The effect of saccades on threshold perception — A model study

The effect of saccadic eye movements on threshold perception is investigated theoretically. The proposed model considers eye movements by taking into account the shifting of the stimulus pattern on the retina during the occurrence of an eye movement. Saccades are characterized by high velocity and short duration. These motions cause overshoots in the response of linear filters to certain stimulus patterns. Therefore, the model predicts facilitation effects of saccades in the perception of low spatial frequency patterns and patterns flickering with high temporal frequencies. These results agree with experimentally obtained data presented in a subsequent paper. A simple approach is formulated which approximates the complex shifting function of a saccade by a switching of the pattern.     

Attention samples stimuli rhythmically

Overt exploration or sampling behaviors, such as whisking, sniffing, and saccadic eye movements, are often characterized by a rhythm. In addition, the electrophysiologically recorded theta or alpha phase predicts global detection performance. These two observations raise the intriguing possibility that covert selective attention samples from multiple stimuli rhythmically. To investigate this possibility, we measured change detection performance on two simultaneously presented stimuli, after resetting attention to one of them. After a reset flash at one stimulus location, detection performance fluctuated rhythmically. When the flash was presented in the right visual field, a 4 Hz rhythm was directly visible in the time courses of behavioral performance at both stimulus locations, and the two rhythms were in antiphase. A left visual field flash exerted only partial reset on performance and induced rhythmic fluctuation at higher frequencies (6-10 Hz). These findings show that selective attention samples multiple stimuli rhythmically, and they position spatial attention within the family of exploration behaviors.     

A minicomputer program for automatic saccade detection and linear analysis of eye movement systems using sine wave stimulus

A FORTRAN IV program is described, which may be run interactively or in batch and which allows a user to obtain the frequency response amplitude ratio and phase resulting from the linear analysis of an eye movement system using sine wave stimuli. The response (eye position) signal may contain components contributed by the saccadic eye movements. The program can digitize analog signals and store data on a magnetic tape. With the aid of digital filters, the program can detect saccades without requiring any input parameters from the user. The program interpolates the saccade interval using a method of least square curve fitting with a sine wave. The interpolation is relatively noise immune and works well regardless of the stimulus frequencies and the width of a saccade interval. Moreover, the program can handle long duration of signals such as 90 min of data which covers about 5 cycles of a 0.001 Hz sine wave signal. Sample runs for the cases of 0.001 and 0.1 Hz are given. The resident driver and the overlayable segments of the program have been implemented on a DEC (Digital Equipment Corp.) LAB-11 minicomputer (PDP 11/20).     

Neural correlates of saccadic suppression in humans

When you look into a mirror and move your eyes left to right, you will see that you cannot observe your own eye movements. This demonstrates the phenomenon of saccadic suppression: during saccadic eye movements, visual sensitivity is much reduced. Given that humans make more than 100,000 eye movements each day, it is clear why suppression is needed: without it, the motion on the retina would prevent us from seeing anything at all. Psychophysical data show that suppression is stimulus selective: it is strongest for the kind of stimuli that preferentially activate magnocellular thalamic neurons. This has led to the hypothesis that saccadic suppression selectively targets the magnocellular stream. We used fMRI to find brain areas with a stimulus-selective suppression of the BOLD signal that matches the psychophysical data. We found such a neural correlate of saccadic suppression in the dorsal stream (hMT+, V7) and in ventral area V4. These areas receive magnocellular input; hence our findings are consistent with the magnocellular hypothesis. The range of effects in our data and in single cell data, however, argues against a single thalamic mechanism that suppresses all cortical input. Instead, we speculate that saccadic suppression relies on multiple mechanisms operating in different cortical areas.     

Negative Potentials of the Human Brain Elicited in Saccadic Latency during Stimulation of the Leading and Nonleading Eyes with an Overlap

Fast presaccadic EEG potentials in saccadic latency were studied with the use of inverse averaging during monocular stimulation of the leading or nonleading eye. Two paradigms were followed, with presentation of visual stimuli consecutively or with a 200-ms overlap. Irrespective of the paradigm and the stimulated eye, the negative N ^–1 potential in the interval of 50–20 ms preceding the beginning of the saccade predominated in the hemisphere contralateral to the saccade direction, reflecting the command processes of saccadic initiation. The N ^–2 potential was more pronounced in the case of direct averaging, starting from the stimulus. Its amplitude increased with increasing concentration of attention on the fixation stimulus under the overlap conditions, and its foci predominated in the left hemisphere, in the frontal, central, and parietosagittal regions. Hence, the N ^–2 potential was assumed to reflect spatial perception and attention as initial stages of saccadic programming. The findings testify to the priority of the leading eye both in fixation and in spatial attention.     

Human fast negative EEG potentials before express saccades

Fast negative EEG potentials preceding fast regular saccades and express saccades were studied by the method of backward averaging under conditions of monocular stimulation of the right and left eye. "Step" and "gap" experimental paradigms were used for visual stimulation. Analysis of parameters of potentials and their spatiotemporal dynamics suggests that, under conditions of the increased attention and optimal readiness of the neural structures, express saccades appear when the previously chosen program of the future eye movement coincides with the actual target coordinates. We assumed that the saccade latency decreases at the expense of the involvement of the main oculomotor areas of motor and saccadic planning in its initiation; an express saccade can be initiated also by means of direct transmission of the signal from the cortex to the brainstem saccadic generator passing by the superior colliculus. Moreover, anticipating release from the central fixation and attention distraction are necessary for the successful initiation of an express saccade.     

Differentiation between vergence and saccadic functional activity within the human frontal eye fields and midbrain revealed through fMRI

Purpose

Eye movement research has traditionally studied solely saccade and/or vergence eye movements by isolating these systems within a laboratory setting. While the neural correlates of saccadic eye movements are established, few studies have quantified the functional activity of vergence eye movements using fMRI. This study mapped the neural substrates of vergence eye movements and compared them to saccades to elucidate the spatial commonality and differentiation between these systems.

Methodology

The stimulus was presented in a block design where the ‘off’ stimulus was a sustained fixation and the ‘on’ stimulus was random vergence or saccadic eye movements. Data were collected with a 3T scanner. A general linear model (GLM) was used in conjunction with cluster size to determine significantly active regions. A paired t-test of the GLM beta weight coefficients was computed between the saccade and vergence functional activities to test the hypothesis that vergence and saccadic stimulation would have spatial differentiation in addition to shared neural substrates.

Results

Segregated functional activation was observed within the frontal eye fields where a portion of the functional activity from the vergence task was located anterior to the saccadic functional activity (z>2.3; p<0.03). An area within the midbrain was significantly correlated with the experimental design for the vergence but not the saccade data set. Similar functional activation was observed within the following regions of interest: the supplementary eye field, dorsolateral prefrontal cortex, ventral lateral prefrontal cortex, lateral intraparietal area, cuneus, precuneus, anterior and posterior cingulates, and cerebellar vermis. The functional activity from these regions was not different between the vergence and saccade data sets assessed by analyzing the beta weights of the paired t-test (p>0.2).

Conclusion

Functional MRI can elucidate the differences between the vergence and saccade neural substrates within the frontal eye fields and midbrain.     

On identification of saccades from sinusoidal eye movement signals

Sinusoidal eye movements and potential saccadic eye movements are examined using the syntactic pattern recognition method presented previously. A few computer tests are presented for the verification of potential saccades from signals of sinusoidal eye movements. The technique was developed and tested with electro-oculographic signals. The verification of saccades consists of three tests: the estimation of average noise peaks in an eye movement signal; an angular velocity threshold; and the comparison between a sinusoidal eye movement signal and the corresponding stimulus signal. The technique is also efficient for noisy signals of eye movements, which were stimulated by both predictive and non-predictive sinusoidal stimulus movements.     

Latency of visually evoked saccadic eye movements

The paper deals with the initiation of visually guided saccades, in order to break down the saccadic reaction time into functionally different periods of time. It takes into account that spatial processing of information is so basic that modelling of saccadic control properties should include spatio-temporal arrangements. The output signal of the saccadic system was measured in response to visual stimuli in which the time between the appearance of a visual stimulus in the peripheral field and the disappearance of the central fixation point was varied. The variation of the mean saccadic latency time, measured with respect to the onset of the peripheral stimulus, as a function of stimulus asynchrony was highly significant. This variation may be represented by a so-called gap-overlap curve, which is characterized here by means of five parameters. A facilitation model is introduced to fit the results of the gap-overlap experiments. The facilitation model for the initiation of visually evoked saccades incorporates a mechanism which governs the efficiency of processing of signals that arise from a stimulus presented at a particular position in space. It explains how visual information may be affected by other sensory information before it is used to command further saccades. It allows determination of saccadic system parameters, such as the peripheral and the foveal afferent processing time, the central processing time for a saccade and the degree of facilitation. These quantities were found to be characteristic for the given test subjects, and where these data could be compared with neurophysiological data, the agreement was within the experimental error.     

Computer simulation of overshoot in saccadic eye movements.

The human horizontal eye movement system produces quick, precise, conjugate eye movements called saccades. These are important in normal vision. For example, reading tasks exclusively utilize saccadic eye movements. The majority of saccades have dynamic overshoot. The amplitude of this overshoot is independent of saccadic amplitude, and is such that it places the image of the stimulus within the retinal region of maximum acuity within a minimum of time. A computer based model of the saccadic mechanisms was used to study the origin of this overshoot. It was discussed that dynamic overshoot cannot be attributed to biomechanism properites of the eye movement mechanism, but must instead be explained by variations in the controlling nervous activity. The form of this neural controller signal is very similar to that required for a time optimal response of an inertial system.     

Stability of the saccadic oculomotor system

A variety of different types of instability has been found in the saccadic system of humans. Some of the instabilities correspond to clinical conditions, whereas others are inherent in the normal saccadic system. How can these instabilities arise within the mechanism of normal saccadic eye movements? A physiologically-based model of the saccadic system predicts that horizontal saccadic oscillations will occur with excessive mutual inhibition between the left and right burst cells and with underaction of the pause cells. The amplitudes and frequencies of the oscillations had ranges of 0–6° and 6–20 cycles per second, respectively. Application of stability analysis techniques to the model reveals that development of the oscillations can be explained by the Hopf bifurcation mechanism. Future development of this approach will involve classifying pathological instabilities of the saccadic system according to the bifurcation involved in their generation.     

Saccadic Eye Movement Characteristics in Adult Niemann-Pick Type C Disease: Relationships with Disease Severity and Brain Structural Measures

Niemann-Pick Type C disease (NPC) is a rare genetic disorder of lipid metabolism. A parameter related to horizontal saccadic peak velocity was one of the primary outcome measures in the clinical trial assessing miglustat as a treatment for NPC. Neuropathology is widespread in NPC, however, and could be expected to affect other saccadic parameters. We compared horizontal saccadic velocity, latency, gain, antisaccade error percentage and self-paced saccade generation in 9 adult NPC patients to data from 10 age-matched controls. These saccadic measures were correlated with appropriate MRI-derived brain structural measures (e.g., dorsolateral prefrontal cortex, frontal eye fields, supplemental eye fields, parietal eye fields, pons, midbrain and cerebellar vermis) and with measures of disease severity and duration. The best discriminators between groups were reflexive saccade gain and the two volitional saccade measures. Gain was also the strongest correlate with disease severity and duration. Most of the saccadic measures showed strongly significant correlations with neurophysiologically appropriate brain regions. While our patient sample is small, the apparent specificity of these relationships suggests that as new diagnostic methods and treatments become available for NPC, a broader range of saccadic measures may be useful tools for the assessment of disease progression and treatment efficacy.     

Lateralization of saccadic latency during monocular presentation of stimuli to a leading and non-leading eye

Saccadic latencies were studied in ten healthy subjects. Peripheral targets were presented monocularly to a leading and nonleading eyes in the right and left hemifields. SS (single step) and OVERLAP (200 ms) schemes of visual stimulation were used. Under OVERLAP conditions, the saccadic latency was longer by 30-39 ms and the number of long-latency saccades was higher than under SS conditions, especially in subjects with mixed asymmetry profiles. In the majority of subjects with right asymmetry profile, the latencies of saccades during stimulation of the leading eye were by 12 ms shorter than during stimulation of the nonleading eye, and the latencies of right saccades were by 24 ms shorter than that of the left saccades independently of the stimulated eye. The obtained results explain some characteristic features of hemyspheric asymmetry in organization of saccadic movements.     

Latency of visually evoked saccadic eye movements

The validness of a model describing the relation between mean saccadic latency and stimulus asynchrony based on facilitation instead of suppression was tested experimentally. As a result, suppression of signals generated by the onset of a peripheral stimulus due to fixation of another target, giving rise to an increase of mean saccadic latency, does not seem very likely. The influence of the intensity of the fixation target on the latency of visually evoked saccades was studied. According to the facilitation model, the offset of the fixation target induces after an afferent delay, a transition of the state of the facilitation mechanism from the unfacilitated condition into a mode of maximal facilitation. The time-period during which this change is accomplished is called Facilitation-Rise-Time (FRT). An interpretation within the context of the facilitation model of gap-overlap latency data for different values of the intensity of the fixation stimulus suggests, in combination with computer-computations of the model, that lowering of this intensity causes an increase in FRT. The results in normal subjects of step stimulus experiments with a dim fixation point substantiate the hypothesis of a facilitation mechanism, which is triggerable not only by an external signal such as the offset of the fixation point, but also by some internal stimulus independent signal. Moreover, data for tracking by an amblyopic eye seem to support this conclusion. The findings of increased saccadic latencies in amblyopic and Optic Neuritis (ON) eyes suggest a slowing of processing of visual information in the sensory pathways from the central retina, subsequently utilized by the oculomotor system in the generation of saccades.(ABSTRACT TRUNCATED AT 250 WORDS)     

用于研究快速扫视对瞳孔光反射调制作用的新方法

瞳孔对光反射系统和快速扫视系统在解剖学和功能上都有着紧密的联系,但是快速扫视系统对瞳孔的光反射系统是否有调制作用尚无报道。研究这两个系统间的调制作用,必须了解光刺激不均匀和近反应对瞳孔直径变化是否有影响。该研究以人为被试,设计了一种全新的实验方法,研究光刺激不均匀和近反应对瞳孔直径变化的影响。实验方法：将被试的一只眼用密闭的眼罩罩住给予脉冲光刺激,刺激由位于眼罩内全视野范围水平排列的一排发光二极管(light emitting diodes,LEDs)给出,被试的另一只眼用来记录眼动和瞳孔直径的变化,研究水平方向的快速扫视对瞳孔对光反射时瞳孔直径变化的影响。实验结果：比较被试注视视野内不同位置的瞳孔对光反射相对收缩率无显著差异（P=0.148, 非配对样本t检验）。结论：本方法消除了光刺激不均匀和近反应对瞳孔直径变化的影响,可用于研究快速扫视系统对瞳孔光反射系统间的调制作用。     

Probability of Seeing Increases Saccadic Readiness

Associating movement directions or endpoints with monetary rewards or costs influences movement parameters in humans, and associating movement directions or endpoints with food reward influences movement parameters in non-human primates. Rewarded movements are facilitated relative to non-rewarded movements. The present study examined to what extent successful foveation facilitated saccadic eye movement behavior, with the hypothesis that foveation may constitute an informational reward. Human adults performed saccades to peripheral targets that either remained visible after saccade completion or were extinguished, preventing visual feedback. Saccades to targets that were systematically extinguished were slower and easier to inhibit than saccades to targets that afforded successful foveation, and this effect was modulated by the probability of successful foveation. These results suggest that successful foveation facilitates behavior, and that obtaining the expected sensory consequences of a saccadic eye movement may serve as a reward for the oculomotor system.     

Influence of Retinal Image Shifts and Extra-Retinal Eye Movement Signals on Binocular Rivalry Alternations

Previous studies have indicated that saccadic eye movements correlate positively with perceptual alternations in binocular rivalry, presumably because the foveal image changes resulting from saccades, rather than the eye movement themselves, cause switches in awareness. Recently, however, we found evidence that retinal image shifts elicit so-called onset rivalry and not percept switches as such. These findings raise the interesting question whether onset rivalry may account for correlations between saccades and percept switches.We therefore studied binocular rivalry when subjects made eye movements across a visual stimulus and compared it with the rivalry in a ‘replay’ condition in which subjects maintained fixation while the same retinal displacements were reproduced by stimulus displacements on the screen. We used dichoptic random-dot motion stimuli viewed through a stereoscope, and measured eye and eyelid movements with scleral search-coils.Positive correlations between retinal image shifts and perceptual switches were observed for both saccades and stimulus jumps, but only for switches towards the subjects'' preferred eye at stimulus onset. A similar asymmetry was observed for blink-induced stimulus interruptions. Moreover, for saccades, amplitude appeared crucial as the positive correlation persisted for small stimulus jumps, but not for small saccades (amplitudes < 1°). These findings corroborate our tenet that saccades elicit a form of onset rivalry, and that rivalry is modulated by extra-retinal eye movement signals.     

Saccadic Reaction Times to Audiovisual Stimuli Show Effects of Oscillatory Phase Reset

Initiating an eye movement towards a suddenly appearing visual target is faster when an accessory auditory stimulus occurs in close spatiotemporal vicinity. Such facilitation of saccadic reaction time (SRT) is well-documented, but the exact neural mechanisms underlying the crossmodal effect remain to be elucidated. From EEG/MEG studies it has been hypothesized that coupled oscillatory activity in primary sensory cortices regulates multisensory processing. Specifically, it is assumed that the phase of an ongoing neural oscillation is shifted due to the occurrence of a sensory stimulus so that, across trials, phase values become highly consistent (phase reset). If one can identify the phase an oscillation is reset to, it is possible to predict when temporal windows of high and low excitability will occur. However, in behavioral experiments the pre-stimulus phase will be different on successive repetitions of the experimental trial, and average performance over many trials will show no signs of the modulation. Here we circumvent this problem by repeatedly presenting an auditory accessory stimulus followed by a visual target stimulus with a temporal delay varied in steps of 2 ms. Performing a discrete time series analysis on SRT as a function of the delay, we provide statistical evidence for the existence of distinct peak spectral components in the power spectrum. These frequencies, although varying across participants, fall within the beta and gamma range (20 to 40 Hz) of neural oscillatory activity observed in neurophysiological studies of multisensory integration. Some evidence for high-theta/alpha activity was found as well. Our results are consistent with the phase reset hypothesis and demonstrate that it is amenable to testing by purely psychophysical methods. Thus, any theory of multisensory processes that connects specific brain states with patterns of saccadic responses should be able to account for traces of oscillatory activity in observable behavior.     

Neural Activity in the Macaque Putamen Associated with Saccades and Behavioral Outcome

It is now widely accepted that the basal ganglia nuclei form segregated, parallel loops with neocortical areas. The prevalent view is that the putamen is part of the motor loop, which receives inputs from sensorimotor areas, whereas the caudate, which receives inputs from frontal cortical eye fields and projects via the substantia nigra pars reticulata to the superior colliculus, belongs to the oculomotor loop. Tracer studies in monkeys and functional neuroimaging studies in human subjects, however, also suggest a potential role for the putamen in oculomotor control. To investigate the role of the putamen in saccadic eye movements, we recorded single neuron activity in the caudal putamen of two rhesus monkeys while they alternated between short blocks of pro- and anti-saccades. In each trial, the instruction cue was provided after the onset of the peripheral stimulus, thus the monkeys could either generate an immediate response to the stimulus based on the internal representation of the rule from the previous trial, or alternatively, could await the visual rule-instruction cue to guide their saccadic response. We found that a subset of putamen neurons showed saccade-related activity, that the preparatory mode (internally- versus externally-cued) influenced the expression of task-selectivity in roughly one third of the task-modulated neurons, and further that a large proportion of neurons encoded the outcome of the saccade. These results suggest that the caudal putamen may be part of the neural network for goal-directed saccades, wherein the monitoring of saccadic eye movements, context and performance feedback may be processed together to ensure optimal behavioural performance and outcomes are achieved during ongoing behaviour.     

Reduced variability of ongoing and evoked cortical activity leads to improved behavioral performance

Sensory responses of the brain are known to be highly variable, but the origin and functional relevance of this variability have long remained enigmatic. Using the variable foreperiod of a visual discrimination task to assess variability in the primate cerebral cortex, we report that visual evoked response variability is not only tied to variability in ongoing cortical activity, but also predicts mean response time. We used cortical local field potentials, simultaneously recorded from widespread cortical areas, to gauge both ongoing and visually evoked activity. Trial-to-trial variability of sensory evoked responses was strongly modulated by foreperiod duration and correlated both with the cortical variability before stimulus onset as well as with response times. In a separate set of experiments we probed the relation between small saccadic eye movements, foreperiod duration and manual response times. The rate of eye movements was modulated by foreperiod duration and eye position variability was positively correlated with response times. Our results indicate that when the time of a sensory stimulus is predictable, reduction in cortical variability before the stimulus can improve normal behavioral function that depends on the stimulus.