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D Guitton

Publications and source records attributed to D Guitton.

At least 19 recordsLinked to original sources
PubMed2001

The superior colliculus and its control of fixation behavior via projections to brainstem omnipause neurons.

Evidence, obtained in the animal whose head is unrestrained, has shown that the superior colliculus (SC) controls, not specifically eye-in-head motion, but rather saccadic shifts of the visual axis (gaze) composed of coordinated eye and head movements. The SC has also been implicated in fixation control. In a current hypothesis, activity on the SC motor map reflects two conflictual behavioral states: 'Orient!' versus 'Don't orient!' The latter behavior is thought to be commanded from a 'fixation zone' in the rostral SC that includes the foveal representation of the retinotopic map. 'Fixation neurons' (SCFNs) in this zone project to brainstem 'omnipause neurons' (OPNs) that inhibit the gaze-saccade generating circuits. It has been proposed that activity in SCFNs during active fixation, drives OPNs which in turn inhibit the gaze saccade generator, thereby maintaining the visual axis stable on a target of interest. Cats with head unrestrained frequently orient in the dark, to a briefly visible target, using multiple-step gaze shifts with a variable number of gaze saccades interspersed with periods of steady fixation. We found that SCFNs are not always active during the inter-step fixation periods when the visual axis is immobile--whereas OPNs are tonically active--thereby implying that activity in FNs and OPNs can be decoupled. In cats with head unrestrained, SCFNs encode the error between desired and actual gaze positions, not necessarily that the visual axis is immobile. By comparison, OPNs are tonically active when the visual axis is immobile, but some also encode gaze position error. Thus, the discharge of an OPN may reflect combined inputs from SCFNs combined with a steady bias.

Animals↗
PubMed2000

Cross-correlated and oscillatory visual responses of superficial-layer and tecto-reticular neurones in cat superior colliculus.

The present study examined, in the superior colliculus (SC) of anaesthetised cats, the functional connectivity between superficial-layer neurones (SLNs) and tectoreticular neurones (TRNs: collicular output cells). TRNs were antidromically identified by electrical stimulation of the predorsal bundle. The auto- and cross-correlation histograms of visual responses of both types of neurones were recorded and analysed. A delayed, sharp peak in cross-correlograms allowed us to verify whether SLN and TRN cells were coupled; in addition, oscillatory activities were compared to verify if rhythmic responses of SLN sites were transmitted to TRN sites. We found that oscillatory activity was rarely observed in spontaneous activity of superficial (1/74) and TRN sites (1/48). Moving light bars induced oscillation in 31% (23/74) of the superficial-layer and in 23% (11/48) of the TRN sites. The strength of the rhythmic responses was determined by specific ranges of stimulus velocity in 83% (19/23) and 64% (7/11) of oscillating SLN and TRN sites, respectively. Frequencies of oscillations ranged between 5 and 125 Hz and were confined, for 53% of the cells, to the 5-20 Hz band. Thus, the band-width of frequencies of the stimulus-related oscillations in the superior colliculus was broader than the gamma range. Analysis of cross-correlation histograms revealed a significant predominant peak with a mean delay of 2.7+/-0.9 ms in 46% (17/37) of SLN-TRN pairs. Most correlated SLN-TRN pairs (88%: 15/17) had superimposed receptive fields, suggesting they were functionally interconnected. However, individual oscillatory frequencies of correlated and oscillatory SLN and TRN cells were never the same (0/8). Together, these results suggest that the neurones in collicular superficial layer contact TRNs and, consequently, support the idea that the superficial layers contribute to collicular outputs producing eye- and head-orienting movements.

Animals↗
PubMed2000

Fixation neurons in the superior colliculus encode distance between current and desired gaze positions.

A visual scene is scrutinized during sequential periods of steady fixation, connected by saccades that shift the visual axis (gaze) to new positions. During such exploratory scan paths, gaze frequently strays from and then returns to salient features. How the brain keeps track of major end-goals and intermediate subgoals is not understood. We studied the discharge of fixation neurons of the brainstem's superior colliculus during multiple-step gaze shifts composed of a sequence of saccades made in the dark and separated by short periods of steady fixation. Cells were initially silent. As sequential gaze saccades approached the goal, firing began; its frequency increased progressively and peaked when gaze was on the remembered target location. We conclude that these fixation neurons encode the error between desired and actual gaze positions, irrespective of trajectory characteristics.

Action Potentials↗
PubMed1999

Three-dimensional eye-head coordination during gaze saccades in the primate.

The purpose of this investigation was to describe the neural constraints on three-dimensional (3-D) orientations of the eye in space (Es), head in space (Hs), and eye in head (Eh) during visual fixations in the monkey and the control strategies used to implement these constraints during head-free gaze saccades. Dual scleral search coil signals were used to compute 3-D orientation quaternions, two-dimensional (2-D) direction vectors, and 3-D angular velocity vectors for both the eye and head in three monkeys during the following visual tasks: radial to/from center, repetitive horizontal, nonrepetitive oblique, random (wide 2-D range), and random with pin-hole goggles. Although 2-D gaze direction (of Es) was controlled more tightly than the contributing 2-D Hs and Eh components, the torsional standard deviation of Es was greater (mean 3.55 degrees ) than Hs (3.10 degrees ), which in turn was greater than Eh (1.87 degrees ) during random fixations. Thus the 3-D Es range appeared to be the byproduct of Hs and Eh constraints, resulting in a pseudoplanar Es range that was twisted (in orthogonal coordinates) like the zero torsion range of Fick coordinates. The Hs fixation range was similarly Fick-like, whereas the Eh fixation range was quasiplanar. The latter Eh range was maintained through exquisite saccade/slow phase coordination, i.e., during each head movement, multiple anticipatory saccades drove the eye torsionally out of the planar range such that subsequent slow phases drove the eye back toward the fixation range. The Fick-like Hs constraint was maintained by the following strategies: first, during purely vertical/horizontal movements, the head rotated about constantly oriented axes that closely resembled physical Fick gimbals, i.e., about head-fixed horizontal axes and space-fixed vertical axes, respectively (although in 1 animal, the latter constraint was relaxed during repetitive horizontal movements, allowing for trajectory optimization). However, during large oblique movements, head orientation made transient but dramatic departures from the zero-torsion Fick surface, taking the shortest path between two torsionally eccentric fixation points on the surface. Moreover, in the pin-hole goggle task, the head-orientation range flattened significantly, suggesting a task-dependent default strategy similar to Listing's law. These and previous observations suggest two quasi-independent brain stem circuits: an oculomotor 2-D to 3-D transformation that coordinates anticipatory saccades with slow phases to uphold Listing's law, and a flexible "Fick operator" that selects head motor error; both nested within a dynamic gaze feedback loop.

Algorithms↗
PubMed1998

Brain stem omnipause neurons and the control of combined eye-head gaze saccades in the alert cat.

When the head is unrestrained, rapid displacements of the visual axis-gaze shifts (eye-re-space)-are made by coordinated movements of the eyes (eye-re-head) and head (head-re-space). To address the problem of the neural control of gaze shifts, we studied and contrasted the discharges of omnipause neurons (OPNs) during a variety of combined eye-head gaze shifts and head-fixed eye saccades executed by alert cats. OPNs discharged tonically during intersaccadic intervals and at a reduced level during slow perisaccadic gaze movements sometimes accompanying saccades. Their activity ceased for the duration of the saccadic gaze shifts the animal executed, either by head-fixed eye saccades alone or by combined eye-head movements. This was true for all types of gaze shifts studied: active movements to visual targets; passive movements induced by whole-body rotation or by head rotation about stationary body; and electrically evoked movements by stimulation of the caudal part of the superior colliculus (SC), a central structure for gaze control. For combined eye-head gaze shifts, the OPN pause was therefore not correlated to the eye-in-head trajectory. For instance, in active gaze movements, the end of the pause was better correlated with the gaze end than with either the eye saccade end or the time of eye counterrotation. The hypothesis that cat OPNs participate in controlling gaze shifts is supported by these results, and also by the observation that the movements of both the eyes and the head were transiently interrupted by stimulation of OPNs during gaze shifts. However, we found that the OPN pause could be dissociated from the gaze-motor-error signal producing the gaze shift. First, OPNs resumed discharging when perturbation of head motion briefly interrupted a gaze shift before its intended amplitude was attained. Second, stimulation of caudal SC sites in head-free cat elicited large head-free gaze shifts consistent with the creation of a large gaze-motor-error signal. However, stimulation of the same sites in head-fixed cat produced small "goal-directed" eye saccades, and OPNs paused only for the duration of the latter; neither a pause nor an eye movement occurred when the same stimulation was applied with the eyes at the goal location. We conclude that OPNs can be controlled by neither a simple eye control system nor an absolute gaze control system. Our data cannot be accounted for by existing models describing the control of combined eye-head gaze shifts and therefore put new constraints on future models, which will have to incorporate all the various signals that act synergistically to control gaze shifts.

Animals↗
PubMed1998

Human head-free gaze saccades to targets flashed before gaze-pursuit are spatially accurate.

Previous studies have shown that accurate saccades can be generated, in the dark, that compensate for movements of the visual axis that result from movements of either the eyes alone or the head alone that intervene between target presentation and saccade onset. We have carried out experiments with human subjects to test whether gaze saccades (gaze = eye-in-space = eye-in-head + head-in-space) can be generated that compensate for smooth pursuit movements of gaze that intervene between target onset and gaze-saccade onset. In both head-unrestrained (head-free) and -restrained (head-fixed) conditions, subjects were asked to make gaze shifts, in the dark, to the remembered location of a briefly flashed target. On most trials, during the memory period, the subjects carried out intervening head-free gaze pursuit or head-fixed ocular pursuit along the horizontal meridian. On the remaining (control) trials, subjects did not carry out intervening pursuit movements during the memory period; this was the classical memory-guided saccade task. We found that the subjects accurately compensated for intervening movements of the visual axis in both the head-free and head-fixed conditions. We conclude that the human gaze-motor system is able to monitor on-line changes in gaze position and add them to initial retinal error, to program spatially accurate gaze saccades.

Adult↗
PubMed1997

Visual-motor transformations required for accurate and kinematically correct saccades.

The goal of this study was to identify and model the three-dimensional (3-D) geometric transformations required for accurate saccades to distant visual targets from arbitrary initial eye positions. In abstract 2-D models, target displacement in space, retinal error (RE), and saccade vectors are trivially interchangeable. However, in real 3-D space, RE is a nontrivial function of objective target displacement and 3-D eye position. To determine the physiological implications of this, a visuomotor "lookup table" was modeled by mapping the horizontal/vertical components of RE onto the corresponding vector components of eye displacement in Listing's plane. This provided the motor error (ME) command for a 3-D displacement-feedback loop. The output of this loop controlled an oculomotor plant that mechanically implemented the position-dependent saccade axis tilts required for Listing's law. This model correctly maintained Listing's law but was unable to correct torsional position deviations from Listing' s plane. Moreover, the model also generated systematic errors in saccade direction (as a function of eye position components orthogonal to RE), predicting errors in final gaze direction of up to 25 degrees in the oculomotor range. Plant modifications could not solve these problems, because the intrisic oculomotor input-output geometry forced a fixed visuomotor mapping to choose between either accuracy or Listing's law. This was reflected internally by a sensorimotor divergence between input-defined visual displacement signals (inherently 2-D and defined in reference to the eye) and output-defined motor displacement signals (inherently 3-D and defined in reference to the head). These problems were solved by rotating RE by estimated 3-D eye position (i.e., a reference frame transformation), inputting the result into a 2-D-to-3-D "Listing's law operator," and then finally subtracting initial 3-D eye position to yield the correct ME. This model was accurate and upheld Listing's law from all initial positions. Moreover, it suggested specific experiments to invasively distinguish visual and motor displacement codes, predicting a systematic position dependence in the directional tuning of RE versus a fixed-vector tuning in ME. We conclude that visual and motor displacement spaces are geometrically distinct such that a fixed visual-motor mapping will produce systematic and measurable behavioral errors. To avoid these errors, the brain would need to implement both a 3-D position-dependent reference frame transformation and nontrivial 2-D-to-3-D transformation. Furthermore, our simulations provide new experimental paradigms to invasively identify the physiological progression of these spatial transformations by reexamining the position-dependent geometry of displacement code directions in the superior colliculus, cerebellum, and various cortical visuomotor areas.

Animals↗
PubMed1997

Primate head-free saccade generator implements a desired (post-VOR) eye position command by anticipating intended head motion.

Primate head-free saccade generator implements a desired (post-VOR) eye position command by anticipating intended head motion. J. Neurophysiol. 78: 2811-2816, 1997. When we glance between objects, the brain ultimately controls gaze direction in space. However, it is currently unclear how this is allocated into separate commands for eye and head movement. To determine the role of desired final eye position commands, and their coordination with intended head movement, we trained three monkeys to make large gaze shifts while wearing opaque goggles with a monocular 8 degrees aperture. Animals eventually developed a new set of context-dependent eye-head coordination strategies, in particular expanding the head range and compressing the eye-in-head range toward the aperture (while wearing the goggles). However, when we shifted the location of the aperture to a different subsection of the normal head-free oculomotor range (by covering the original aperture and creating a new one), eye-head saccades failed to acquire visual targets, because they continued to drive the eye ultimately toward the now occluded original aperture. Even when a head-stationary saccade acquired the new aperture, subsequent head-free saccades drove the eye eccentrically toward a point that anticipated the intended head movement, such that the subsequent vestibuloocular reflex slow phase brought the eye onto the location of the original aperture. Animals could only acquire the new aperture consistently after several days of retraining. These results suggest that 1) eye-head coordination is achieved by a plastic, context-dependent neural operator that uses information about initial eye/head position and intended movement to compute desired combinations of final eye/head position and 2) acquisition of these positions involves sophisticated anticipatory compensations for subsequent movement components, akin to those observed previously in complex oral and manual behaviors.

Animals↗
PubMed1997

Analysis of primate IBN spike trains using system identification techniques. I. Relationship To eye movement dynamics during head-fixed saccades.

The dynamic behavior of primate (Macaca fascicularis) inhibitory burst neurons (IBNs) during head-fixed saccades was analyzed by using system identification techniques. Neurons were categorized as IBNs on the basis of their anatomic location as well as by their activity during horizontal head-fixed saccadic and smooth pursuit eye movements and vestibular nystagmus. Each IBN's latency or "dynamic lead time" (td) was determined by shifting the unit discharge in time until an optimal fit to the firing rate frequency B(t) profile was obtained by using the simple model based on eye movement dynamics,B(t) = r + b1(t); where is eye velocity. For the population of IBNs, the dynamic estimate of lead time provided a significantly lower value than a method that used the onset of the first spike. We then compared the relative abilities of different eye movement-based models to predict B(t) by using objective optimization algorithms. The most important terms for predicting B(t) were eye velocity gain (b1) and bias terms (r) mentioned above. The contributions of higher-order velocity, acceleration, and/or eye position terms to model fits were found to be negligible. The addition of a pole term [the time derivative of B(t)] in conjunction with an acceleration term significantly improved model fits to IBN spike trains, particularly when the firing rates at the beginning of each saccade [initial conditions (ICs)] were estimated as parameters. Such a model fit the data well (a fit comparable to a linear regression analysis with a R2 value of 0.5, or equivalently, a correlation coefficient of 0.74). A simplified version of this model [B(t) = rk + b1(t)], which did not contain a pole term, but in which the bias term (rk) was estimated separately for each saccade, provided nearly equivalent fits of the data. However, models in which ICs or rks were estimated separately for each saccade contained too many parameters to be considered as useful models of IBN discharges. We discovered that estimated ICs and rks were correlated with saccade amplitude for the majority of short-lead IBNs (SLIBNs; 56%) and many long-lead IBNs (LLIBNs; 42%). This observation led us to construct a more simple model that included a term that was inversely related to the amplitude of the saccade, in addition to eye velocity and constant bias terms. Such a model better described neuron discharges than more complex models based on a third-order nonlinear function of eye velocity. Given the small number of parameters required by this model (only 3) and its ability to fit the data, we suggest that it provides the most valuable description of IBN discharges. This model emphasizes that the IBN discharges are dependent on saccade amplitude and implies further that a mechanism must exist, at the motoneuron (MN) level, to offset the effect of the bias and amplitude-dependent terms. In addition, we did not find a significant difference in the variance accounted for by any of the downstream models tested for SLIBNs versus LLIBNs. Therefore we conclude that the eye movement signals encoded dynamically by SLIBNs and LLIBNs are similar in nature. Put another way, SLIBNs are not closer, dynamically, to MNs than LLIBNs.

Action Potentials↗
PubMed1997

Analysis of primate IBN spike trains using system identification techniques. II. Relationship to gaze, eye, and head movement dynamics during head-free gaze shifts.

We have investigated the relationships among the firing frequency B(t) of inhibitory burst neurons (IBNs) and the metrics and dynamics of the eye, head, and gaze (eye + head) movements generated during voluntary combined eye-head gaze shifts in monkey. The same IBNs were characterized during head-fixed saccades in our first of three companion papers. In head-free gaze shifts, the number of spikes (NOS) in a burst was, for 82% of the neurons, better correlated with gaze amplitude than with the amplitude of either the eye or head components of the gaze shift. A multiple regression analysis confirmed that NOS was well correlated to the sum of head and eye amplitudes during head-free gaze shifts. Furthermore, the mean slope of the relationship between NOS and gaze amplitude was significantly less for head-free gaze shifts than for head-fixed saccades. NOS is a global parameter. To refine we used system identification techniques to evaluate a series of dynamic models in which IBN spike trains were related to gaze or eye movements. We found that gaze- and eye-based models predicted the discharges of IBNs equally well. However, the bias values required by gaze-based models were comparable to those required in our head-fixed models whereas those required by eye-based models were significantly larger. The difference in biases between gaze- and eye-based models was very strongly correlated to the mean head velocity () during gaze shifts [R = -0.93 +/- 0.15 (SD)]. This result suggested that the increased bias required by the eye-based models reflected an unmodeled input onto these cells. To pursue this argument further we investigated a series of dynamic models that included both eye velocity () and terms and this confirmed the importance of these two terms. As in our head-fixed analysis of companion paper I, the most valuable model formulation also included an eye saccade amplitude term (DeltaE) and was given by B(t) = r0 + r1DeltaE + b1 + g1 where r0, r1, b1, and g1 are constants. The amplitude of the head velocity coefficient was significantly less than that of the eye velocity coefficient. Furthermore, in our population long-lead IBNs tended to have a smaller head velocity coefficients than short-lead IBNs. We conclude that during head-free gaze shifts, the head velocity signal carried to the abducens nucleus by primate excitatory burst neurons (EBNs; if EBNs and IBNs carry similar signals) must be offset by other premotor cells.

Action Potentials↗
PubMed1997

Analysis of primate IBN spike trains using system identification techniques. III. Relationship To motor error during head-fixed saccades and head-free gaze shifts.

The classic model of saccade generation assumes that the burst generator is driven by a motor-error signal, the difference between the actual eye position and the final "desired" eye position in the orbit. Here we evaluate objectively, using system identification techniques, the dynamic relationship between motor-error signals and primate inhibitory burst neuron (IBN) discharges (upstream analysis). The IBNs presented here are the same neurons whose downstream relationships were characterized during head-fixed saccades and head-free gaze shifts in our companion papers. In our analysis of head-fixed saccades we determined how well IBN discharges encode eye motor error (epsilone) compared with downstream saccadic eye movement dynamics and whether long-lead IBN (LLIBN) discharges encode epsilone better than short-lead IBNs (SLIBNs), given that it is commonly assumed that short-lead burst neurons (BNs) are closer than long-lead BNs to the motor output and thus further from the epsilone signal. In the epsilone-based models tested, IBN firing frequency B(t) was represented by one of the following: 1) model 1u, a nonlinear function of epsilone; 2) model 2u, a linear function of epsilone [B(t) = rk + a1epsilone(t)] where the bias term rk was estimated separately for each saccade; 3) model 3u, a version of model 2u wherein the bias term was a function of saccade amplitude; or 4) model 4u, a linear function of epsilone with an added pole term (the derivative of firing rate). Models based on epsilone consistently produced worse predictions of IBN activity than models of comparable complexity based on eye movement dynamics (e.g., eye velocity). Hence, the simple two parameter downstream model 2d [B(t) = r + b1(t)] was much better than any upstream (epsilone-based) model with a comparable number of parameters. The link between B(t) and epsilone is due primarily to the correlation between the declining phases of B(t) and epsilone; motor-error models did not predict well the rising phase of the discharge. We could improve substantially the performance of upstream models by adding an e term. Because e = -, this process was equivalent to incorporating terms into upstream models thereby erasing the distinction between upstream and downstream analyses. Adding an e term to the upstream models made them as good as downstream ones in predicting B(t). However, the epsilone term now became redundant because its removal did not affect model accuracy. Thus, when is available as a parameter, epsilone becomes irrelevant. In the head-free monkey the ability of upstream models to predict IBN firing during head-free gaze shifts when gaze, eye, or head motor-error signals were model inputs was poor and similar to the upstream analysis of the head-fixed condition. We conclude that during saccades (head-fixed) or gaze shifts (head-free) the activity of both SLIBNs and LLIBNs is more closely linked to downstream events (i.e., the dynamics of ongoing movements) than to the coincident upstream motor-error signal. Furthermore, SLIBNs and LLIBNs do not differ in their characteristics; the latter are not, as is usually hypothesized, closer to a motor-error signal than the former.

Animals↗
PubMed1996

The use of system identification techniques in the analysis of oculomotor burst neuron spike train dynamics.

The objective of system identification methods is to construct a mathematical model of a dynamical system in order to describe adequately the input-output relationship observed in that system. Over the past several decades, mathematical models have been employed frequently in the oculomotor field, and their use has contributed greatly to our understanding of how information flows through the implicated brain regions. However, the existing analyses of oculomotor neural discharges have not taken advantage of the power of optimization algorithms that have been developed for system identification purposes. In this article, we employ these techniques to specifically investigate the "burst generator" in the brainstem that drives saccadic eye movements. The discharge characteristics of a specific class of neurons, inhibitory burst neurons (IBNs) that project monosynaptically to ocular motoneurons, are examined. The discharges of IBNs are analyzed using different linear and nonlinear equations that express a neuron's firing frequency and history (i.e., the derivative of frequency), in terms of quantities that describe a saccade trajectory, such as eye position, velocity, and acceleration. The variance accounted for by each equation can be compared to choose the optimal model. The methods we present allow optimization across multiple saccade trajectories simultaneously. We are able to investigate objectively how well a specific equation predicts a neuron's discharge pattern as well as whether increasing the complexity of a model is justifiable. In addition, we demonstrate that these techniques can be used both to provide an objective estimate of a neuron's dynamic latency and to test whether a neuron's initial firing rate (expressed as an initial condition) is a function of a quantity describing a saccade trajectory (such as initial eye position).

Action Potentials↗
PubMed1995

On-line compensation of gaze shifts perturbed by micro-stimulation of the superior colliculus in the cat with unrestrained head.

Prior studies have led to the gaze feedback hypothesis, which states that quick orienting movements of the visual axis (gaze shifts) are controlled by a feedback system. We have previously provided evidence for this hypothesis by extending the original study of Mays and Sparks (1980) to the cat with unrestrained head (Pélisson et al. 1989). We showed that cats compensated for a stimulation-induced perturbation of initial gaze position by generating, in the dark, an accurate gaze shift towards the remembered location of a flashed target. In the present study, we investigate goal-directed gaze shifts perturbed "in flight" by a brief stimulation of the superior colliculus. The microstimulation parameters were tuned such that significant perturbations were induced without halting the movement. The ambient light was turned off at the onset of the gaze shift, suppressing any visual feedback. We observed that, following stimulation offset, the gaze shift showed temporal and spatial changes in its trajectory to compensate for the transient perturbation. Such compensations, which occurred "on-line" before gaze shift termination, involved both eye and head movements and had dynamic characteristics resembling those of unperturbed saccadic gaze shifts. These on-line compensations maintained gaze accuracy when the stimulation was applied during the early phase of large and medium (about 60 and 40 degrees) movements. These results are compatible with the notion of a gaze feedback loop providing a dynamic gaze error signal.

Animals↗
PubMed1994

The fixation area of the cat superior colliculus: effects of electrical stimulation and direct connection with brainstem omnipause neurons.

The superior colliculus has long been recognized as an important structure in the generation of saccadic displacements of the visual axis. Neurons with presaccadic activity encoding saccade vectors are topographically organized and form a "motor map." Recently, neurons with fixation-related activity have been recorded at the collicular rostral pole, at the area centralis representation or fixation area. Another collicular function which deals with the maintenance of fixation behavior by means of active inhibition of orientation commands was then suggested. We tested that hypothesis as it relates to the suppression of gaze saccades (gaze = eye in space = eye in head + head in space) in the head-free cat by increasing the activity of the fixation cells at the rostral pole with electrical microstimulation. Long stimulation trains applied before gaze saccades delayed their initiation. Short stimuli, delivered during the gaze saccades, transiently interrupted both eye and head components. These results provide further support for a role in fixation behavior for collicular fixation neurons. Brainstem omnipause neurons also exhibit fixation-related activity and have been shown to receive a direct excitatory input from the superior colliculus. To determine whether the collicular projection to omnipause neurons arises from the fixation area, the deep layers of the superior colliculus were electrically stimulated either at the rostral pole including the fixation area or in more caudal regions where stimulation evokes orienting responses. Forty-nine neurons were examined in three cats. 61% of the neurons were found to be orthodromically excited by single-pulse stimulation of the rostral pole, whereas only 29% responded to caudal stimulation. In addition, stimuli delivered to the rostral pole activated, on average, omnipause neurons at shorter latencies and with lower currents than those applied in caudal regions. These results suggest that excitatory inputs to omnipause neurons from the superior colliculus are principally provided by the fixation area, via which the superior colliculus could play a role in suppression of gaze shifts.

Animals↗
PubMed1994

Gaze shifts evoked by stimulation of the superior colliculus in the head-free cat conform to the motor map but also depend on stimulus strength and fixation activity.

In our previous paper we demonstrated that electrical microstimulation of the fixation area at the rostral pole of the cat superior colliculus (SC) elicits no gaze movement but, rather, transiently suppresses eye-head gaze saccades. In this paper, we investigated the more caudal region of the SC and its interaction with the fixation area. In the alert head-free cat, suprathreshold stimulation in the anterior portion of the SC but outside the fixation area evoked small saccadic shifts of gaze consisting mainly of an eye movement, the head's contribution being small. Stimulating more posteriorly elicited large gaze saccades consisting of an ocular saccade combined with a rapid head movement. At these latter stimulation sites, craniocentric (goal-directed) eye movements were evoked when the cat's head was restrained. The amplitude of eye-head gaze saccades elicited at a particular stimulation site increased with stimulus duration, current strength, and pulse rate, until a constant or "unit" value was reached. The peak velocity of gaze shifts depended on both pulse rate and current strength. The movement direction was not affected by stimulus parameters. The unit gaze vector evoked, in the head-free condition, by stimulating one collicular site was similar to that coded by efferent neurons recorded at that site, thereby indicating a retinotopically coded gaze error representation on the collicular motor map which is not revealed by stimulating the head-fixed animal. Evoked gaze saccades were found to be influenced by fixation behavior. The amplitude of evoked gaze shifts was reduced if stimulation occurred when the hungry animal fixated a food target. Electrical activation of the collicular fixation area was found to mimic well the effects of natural fixation on evoked gaze shifts. Taken together, our results support the view that the overall distribution and level of collicular activity contributes to the encoding of the metrics of gaze saccades. We suggest that the combined levels of activity at the site being stimulated and at the fixation area influence the amplitude of evoked gaze saccades through competition. When stimulation is at low intensities, fixation-related activity reduces the amplitude of evoked gaze saccades. At high activation levels, the site being stimulated dominates and the gaze vector is specified only by that site's collicular output neurons, from which arises the close correspondence between the unit-evoked gaze saccades and the neurally coded gaze vector at that site.

Animals↗
PubMed1993

Human gaze shifts in which head and eyes are not initially aligned.

Most studies of rapid orienting gaze shifts generated by combined eye and head movements have focused on an experimental condition in which gaze displacements are started with the subject's eyes in the normal straight-ahead position in the orbit. Such an experimental approach does not permit a clear identification of the input signal to the head motor system, because target offset angle is the same for both the eye and head. We have studied gaze shifts in human subjects which began with the visual axis straight ahead relative to the body (i.e., gaze or line of sight aligned with body sagittal plane) and with head offset from straight ahead at various angular positions. In our experimental conditions, the amplitude of head movement during a gaze shift was nearly equal to the angular distance between the target position and the starting head position (target-re-head), even though subjects were not specifically instructed to move their heads. This observation contrasts with other published reports in the literature showing considerable variability amongst subjects in the amplitude of head rotation within a given task and between tasks. The difference may be related to the initial conditions which required subjects to align the eye and head on specific starting targets, since others have shown that requiring head alignment enhances head displacement. The amplitude of the saccadic eye movement was not determined by either the target's position relative to the starting eye or head positions. The value that best described the eye movement amplitude was the eye position in the orbit at the end of the saccade.(ABSTRACT TRUNCATED AT 250 WORDS)

Adult↗