend on the type of saccades and that, in the case of voluntary saccades, they are shared by hand pointing movements. Implications for the neurophysiological substrates of the adaptation of reactive and voluntary saccades are discussed.
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de Brouwer, Sophie, Marcus Missal, Graham Barnes i Philippe Lefèvre. "Quantitative Analysis of Catch-Up Saccades During Sustained Pursuit". Journal of Neurophysiology 87, nr 4 (1.04.2002): 1772–80. http://dx.doi.org/10.1152/jn.00621.2001.
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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°/s) in the contribution of retinal slip to the amplitude of catch-up saccades.
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Krebs, Ruth M., C. Nicolas Boehler, Helen H. Zhang, Mircea A. Schoenfeld i Marty G. Woldorff. "Electrophysiological recordings in humans reveal reduced location-specific attentional-shift activity prior to recentering saccades". Journal of Neurophysiology 107, nr 5 (1.03.2012): 1393–402. http://dx.doi.org/10.1152/jn.00912.2010.
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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.
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Kustov, A. A., i D. L. Robinson. "Modified saccades evoked by stimulation of the macaque superior colliculus account for properties of the resettable integrator". Journal of Neurophysiology 73, nr 4 (1.04.1995): 1724–28. http://dx.doi.org/10.1152/jn.1995.73.4.1724.
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1. Models of the saccadic system propose that there is an integration of the pulse signal, and there is good evidence that the integrator is reset gradually (Nichols and Sparks 1994, 1995). Other studies of the superior collicular contribution to the saccadic system have proposed a sensory, not motor, nature for its signal. 2. To test experimentally the resetting of the integrator and the nature of the collicular signal, we electrically stimulated the superior colliculus during periods of fixation and during the course of visually guided saccades. Trains of stimuli which were presented during periods of fixation evoked saccades with fixed vectors. Identical stimulation at the beginning of a visually guided saccade evoked saccades whose direction was rotated and amplitude extended from the fixed vector. The direction of the rotation was opposite that of the visually guided saccade, and the magnitude of this rotation could be as large as 80 degrees. 3. Stimulation which was applied at progressively later times during the visually guided saccade, evoked saccades with progressively smaller rotations and progressively less elongations. The time period during which saccades were modified persisted beyond the end of the visually guided saccade, when the eyes were stationary. Thus, we confirm the previous findings (Nichols and Sparks 1994, 1995; Robinson, 1972), that the end of the saccade is not a period of quiescence within the oculomotor pathways. 4. Our results confirm that the resetting of the integration of the saccade signal is gradual rather than abrupt. Furthermore, these data suggest that the superior colliculus signals a motor error.
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Munoz, D. P., i R. H. Wurtz. "Fixation cells in monkey superior colliculus. II. Reversible activation and deactivation". Journal of Neurophysiology 70, nr 2 (1.08.1993): 576–89. http://dx.doi.org/10.1152/jn.1993.70.2.576.
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1. We tested the hypothesis that a subset of neurons, which we have referred to as fixation cells, located within the rostral pole of the monkey superior colliculus (SC) controls the generation of saccadic eye movements. We altered the activity of these neurons with either electrical stimulation or GABAergic drugs. 2. An increase in the activity of fixation cells in the rostral SC, induced by a train of low-frequency electrical stimulation, delayed the initiation of saccades. With bilateral stimulation the monkey was able to make saccades only after stimulation ceased. 3. Pulses of stimulation delivered during the saccade produced an interruption of the saccade in midflight. The latency to the onset of this perturbation was as short as 12 ms. 4. Injection of the gamma-aminobutyric acid (GABA) antagonist bicuculline into the rostral pole of the SC, which decreases normal GABA inhibition and increases cell activity, increased the latency of saccades to both visual and remembered targets. 5. Injection of the GABA agonist muscimol into the rostral SC, which increases normal GABA inhibition and decreases activity, reduced the latency for saccades to visual targets. The monkey also had difficulty maintaining visual fixation and suppressing unwanted saccades. 6. After muscimol injections, monkeys frequently made very short-latency saccades forming a peak in the saccade latency histogram at < 100 ms. These saccades are similar to express saccades made by normal monkeys. This finding suggests that the fixation cells in the rostral SC are critical for controlling the frequency of express saccades. 7. These results support the hypothesis that fixation cells in the rostral SC inhibit the generation of saccadic eye movements and that they form part of a system of oculomotor control, that of visual fixation.
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Goossens, H. H. L. M., i A. J. Van Opstal. "Local Feedback Signals Are Not Distorted By Prior Eye Movements: Evidence From Visually Evoked Double Saccades". Journal of Neurophysiology 78, nr 1 (1.07.1997): 533–38. http://dx.doi.org/10.1152/jn.1997.78.1.533.
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Goossens, H.H.L.M. and A. J. Van Opstal. Local feedback signals are not distorted by prior eye movements: evidence from visually evoked double saccades. J. Neurophysiol. 78: 533–538, 1997. Recent experiments have shown that the amplitude and direction of saccades evoked by microstimulation of the monkey superior colliculus depend systematically on the amplitude and direction of preceding visually guided saccades as well as on the postsaccade stimulation interval. The data are consistent with the hypothesis that an eye displacement integrator in the local feedback loop of the saccadic burst generator is gradually reset with a time constant of ∼45 ms. If this is true, similar effects should occur during naturally evoked saccade sequences, causing systematic interval-dependent errors. To test this prediction in humans, saccades toward visual single- and double-step stimuli were elicited, and the properties of the second saccades were investigated as a function of the intersaccadic interval (ISI). In 15–20% of the saccadic responses, ISIs fell well below 100 ms. The errors of the second saccades were not systematically affected by the preceding primary saccade, irrespective of the ISI. Only a slight increase in the endpoint variability of second saccades was observed for the shortest ISIs. These results are at odds with the hypothesis that the putative eye displacement integrator has a reset time constant >10 ms. Instead, it is concluded that the signals involved in the internal feedback control of the saccadic burst generator reflect eye position and/or eye displacement accurately, irrespective of preceding eye movements.
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Phillips, Adam N., i Mark A. Segraves. "Predictive Activity in Macaque Frontal Eye Field Neurons During Natural Scene Searching". Journal of Neurophysiology 103, nr 3 (marzec 2010): 1238–52. http://dx.doi.org/10.1152/jn.00776.2009.
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Generating sequences of multiple saccadic eye movements allows us to search our environment quickly and efficiently. Although the frontal eye field cortex (FEF) has been linked to target selection and making saccades, little is known about its role in the control and performance of the sequences of saccades made during self-guided visual search. We recorded from FEF cells while monkeys searched for a target embedded in natural scenes and examined the degree to which cells with visual and visuo-movement activity showed evidence of target selection for future saccades. We found that for about half of these cells, activity during the fixation period between saccades predicted the next saccade in a sequence at an early time that precluded selection based on current visual input to a cell's response field. In addition to predicting the next saccade, activity during the fixation prior to two successive saccades also predicted the direction and goal of the second saccade in the sequence. We refer to this as advanced predictive activity. Unlike activity indicating the upcoming saccade, advanced predictive activity occurred later in the fixation period, mirroring the order of the saccade sequence itself. The remaining cells without advanced predictive activity did not predict future saccades but reintroduced the signal for the upcoming saccade at an intermediate time in the fixation period. Together these findings suggest that during natural visual search the timing of FEF cell activity is consistent with a role in specifying targets for one or more future saccades in a search sequence.
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Wulff, Svenja, Annalisa Bosco, Katharina Havermann, Giacomo Placenti, Patrizia Fattori i Markus Lappe. "Eye position effects in saccadic adaptation in macaque monkeys". Journal of Neurophysiology 108, nr 10 (15.11.2012): 2819–26. http://dx.doi.org/10.1152/jn.00212.2012.
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The saccadic amplitude of humans and monkeys can be adapted using intrasaccadic target steps in the McLaughlin paradigm. It is generally believed that, as a result of a purely retinal reference frame, after adaptation of a saccade of a certain amplitude and direction, saccades of the same amplitude and direction are all adapted to the same extent, independently from the initial eye position. However, recent studies in humans have put the pure retinal coding in doubt by revealing that the initial eye position has an effect on the transfer of adaptation to saccades of different starting points. Since humans and monkeys show some species differences in adaptation, we tested the eye position dependence in monkeys. Two trained Macaca fascicularis performed reactive rightward saccades from five equally horizontally distributed starting positions. All saccades were made to targets with the same retinotopic motor vector. In each session, the saccades that started at one particular initial eye position, the adaptation position, were adapted to shorter amplitude, and the adaptation of the saccades starting at the other four positions was measured. The results show that saccades that started at the other positions were less adapted than saccades that started at the adaptation position. With increasing distance between the starting position of the test saccade and the adaptation position, the amplitude change of the test saccades decreased with a Gaussian profile. We conclude that gain-decreasing saccadic adaptation in macaques is specific to the initial eye position at which the adaptation has been induced.
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Izawa, Yoshiko, Hisao Suzuki i Yoshikazu Shinoda. "Suppression of Visually and Memory-Guided Saccades Induced by Electrical Stimulation of the Monkey Frontal Eye Field. I. Suppression of Ipsilateral Saccades". Journal of Neurophysiology 92, nr 4 (październik 2004): 2248–60. http://dx.doi.org/10.1152/jn.01021.2003.
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When a saccade occurs to an interesting object, visual fixation holds its image on the fovea and suppresses saccades to other objects. Electrical stimulation of the frontal eye field (FEF) has been reported to elicit saccades, and recently also to suppress saccades. This study was performed to characterize properties of the suppression of visually guided (Vsacs) and memory-guided saccades (Msacs) induced by electrical stimulation of the FEF in trained monkeys. For any given stimulation site, we determined the threshold for electrically evoked saccades (Esacs) at ≤50 μA and then examined suppressive effects of stimulation at the same site on Vsacs and Msacs. FEF stimulation suppressed the initiation of both Vsacs and Msacs during and about 50 ms after stimulation at stimulus intensities lower than those for eliciting Esacs, but did not affect the vector of these saccades. Suppression occurred for ipsiversive but not contraversive saccades, and more strongly for saccades with larger amplitudes and those with initial eye positions shifted more in the saccadic direction. The most effective stimulation timing for suppression was about 50 ms before saccade onset, which suggests that suppression occurred in the efferent pathway for generating Vsacs at the premotor rather than the motoneuronal level, most probably in the superior colliculus and/or the paramedian pontine reticular formation. Suppression sites of ipsilateral saccades were distributed over the classical FEF where saccade-related movement neurons were observed. The results suggest that the FEF may play roles in not only generating contraversive saccades but also maintaining visual fixation by suppressing ipsiversive saccades.
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Grujic, Nikola, Nils Brehm, Cordula Gloge, Weijie Zhuo i Ziad M. Hafed. "Perisaccadic perceptual mislocalization is different for upward saccades". Journal of Neurophysiology 120, nr 6 (1.12.2018): 3198–216. http://dx.doi.org/10.1152/jn.00350.2018.
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Saccadic eye movements, which dramatically alter retinal images, are associated with robust perimovement perceptual alterations. Such alterations, thought to reflect brain mechanisms for maintaining perceptual stability in the face of saccade-induced retinal image disruptions, are often studied by asking subjects to localize brief stimuli presented around the time of horizontal saccades. However, other saccade directions are not usually explored. Motivated by recently discovered asymmetries in upper and lower visual field representations in the superior colliculus, a structure important for both saccade generation and visual analysis, we observed significant differences in perisaccadic perceptual alterations for upward saccades relative to other saccade directions. We also found that, even for purely horizontal saccades, perceptual alterations differ for upper vs. lower retinotopic stimulus locations. Our results, coupled with conceptual modeling, suggest that perisaccadic perceptual alterations might critically depend on neural circuits, such as superior colliculus, that asymmetrically represent the upper and lower visual fields. NEW & NOTEWORTHY Brief visual stimuli are robustly mislocalized around the time of saccades. Such mislocalization is thought to arise because oculomotor and visual neural maps distort space through foveal magnification. However, other neural asymmetries, such as upper visual field magnification in the superior colliculus, may also exist, raising the possibility that interactions between saccades and visual stimuli would depend on saccade direction. We confirmed this behaviorally by exploring and characterizing perisaccadic perception for upward saccades.
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Soetedjo, Robijanto. "Signals driving the adaptation of saccades that require spatial updating". Journal of Neurophysiology 120, nr 2 (1.08.2018): 525–38. http://dx.doi.org/10.1152/jn.00075.2018.
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Saccades adapt to persistent natural or artificially imposed dysmetrias. The characteristics and circuitry of saccade adaptation have been revealed using a visually guided task (VGT) where the vectors of the target step and the intended saccade command are the same. However, in real life, another saccade occasionally intervenes before the saccade to the target occurs. This necessitates an updating of the intended saccade to account for the intervening saccadic displacement, which dissociates the visual target signal and the intended saccade command. We determined whether the adaptation process is similar for VGT and updated saccades by studying the transfer of adaptation between them. The ultimate visual target was dissociated from the intended saccade command with double-step saccade tasks (DSTs) in which two targets are flashed sequentially at different locations while the monkey maintains fixation. The resulting saccades toward the first and second targets occur in the dark. The transfer of visually guided saccade adaptation to the second saccades of a DST and vice versa depended on the eccentricity of the second visual target, and not the second saccade command. If a target with the same eccentricity as the adapted target appears briefly during the intersaccadic interval of a DST, more adaptation transfers. Because a brief appearance of the visual target either before the first saccade or during the intersaccadic interval influences how much adaptation transfer the second saccade will express, the processing of adaptation and DST updating may overlap. NEW & NOTEWORTHY Adaptation and the spatial updating of saccades are thought to be independent processes. When we dissociate the visual target and the intended saccade command, the transfer of visually guided saccade adaptation to the saccades of the double-step saccade tasks (DST) and vice versa is driven by a visual not motor error. The visual target has an effect until the second saccade of a DST occurs. Therefore, the processing of adaptation and the spatial updating of saccades may overlap.
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Schmidt, M. "Neurons in the cat pretectum that project to the dorsal lateral geniculate nucleus are activated during saccades". Journal of Neurophysiology 76, nr 5 (1.11.1996): 2907–18. http://dx.doi.org/10.1152/jn.1996.76.5.2907.
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1. Neurons in the pretectal nuclear complex that project to the ipsilateral dorsal lateral geniculate nucleus (LGNd) were identified by antidromic activation after electrical LGNd stimulation in awake cats, and their response properties were characterized to retinal image shifts elicited either by external visual stimulus movements or during spontaneous saccadic eye movements on a stationary visual stimulus, and to saccades in darkness. Eye position was monitored with the use of a scleral search coil and care was taken to assure stability of the eyes during presentation of moving visual stimuli. 2. Of a total sample of 134 cells recorded, 27 neurons were antidromically activated by electrical LGNd stimulation. In addition, responses from neurons that were not activated from the LGNd were also analyzed, including 19 “retinal slip” cells, which selectively respond to slow horizontal stimulus movements, and 21 “jerk” cells, which are specifically activated by rapid stimulus shifts. All recorded neurons were located in the nucleus of the optic tract and in the posterior pretectal nucleus. 3. In the light, neurons identified as projecting to the LGNd responded maximally to saccadic eye movements and to externally generated sudden shifts of large visual stimuli. Slow stimulus drifts did not activate these neurons. Response latencies were shorter and peak activities were increased during saccades compared with pure visual stimulation. No systematic correlation between response latency, response duration, or the number of spikes in the response and saccade direction, saccade amplitude, or saccade duration was found. Saccades and rapid stimulus shifts in the light also activated jerk cells but not retinal slip cells. 4. All 27 antidromically activated neurons also responded to spontaneous saccadic eye movements in complete darkness. Responses to saccades in the dark, however, had longer response latencies and lower peak activities than responses to saccades in light. As in the light, response parameters in darkness seemed not to code specific saccade parameters. Cells that were not activated from LGNd were found to be unresponsive to saccades in the dark. 5. According to their specific activation by saccades in darkness, LGNd-projecting pretectal neurons are termed “saccade neurons” to distinguish them from other pretectal cell populations, in particular from jerk neurons, which show similar response properties in light. 6. The saccade-related activation of pretectal saccade neurons may be used to modulate visual responses of LGNd relay cells following saccadic eye movements. Because the pretectogeniculate projection in cat most likely is GABAergic and terminates on inhibitory LGNd interneurons, its activation may lead to a saccade-locked disinhibition of relay cells. This input could counter the strong inhibition induced in the LGNd after shifts of gaze direction and lead to a resetting of LGNd cell activity.
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Russo, G. S., i C. J. Bruce. "Frontal eye field activity preceding aurally guided saccades". Journal of Neurophysiology 71, nr 3 (1.03.1994): 1250–53. http://dx.doi.org/10.1152/jn.1994.71.3.1250.
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1. We studied neuronal activity in the monkey's frontal eye field (FEF) in conjunction with saccades directed to auditory targets. 2. All FEF neurons with movement activity preceding saccades to visual targets also were active preceding saccades to auditory targets, even when such saccades were made in the dark. Movement cells generally had comparable bursts for aurally and visually guided saccades; visuomovement cells often had weaker bursts in conjunction with aurally guided saccades. 3. When these cells were tested from different initial fixation directions, movement fields associated with aurally guided saccades, like fields mapped with visual targets, were a function of saccade dimensions, and not the speaker's spatial location. Thus, even though sound location cues are chiefly craniotopic, the crucial factor for a FEF discharge before aurally guided saccades was the location of auditory target relative to the current direction of gaze. 4. Intracortical microstimulation at the sites of these cells evoked constant-vector saccades, and not goal-directed saccades. The direction and size of electrically elicited saccades generally matched the cell's movement field for aurally guided saccades. 5. Thus FEF activity appears to have a role in aurally guided as well as visually guided saccades. Moreover, visual and auditory target representations, although initially obtained in different coordinate systems, appear to converge to a common movement vector representation at the FEF stage of saccadic processing that is appropriate for transmittal to saccade-related burst neurons in the superior colliculus and pons.
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Heinen, Stephen J., Jeremy B. Badler i Scott N. J. Watamaniuk. "Choosing a foveal goal recruits the saccadic system during smooth pursuit". Journal of Neurophysiology 120, nr 2 (1.08.2018): 489–96. http://dx.doi.org/10.1152/jn.00418.2017.
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Models of smooth pursuit eye movements stabilize an object’s retinal image, yet pursuit is peppered with small, destabilizing “catch-up” saccades. Catch-up saccades might help follow a small, spot stimulus used in most pursuit experiments, since fewer of them occur with large stimuli. However, they can return when a large stimulus has a small central feature. It may be that a central feature on a large object automatically recruits the saccadic system. Alternatively, a cognitive choice is made that the feature is the pursuit goal, and the saccadic system is then recruited to pursue it. Observers pursued a 5-dot stimulus composed of a central dot surrounded by four peripheral dots arranged as a diamond. An attention task specified the pursuit goal as either the central element, or the diamond gestalt. Fewer catch-up saccades occurred with the Gestalt goal than with the central goal, although the additional saccades with the central goal neither enhanced nor impeded pursuit. Furthermore, removing the central element from the diamond goal further reduced catch-up saccade frequency, indicating that the central element automatically triggered some saccades. Higher saccade frequency was not simply due to narrowly focused attention, since attending a small peripheral diamond during pursuit elicited fewer saccades than attending the diamond positioned foveally. The results suggest some saccades are automatically elicited by a small central element, but when it is chosen as the pursuit goal the saccadic system is further recruited to pursue it. NEW & NOTEWORTHY Smooth-pursuit eye movements stabilize retinal image motion to prevent blur. Curiously, smooth pursuit is frequently supplemented by small catchup saccades that could reduce image clarity. Catchup saccades might only be needed to pursue small laboratory stimuli, as they are infrequent during large object pursuit. Yet large objects with central features revive them. Here, we show that voluntarily selecting a feature as the pursuit goal elicits saccades that do not help pursuit.
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SCHILLER, PETER H., JOHANNES HAUSHOFER i GEOFFERY KENDALL. "An examination of the variables that affect express saccade generation". Visual Neuroscience 21, nr 2 (marzec 2004): 119–27. http://dx.doi.org/10.1017/s0952523804042038.
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The frequency with which express saccades are generated under a variety of conditions in rhesus monkeys was examined. Increasing the gap time between fixation spot termination and target onset increased express saccade frequency but was progressively less effective in doing so as the number of target positions in the sample was increased. Express saccades were rarely produced when two targets were presented simultaneously and the choice of either of which was rewarded; a temporal asynchrony of only 17 ms between the targets reinstated express saccade generation. Express saccades continued to be generated when the vergence or pursuit systems was coactivated with the saccadic system.