Eye-head coordination during large gaze shifts.

In addition to many other symptoms, Huntington's Disease (HD) also causes an impairment of oculomotor functions. In particular, saccadic eye movements become progressively slower and more difficult to initiate; ultimately, patients are forced to recur to large head thrusts as means to initiate gaze shifts. We wondered whether, as a precursor of this condition, head movements would facilitate gaze shifts already in early stages of the disease. We studied horizontal head movements and eye-head coordination in 29 early stage HD patients (Ps) and 24 age matched controls (Cs). Subjects tracked random horizontal steps of visual or auditory targets while their heads were either stabilised (saccade amplitudes <or=40 degrees) or free to move (amplitudes <or=160 degrees). Subjects were to react either immediately (reactive mode), or wait until a go signal was sounded (delayed mode), or by antisaccades. Ps' head velocity was found to depend on the age of disease onset in a similar way as their saccadic eye velocity does, being clearly reduced in early affected Ps, but increasing to normal levels in lately affected Ps. Yet, saccade and head velocity were only loosely correlated although both exhibited a negative correlation with the severity of Ps' genetic condition (number of Ps' CAG repeats). Eye-head coordination turned out to be identical in Ps and Cs except for quantitative differences caused by the lower saccade and head velocities of Ps. Specifically, the timing between head and eyes and the head contribution to gaze shifts were similar in both groups. Moreover, preventing head movements did not affect the saccade latency or accuracy of Ps. Although Ps made more small involuntary head movements in this condition than Cs, these movements were not instrumental in generating saccades since they occurred only late after saccade onset. Thus, the head manoeuvres of severely affected patients must be considered a late adaptive behaviour. Finally, the ability of both Ps and Cs to suppress immediate reactions in the delayed and antisaccade conditions diminished as target distance decreased, with failure rates in Ps being much larger than in Cs. Unlike eye and head velocity, these failure rates were not correlated with age and, by the same token, neither with the variations in head and eye velocity nor with the number of CAG repeats. Hence, the pattern of brain areas prominently affected by HD is likely to vary significantly among individuals.

The mechanics of the eyeball and its surrounding tissues, which together form the oculomotor plant, have been shown to be the same for smooth pursuit and saccadic eye movements. Hence it was postulated that similar signals would be carried by motoneurons during slow and rapid eye movements. In the present study, we directly addressed this proposal by determining which eye movement-based models best describe the discharge dynamics of primate abducens neurons during a variety of eye movement behaviors. We first characterized abducens neuron spike trains, as has been classically done, during fixation and sinusoidal smooth pursuit. We then systematically analyzed the discharge dynamics of abducens neurons during and following saccades, during step-ramp pursuit and during high velocity slow-phase vestibular nystagmus. We found that the commonly utilized first-order description of abducens neuron firing rates (FR = b + kE + r, where FR is firing rate, E and are eye position and velocity, respectively, and b, k, and r are constants) provided an adequate model of neuronal activity during saccades, smooth pursuit, and slow phase vestibular nystagmus. However, the use of a second-order model, which included an exponentially decaying term or "slide" (FR = b + kE + r + uE - c), notably improved our ability to describe neuronal activity when the eye was moving and also enabled us to model abducens neuron discharges during the postsaccadic interval. We also found that, for a given model, a single set of parameters could not be used to describe neuronal firing rates during both slow and rapid eye movements. Specifically, the eye velocity and position coefficients (r and k in the above models, respectively) consistently decreased as a function of the mean (and peak) eye velocity that was generated. In contrast, the bias (b, firing rate when looking straight ahead) invariably increased with eye velocity. Although these trends are likely to reflect, in part, nonlinearities that are intrinsic to the extraocular muscles, we propose that these results can also be explained by considering the time-varying resistance to movement that is generated by the antagonist muscle. We conclude that to create realistic and meaningful models of the neural control of horizontal eye movements, it is essential to consider the activation of the antagonist, as well as agonist motoneuron pools.

The superior colliculus (SC) is a multilayered structure in the mammalian midbrain. Even though the behavioral function of the SC has been studied for well over 30 yr, it continues to be a source of surprising new findings. Traditionally it has been assumed that the primate SC selects and generates

Oculomotor related interaction of vestibular and visual stimulation in vestibular nucleus cells in alert monkey

Medial rectus motoneurons mediate nasally directed horizontal eye movements. These motoneurons receive two major excitatory inputs, from the abducens internuclear neurons (ABD Ints) and neurons of the lateral vestibular nucleus whose axons course through the ascending tract of Deiters (ATD). In the present work, we have recorded in the alert chronic cat preparation the discharge activity of these two premotor neurons simultaneously with eye movements, to discern their relative contribution to the firing pattern of medial rectus motoneurons. ABD Int discharge was accurately correlated with eye movements, displaying high sensitivities to eye position and eye velocity. ATD neurons also discharged in relation to spontaneous and vestibular eye movements but showed significantly lower eye position and eye velocity sensitivities. Outstandingly, ATD neurons presented a significantly lower eye position threshold for recruitment compared to both ABD Ints and medial rectus motoneurons. Therefore, ATD neurons exhibited eye position and velocity signals during spontaneous and vestibular eye movements, which were of lower magnitude than those of ABD Ints, but due to their low recruitment threshold, they could play a significant role in facilitating ABD Int signal transmission onto medial rectus motoneurons.

1. The spike activity of deep cerebellar nuclear neurons was recorded in the alert cat during spontaneous and during vestibularly and visually induced eye movements. 2. Neurons were classified according to their location in the nuclei, their antidromic activation from projection sites, their sensitivity to eye position and velocity during spontaneous eye movements, and their responses to vestibular and optokinetic stimuli. 3. Type I EPV (eye position and velocity) neurons were located mainly in the posterior part of the fastigial nucleus and activated antidromically almost exclusively from the medial longitudinal fasciculus close to the oculomotor complex. These neurons, reported here for the first time, increased their firing rate during saccades and eye fixations towards the contralateral hemifield. Their position sensitivity to eye fixations in the horizontal plane was 5.3 +/- 2.6 spikes s-1 deg-1 (mean +/- S.D.). Eye velocity sensitivity during horizontal saccades was 0.71 +/- 0.52 spikes s-1 deg-1 s-1. Variability of their firing rate during a given eye fixation was higher than that shown by abducens motoneurons. 4. Type I EPV neurons increased their firing rate during ipsilateral head rotations at 0.5 Hz with a mean phase lead over eye position of 95.3 +/- 9.5 deg. They were also activated by contralateral optokinetic stimulation at 30 deg s-1. Their sensitivity to eye position and velocity in the horizontal plane during vestibular and optokinetic stimuli yielded values similar to those obtained for spontaneous eye movements. 5. Type II neurons were located in both fastigial and dentate nuclei and were activated antidromically from the restiform body, the medial longitudinal fasciculus close to the oculomotor complex, the red nucleus and the pontine nuclei. Type II neurons were not related to spontaneous eye movements. These neurons increased their firing rate in response to contralateral head rotation and during ipsilateral optokinetic stimulation, and decreased it with the oppositely directed movements. 6. Saccade-related neurons were located mostly in the fastigial and dentate nuclei. Fastigial neurons were activated antidromically from the medial longitudinal fasciculus, while dentate neurons were activated from the red nucleus. These neurons fired a burst of spikes whose duration was significantly related to saccade duration. Dentate neurons responded during the omni-directional saccades, while some fastigial neurons fired more actively during contralateral saccades. 7. These three types of neuron represent the output channel for oculomotor signals of the posterior vermis and paravermis. It is proposed that type I EPV neurons correspond to a group of premotor neurons directly involved in oculomotor control.(ABSTRACT TRUNCATED AT 400 WORDS)

1. This series of three papers aims to describe the three-dimensional, kinematic input-output relations of the rotational vestibuloocular reflex (VOR) in humans, and to identify the functional advantages of these relations. In this first paper the response to sinusoidal rotation in darkness at 0.3 Hz, maximum speed 37.5%/s, was quantified by the use of the three-dimensional analogue of VOR gain: a 3 x 3 matrix where each element describes the dependence of one component (torsional, vertical, or horizontal) of eye velocity on one component of head velocity. 2. The three matrix elements indicating collinear gains (i.e., dependence of torsional eye velocity on torsional head velocity, vertical on vertical, and horizontal on horizontal) were smaller than the -1's required for optimal retinal image stabilization. Of these three the torsional gain was weakest: -0.37 for rotation about an earth-vertical axis, versus -0.73 and -0.64 for vertical and horizontal gains. Matrix elements indicating cross talk were mostly negligible. There was a tendency to leftward eye rotation in response to clockwise head motion, but this was not statistically significant. 3. VOR responses were compared for rotation about earth-vertical and earth-horizontal axes. The varying otolith input due to the rotation of the gravity vector relative to the head during earth-horizontal axis rotation made no difference to the collinear gains. 4. There were no consistent phase leads or lags except for a torsional phase lead of up to 10 degrees, usually more marked for clock-wise head rotation versus counterclockwise, and for oblique axis rotations versus purely torsional. 5. Torsional gain was magnified, averaging -0.52, when the torsional component of head rotation was only a small part of a predominantly vertical or horizontal rotation, i.e., when the axis of head rotation was near the frontal plane. Because most natural head rotations occur about such axes, the torsional VOR is probably somewhat stronger than the response to pure torsion would suggest. 6. The speed of eye rotation in response to a given stimulus varied widely among subjects, but the direction of rotation was much more uniform. For head rotations about oblique axes out of the frontal plane, there was a systematic misalignment of eye and head axes, with eye axes tilted toward the frontal plane. These findings can be explained on the basis of a strategy where the VOR balances the muscular effort of rotating the eyes against the cost of retinal slip.

The goal of this study was to test whether a superposition model of smooth-pursuit and vestibuloocular reflex (VOR) eye movements could account for the stability of gaze that subjects show as they view a stationary target, during head rotations at frequencies that correspond to natural movements. Horizontal smooth-pursuit and the VOR were tested using sinusoidal stimuli with frequencies in the range 1.0-3.5 Hz. During head rotation, subjects viewed a stationary target either directly or through an optical device that required eye movements to be approximately twice the amplitude of head movements in order to maintain foveal vision of the target. The gain of compensatory eye movements during viewing through the optical device was generally greater than during direct viewing or during attempted fixation of the remembered target location in darkness. This suggests that visual factors influence the response, even at high frequencies of head rotation. During viewing through the optical device, the gain of compensatory eye movements declined as a function of the frequency of head rotation (P < 0.001) but, at any particular frequency, there was no correlation with peak head velocity 9P > 0.23), peak head acceleration (P > 0.22) or retinal slip speed (P > 0.22). The optimal values of parameters of smooth-pursuit and VOR components of a simple superposition model were estimated in the frequency domain, using the measured responses during head rotation, as each subject viewed the stationary target through the optical device. We then compared the model's prediction of smooth-pursuit gain and phase, at each frequency, with values obtained experimentally. Each subject's pursuit showed lower gain and greater phase lag than the model predicted. Smooth-pursuit performance did not improve significantly if the moving target was a 10 deg x 10 deg Amsler grid, or if sinusoidal oscillation of the target was superimposed on ramp motion. Further, subjects were still able to modulate the gain of compensatory eye movements during pseudo-random head perturbations, making improved predictor performance during visual-vestibular interactions unlikely. We conclude that the increase in gain of eye movements that compensate for head rotations when subjects view, rather than imagine, a stationary target cannot be adequately explained by superposition of VOR and smooth-pursuit signals. Instead, vision may affect VOR performance by determining the context of the behavior.

Single-unit activities were recorded from floccular Purkinje cells in 29 alert pigmented rabbits. The floccular areas specifically related to horizontal eye movement (H-zone) were identified by the effects of local stimulation and later confirmed histologically. Most of the 53 H-zone Purkinje cells responded to both vestibular stimulation with turntable oscillation (5 degrees peak-to-peak, 0.1 Hz) in darkness and optokinetic stimulation with dot screen oscillation (2.5 degrees, 0.33 Hz), often showing a preference either for turntable oscillation or screen oscillation. Since these responses were not correlated with the velocity of evoked eye movements or eye position shift, it is concluded that the major inputs to flocculus H-zone Purkinje cells are head velocity or retinal slip signals rather than eye velocity or position signals.

Purkinje cell activity in the middle zone of the cerebellar flocculus during optokinetic and vestibular eye movement in cats.

To investigate the characteristics of the head movements and the mechanisms of gaze, stabilization during walking and running, eye and head movements were recorded in 10 normal subjects and 8 patients with labyrinthine lesions, who repeated stepping (1.OHz) and running (1.5Hz) on the same spot. 1. In normal subjects, head displacements during stepping were limited in the vertical, right-left and anteroposterior directions (0.8, 1.5 and 1.5cm on average, respectively), and the head rotations and the eye movements were also small. During running, the increase in the vertical head displacement (5.5cm on average) induced rhythmical pitching motions of the head and vertical eye movements. 2. In patients with labyrinthine lesions, both the horizontal head displacement and the rotation amplitude became twice as large as those in normal subjects during either locomotion, however, horizontal eye movements did not correspondingly increase. The vertical head displacements during running resembled those of normal subjects, but the amplitude of pitching motion of the head significantly decreased. 3) The spatial stabilization of the head during stepping in normal subjects indicated that walking is a suitable locomotion for gaze stabilization. On the other hand, the head was more unstable during running, however, the pitching motions of the head resulting from vertical head displacements seemed to reduce the concussion of the head and produce vertical compensatory eye movements which maintained the gaze stabilization. 4) In patients with labyrinthine lesions, the horizontal eye movements did not compensate the horizontal head movements during walking and running. Furthermore, since the pitching motions of the head could not be produced rhythmically during running, the vertical body oscillations were directly conducted to the head. The abnormalities of the head movements in addition to the insufficient compensations may cause oscillopsia in these patients. 5) The present study indicated that the head stabilization is essential to maintain the gaze stabilization during walking and running and that the semicircular canals-ocular reflexes act a major role in these mechanisms.

Passive head rotation in darkness produces vestibular nystagmus, consisting of slow and quick phases. The vestibulo-ocular reflex produces the slow phases, in the compensatory direction, while the fast phases, in the same direction as head rotation, are of saccadic origin. We have investigated how the saccadic components of the ocular motor responses evoked by active head rotation in darkness are generated, assuming the only available sensory information is that provided by the vestibular system. We recorded the eye and head movements of nine normal subjects during active head rotation in darkness. Subjects were instructed to rotate their heads in a sinusoidal-like manner and to focus their attention on producing a smooth head rotation. We found that the desired eye position signal provided to the saccadic mechanism by the vestibular system may be modeled as a linear combination of head velocity and head displacement information. Here we present a mathematical model for the generation of both the slow and quick phases of vestibular nystagmus based on our findings. Simulations of this model accurately fit experimental data recorded from subjects.

The study of the earth mass movement has long been regarded as one of the most important and interesting aspect of engineering geology and geotechnical engineering, which the designers and planners from the private and public sectors address when implementing the initial stage of urban and rural development projects. This involves highways and infrastructures construction and land use planning among the others. Failure to appreciate the problems relating to mass movements of earth material could lead to damage of man made structures and even the loss of lives. These studies focused on the mass movement in Bundu Tuhan to Kundasang highway area approximately 84 km to 96 km from Kota Kinabalu city, Sabah, one of the most vulnerable to mass movements occurrence in west coast of Sabah. It is bounded by longitude line E 116o 31.592’ to E 116o 36.183’ and latitude line N 06o 00.269’ to N 05o 57.610’. The main objectives of this study are; 1) to map and locate the landslides in the study area; and 2) to study the mechanism and the influence of geological factors causing the mass movement. Geology of the study area and its surrounding is hosted mainly by three sedimentary rock formations: Trusmadi Formation (Palaeocene to Eocene age), Crocker Formation (Late Eocene to Early Miocene age) and Pinousuk Gravel (Upper Pleistocene to Holocene age). These three geologic formations dissected by numerous geological lineaments structural produced by a complex tectonic history of multi phase folding and thrust, normal and reverse faulting. These tectonic setting reduce the physical and mechanical properties of the soil and produced intensive displacement in substrata resulting in intensive high degree of weathering processes. The weathered materials are weak and cause sinking, subsidence and sliding due to high pore pressure subjected by both shallow and deep groundwater. Evaluation 10 boreholes data in study area indicated that the groundwater table in study area is shallow and range 1.9 meter to about 11.3 meters. The groundwater in study area fluctuate drastically even within short period. Sand and gravel layer with variable thickness defined the major shallow aquifers within the top weathered materials while the highly fracture sedimentary rocks defined the major deep aquifers. Most of the aquifer within top unconsolidated weathered material is under unconfined condition. Most of significant aquifers within the sedimentary rocks are sandstones. The sandstones generally fracture and contain coarse sediments, which increase the permeability. Geologic and geotechnic evaluation of the study area indicates that the mass movement take place when slope materials are no longer able to resist the force of gravity. These decrease the shear resistance resulting mass movement, which is due to internal and external factors. Internal factors involve some change in either physical or chemical properties of the rock and soil. External factors involve increase of shear stress on slope, which usually involves a form of disturbance that is induced by man. The triggering mechanism in the study area most likely involves heavy rainfall causing water saturation of the slope material and loss of cohesion along rapture planes. The sheared shale, bedding and fault planes, and opening fractures are all structural weaknesses, which acting as pathways for water seepage, hastening the weakening and eventual mass movement in the study area. Development planning has to consider these hazards in order to counter their effect. An environmental management program should be implemented to prevent these losses. Geological and geotechnical studies will play a vital role in ground stability assessment that critical in public safety.

Heading detection using motion templates and eye velocity gain fields.

Analysis and Algebra on Differentiable Manifolds