On the production and correction of involuntary prosaccades in a gap antisaccade task.

In an antisaccade task, where saccades in the direction opposite of a suddenly presented stimulus are required, certain numbers of prosaccades can occur. The hypothesis is put forward that poor fixation and poor voluntary saccade control constitute two independent sources for the errors. This possibility is investigated by including the corrections of the errors in the analysis. First, the eye movements of 346 normal subjects (group N) performing a gap antisaccade and an overlap prosaccade task were measured. For each subject the proportion of express saccades in the overlap prosaccade task and the proportion of prosaccades in the gap antisaccade task were determined. The data of 150 subjects with more than 20% proerrors were divided into two groups: group A with relatively many, group B with relatively few express saccades in the overlap prosaccade task. Group A subjects produced their errors after significantly shorter reaction times and they corrected their errors significantly faster and more often than group B subjects. Second, we analysed the data of three groups of subjects: the complete normal group N, a group D of dyslexic subjects (n=343), and a group T containing all subjects irrespective of their cognitive achievements (n=780). A highly significant negative correlation exists between the correction rates and the error rates. A factor analysis of the variables performed for each group separately results in only two factors, one describing prosaccade the other antisaccade control. Only the error rate contributes significantly to both factors indicating that high errors may have two independent reasons.

Poor saccadic control correlates with dyslexia.

A large group of subjects, either average readers or reading/spelling disabled subjects (n = 185; age between 8-25 years; M = 13 +/- 4 years), were tested in various standardized cognitive tasks including reading/spelling assessment and in non-cognitive saccadic eye movement tasks. Dyslexics were separated into a subgroup (D1) with deficits in the serial auditory short-term memory and a subgroup (D2) with an isolated low achievement in reading/writing. Control subjects had no relevant cognitive deficit of any type. Saccadic eye movements were measured in a single target and in a sequential-target task. A significant correlation was found between abnormal saccadic control and reading disability. The two dyslexic groups showed only slight differences. As compared to the control group, the mean values of the standard deviations of the saccadic reaction times (SRT) and the amount of late saccades (SRT > 700) were significantly increased in both dyslexic groups and especially in group D1 who also showed an increased amount of anticipatory saccades. The number of express saccades (SRT = 80-134 ms) was increased, but not significantly, in D2 dyslexics. Both dyslexic groups produced significantly more regressive saccades in the sequential-target task. The correlation between saccadic variables and "reading factor" was 0.4. Significant deviations from normal performance of the saccadic variables were found in an estimated 50% of the dyslexics as compared to 20% of the control subjects. In spite of their worse level in saccadic control, dyslexics also developed with age in the eye movement performance as the control subjects did. Yet, the development was slower in group D1. It is suggested that reading process and saccade system are both controlled by visuo spatial attention and fixation systems that maybe impaired or develop slowly in many dyslexic subjects.

Detecting multimodality in saccadic reaction time distributions in gap and overlap tasks.

In many cases the distribution of saccadic reaction times (SRT) deviates considerably from a unimodal distribution and may often exhibit several peaks. We present a statistical approach to determining the number and form of the individual peaks. The overall density of the reaction times fi(t), i = 1...M obtained in M different experiments with the same subject is described as the sum of K basis functions xk(t), k = 1...K with different weights and an error term. A change in the experimental conditions is assumed to cause a change in the weights, not in the basis functions. We minimize the square of the difference (measured data minus approximation), divided by the error of the data. Incrementing K step by step we determine the necessary number of basis functions. This method is applied to data of six subjects tested in different saccade tasks. We detect five different modes: two in the range 80-140 ms (express modes), two in the range 145-190 ms (fast-regular mode) and one at about 230 ms (slow-regular mode). These modes are located at about the same positions for different subjects. The method presented here not only proves statistically the existence of several modes in SRT distributions but also allows the distributions to be described by a few characteristic numbers that go beyond the mean values and standard deviations.

Saccadic reaction times: a statistical analysis of multimodal distributions.

The distributions of saccadic reaction times (SRT) often deviate from unimodal normal distributions. An excess-mass procedure was used to detect peaks in 963 data sets containing 90,927 reaction times from 170 subjects. About 55% showed one, 30% two, 12% three and 3% four peaks. According to their clustering along the reaction time scale the modes could be classified into express (90-120 msec), fast regular (135-170 msec) and slow regular (200-220 msec) modes. Among the unimodal distributions 29% had peaks in the range of the express mode and 46% had peaks in the range of the fast regular mode. Therefore, 87% of the data sets support the notion of saccadic reaction time distributions being the superposition of three modes. All experimental distributions were fitted by as many gamma distributions as determined by the excess-mass test. The significance of the multimodality for saccade generation processes is discussed.

On the development of voluntary and reflexive components in human saccade generation.

The saccadic performance of a large number (n = 281) of subjects of different ages (8-70 years) was studied applying two saccade tasks: the prosaccade overlap (PO) task and the antisaccade gap (AG) task. From the PO task, the mean reaction times and the percentage of express saccades were determined for each subject. From the AG task, the mean reaction time of the correct antisaccades and of the erratic prosaccades were measured. In addition, we determined the error rate and the mean correction time, i.e. the time between the end of the first erratic prosaccade and the following corrective antisaccade. These variables were measured separately for stimuli presented (in random order) at the right or left side. While strong correlations were seen between variables for the right and left sides, considerable side asymmetries were obtained from many subjects. A factor analysis revealed that the seven variables (six eye movement variables plus age) were mainly determined by only two factors, V and F. The V factor was dominated by the variables from the AG task (reaction time, correction time, error rate) the F factor by variables from the PO task (reaction time, percentage express saccades) and the reaction time of the errors (prosaccades!) from the AG task. The relationship between the percentage number of express saccades and the percentage number of errors was completely asymmetric: high numbers of express saccades were accompanied by high numbers of errors but not vice versa. Only the variables in the V factor covaried with age. A fast decrease of the antisaccade reaction time (by 50 ms), of the correction times (by 70 ms) and of the error rate (from 60 to 22%) was observed between age 9 and 15 years, followed by a further period of slower decrease until age 25 years. The mean time a subject needed to reach the side opposite to the stimulus as required by the antisaccade task decreased from approximately 350 to 250 ms until age 15 years and decreased further by 20 ms before it increased again to approximately 280 ms. At higher ages, there was a slight indication for a return development. Subjects with high error rates had long antisaccade latencies and needed a long time to reach the opposite side on error trials. The variables obtained from the PO task varied also significantly with age but by smaller amounts. The results are discussed in relation to the subsystems controlling saccade generation: a voluntary and a reflex component the latter being suppressed by active fixation. Both systems seem to develop differentially. The data offer a detailed baseline for clinical studies using the pro- and antisaccade tasks as an indication of functional impairments, circumscribed brain lesions, neurological and psychiatric diseases and cognitive deficits.

The analysis of saccadic eye movements from gap and overlap paradigms.

This protocol describes the acquisition and evaluation of saccadic eye movement data for use in basic neuroscience research and clinical application. The experimental protocol requires the subject to make saccadic eye movements in response to visual stimuli presented, in random order, on consecutive trials. The gap and overlap paradigms are described together with the instruction to generate pro- or antisaccades. The protocol includes the description of saccade detection, the determination of the beginning, the end, the size, and the velocity of a saccade, the exact way of calculating the proportion of different kinds of trials, and the treatment of erratic or artifact trials. Relevant variables are defined. The results obtained from a large number (300) of subjects of different ages (8-65 years) are described and analysed with respect to their development with age. The protocol allows to test a subject's saccadic status in many different circumstances in particular with respect to diagnostic help in neurology, psychiatry and psychology.

The three-loop model: a neural network for the generation of saccadic reaction times.

This paper presents a computer simulation of the three-loop model for the temporal aspects of the generation of visually guided saccadic eye movements. The intention is to reproduce complex experimental reaction time distributions by a simple neural network. The operating elements are artificial but realistic neurones. Four modules are constructed, each consisting of 16 neural elements. Within each module, the elements are connected in an all-to-all manner. The modules are working parallel and serial according to the anatomically and physiologically identified visuomotor pathways including the superior colliculus, the frontal eye fields, and the parietal cortex. Two transient-sustained input lines drive the network: one represents the visual activity produced by the onset of the saccade target, the other represents a central activity controlling the preparation of saccades, e.g. the end of active fixation. The model works completely deterministically; its stochastic output is a consequence of the stochastic properties of the input only. Simulations show how multimodal distributions of saccadic reaction times are produced as a natural consequence of the model structure. The gap effect on saccadic reaction times is correctly produced by the model: depending only on the gap duration (all model parameters unchanged) express, fast-regular, and slow-regular saccades are obtained in different numbers. In agreement with the experiments, bi- or trimodal distributions are produced only for medium gap durations (around 200 ms), while for shorter or longer gaps the express mode disappears and the distributions turn bi- or even unimodal. The effect of varying the strength of the transient-sustained components and the ongoing activity driving the hierarchically highest module are considered to account for the interindividual variability of the latency distributions obtained from different subjects, effects of different instructions to the same subject, and the observation of express makers (subjects who produce exclusively express saccades). How the model can be extended to describe the spatial aspects of the saccade system will be discussed as well as the effects of training and/or rapid adaptation to experimental conditions.