Centripetal Saccades Research Articles

Recentering bias for temporal saccades only: Evidence from binocular recordings of eye movements.

It is well known that the saccadic system presents multiple asymmetries. Notably, temporal (as opposed to nasal) saccades, centripetal (as opposed to centrifugal) saccades (i.e., the recentering bias) and saccades from the abducting eye (as opposed to the concomitant saccades from the adducting eye) exhibit higher peak velocities. However, these naso-temporal and centripetal-centrifugal asymmetries have always been studied separately. It is thus unknown which asymmetry prevails when there is a conflict between both asymmetries, i.e., in case of centripetal nasal saccades or centrifugal temporal saccades. This study involved binocular recordings of eye movements to examine both the naso-temporal and centripetal-centrifugal asymmetries so as to determine how they work together. Twenty-eight participants had to make saccades toward stimuli presented either centrally or in the periphery in binocular conditions. We found that temporal and abducting saccades always exhibit higher peak velocities than nasal and adducting saccades, irrespective of their centripetal or centrifugal nature. However, we showed that the velocity advantage for centripetal saccades is only found for temporal and not for nasal saccades. Such a result is of importance as it could provide new insights about the physiological origins of the asymmetries found in the saccadic system.

Visual straight-ahead preference in saccadic eye movements.

Ocular saccades bringing the gaze toward the straight-ahead direction (centripetal) exhibit higher dynamics than those steering the gaze away (centrifugal). This is generally explained by oculomotor determinants: centripetal saccades are more efficient because they pull the eyes back toward their primary orbital position. However, visual determinants might also be invoked: elements located straight-ahead trigger saccades more efficiently because they receive a privileged visual processing. Here, we addressed this issue by using both pro- and anti-saccade tasks in order to dissociate the centripetal/centrifugal directions of the saccades, from the straight-ahead/eccentric locations of the visual elements triggering those saccades. Twenty participants underwent alternating blocks of pro- and anti-saccades during which eye movements were recorded binocularly at 1 kHz. The results confirm that centripetal saccades are always executed faster than centrifugal ones, irrespective of whether the visual elements have straight-ahead or eccentric locations. However, by contrast, saccades triggered by elements located straight-ahead are consistently initiated more rapidly than those evoked by eccentric elements, irrespective of their centripetal or centrifugal direction. Importantly, this double dissociation reveals that the higher dynamics of centripetal pro-saccades stem from both oculomotor and visual determinants, which act respectively on the execution and initiation of ocular saccades.

High-Field fMRI Reveals Brain Activation Patterns Underlying Saccade Execution in the Human Superior Colliculus

BackgroundThe superior colliculus (SC) has been shown to play a crucial role in the initiation and coordination of eye- and head-movements. The knowledge about the function of this structure is mainly based on single-unit recordings in animals with relatively few neuroimaging studies investigating eye-movement related brain activity in humans.Methodology/Principal FindingsThe present study employed high-field (7 Tesla) functional magnetic resonance imaging (fMRI) to investigate SC responses during endogenously cued saccades in humans. In response to centrally presented instructional cues, subjects either performed saccades away from (centrifugal) or towards (centripetal) the center of straight gaze or maintained fixation at the center position. Compared to central fixation, the execution of saccades elicited hemodynamic activity within a network of cortical and subcortical areas that included the SC, lateral geniculate nucleus (LGN), occipital cortex, striatum, and the pulvinar.Conclusions/SignificanceActivity in the SC was enhanced contralateral to the direction of the saccade (i.e., greater activity in the right as compared to left SC during leftward saccades and vice versa) during both centrifugal and centripetal saccades, thereby demonstrating that the contralateral predominance for saccade execution that has been shown to exist in animals is also present in the human SC. In addition, centrifugal saccades elicited greater activity in the SC than did centripetal saccades, while also being accompanied by an enhanced deactivation within the prefrontal default-mode network. This pattern of brain activity might reflect the reduced processing effort required to move the eyes toward as compared to away from the center of straight gaze, a position that might serve as a spatial baseline in which the retinotopic and craniotopic reference frames are aligned.

Long Time-Constant Behavior of the Oculomotor Plant in Barbiturate-Anesthetized Primate

The mechanics of the extraocular muscles and orbital tissue ("oculomotor plant") can be approximated by a small number of viscoelastic (Voigt) elements in series. Recent analysis of the eye's return from displacement in lightly anesthetized rhesus monkeys has suggested a four-element plant model with time constants (TCs) of approximately 0.01, 0.1, 1, and 10 s. To demonstrate directly the presence of long (1,10 s) TC elements and to assess their contribution quantitatively, horizontal eye displacement was induced in Cynomolgus monkeys under deep barbiturate anesthesia that prevented interference from spontaneous eye movements. The displacement was maintained for either a prolonged (30 s) or brief (0.2 s) period before release. Return to resting position took 20-30 s after prolonged displacement but only 1-2 s after brief displacement, consistent with the presence of long TC elements that would only be substantially stretched in the former condition. Quantitative fitting of the release curves after prolonged displacement indicated that the two long TC elements contribute a substantial proportion (approximately 30%) of the total plant compliance. A model based on the estimated compliance values is shown to account quantitatively both for our release data and for Goldstein and Robinson's data on hysteresis of ocular motoneuron firing rates measured after centripetal saccades following prolonged eccentric fixation. Long time-constant elements in the plant thus make a substantial contribution to some types of eye movement, and their inclusion in plant models can help interpret the firing patterns of single units in the oculomotor system.

Saccade-related neurons in the primate fastigial nucleus: what do they encode?

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 degrees ) 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 approximately 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.

Cause of kinematic differences during centrifugal and centripetal saccades

Measurements of eye movements have shown that centrifugal movements (i.e. away from the primary position) have a lower maximum velocity and a longer duration than centripetal movements (i.e. toward the primary position) of the same size. In 1988 Pelisson proposed that these kinematic differences might be caused by differences in the neural command signals, oculomotor mechanics or a combination of the two. By using the result of muscle force measurements that were made in recent years (Orbit TM1.8 Gaze mechanics simulation, Eidactics, San Francisco, 1999) we simulated the muscle forces during centrifugal and centripetal saccades. Based on these simulations we show that the cause of the kinematic differences between the centrifugal and centripetal saccades is the non-linear force–velocity relationship (i.e. muscle viscosity) of the muscles.

Expression of a re-centering bias in saccade regulation by superior colliculus neurons.

In previous studies of saccadic eye movement reaction time, the manipulation of initial eye position revealed a behavioral bias that facilitates the initiation of movements towards the central orbital position. An interesting hypothesis for this re-centering bias suggests that it reflects a visuo-motor optimizing strategy, rather than peripheral muscular constraints. Given that the range of positions that the eyes can take in the orbits delimits the extent of visual exploration by head-fixed subjects, keeping the eyes centered in the orbits may indeed permit flexible orienting responses to engaging stimuli. To investigate the influence of initial eye position on central processes such as saccade selection and initiation, we examined the activity of saccade-related neurons in the primate superior colliculus (SC). Using a simple reaction time paradigm wherein an initially fixated visual stimulus varying in position was extinguished 200 ms before the presentation of a saccadic target, we studied the relationship between initial eye position and neuronal activation in advance of saccade initiation. We found that the magnitude of the early activity of SC neurons, especially during the immediate pre-target period that followed the fixation stimulus disappearance, was correlated with changes in initial eye position. For the great majority of neurons, the pre-target activity increased with changes in initial eye position in the direction opposite to their movement fields, and it was also strongly correlated with the concomitant reduction in reaction time of centripetal saccades directed within their movement fields. Taking into account the correlation with saccadic reaction time, the relationship between neuronal activity and initial eye position remained significant. These results suggest that eye-position-dependent changes in the excitability of SC neurons could represent the neural substrate underlying a re-centering bias in saccade regulation. More generally, the low frequency SC pre-target activity could use eccentric eye position signals to regulate both when and which saccades are produced by promoting the emergence of a high frequency burst of activity that can act as a saccadic command. However, only saccades initiated within approximately 200 ms of target presentation were associated with SC pre-target activity. This eye-dependent pre-target activation mechanism therefore appears to be restricted to the initiation of saccades with relatively short reaction times, which specifically require the integrity of the SC.

Disconjugate adaptation of saccades: contribution of binocular and monocular mechanisms

We studied the effects of prism–induced disparity on static and intrasaccadic alignment in six normal human subjects. A ten diopter base–out prism, calling for convergence, was placed in front of the central field of the right eye, so that at the center the eye viewed through the prism; at left and right, outside the prism. During 15 min of training, subjects made repetitive saccades solely in the right field of vision (C-R-C sequence). This paradigm required relative divergence for centrifugal (C-R) saccades and relative convergence for centripetal (R-C) saccades, as well as increase of the amplitude for all saccades made by the right eye. We found that during training, all subjects incorporated the necessary change in alignment into the saccades. After training the resultant intrasaccadic disconjugacy persisted when tested during monocular viewing, indicating that motor learning had occurred. Subjects demonstrated increased divergence for C-R and increased convergence for R-C saccades, in accordance with the change acquired during adaptation to the prism. In addition, five subjects developed increased divergence for C-L saccades, for which they did not train. Smaller and less consistent divergence was also observed for L-C saccades. Changes in intrasaccadic alignment were accompanied by changes in the relative velocities of the two eyes’ saccades and slowing of the peak velocities in both eyes during training. Static alignment showed a general tendency toward convergence that did not parallel the changes in the intrasaccadic alignment, suggesting that saccade adaptation is system–specific. The pattern of transfer of the intrasaccadic disconjugacy to saccades in the untrained field and the changes in the relative speeds of the two eyes cannot be explained by monocular adjustment of the saccades. Our results indicate that both a binocular mechanism—saccade–vergence interaction—and monocular adaptation contribute to disconjugate adaptation of saccades.

Effects of lesions of the oculomotor vermis on eye movements in primate: saccades.

We studied the effects on saccades of ablation of the dorsal cerebellar vermis (lesions centered on lobules VI and VII) in three monkeys in which the deep cerebellar nuclei were spared. One animal, with a symmetrical lesion, showed bilateral hypometric horizontal saccades. Two animals, with asymmetrical lesions, showed hypometric ipsilateral saccades, and saccades to vertically positioned targets were misdirected, usually deviating away from the side to which horizontal saccades were hypometric. Postlesion, all animals showed an increase (2- to 5-fold) in trial-to-trial variability of saccade amplitude. They also showed a change in the ratio of the amplitudes of centripetal to centrifugal saccades (orbital-position effect); usually centrifugal saccades became smaller. In the two animals with asymmetrical lesions, for saccades in the hypometric direction, latencies were markedly increased (up to approximately 500 ms). There was also an absence of express and anticipatory saccades in the hypometric direction. When overall saccade latency was increased, centrifugal saccades became relatively more delayed than centripetal saccades. The dynamic characteristics of saccades were affected to some extent in all monkeys with changes in peak velocity, eye acceleration, and especially eye deceleration. There was relatively little effect of orbital position on saccade dynamics, however, with the exception of one animal that showed an orbital position effect for eye acceleration. In a double-step adaptation paradigm, animals showed an impaired ability to adaptively adjust saccade amplitude, though increased amplitude variability postlesion may have played a role in this deficit. During a single training session, however, the latency to corrective saccades-which had been increased postlesion-gradually decreased and so enabled the animal to reach the final position of the target more quickly. Overall, both in the early postlesion period and during recovery, changes in saccade amplitude and latency tended to vary together but not with changes in saccade dynamics or adaptive capability, both of which behaved relatively independently. These findings suggest that the cerebellum can adjust saccade amplitude and saccade dynamics independently. Our results implicate the cerebellar vermis directly in every aspect of the on-line control of saccades: initiation (latency), accuracy (amplitude and direction), and dynamics (velocity and acceleration) and also in the acquisition of adaptive ocular motor behavior.

Properties of Horizontal Saccades Accompanied by Blinks

Using the magnetic search coil technique to record eye and lid movements, we investigated the effect of voluntary blinks on horizontal saccades in five normal human subjects. The main goal of the study was to determine whether changes in the dynamics of saccades with blinks could be accounted for by a superposition of the eye movements induced by blinks as subjects fixated a stationary target and saccadic movements made without a blink. First, subjects made voluntary blinks as they fixed on stationary targets located straight ahead or 20 degrees to the right or left. They then made saccades between two continuously visible targets 20 or 40 degrees apart, while either attempting not to blink, or voluntarily blinking, with each saccade. During fixation of a target located straight ahead, blinks induced brief downward and nasalward deflections of eye position. When subjects looked at targets located at right or left 20 degrees, similar initial movements were made by four of the subjects, but the amplitude of the adducted eye was reduced by 65% and was followed by a larger temporalward movement. Blinks caused substantial changes in the dynamic properties of saccades. For 20 degrees saccades made with blinks, peak velocity and peak acceleration were decreased by approximately 20% in all subjects compared with saccades made without blinks. Blinks caused the duration of 20 degrees saccades to increase, on average, by 36%. On the other hand, blinks had only small effects on the gain of saccades. Blinks had little influence on the relative velocities of centrifugal versus centripetal saccades, and abducting versus adducting saccades. Three of five subjects showed a significantly increased incidence of dynamic overshoot in saccades accompanied by blinks, especially for 20 degrees movements. Taken with other evidence, this finding suggests that saccadic omnipause neurons are inhibited by blinks, which have longer duration than the saccades that company them. In conclusion, the changes in dynamic properties of saccades brought about by blinks cannot be accounted for simply by a summation of gaze perturbations produced by blinks during fixation and saccadic eye movements made without blinks. Our findings, especially the appearance of dynamic overshoots, suggest that blinks affect the central programming of saccades. These effects of blinks need to be taken into account during studies of the dynamic properties of saccades.

Abnormalities of ocular motility in myotonic dystrophy.

Are the oculomotor disturbances in myotonic dystrophy (MD), i.e. reduced smooth pursuit (SP) gain and reduced saccadic peak velocity (PV), of muscular or central origin? To answer this question the following two approaches were used. (i) The performance of SP was compared with the patient's ability to suppress the vestibulo-ocular reflex (VOR) visually (VOR suppression; VOR-S). In the latter task the SP system is involved, but the eyes hardly move within the orbits. A parallel impairment of SP and VOR-S would indicate a central dysfunction. (ii) Peak saccadic velocity was compared between two saccades performed to and fro in rapid succession. The intention was to measure any myotonic effect which might build up after the first saccade and slow down the second saccade. We studied 15 MD patients and 15 age-matched controls. Stimuli for slow eye responses consisted of sinusoidal horizontal rotations of the SP target and/or the vestibular rotation chair at frequencies between 0.1 and 0.8 Hz. Saccades were analysed in terms of PV. accuracy, duration and latency, comparing centripetal versus centrifugal saccades at short and long intersaccadic intervals (ISI; 400 ms and 900 ms, respectively). The SP gain was reduced in patients compared with the controls, the effect being most pronounced (32% less) at the highest stimulus frequency. Whereas VOR was normal in the patients, VOR-S was clearly impaired (50% worse at 0.8 Hz). Despite normal saccadic accuracy, peak saccadic velocity was significantly lower in the patient group (23% less for saccades of 12 degrees amplitude), similarly for centrifugal and centripetal saccades; all these differences were independent of the ISI. Latency was normal with centrifugal saccades, but was considerably increased with centripetal saccades at short ISI (67% longer compared with controls). The observation of a parallel degradation of SP and VOR-S in the patients is interpreted in terms of a central deficit in the SP pathways. Thus, it appears that slow eye movements were not impaired by muscle dystrophy and myotonia to a considerable degree in our patients. The increase in saccadic latency for centripetal saccades at the short ISI also reflects a central deficit. However, the observed slowing of saccades might have a myopathic or neural origin; a distinction was not possible at present. A myotonic origin of the saccade slowing seems unlikely, because the effect was independent of the presaccadic activation of the relaxing (antagonistic) eye muscle.

Immediate saccade amplitude disconjugacy induced by unequal images

We tested the ability of normal subjects to make changes in the conjugacy of their saccades. Subjects dichoptically viewed a grid the size of which was 10% larger in one eye. The grids were centred onto a flat screen at 57 cm or 1 m from the subject. Horizontal saccades immediately became larger in the eye viewing the larger grid. For some subjects this disconjugacy persisted even under subsequent monocular viewing. Such persistent changes occurred mainly in the field where the required disconjugacy was divergent for centrifugal saccades, convergent for centripetal saccades. Vertical saccades also developed compensatory disconjugacy; its amplitude was smaller but less variable. To explain these results we propose a fast associative learning mechanism that pairs peripheral disparity with saccades and is capable of producing saccade disconjugacy even in the absence of disparity. For horizontal saccades a secondary conditioning of monocular depth cues by the disparity would also be involved.

Saccadic burst neurons in the fastigial nucleus are not involved in compensating for orbital nonlinearities

1. To test whether the cerebellum is involved in compensating for orbital nonlinearities, we recorded discharges of saccadic burst neurons in the fastigial oculomotor region (FOR) during both centrifugal and centripetal saccades in two trained macaque monkeys. 2. We also investigated the metrics of centrifugal and centripetal saccades with the same amplitude while output signals from the fastigial nucleus were blocked by muscimol, an agonist of inhibitory neurotransmitter gamma-aminobutyric acid (GABA), injected into the FOR. 3. We studied discharges of 32 fastigial neurons that were inhibited by microstimulation of the oculomotor vermis. These neurons exhibited unique burst responses during contralaterally and ipsilaterally directed saccades, as shown in our previous report. The discharge pattern, the peak burst frequency, the burst lead time, and the burst duration were consistent, regardless of initial eye position. 4. The muscimol injection produced hypermetric saccades toward the injection side and hypometric saccades to the opposite side, regardless of whether saccade was centripetally or centrifugally directed. 5. These findings suggest that the FOR is not involved in the compensation of saccade metrics for orbital nonlinearities.

Role of the caudal fastigial nucleus in saccade generation. II. Effects of muscimol inactivation.

1. We studied the effect of temporarily inhibiting neurons in the caudal fastigial nucleus in two rhesus macaques trained to make saccades to jumping targets. We placed injections of the gamma-aminobutyric acid (GABA) agonist muscimol unilaterally or bilaterally at sites in the caudal fastigial nucleus where we had recorded saccade-related neurons a few minutes earlier. 2. Unilateral injections (n = 9) made horizontal saccades to the injected side hypermetric and those to the other side hypometric (mean gain of 1.37 and 0.61, respectively, for 10 degrees target steps, and 1.26 and 0.81 for 20 degrees target steps; normal saccade gain was 0.96). Saccades to vertical targets showed a small but significant hypermetria and curved strongly toward the side of the injection. The trajectories and end points of all targeted saccades were more variable than normal. 3. After unilateral injections, centripetal saccades were slightly larger than centrifugal saccades (mean gains for ipsilateral saccades were 1.42 and 1.31, respectively, for 10 degrees target steps, and 1.37 and 1.15 for 20 degrees target steps). 4. Unilateral injections increased the average acceleration of ipsilateral saccades and decreased the acceleration of contralateral saccades. Injections decreased both the acceleration and deceleration of vertical saccades. 5. After dysmetric saccades, monkeys acquired the target with an abnormally high number of hypometric corrective saccades. Injection increased the average number of corrective saccades from 0.6 to 2.1 after 10 degrees horizontal target steps and from 0.8 to 2.1 after 20 degrees steps. The size of each successive corrective saccade in a series decreased, and the latency from the previous corrective saccade increased. 6. Bilateral injections (n = 2) of muscimol, in which we injected first into the left caudal fastigial nucleus and then, within 30 min, into the right, made all saccades hypermetric (mean gain for 10 degrees right, left, up, and down saccades was 1.18, 1.49, 1.43, and 1.10, respectively). Paradoxically, bilateral injection decreased both saccade acceleration and deceleration. Saccade trajectories and end points were more variable than normal. 7. To account for the effects of our injections, we propose that the activity of caudal fastigial neurons on one side normally helps to decelerate ipsilateral saccades and helps to accelerate contralateral saccades by influencing the feedback loop of the saccade burst generator in the brain stem. Without caudal fastigial activity the brain stem burst generator produces hypermetric, variable saccades. We therefore also propose that the influence of caudal fastigial neurons on the burst generator makes saccades more consistent and accurate.(ABSTRACT TRUNCATED AT 400 WORDS)

Normal and pathological saccadic dysmetria.

Initial saccades to visual targets are generally not precise in either normal subjects or patients with saccadic dysmetria. Quantitative criteria have to be applied to clearly distinguish between normal and pathological saccadic dysmetria, which is often found in patients with cerebellar lesions. To establish these criteria, the accuracy of visually guided horizontal saccades (10 degrees and 20 degrees target steps) was studied in a group of 24 patients with lesions affecting the cerebellum or its connections, and compared with data from 17 normal subjects. Eye movements were recorded with infrared oculography. Saccades of normal subjects had an average gain of 0.92-0.95 depending on the stimulus condition. Centripetal saccades were significantly larger than centrifugal saccades, for 20 degrees target steps. Most patients (n = 15) had significantly larger saccadic amplitudes than normal subjects (hypermetria), at least in one direction. Saccades in the opposite direction could be either hypometric, hypermetric or normal. Two patients had hypometric saccades in both directions. For one of the patients with hypermetria, in addition, the amplitude difference between centrifugal and centripetal saccades was significantly larger than in the normal subjects. Five patients had no significant pathology of the initial (primary) saccade, but a pathological pattern of corrective saccades. Two patients had normal saccades under all conditions. The quantitative comparison with normal subjects allows the detection even of mild pathology. According to our results, a pathology can be assumed when the average gain of saccades in at least one direction is 1.0 or more, or when more than two out of 20 saccades are followed by two corrective saccades of which the last is in the direction opposite to the initial saccade (pathological pattern of corrective saccades). Target steps of 20 degrees reveal a pathological condition more often than 10 degrees target steps. The application of quantitative criteria might be useful to establish a diagnosis of pathologic saccadic dysmetria even in instances in which it is clinically not obvious.

Cortical potentials preceding centrifugal and centripetal self-paced horizontal saccades

Cortical potentials preceding self-paced centrifugal and centripetal saccades were recorded in 15 subjects from F3, Fz, F4, C3, Cz, C4, P3, Pz and P4 versus linked mastoid electrodes. A negative potential starting about 1.0 sec prior to saccade onset, reaching a peak amplitude of 6.8 μV on average, preceded centrifugal saccades. In contrast the negativity preceded centripetal saccades by only 500 msec, and its peak amplitude was smaller (4.6 μV). We conclude that these, differences reflect the fact that less ‘effort’ is needed with centriperal as compared to centrifugal saccades.

Kinematics of centrifugal and centripetal saccadic eye movements in man

The kinematics of centrifugal and centripetal saccadic eye movements were quantified in human subjects. The maximum velocity of centripetal saccades increased with the eccentricity of the orbital starting point and was systematically higher than that of centrifugal saccades starting from primary orbital position. The slope of this linear increase was related to target step amplitude (2.6 and 3.9 deg/sec/deg for 20 and 30 deg, respectively). Despite these velocity changes, saccade amplitude was maintained by corresponding variations of the duration of deceleration. These findings, which are relevant with respect to saccadic control theories, indicate that initial eye position must be considered before comparing saccades based on their kinematic properties.

Saccadic undershoot is not inevitable: Saccades can be accurate

Saccades normally take the eye 90% of the way to a target, followed by a 10% corrective saccade. An exception to this rule occurs with the range effect. When targets appear in a set of positions, saccades overshoot the near positions and undershoot the far. This phenomenon, previously reported, was confirmed with more accurate methods. The range effect increases if a visual discrimination task is added. It is established rapidly in only a few trials. Latencies of corrective saccades from overshoots and undershoots were the same. Centripetal saccades were more accurate than centrifugal. Thus, undershooting is not inevitable.