Visual fixation offsets affect both the initiation and the kinematic features of saccades

It is well known that the removal of a fixation point prior to the presentation of a peripheral target dramatically reduces saccadic reaction time (SRT). This effect has become known as the “gap effect”. The present study examined several detailed kinematic variables to determine whether the removal of the fixation point also affects the manner in which saccades are produced. The findings indicate that saccades that were initiated after the removal of the fixation point had higher average velocities and reached greater peak velocities, accelerations, and decelerations than did saccades produced in the presence of the fixation point. The results suggest that the removal of the fixation point may affect the force-time curves of saccades in addition to affecting the time needed to initiate the saccades. Received: 21 February 1997 / Accepted: 24 July 1997     

Proportional Jerk: A New Measure of Motion as Applied to Eye Movements in Sleep and Waking

The third derivative of motion or the change in the rate of acceleration (also known as jerk) is examined in terms of its applicability to the study of psychophysiological function. An algorithm of this third derivative is presented to show that jerk can be derived from the arithmetic difference of two slopes which constitute the portion of motion being differentiated. By modifying the algorithm, a new parameter termed “proportional jerk’ or PJ is formulated whereby one slope is measured relative to the other slope; this PJ provides information about the smoothness of movement without being influenced by the velocity as is the traditional jerk measure. A practical application of the PJ to waking saccades and REMs in 11 human subjects revealed that REMs are significantly “jerkier’ or less smooth than waking calibration eye movements. Whereas the waking eye movements had a well-defined negative phase of the PJ, the REMs did not show such stereotypical behavior. This is in accord with previous work which showed that waking saccade velocity increases to a maximum and then decreases whereas REMs maintain their peak or near peak velocities for varying periods. It was suggested that PJ can be useful in detecting subtle temporal changes in the course of movements and may be used as a parameter of motion even when the absolute amplitude is unknown.     

Accuracies of saccades to moving targets during pursuit initiation and maintenance

The overall goals of the studies presented here were to compare (1) the accuracies of saccades to moving targets with either a novel or a known target motion, and (2) the relationships between the measures of target motion and saccadic amplitude during pursuit initiation and maintenance. Since resampling of position error just prior to saccade initiation can confound the interpretation of results, the target ramp was masked during the planning and execution of the saccade. The results suggest that saccades to moving targets were significantly more accurate if the target motion was known from the early part of the trial (e.g., during pursuit maintenance) than in the case of novel target motion (e.g., during pursuit initiation); both these types of saccades were more accuate than those when target motion information was not available. Using target velocity in space as a rough estimate of the magnitude of the extra-retinal signal during pursuit maintenance, the saccadic amplitude was significantly associated with the extra-retinal target motion information after accounting for the position error. In most subjects, this association was stronger than the one between retinal slip velocity and saccadic amplitude during pursuit initiation. The results were similar even when the smooth eye motion prior to the saccade was controlled. These results suggest that different sources of target motion information (retinal image velocity vs internal representation of previous target motion in space) are used in planning saccades during different stages of pursuit. The association between retinal slip velocity and saccadic amplitude is weak during initiation, thus explaining poor saccadic accuracy during this stage of pursuit.     

Eye movement disorders after frontal eye field lesions in humans

Eye movements were recorded electro-oculographically in three patients with a small ischemic lesion affecting the left frontal eye field (FEF) and in 12 control subjects. Reflexive visually guided saccades (gap and overlap tasks), antisaccades, predictive saccades, memory-guided saccades, smooth pursuit and optokinetic nystagmus (OKN) were studied in the three patients. Staircase saccades and double step saccades were also studied in one of the three patients. For both leftward and rightward saccades, latency in the overlap task (but not in the gap task) and that of correct antisaccades and of memory-guided saccades was significantly increased, compared with the results of controls. There was a significant decrease in the amplitude gain of all rightward saccades programmed using retinotopic coordinates (gap and overlap tasks, predictive and memory-guided saccades), whereas the amplitude gain of corresponding leftward saccades was preserved. Such an asymmetry between leftward and rightward saccades was significant. In the staircase paradigm as well as for the first saccade in the double step paradigm (with the use of retinotopic coordinates in both cases), the amplitude gain of rightward saccades was also significantly lower than that of leftward saccades. Moreover, in the double step paradigm, the amplitude gain of the first rightward saccade was significantly lower than that of the second rightward saccade (programmed using extraretinal signals), which was preserved. The percentage of errors in the antisaccade task did not differ significantly from that of normal subjects. In the predictive saccade paradigm, the percentage of predictive rightward saccades was significantly decreased. The left smooth pursuit gain for all tested velocities, the right smooth pursuit gain for higher velocities, and the left OKN gain were significantly decreased. The results show, for the first time in humans, that the FEF plays an important role in (1) the disengagement from central fixation, (2) the control of contralateral saccades programmed using retinotopic coordinates, (3) saccade prediction and (4) the control of smooth pursuit and OKN, mainly ipsilaterally. In contrast, the left FEF did not appear to be crucial for the control of the only type of saccades programmed using extraretinal signals studied here.     

Saccades and multisaccadic gaze shifts are gated by different pontine omnipause neurons in head-fixed cats

 Pontine omnipause neurons (OPNs) have so far been considered as forming a homogeneous group of neurons whose tonic firing stops during the duration of saccades, when the head is immobilized. In cats, they pause for the total duration of gaze shifts, when the head is free to move. In the present study, carried out on alert cats with fixed heads, we present observations made during self-initiated saccades and during tracking of a moving target which show that the OPN population is not homogeneous. Of the 76 OPNs we identified, 39 were found to have characteristics similar to those of previously described neurons, ”saccade” (S-) OPNs: (1) the durations of their pauses were significantly correlated with the durations of saccades; (2) the discharge ceased shortly before saccade onset and resumed before saccade end; (3) visual responses to target motion were excitatory; and (4) during tracking, S-OPNs interrupted the discharge for the duration of saccades and resumed firing during perisaccadic ”drifts”. However, the characteristics of 37 neurons (”complex” (C-) OPNs) were different: (1) the pause duration was not correlated with the duration of self-initiated saccades; (2) time lead of pause onsets relative to saccades was, on average, longer than in the group of S-OPNs, and firing resumed after the saccade end; (3) visual target motion suppressed tonic discharges; and (4) during tracking, firing was interrupted for the total duration of gaze shifts, including not only saccades but also perisaccadic ”drifts”. We conclude that cat OPNs can be subdivided into two main groups. The first comprises neurons whose firing patterns are compatible with gating individual saccades (”saccade” OPNs). The second group consists of ”complex” OPNs whose firing characteristics are appropriate to gate total gaze displacements rather than individual saccades. The function of these neurons may be to disinhibit pontobulbar circuits participating in the generation of saccade sequences and associated perisaccadic drifts. Received: 20 January 1998 / Accepted: 22 October 1998     

Independent contributions of the orienting of attention,fixation offset and bilateral stimulation on human saccadic latencies

In a series of experiments we examined the effects of the endogenous orienting of visual attention on human saccade latency. Three separate manipulations were performed: the orienting of visual attention, the prior offset of fixation (gap paradigm) and the bilateral presentation of saccade targets. Each of these manipulations was shown to make an independent contribution to saccade latency. In experiments 1 and 2 subjects were instructed to orient their attention covertly to a location by a verbal pre-cue; targets could appear in the attended hemifield (valid) or in the non-attended hemifield (invalid) together with a no-instruction (neutral) condition. Saccades were made under fixation gap and overlap conditions, to either single targets or two bilaterally presented targets which appeared at equal and opposite eccentricities in both hemifields. The results showed a large increase (cost) of saccade latency to invalid targets and a small non-significant decrease (benefit) of saccade latency to valid targets. The cost associated with invalid targets replicates the meridian crossing effect shown in manual reaction time experiments and is consistent with the hemifield inhibition and premotor models of attentional orienting. The use of a gap procedure produced a generalised facilitation of saccade latency, which was not modified by the prior orienting of visual attention. The magnitude of the gap effect was similar for saccades made to attended and non-attended stimulis. This suggests that the gap effect may be due to ocular motor disengagement, or a warning signal effect, rather than to the prior disengagement of visual attention. When two targets were presented simultaneously, one in each hemifield, saccade latency was slowed compared with the single target condition. The magnitude of this slowing was unaffected by the prior orienting of visual attention or by the fixation condition. The slowing was examined in more detail in experiment 3, by presenting targets with brief offset delays. The latency increase was maximal if the two targets were presented simultaneously and decreased if the distractor appeared at short intervals (20–80 ms) before or after the saccade target onset. If the non-attended stimulus was presented at greater intervals (160, 240 ms) before the saccade target, then a facilitation effect was observed. This demonstrates that the onset of a distractor in the non-attended hemifield can have both an inhibitory and a facilitatory effect on a saccade production.     

Differentiating early Parkinson's disease and multiple system atrophy with parkinsonism by saccade velocity profiles

ObjectivePatients with Parkinson's disease (PD) and multiple system atrophy both present predominantly with parkinsonism at early stages, whereas cerebellar symptoms are largely masked in multiple system atrophy with parkinsonism (MSAP). We sought to determine whether the velocity profiles of saccades could be used to differentiate between these two disorders, revealing the underlying basal ganglia and/or cerebellar dysfunction and brainstem pathology in these disorders.MethodsSixteen MSA-P patients, 63 PD patients, and 36 age-matched normal subjects performed the visually guided (VGS) and memory-guided saccade (MGS) tasks. Targets were presented at eccentricities of 5, 10, 20, and 30 degrees. The amplitude, peak velocity, and duration of saccades were compared among subject groups. Duration was further subdivided into acceleration and deceleration periods, corresponding to the times before and after peak velocity. These parameters correlated with the severity of Parkinsonism as assessed by the UPDRS motor score.ResultsHypometria predominated in both PD and MSAP patients, whereas hypermetria, frequently noted in cerebellar ataxia, was rarely observed. Saccades in MSAP were characterized both by prolonged acceleration and deceleration periods with reduced peak velocity. In contrast, the velocity profile of PD patients was characterized mainly by the prolonged deceleration period. The changes observed in velocity profiles of MGS deteriorated with advancing severity of parkinsonism in MSAP and PD patients.ConclusionSaccade profiles provide useful information for differentiating between PD and MSAP at early stages. While the changes in velocity profiles may be explained by the cerebellar and brainstem pathology in MSAP, the changes in velocity profile in both PD and MSAP correlated significantly with increasing severity of Parkinsonism in both disorders, suggesting a link with striatonigral pathology.SignificanceThe differential changes in saccade velocity profiles of MSAP and PD may be used as a measure indexing the progression of cerebellar and basal ganglia dysfunction as well as for assessing the functional improvement when clinical treatment becomes available.     

Behavioral analysis of predictive saccade tracking as studied by countermanding

The ability to make predictive saccadic eye movements is dependent on neural signals that anticipate the onset of a visual target. We used a novel paradigm—based on the saccade-countermanding task—as a tool to investigate rhythm saccade pacing and to provide information on the mechanisms of predictive timing. In particular, we examined the ability of normal subjects to stop a sequence of periodically paced eye movements when cued by a stop signal that was presented at different times with respect to the last target of the sequence (stop signal delay, SSD). The timing of the stop signal affected the ability to stop the saccadic sequence (make a saccade to a central target rather than to the peripheral alternating targets) in different ways, depending on the preceding tracking behavior. For the same SSD, subjects cancelled fewer trials during predictive tracking (promoted by tracking targets alternating at a fast pacing rate, 1.0 Hz) than during reactive tracking (tracking alternating targets at a low pacing rate, 0.2 Hz). In addition, on non-canceled trials, there was an increase in the delay of the corrective saccade to the central target with increasing SSD for pacing at 0.2 Hz, but the timing of the corrective saccade remained near constant for 1.0 Hz pacing. In examining the timing between movements, we estimate that the repetitive GO process that drives the saccades during predictive tracking begins earlier and has a shorter duration than the repetitive GO process during reactive tracking. These behavioral results provide further insight into the initiation process of predictive responses. In particular, the reduced reaction time and the corresponding short duration of the predictive process may result from a faster accumulation of neuronal discharge to a relatively fixed threshold.     

Sensory versus motor information in the control of predictive saccade timing

Humans readily make predictive saccades to periodic alternating targets. This predictive behavior depends on internal monitoring of timing error of past saccades in order to determine the time of initiation of future saccades; our earlier studies have confirmed this by finding correlations between latencies of consecutive predictive saccades. It is natural to consider that timing error is determined by visual detection of the difference between the time the target appears and the time the eyes arrive at the target; this in turn implies that saccades must actually be produced in order for their timing errors to be determined and predictive saccade timing to be established. We tested this hypothesis by having subjects view alternating visual targets while fixating a central target in order to eliminate saccade production. After six alternating target presentations, subjects began tracking the alternating targets. Tracking performance was assessed with an error measure that compared saccade latency and inter-saccade interval with desired values (zero and inter-stimulus interval, respectively). Errors in this Prior Viewing paradigm were compared to those from a conventional De Novo paradigm in which saccades began as soon as the alternating targets were presented. Saccades under Prior Viewing reached a low-error steady predictive state more rapidly than under De Novo tracking. The initial saccade under Prior Viewing had a higher latency than the others, suggesting that this saccade was reactive even though the paradigm is predictable; other reasons for this higher latency include time to disengage from the fixation target and time required to pre-program the initial set of saccades. The results show that visual detection of timing error from an actual motor act (saccades) is not necessary to establish predictive saccadic pacing: sensory-only information from viewing the moving targets can help to establish this predictive state.     

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

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