Combined eye-head gaze shifts produced by electrical stimulation of the superior colliculus in rhesus monkeys

1. We electrically stimulated the intermediate and deep layers of the superior colliculus (SC) in two rhesus macaques free to move their heads both vertically and horizontally (head unrestrained). Stimulation of the primate SC can elicit high-velocity, combined, eye-head gaze shifts that are similar to visually guided gaze shifts of comparable amplitude and direction. The amplitude of gaze shifts produced by collicular stimulation depends on the site of stimulation and on the parameters of stimulation (frequency, current, and duration of the stimulation train). 2. The maximal amplitude gaze shifts, produced by electrical stimulation at 56 sites in the SC of two rhesus monkeys, ranged in amplitude from approximately 7 to approximately 80 deg. Because the head was unrestrained, stimulation-induced gaze shifts often included movements of the head. Head movements produced at the 56 stimulation sites ranged in amplitude from 0 to approximately 70 deg. 3. The relationships between peak velocity and amplitude and between duration and amplitude of stimulation-induced head movements and gaze shifts were comparable with the relationships observed during visually guided gaze shifts. The relative contributions of the eyes and head to visually guided and stimulation-induced gaze shifts were also similar. 4. As was true for visually guided gaze shifts, the head contribution to stimulation-induced gaze shifts depended on the position of the eyes relative to the head at the onset of stimulation. When the eyes were deviated in the direction of the ensuing gaze shift, the head contribution increased and the latency to head movement onset was decreased. 5. We systematically altered the duration of stimulation trains (10-400 ms) while stimulation frequency and current remained constant. Increases in stimulation duration systematically increased the amplitude of the evoked gaze shift until a site specific maximal amplitude was reached. Further increases in stimulation duration did not increase gaze amplitude. There was a high correlation between the end of the stimulation train and the end of the evoked gaze shift for movements smaller than the site-specific maximal amplitude. 6. Unlike the effects of stimulation duration on gaze amplitude, the amplitude and duration of evoked head movements did not saturate for the range of durations tested (10-400 ms), but continued to increase linearly with increases in stimulation duration. 7. The frequency of stimulation was systematically varied (range: 63-1,000 Hz) while other stimulation parameters remained constant. The velocity of evoked gaze shifts was related to the frequency of stimulation; higher stimulation frequencies resulted in higher peak velocities. The maximal, site-specific amplitude was independent of stimulation frequency. 8. When stimulating a single collicular site using identical stimulation parameters, the amplitude and direction of stimulation-induced gaze shifts, initiated from different initial positions, were relatively constant. In contrast, the amplitude and direction of the eye component of these fixed vector gaze shifts depended upon the initial position of the eyes in the orbits; the endpoints of the eye movements converged on an orbital region, or "goal," that depended on the site of collicular stimulation. 9. When identical stimulation parameters were used and when the eyes were centered initially in the orbits, the gaze shifts produced by caudal collicular stimulation when the head was restrained were typically smaller than those evoked from the same site when the head was unrestrained. This attenuation occurred because stimulation drove the eyes to approximately the same orbital position when the head was restrained or unrestrained. Thus movements produced when the head was restrained were reduced in amplitude by approximately the amount that the head would have contributed if free to move. 10. When the head was restrained, only the eye component of the intended gaze shift     

Morphological studies on Mn-SOD, NOS and calcium binding proteins in the rat hippocampus

Head movement propensity-the pattern of head saccades dependent on methods of target presentation-varies among individuals. The present group of 9 young adults was previously ranked in a visual saccadic task according to this propensity. The present report examines how and why this propensity changes if the saccades are made to auditory targets. 1) Spatially identical, interleaved, auditorily and visually elicited horizontal saccadic gaze shifts (jumps) differed in amplitude and in starting and/or ending position. The jumps were executed in two head movement modes: first, the non-aligned mode was a standard reaction-time single gaze step between two points. Second, the head-aligned mode required alignment of the head with the fixation (starting) point; thereafter both modes were identical. All results in the auditory task are expressed relative to the visual results. 2) In the non-aligned mode, head movement amplitudes were increased on average by 15% (for example, an 80 degrees jump elicited a 12 degrees larger head movement), and velocity decreased by 12%, reflecting the increased demands of the auditory task. More importantly, the differences between subjects was narrowed; that is, head movement propensity was homogenized in the auditory task. In the visual task, head-movers willingly move their heads off and across the midline, whereas non-movers are unwilling to leave the midline from eccentric starting points or to eccentric ending points. This is called the midline attraction effect and was previously linked to spatial reference frames. The homogenization in the auditory task was characterized by head-movers increasing, and non-movers decreasing, their midline attraction, suggesting altered spatial reference frames. 3) For heuristic purposes, the ideal head-mover is defined by a gain of 1.0 in the visual task, and by external earth-fixed reference frames. Similarly, the ideal non-mover has a gain of 0.0 and has a bias toward body (or some par of the body)-fixed reference frames. In the auditory task these gains (and reference frames) in head movers and non-movers are homogenized (close to 0.5), either by the participation of the head (movement of the ears in space) in sensory acquisition or by differences in central nervous processing of the two modalities, or both.     

Gain adaptation of eye and head movement components of simian gaze shifts

Gain adaptation of eye and head movement components of simian gaze shifts. J. Neurophysiol. 78: 2817-2821, 1997. To investigate the site of gaze adaptation in primates, we reduced the gain of large head-restrained gaze shifts made to 50 degrees target steps by jumping the target 40% backwards during a targeting saccade and then tested gain transfer to the eye- and head-movement components of head-unrestrained gaze shifts. After several hundred backstep trials, saccadic gain decreased by at least 10% in 8 of 13 experiments, which were then selected for further study. The minimum saccadic gain decrease in these eight experiments was 12.8% (mean = 18.4%). Head-unrestrained gaze shifts to ordinary 50 degrees target steps experienced a gain reduction of at least 9.3% (mean = 14.9%), a mean gain transfer of 81%. Both the eye and head components of the gaze shift also decreased. However, average head movement gain decreased much more (22.1%) than eye movement gain (9.2%). Also, peak head velocity generally decreased significantly (20%), but peak eye velocity either increased or remained constant (average increase of 5.6%). However, the adapted peak eye and head velocities were appropriate for the adapted, smaller gaze amplitudes. Similar dissociations in eye and head metrics occurred when head-unrestrained gaze shifts were adapted directly (n = 2). These results indicated that head-restrained saccadic gain adaptation did not produce adaptation of eye movement alone. Nor did it produce a proportional gain change in both eye and head movement. Rather, normal eye and head amplitude and velocity relations for a given gaze amplitude were preserved. Such a result could be explained most easily if head-restrained adaptation were realized before the eye and head commands had been individualized. Therefore, gaze adaptation is most likely to occur upstream of the creation of separate eye and head movement commands.     

A modification of eye-head coordination by CNS disease

When a subject, seated and facing ahead, was asked to look toward one side, the result was a combined movement of the eyes and head. Normal subjects began the eye movement just before the onset of head movement; 4 neurologic patients who showed abnormalities in eye movements (saccades that tended to be smaller in amplitude and lower in velocity than those of the control subjects) regularly began eye movement after the onset of head movement. Thus the initiation of the head rotation was not as much retarded in these patients as that of eye movement. Amplitudes of the movements were reduced in the patients, but this change too was less for the head than for the eyes. Because the amplitude and velocity of the head movement were less affected in the patient group, the relative contribution of the head to the total gaze shift was increased. It appears as if, when the oculomotor system is affected, the head can assume a leading role in the initiation and execution of gaze shifts.     

Gaze-related activity of putative inhibitory burst neurons in the head-free cat

1. Previous studies in the cat have demonstrated that output neurons of the superior collicular as well as brain stem omnipause neurons have discharges that are best correlated, not with the trajectory of the eye in the head but, with the trajectory of the visual axis in space (gaze = eye-in-head + head-in-space) during rapid orienting coordinated eye and head movements. In this study, we describe the gaze-related activity of cat premotor "inhibitory burst neurons" (IBNs) identified on the basis of their position relative to the abducens nucleus. 2. The firing behavior of IBNs was studied during 1) saccades made with the head stationary, 2) active orienting combined eye-head gaze shifts, and 3) passive movements of the head on the body. IBN discharges were well correlated with the duration and amplitude of saccades made when the head was stationary. In both head-free paradigms, the behavior of cat IBNs differed from that of previously described primate "saccade bursters". The duration of their burst was better correlated with gaze than saccade duration, and the total number of spikes in a burst was well correlated with gaze amplitude and generally poorly correlated with saccade amplitude. The behavior of cat IBNs also differed from that of previously described primate "gaze bursters". The slope of the relationship between the total number of spikes and gaze amplitude observed during head-free gaze shifts was significantly lower than that observed during head-fixed saccades. 3. These studies suggest that cat IBNs do not fit into the categories of gaze-bursters or saccade-bursters that have been described in primate studies.(ABSTRACT TRUNCATED AT 250 WORDS)     

Contribution of the rostral fastigial nucleus to the control of orienting gaze shifts in the head-unrestrained cat

The implication of the caudal part of the fastigial nucleus (cFN) in the control of saccadic shifts of the visual axis is now well established. In contrast a possible involvement of the rostral part of the fastigial nuceus (rFN) remains unknown. In the current study we investigated in the head-unrestrained cat the contribution of the rFN to the control of visually triggered saccadic gaze shifts by measuring the deficits after unilateral muscimol injection in the rFN. A typical gaze dysmetria was observed: gaze saccades directed toward the inactivated side were hypermetric, whereas those with an opposite direction were hypometric. For both movement directions, gaze dysmetria was proportional to target retinal eccentricity and could be described as a modified gain in the translation of visual signals into eye and head motor commands. Correction saccades were triggered when the target remained visible and reduced the gaze fixation error to 2.7 +/- 1.3 degrees (mean +/- SD) on average. The hypermetria of ipsiversive gaze shifts resulted predominantly from a hypermetric response of the eyes, whereas the hypometria of contraversive gaze shifts resulted from hypometric responses of both eye and head. However, even in this latter case, the eye saccade was more affected than the motion of the head. As a consequence, for both directions of gaze shift the relative contributions of the eye and head to the overall gaze displacement were altered by muscimol injection. This was revealed by a decreased contribution of the head for ipsiversive gaze shifts and an increased head contribution for contraversive movements. These modifications were associated with slight changes in the delay between eye and head movement onsets. Inactivation of the rFN also affected the initiation of eye and head movements. Indeed, the latency of ipsiversive gaze and head movements decreased to 88 and 92% of normal, respectively, whereas the latency of contraversive ones increased to 149 and 145%. The deficits induced by rFN inactivation were then compared with those obtained after muscimol injection in the cFN of the same animals. Several deficits differed according to the site of injection within the fastigial nucleus (tonic orbital eye rotation, hypermetria of ipsiversive gaze shifts and fixation offset, relationship between dysmetria and latency of contraversive gaze shifts, postural deficit). In conclusion, the present study demonstrates that the rFN is involved in the initiation and the control of combined eye-head gaze shifts. In addition our findings support a functional distinction between the rFN and cFN for the control of orienting gaze shifts. This distinction is discussed with respect to the segregated fastigiofugal projections arising from the rFN and cFN.     

Microbial dehalogenases: enzymes recruited to convert xenobiotic substrates

Ocular torsion was measured in five subjects during sinusoidal lateral tilt (amplitude 25 degrees, 0.2 Hz). The cervical contribution to ocular torsion was best visible as the difference between the signals obtained in conditions with only head tilt and conditions with whole body tilt. Contribution of the neck did not affect the slow component, but produced an anticompensatory modulation of the beating field offset by means of saccades (analogous to gaze shift). Static tilt conditions (25 degrees tilt) of the trunk only, the head only or the whole body showed similar data, although of smaller amplitude. The results from patients suffering from post-whiplash syndrome were similar to those of healthy subjects, showing large intersubject variability. The reduced tolerance to head tilt of whiplash patients restricts useful implementation of this sort of test in the clinic.     

Influence of joint interactional effects on the coordination of planar two-joint arm movements

We have examined EMG-movement relations in two-joint planar arm movements to determine the influence of interactional torques on movement coordination. Explicitly defined combinations of elbow movements (ranging from 20 to 70 degrees) and wrist movements (ranging from 20 to 40 degrees) were performed during a visual, step-tracking task in which subjects were specifically required to attend to the initial and final angles at each joint. In all conditions the wrist and elbow rotated in the same direction, that is, flexion-flexion or extension-extension. Elbow movement kinematics were only slightly influenced by motion about the wrist. In contrast, the trajectory of the wrist movement was significantly influenced by uncompensated reaction torques resulting from movement about the elbow joint. At any given wrist amplitude, wrist movement duration increased and peak velocity decreased as elbow amplitude increased. In addition, as elbow amplitude increased, wrist movement onset was progressively delayed relative to this elbow movement. Surprisingly, the changes between joint movement onsets were not accompanied by corresponding changes between agonist EMG onsets at the elbow and wrist joints. The mean difference in onset times between elbow and wrist agonists (22-30 ms) remained unchanged across conditions. In addition, a basic pattern of muscle activation that scaled with movement amplitude was observed at each joint. Phasic agonist activity at the wrist and elbow joints remained remarkably similar across conditions and thus the changes in joint movement onset could not be attributed to changes in the motor commands.(ABSTRACT TRUNCATED AT 250 WORDS)     

Learning and limits of use of eye gaze by capuchin monkeys (Cebus apella) in an object-choice task.

The ability of 3 capuchin monkeys (Cebus apella) to use experimenter-given cues to solve an object-choice task was assessed. The monkeys learned to use explicit gestural and postural cues and then progressed to using eye-gaze-only cues to solve the task, that is, to choose the baited 1 of 2 objects and thus obtain a food reward. Increasing cue-stimulus distance and introducing movement of the eyes impeded the establishment of effective eye-gaze reading. One monkey showed positive but imperfect transfer of use of eye gaze when a novel experimenter presented the cue. When head and eye orientation cues were presented simultaneously and in conflict, the monkeys showed greater responsiveness to head orientation cues. The results show that capuchin monkeys can learn to use eye gaze as a discriminative cue, but there was no evidence for any underlying awareness of eye gaze as a cue to direction of attention. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

The effect of eye position on auditory lateralization

The present study examines whether the direction of gaze can influence sound lateralization. For this purpose, dichotic stimuli with variable interaural level difference (ILD) were presented under different conditions of visual fixation. In experiment 1, subjects with their head fixed directed their gaze to a given target, simultaneously adjusting the ILD of continuous pure tone or noise stimuli so that their location was perceived in the median plane of the head. The auditory adjustments were significantly correlated with gaze direction. During eccentric fixation, the psychophysical adjustments to the median plane shifted slightly toward the direction of gaze. The magnitude of the shift was about 1-3 dB, over a range of fixation angles of 45 degrees to either side. The eye position effect, measured as a function of pure-tone frequency, was most pronounced at 2 kHz and showed a tendency to decrease at lower and higher frequencies. The effect still occurred, although weaker, even when the eyes were directed to eccentric positions in darkness and without a fixation target. In experiment 2, the adjustment method was replaced by a two-alternative forced-choice method. Subjects judged whether sound bursts, presented with variable ILDs, were perceived on the left or right of the median plane during fixation of targets in various directions. Corresponding to experiment 1, the psychometric functions shifted significantly with gaze direction. However, the shift was only about half as large as that found in experiment 1. The shift of the subjective auditory median plane in the direction of eccentric gaze, observed in both experiments, indicates that dichotic sound is localized slightly to the opposite side, i.e., to the left when the gaze is directed to the right and vice versa. The effect may be related to auditory neurons which exhibit spatially selective receptive fields that shift with eye position.     

Eye movement guidance in reading: The role of parafoveal letter and space information.

The relation between parafoveal letter and space information in eye movement guidance during reading was investigated in 2 experiments. Contingent upon the reader's fixation, the type of parafoveal information available to the right of fixation was varied by (a) space information only, (b) space information with letter information added at some delay, or (c) letter and space information simultaneously. In addition, the onset of the relevant parafoveal information was delayed between 0 and 250 ms into the fixation. The time course of processing the 2 types of information (letters or spaces) differed, as did the nature of their impact on the eye movement record. Although both letter and space information influenced saccade length and initial landing positions within words, only letter information had an effect on fixation duration. In addition, fixation duration was affected only by information entering within the first 50 ms of the fixation, whereas saccade length was affected by information arriving at any time during the fixation. The results are consistent with a model of eye movement control (A. W. Inhoff and K. Raner [see PA, Vol 75:6513] and A. Pollatsek et al [see PA, Vol 74:27114]) in which 2 independent processes are operating in tandem to determine when and where to move the eyes during reading. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Eye- and head movements in freely moving rabbits

1. Eye- and head movements were recorded in unrestrained, spontaneously behaving rabbits with a new technique, based upon phase detection of signals induced in implanted coils by a rotating magnetic field. 2. Movements of the eye in space were exclusively saccadic. In the intersaccadic intervals the eyes were stabilized in space, even during vigorous head movements. Most of this stability was maintained in darkness, except for the occurrence of slow drift. 3. Many saccades were initiated while the head was stationary. They were accompanied by a similar, but slower head rotation with approximately the same amplitude. The displacement of the eye in space was a pure step without appreciable under- or over-shoot. The deviation of the eye in the head was mostly transient. 4. Other saccades were started while the head was moving and were possibly fast phases of a vestibulo-ocular reflex. The time course of the eye movement in space was identical for all saccades, whether the head was moving prior to the saccade or not. Eye movements without any head movement were not observed. 5. Saccades were mostly large (average 20-6 +/- 12-4 degrees S.D.) and never smaller than 1 degree. The relations of maximal velocity and duration to amplitude were similar to those reported for man. 6. Visual pursuit of moving objects, when elicited, was only saccadic and never smooth. 7. It is concluded that the co-ordination and dynamics of the rabbit's head- and eye movements are similar to those of primates. In the absence of foveal specilization, the eye movements are restricted to a rather global redirection of the visual field, possibly in particular of the binocular area.     

Amplitude and period discrimination of haptic stimuli

As part of a project to examine the ability of the hand to receive speech information, the present study examined subjects' ability to discriminate finger movements along the dimensions of amplitude and period (movement duration). The movements consisted of single-cycle, sinewave movements and single-cycle, cosine movements presented to the index finger. Difference thresholds were collected using an adaptive, two-interval, temporal forced-choice procedure. Amplitudes from 6 to 19 mm were examined, and the difference thresholds ranged from 10% to 18%. The thresholds were unaffected by the period of the movement. Periods from 3000 to 111 ms (0.33-9 Hz) were examined, and thresholds ranged from 6% to 16%. The thresholds were unaffected by the amplitude of the movement. Further measurements in which period was varied in the amplitude discrimination task and amplitude was varied in the period discrimination task indicated that subjects were not using peak velocity as the basis for discrimination. These measurements were collected using a display specifically designed for the examination of haptic stimulation and capable of presenting controlled patterns of movement and vibration to the fingers.     

Vestibular and somatosensory contributions to responses to head and body displacements in stance

The relative contribution of vestibular and somatosensory information to triggering postural responses to external body displacements may depend on the task and on the availability of sensory information in each system. To separate the contribution of vestibular and neck mechanisms to the stabilization of upright stance from that of lower body somatosensory mechanisms, responses to displacements of the head alone were compared with responses to displacements of the head and body, in both healthy subjects and in patients with profound bilateral vestibular loss. Head displacements were induced by translating two 1-kg weights suspended on either side of the head at the level of the mastoid bone, and body displacements were induced translating the support surface. Head displacements resulted in maximum forward and backward head accelerations similar to those resulting from body displacements, but were not accompanied by significant center of body mass, ankle, knee, or hip motions. We tested the effect of disrupting somatosensory information from the legs on postural responses to head or body displacements by sway-referencing the support surface. The subjects' eyes were closed during all testing to eliminate the effects of vision. Results showed that head displacements alone can trigger medium latency (48-84 ms) responses in the same leg and trunk muscles as body displacements. Nevertheless, it is unlikely that vestibular signals alone normally trigger directionally specific postural responses to support surface translations in standing humans because: (1) initial head accelerations resulting from body and head displacements were in opposite directions, but were associated with activation of the same leg and trunk postural muscles; (2) muscle responses to displacements of the head alone were only one third of the amplitude of responses to body displacements with equivalent maximum head accelerations; and (3) patients with profound bilateral vestibular loss showed patterns and latencies of leg and trunk muscle responses to body displacements similar to those of healthy subjects. Altering somatosensory information, by sway-referencing the support surface, increased the amplitude of ankle muscle activation to head displacements and reduced the amplitude of ankle muscle activation to body displacements, suggesting context-specific reweighting of vestibular and somatosensory inputs for posture. In contrast to responses to body displacements, responses to direct head displacements appear to depend upon a vestibulospinal trigger, since trunk and leg muscle responses to head displacements were absent in patients who had lost vestibular function as adults.(ABSTRACT TRUNCATED AT 400 WORDS)     

Children's detection of on-face and off-face gazes.

Assessed the accurate detection of gaze direction for 24 6-yr-old boys in 2 groups, using an on-face vs off-face gaze detection task. Compared to adults in a prior study (J. J. Gibson and A. D. Pick, 1963) who were tested with an on-face vs off-face detection task, and compared to children and adults in a prior study (C. Lord, 1974) who were tested with only an on-face detection task, these children were less accurate at detecting deviations from direct eye contact from an adult as well as a child peer. Discussion focuses on the functional equivalence of various degrees of off-face gazing and eye contact for young children. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Use of human visual attention cues by olive baboons (Papio anubis) in a competitive task.

The ability of 4 olive baboons (Papio anubis) to use human gaze cues during a competitive food task was investigated. Three baboons used head orientation as a cue, and 1 individual also used eye direction alone. As the baboons did not receive prior training with gestural cuts, their performance suggests that the competitive paradigm may be more appropriate for testing nonhuman primates than the standard object-choice paradigm. However, the baboons were insensitive to whether the experimenter could actually perceive the food item, and therefore the use of visual orientation cues may not be indicative of visual perspective-taking abilities. Performance was disrupted by the introduction of a screen and objects to conceal food items and by the absence of movement in cues presented. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Bottlenosed Dolphins (Tursiops truncatus) Comprehend the Referent of Both Static and Dynamic Human Gazing and Pointing in an Object-Choice Task.

The authors tested 2 bottlenosed dolphins (Tursiops truncatus) for their understanding of human-directed gazing or pointing in a 2-alternative object-choice task. A dolphin watched a human informant either gazing at or pointing toward 1 of 2 laterally placed objects and was required to perform a previously indicated action to that object. Both static and dynamic gaze, as well as static and dynamic direct points and cross-body points, yielded errorless or nearly errorless performance. Gaze with the informant's torso obscured (only the head was shown) produced no performance decrement, but gaze with eyes only resulted in chance performance. The results revealed spontaneous understanding of human gaze accomplished through head orientation, with or without the human informant's eyes obscured, and demonstrated that gaze-directed cues were as effective as point-directed cues in the object-choice task. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Anxiety and sensitivity to gaze direction in emotionally expressive faces.

This study investigated the role of neutral, happy, fearful, and angry facial expressions in enhancing orienting to the direction of eye gaze. Photographs of faces with either direct or averted gaze were presented. A target letter (T or L) appeared unpredictably to the left or the right of the face, either 300 ms or 700 ms after gaze direction changed. Response times were faster in congruent conditions (i.e., when the eyes gazed toward the target) relative to incongruent conditions (when the eyes gazed away from the target letter). Facial expression did influence reaction times, but these effects were qualified by individual differences in self-reported anxiety. High trait-anxious participants showed an enhanced orienting to the eye gaze of faces with fearful expressions relative to all other expressions. In contrast, when the eyes stared straight ahead, trait anxiety was associated with slower responding when the facial expressions depicted anger. Thus, in anxiety-prone people attention is more likely to be held by an expression of anger, whereas attention is guided more potently by fearful facial expressions. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Eye movements in reading: Perceptual and cognitive processes.

This article discusses the variability of eye movement behavior. The variability exists between readers, so that the average fixation duration for sonic readers my be around 200 ms and for other readers it may be closer to 300 ms; the average saccade length for some readers may be 6 letter spaces, while for others it may be closer lo 10 letter spaces; and some readers rarely regress while others regress quite frequently. These eye movement characteristics can also be influenced by text difficulty; more difficult text leads to longer fixation durations, shorter saccades, and more regressions. More important than the between reader variability is the within reader variability that exists with each of these measures. The article also discusses research on eye movement during reading. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Orienting gaze shifts during muscimol inactivation of caudal fastigial nucleus in the cat. I. Gaze dysmetria

The cerebellar control of orienting behavior toward visual targets was studied in the head-unrestrained cat by analyzing the deficits of saccadic gaze shifts after unilateral injection of muscimol in the caudal part of the fastigial nucleus (cFN). Gaze shifts are rendered strongly inaccurate by muscimol cFN inactivation. The characteristics of gaze dysmetria are specific to the direction of the movement with respect to the inactivated cFN. Gaze shifts directed toward the injected side are hypermetric. Irrespective of their starting position, all these ipsiversive gaze shifts overshoot the target by a constant horizontal error (or bias) to terminate at a "shifted goal" location. In particular, when gaze is directed initially at the future target's location, a response with an amplitude corresponding to the bias moves gaze away from the actual target. Additionally, when gaze is initially in between the target and this shifted goal location, the response again is directed toward the latter. This deficit of ipsiversive gaze shifts is characterized by a consistent increase in the y intercept of the relationship between horizontal gaze amplitude and horizontal retinal error. Slight increases in the slope sometimes are observed as well. Contraversive gaze shifts are markedly hypometric and, in contrast to ipsiversive responses, they do not converge onto a shifted goal but rather underestimate target eccentricity in a proportional way. This is reflected by a decrease in the slope of the relationship between horizontal gaze amplitude and horizontal retinal error, with, for some experiments, a moderate change in the y-intercept value. The same deficits are observed in a different setup, which permits the control of initial gaze position. Correction saccades rarely are observed when visual feedback is eliminated on initiation of the primary orienting response; instead, they occur frequently when the target remains visible. Like the primary contraversive saccades, they are hypometric and the ever-decreasing series of three to five correction saccades reduces the gaze fixation error but often does not completely eliminate it. We measured the position of gaze after the final correction saccade and found that fixation of a visible target is still shifted toward the inactivated cFN by 4.9 +/- 2.4 degrees. This fixation offset is correlated to, but on average 54% smaller than, the hypermetric bias of ipsiversive responses measured in the same experiments. In conclusion, the cFN contributes to the control of saccadic shifts of the visual axis toward a visual target. The hypometria of contraversive gaze shifts suggests a cFN role in adjusting a gain in the translation of retinal signals into gaze motor commands. On the basis of the convergence of ipsiversive gaze shifts onto a shifted goal, the straightness of gaze trajectory during these responses and the production of misdirected or inappropriately initiated responses toward this shifted goal, we propose that the cFN influences the processes that specify the goal of ipsiversive gaze shifts.