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R H Schor

Publications and source records attributed to R H Schor.

At least 19 recordsLinked to original sources

The "practical mathematics" of recording three-dimensional eye position using scleral coils.

The "gold standard" for recording the three-dimensional rotation of the eye involves placing two coils of wire, embedded in a soft plastic ring, on the sclera of the eye, then placing the subject inside a set of orthogonal oscillating magnetic fields, and using the currents induced in the eye coils to deduce the position of the coil, and hence of the eye, in space. Eye movements are actually eye rotations, which can be described mathematically by a special class of matrices, rotation matrices, or, alternatively, by a rotation vector related to the axis of the rotation. This article deals with the mathematical tools needed to implement the signal processing from such a multifield, dual-coil system and compute the precise rotational movement of the eye. One reason for making such careful measurements is to study an interesting constraint on eye movements, called Listing's law, which expresses ocular torsion, or rotation of the eye about its line of sight, in terms of the direction of gaze. Techniques for experimentally quantitating these constraints are also presented. Following a treatment of the "ideal" case, with coils and eye in perfect alignment, the additional techniques for dealing with various departures from ideality that are almost always encountered experimentally are examined. A final section deals with developing a validation protocol for eye movement analysis techniques using mechanical and computer simulations of eye movements.

Algorithms↗

Semicircular canal-otolith organ interaction during off-vertical axis rotation in humans.

The aim of this study was to examine the interaction of signals from the semicircular canals and the otolith organs during off-vertical axis rotation (OVAR). We recorded horizontal eye position using electro-oculography in 22 young normal human subjects and stimulated the vestibulo-ocular reflex with both constant velocity trapezoids and sinusoidal yaw rotations, using both earth-vertical axis rotation (EVAR) and OVAR. We found that per-rotatory long vestibulo-ocular reflex (VOR) time constants during velocity trapezoids were shorter for OVAR than for EVAR, suggesting a reduction in the efficacy of the velocity storage system during OVAR. However, when we tested with very-low-frequency sinusoids (0.01 Hz and below), the phase lead of the VOR re head velocity was smaller during OVAR than EVAR, suggesting a longer time constant and enhanced efficacy of velocity storage during OVAR. These rotational responses can be explained by two competing influences of signals from the otolith organs, one that diminishes the effectiveness of velocity storage and another that contributes to an estimate of head velocity.

Acceleration↗

Otolith function after acoustic neuroma surgery.

The response of the vestibular system after acoustic neuroma surgery was investigated in nine patients. The otolith system was studied by means of ocular counterrolling, assessed by video oculography. Horizontal vestibulo-ocular reflex (VOR) function was tested by the sinusoidal harmonic acceleration test using electronystagmography. The results were compared with those obtained from a normal control population. The response to slow rotation tests was symmetric, but the gain was significantly reduced when compared to the normal population. Phase lag was significantly increased. No difference in ocular torsion was observed with lateroflexion of the head to the ipsilateral side in comparison with lateroflexion to the contralateral side. Moreover, the overall ocular counterrolling was well within normal limits. We conclude that the semicircular canal response differs from the otolith response. The component of the torsional VOR mediated by otolith stimulation appears to be more robust than the horizontal VOR mediated mainly by the horizontal semicircular canal system. Ocular counterrolling induced by lateroflexion does not reveal abnormalities in patients with surgically produced unilateral peripheral loss.

Adult↗

Characteristics of secondary phase post-rotatory nystagmus following off-vertical axis rotation in humans.

The nystagmus following yaw earth-vertical axis rotation often reverses direction, a phenomenon known as the "secondary phase". The purpose of this study was to examine the existence and the spatial and temporal properties of the secondary phase of post-rotatory nystagmus following off-vertical axis rotation (OVAR). Eleven normal human subjects were rotated at 120 or 180 degrees/s about an off-vertical axis and stopped in the left ear down or right ear down lateral position. Horizontal and vertical eye positions were recorded with a scleral search coil, and horizontal and vertical slow component eye velocities were computed. Our results indicate that (a) there is a robust secondary phase nystagmus following OVAR, and (b) the direction of the secondary phase nystagmus tends to align with earth-horizontal. These results can be explained by a minor modification of an existing VOR model that has been shown to produce secondary phase responses.

Adult↗

Cortical influences on the vestibular nuclei of the cat.

Our goal was to study potential substrates for cortical modulation of vestibular reflexes in the cat. In initial experiments, injections of wheat-germ-agglutinate-horseradish-peroxidase into Deiters' nucleus and the rostral descending nucleus revealed bilateral colonies of retrogradely filled neurons in cortical areas 6, 2, and 3a (about 60 cells per colony). In cats anesthetized with chloralose-urethane, we stimulated areas 2 and 3a with trains of pulses while recording from ipsilateral vestibular-nucleus neurons, which were characterized by their responses to sinusoidal tilts and tested for the presence of antidromic responses to stimulation of the upper cervical cord. A majority of the neurons was affected by cortical stimulation, showing either facilitation, inhibition, or a mixture of the two. Stimulation in area 2 was more effective than stimulation in area 3a. Despite the anatomic presence of direct cortico-vestibular projections, properties of facilitation and inhibition suggest that both were evoked by polysynaptic pathways. Cortical effects were broadly distributed to vestibular neurons without regard to responses of these neurons to sinusoidal tilts. There was no significant difference between effects on lateral and medial vestibulospinal tract neurons, but, as a group, vestibulospinal neurons were much more likely to be affected by cortical stimulation than neurons not antidromically activated from the C2 segment. We conclude that, by their influence on vestibulospinal neurons, neurons in cortical areas 2 and 3a should be able to modulate, in behaving animals, vestibular reflexes acting on the neck and limbs.

Animals↗

The neural substrate of the vestibulocollic reflex. What needs to be learned.

The purpose of this review is to assess the role of short-latency pathways in the vestibulocollic reflex (VCR). First the current knowledge about the disynaptic and trisynaptic pathways linking semicircular canal and otolith afferents with cat neck motoneurons is summarized. We then discuss whether these pathways are sufficient or necessary to produce the responses observed in neck muscles by natural vestibular stimulation and conclude that they are neither. Finally, alternate pathways are considered, most likely involving reticulospinal fibers, which are an important part of the neural substrate of the VCR.

Animals↗

Tilt responses of neurons in the caudal descending nucleus of the decerebrate cat: influence of the caudal cerebellar vermis and of neck receptors.

1. In decerebrate cats with intact cerebellums, we studied the responses of neurons in the caudal areas of the vestibular nuclei to natural vestibular stimulation in vertical planes and to neck rotation. The activity of most neurons was recorded in the caudal half of the descending nucleus. 2. One goal of our experiments was to compare the dynamic and spatial properties of responses to sinusoidal vestibular stimulation with those seen in previous experiments in which the caudal cerebellar vermis, including the nodulus and uvula, was removed. This part of the cerebellum receives vestibular input and projects to the caudal areas of the vestibular nuclei, suggesting that it could influence responses to stimulation of the labyrinth. 3. As in our previous experiments, most neurons could be classified as receiving predominant input either from the otoliths or from one vertical semicircular canal. When mean gain and phase and response vector orientations were compared, there were no obvious differences between the behavior of neurons in the partially decerebellate preparation and the one with the cerebellum intact, demonstrating that in the decerebrate cat the nodulus and uvula have little or no influence on the processing of vertical vestibular input in this region of the vestibular nuclei. 4. Only 23 of 74 (31%) of neurons tested responded to neck rotation. This contrasts with the much larger fractions that respond to this stimulus in Deiters' nucleus and in the rostral descending nucleus. We also recorded from neurons near the vestibular nuclei, mainly in the external cuneate nucleus. All of them (9 of 9) responded to neck rotation. 5. Responses to neck rotation also differed in their dynamics from those found more rostrally in the vestibular nuclei. Dynamics of more rostral neurons resemble those of neck muscle spindles, as do those of external cuneate neurons. The dynamics of caudal vestibular neurons, on the other hand, have a steeper gain slope and more advanced phases than do those of neurons in the more rostral vestibular nuclei. This suggests the possibility of involvement of additional receptors in the production of these responses. 6. In the more rostral vestibular nuclei, responses to vestibular and neck rotation are most often antagonistic, so that head rotation results in little or no response. This is not the case in the caudal areas of the vestibular nuclei, where less than half the neurons tested displayed antagonistic behavior. Further experiments are required to put the neck projection to the caudal vestibular nuclei in a functional context.

Animals↗

Off-vertical axis rotational responses in patients with unilateral peripheral vestibular lesions.

Off-vertical axis rotation (OVAR) stimulates the otolith organs in a manner that is suitable for assessment of the otolith-ocular reflex. To further assess the potential clinical usefulness of OVAR, the eye movement responses of seven patients with surgically confirmed unilateral peripheral vestibular lesions were compared with the eye movement responses of a group of age-matched, healthy, asymptomatic control subjects. Patients and controls were tested with constant velocity rotations that followed a brief period of angular acceleration (velocity trapezoid) using either earth-vertical axis (EVA) rotation or OVAR. Both EVA and OVAR sinusoidal velocity profiles were also performed. Results indicated that each patient had 1) an asymmetric OVAR response, ie, a bias component whose direction was opposite normal when rotating toward the lesioned ear, and 2) a normal modulation component. Population data suggested that patients had 1) a more rapid decay of response than normal subjects during OVAR velocity trapezoids, 2) an increased phase lead as compared to normal subjects during sinusoidal OVAR, and 3) like normal subjects, a less rapid decay of response during OVAR velocity trapezoids than during EVA rotational velocity trapezoids. Taken together, these findings suggest that patients with unilateral peripheral vestibular deficits have abnormal otolith-ocular and semicircular canal-ocular reflexes but that a single labyrinth appears to provide an otolithic signal sufficient for qualitatively normal semicircular canal-otolith interaction.

Adult↗

Response of pontomedullary reticulospinal neurons to vestibular stimuli in vertical planes. Role in vertical vestibulospinal reflexes of the decerebrate cat.

1. To investigate the neural substrate of vestibulospinal reflexes in decerebrate cats, we studied the responses of pontomedullary reticulospinal neurons to natural stimulation of the labyrinth in vertical planes. Our principal aim was to determine whether reticulospinal neurons that terminate in, or are likely to give off collaterals to, the upper cervical segments had properties similar to those of the vestibulocollic reflex (VCR). 2. Antidromic stimulation was used to determine whether the neurons projected to the neck, lower cervical, thoracic, or lumbar levels. Dynamics of the responses of spontaneously firing neurons were studied with sinusoidal stimuli delivered at 0.05-1 Hz and aligned to the plane of body rotation, that produced maximal modulation of the neuron (response vector orientation). Each neuron was assigned a vestibular input classification of otolith, vertical canal, otolith + canal, or spatial-temporal convergence (STC). 3. We found, in agreement with previous studies, that the largest fraction of pontomedullary reticulospinal neurons projected to the lumbar cord, and that only a small number ended in the neck segments. Neurons projecting to all levels of the spinal cord had similar responses to labyrinth stimulation. 4. Reticulospinal neurons that received only vertical canal inputs were rare (1 of 67 units). Most reticulospinal neurons (48%) received predominant otolith inputs, 18% received otolith + canal input, and only 9% had STC behavior. These data are in sharp contrast to the results of our previous studies of vestibulospinal neurons. A considerable portion of vestibulospinal neurons receives vertical canal input (38%), fewer receive predominantly otolith input (22%), whereas the proportion that have otolith + canal input or STC behavior is similar to our present reticulospinal data. 5. The response vector orientations of our reticulospinal neurons, particularly those with canal inputs (canal, otolith + canal, STC) were predominantly in the roll quadrants. There was no evidence of convergence of inputs from like canals across the midline (e.g., right anterior + left anterior). 6. Two characteristics of the VCR, STC behavior and bilateral input from symmetric vertical canals (in some muscles), cannot be accounted for by the reticulospinal neurons that we studied. Because these characteristics are also not seen in vestibulocollic neurons, they are likely to be the result of the appropriate convergence of vestibular signals in the spinal cord. 7. Pontomedullary reticulospinal neurons seem particularly well suited to play a role in gravity-dependent postural reflexes of neck and limbs.

Acoustic Stimulation↗

Off-vertical axis rotation: a test of the otolith-ocular reflex.

The vestibulo-ocular reflex was studied via off-vertical axis rotation (OVAR) in the dark. The axis of the turntable could be tilted from vertical by up to 30 degrees. Eye movements were measured with electro-oculography. Results from healthy asymptomatic subjects indicated that 1) a reliable otolith-induced response could be obtained during constant velocity OVAR using a velocity of 60 degrees/s with a tilt of 30 degrees; 2) constant velocity OVAR rotation was nausea-producing and, especially if subjects were rotated in the dark about an earth-vertical axis prior to being tilted, disorienting; and 3) sinusoidal OVAR produced minimal nausea; the eye movement response appeared to be the result of a combination of semicircular canal and otolith components. We conclude that OVAR has the potential of becoming a useful method for clinically assessing both the otolith-ocular reflex and semicircular canal-otolith interaction.

Adult↗

Bilateral labyrinthectomy in the cat: motor behaviour and quiet stance parameters.

The purpose of this study was to examine the effect of bilateral labyrinthectomy on quiet stance in the freely-standing cat. Since loss of the vestibular end organs produces marked deficits in motor behaviour, including ataxia and problems with balance, we hypothesized that labyrinthectomized animals would show impairment in quantitative measures of stance. Stance was quantified in terms of the ground reaction forces under each limb and the tonic electromyographic (EMG) activity of selected muscles. Animals were labyrinthectomized by drilling into the vestibule and removing the vestibular epithelium. Following lesion, animals were able to stand unsupported on the force platform within 2 days. To our surprise, the lesioned animals showed little change in stance parameters from the control, pre-lesion state. Thus, our hypothesis of changes in stance parameters was not supported. There was no change in the distribution of vertical forces under the limbs and no increase in sway, as measured by the area of excursion of the centre of pressure over time. The horizontal plane forces, which were diagonally directed prior to lesion, became more laterally directed and larger in amplitude. The change in direction persisted even after the animals had fully compensated for the lesion, but the force amplitudes returned to control values within 10-12 days. The change in horizontal force direction was similar to that observed in normal animals that were required to stand with their paws closer than preferred in the sagittal plane (unpublished observations). EMG activity changed in some muscles but not others, and usually transiently.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Response of vestibular neurons to head rotations in vertical planes. III. Response of vestibulocollic neurons to vestibular and neck stimulation.

1. To compare the properties of the vestibulocollic reflex (VCR) with those of vestibular neurons projecting to the neck [vestibulocollic (VC) neurons], we have studied the behavior of the latter in the decerebrate cat. Neurons were identified by their antidromic responses to stimulation in C1-C2, but not C5. Responses to stimulation of vestibular and neck receptors were produced by rotation of the body and head in vertical planes. 2. We determined the plane of whole body (vestibular) or body with head counter-rotated (neck) rotation, which produced the maximal modulation of each neuron (response vector orientation). Neuron dynamics were then studied with sinusoidal (0.02-2 Hz) stimuli aligned with this orientation. 3. On the basis of dynamics and vector orientation, the neuron was assigned a vestibular input classification of otolith, vertical canal, otolith + canal, or spatial-temporal convergence (STC). 4. The properties of this sample of VC neurons are similar to those of a larger population of vestibular neurons whose projection was not identified. For example, the distributions of cells with different types of vestibular inputs were roughly the same; in particular, few cells showed STC responses. In addition, there was no evidence of significant convergence of like canals across the midline (e.g., right anterior + left anterior). 5. Also similar to the larger unidentified population, 80% of VC neurons tested for neck input received such an input. The neck and vestibular responses tended to be antagonistic; the vector orientations were usually opposite, and the response gains and phases similar.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Neck-vestibular interaction in the vestibular nuclei. A dynamic, two-dimensional study.

In decerebrate cats we used response dynamics to classify neurons in Deiters' nucleus and the rostral descending nucleus as receiving input from vertical canals, otolith organs (utricle) or canal + otolith. Many of these neurons, particularly those with convergent vestibular input (canal + otolith, canal + canal) and preferred stimulus directions near roll, also received neck input. Neck and vestibular response vectors tended to point in opposite directions. Neck and vestibular response dynamics were often well matched, so that combined stimuli (head rotation) produced no response over a wide frequency range. In some cases differing dynamics produced an output that appeared to code head position.

Animals↗

Three-dimensional sensitivity and caudal projection of neck spindle afferents.

1. We recorded from neck muscle spindle afferents in the C2 dorsal root ganglion of the decerebrate cat using floating electrodes. The afferents presumably innervated mainly ventral and ventrolateral perivertebral muscles, and sternocleidomastoid. Stimuli consisted of combinations of rotatory head movements about the roll/pitch or pitch/yaw axes. An important difference from our earlier experiments (10) was the addition of yaw movement to the stimulus paradigm making possible a three-dimensional analysis of afferent behavior. 2. For each afferent we determined the most effective direction of tilt (orientation of the response vector) in three dimensions by using sinusoidal stimuli that combined pitch and roll, or pitch and yaw, or by measuring the gains to responses to roll, pitch, and yaw rotation. 3. Most afferents were sensitive to rotation around all three axes; pitch and yaw were usually more effective than roll. There was no indication of clustering of response vectors, as might be expected if the receptors were located in a small number of muscles each of which has receptors aligned in a homogeneous direction. 4. The responses of afferents were further studied using sinusoidal and trapezoidal stimuli aligned as closely as possible with the orientation of their response vector. The availability of the yaw stimulus made receptor classification based on response linearity, gain, and dynamic index more reliable than in our earlier experiments (10). 5. Muscle spindle responses were divided into three categories: A, B, and ambiguous. The evidence suggests that category A are probably spindle primary receptors and category B are secondaries. Ambiguous receptors have intermediate properties. 6. The caudal projection of spindle afferents was examined by delivering antidromic stimuli with a movable electrode on the surface of the ipsilateral dorsal column. Eighteen percent of the afferents projected to C4, and 14% as far as C5. Long caudal projections can be found in A, B, and ambiguous receptors with a range of directional sensitivities. 7. The evidence suggests that C2 spindle afferents make synapses in the midcervical segments with interneurons and propriospinal neurons that are part of the intraspinal pathway of the tonic neck reflex.

Action Potentials↗