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C Evinger

Publications and source records attributed to C Evinger.

49 records · Page 3Linked to original sources

Different forms of blinks and their two-stage control.

The purpose of this paper is to examine blink kinematics and the neural basis of blinks evoked reflexively by different kinds of stimuli. The kinematics of the upper lid movement and the electromyographic response of lid muscles levator palpebrae and orbicularis oculi were recorded in the rabbit during trigeminally and visually-evoked blinks. We find that there is a basic, kinematic difference between blinks. A blink in response to an airpuff is more rapidly accomplished and achieves a higher velocity than does an equal amplitude blink in response to a flash of light. The two forms of the reflex blink result from differences in the nature and timing of activity in antagonistic lid muscle motoneurons. Nevertheless, most characteristics of blink neural control are common to both reflex blinks. Most importantly, it appears that blinks are produced by two-stage neural control, an early component that is preprogrammed and a late component that is under stimulus control.

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Retrograde horseradish peroxidase transport after oculomotor nerve injury.

We studied the distribution of somatic motor neurons innervating the cat superior rectus 3-6 months after oculomotor nerve injury using intramuscular horseradish peroxidase (HRP). In normal cats, 98% or more of the labelled superior rectus motoneurons were in the contralateral oculomotor subnucleus. Two experimental cats who exhibited little or no evidence of recovery showed few labelled cells (4% of controls) which were distributed in both the ipsilateral and contralateral oculomotor nucleus. The other three experimental cats demonstrated definite signs of recovery, and HRP injections labelled more cells (20% of controls) also distributed in the ipsilateral and contralateral oculomotor subnuclei. This study shows that, after sectioning, the oculomotor nerve regenerates and anomalous connections develop between the somatic motoneurons of the ipsilateral oculomotor nucleus and the superior rectus. These findings support the hypothesis that acquired oculomotor synkinesis developing after third nerve injury results from misdirection of regenerating axons.

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Blinking and associated eye movements in humans, guinea pigs, and rabbits.

Recordings of upper eyelid movements in humans, guinea pigs, and rabbits demonstrated that all three species displayed qualitatively similar patterns of eyelid movement. The relation between amplitude, duration, and maximum velocity in rabbits and humans was nearly identical. Guinea pig blinks were faster than those of rabbit and man. Electromyographic (EMG) recordings in humans demonstrated that the orbicularis oculis muscle participated in downward movement of the upper eyelid during blinks and eyelid closure but did not participate actively in the downward lid movement occurring with gaze changes. When looking straight ahead, the estimated stiffness and viscosity of the upper eyelid were 10 g/mm and 0.38 g X s X mm-1 for humans and 1.17 g/mm and 0.062 g X s X mm-1 for rabbits. Upward and abducting rotations of the eye accompanied blinks in rabbits and guinea pigs. Simultaneously, the eyeball retracted (translational movement) into the orbit. These translational and rotational eye movements resulted from contraction of the retractor bulbi muscle and cocontraction of antagonistic extraocular muscles. The data suggested that humans also retracted the eye during voluntary blinks. The retraction produced a rotation of the eye toward a "primary position" rather than a rotation in one specific direction. The relationship between the maximum velocity, duration, and amplitude of the down phase of a blink may be expressed as a single equation, maximum velocity = c X average velocity, where c is a constant. The same relationship, with a similar value for c, also describes saccadic eye movements and rapid skeletal movements. This implies that all three movements employ comparable neural mechanisms.

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Electron microscopic observations of axon collateral synaptic endings of cat oculomotor motoneurons stained by intracellular injection of horseradish peroxidase.

Motoneurons in the cat oculomotor nucleus have been identified electrophysiologically and stained by intracellular injection of horseradish peroxidase. Axon collateral arborizations with preterminal and terminal boutons identified by light microscopy corresponded to synaptic endings observed by electron microscopy. Despite variations in size and shape, synaptic endings showed similar ultra-structural features and established asymmetrical predominantly axodendritic synaptic contacts usually characterized by the presence of subjunctional dense bodies underlying the postsynaptic membrane densification.

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Activity of omnipause neurons in alert cats during saccadic eye movements and visual stimuli.

1. In the cats trained to follow a target spot with their eyes, activity was recorded from omnipause neurons (OPNs). OPNs discharge at a relatively high steady tonic rate (50-130 spikes/s) during visual fixation and smooth-pursuit eye movements but exhibit a complete cessation of discharge that begins before saccades in any direction. They are located in a compact region of the dorsal pontine tegmentum near the midline, just rostral to the abducens nucleus. 2. The average duration of the horizontal or vertical component of a saccade increases monotonically with pause duration, but a given pause duration is associated with a large range of individual saccade parameters and the timing of the pause, such as the latency from the pause onset to saccade onset or the interval from the maximum saccade velocity to the end of the pause, is no better. However, OPNs can be divided into two distinct groups on the basis of the timing of the pause relative to the parameters of the saccade. One group ceases discharging 32.4 +/- 4.6 ms, on average, before the saccade, while the second pauses 18.2 +/- 3.4 ms before the saccade. 3. Microstimulation at the site of OPNs affects the occurrence and trajectory of saccades but not smooth pursuit or fixation. Sustained electrical stimulation (20 micro A) lasting several seconds prevents the occurrence of saccades while brief trains (10-60 ms), timed to occur early in the saccade, interrupt it in midflight for the duration of the train. The latency to the interruption is about 26 ms. These data support the view that OPNs tonically inhibit the saccadic machinery between saccades and must be turned off to allow a saccade to occur. 4. Almost every (65 of 69) feline OPN exhibited a brief transient increase in activity for visual stimuli moving in any direction with a wide range of velocities. A moving 1 degree spot was generally more effective than a moving full-field, striped background. All units also showed a transient increase in firing when the spot was turned either on or off. Receptive fields plotted with the spot were greater than 250 deg2 and always included the area centralis. Two-thirds of the cells tested also responded to auditory stimuli. 5. Interaction between the excitatory visual input and the saccade-related pause was tested by comparing OPN activity and the saccadic trajectory during eye movements in the dark versus the light and by presenting brief flashes of light during a saccade. During saccades in the dark, the steady firing of OPNs was less than during saccades in the light. Only by stabilizing a flashed spot of light to occur on the area centralis at the beginning of the saccade was it possible to activate an OPN artificially to interrupt the saccade in midflight. Therefore, rather than being instrumental in specifically controlling the saccade trajectory, the visual input, along with the auditory and other sensory inputs, probably serves, under normal visual conditions, to help establish the tonic rate of OPNs. 6...

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Oblique saccadic eye movements of the cat.

A quantitative study of saccadic eye movements in the cat was undertaken to attempt to account for the high degree of variability in the trajectory of feline saccades compared with the more stereotyped monkey saccades. Cats were trained to make oblique saccades so that a large variety of saccadic component amplitudes, maximum velocities, and durations could be obtained. The horizontal and vertical components of oblique saccades were either stretched or compressed, relative to equal amplitude movements without orthogonal components, so that the two components were nearly equal in duration. On average, the components began approximately synchronously but were more asynchronous in their termination times. Consistent with the stretching or compression of a saccadic component, there was the complementary decrease or increase respectively of its maximum velocity. The product of maximum velocity and duration was constant for saccades of a given size. Furthermore, the constant relating maximum velocity and duration was a linear function of saccade size. Therefore, any saccade size was uniquely determined by knowing both duration and maximum velocity while knowing either alone resulted in considerable ambiguity in specifying saccade size. In addition to uniquely specifying feline saccades with different degrees of obliquity, this two parameter description allowed us to fit not only the monkey data but data obtained from rabbit and human as well. Therefore, specification of both duration and maximum velocity might be a basic organizing principle of the neural mechanisms generating saccades in many species.

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Saccadic, smooth pursuit, and optokinetic eye movements of the trained cat.

1. Cats were trained to track a small target by rewarding them for keeping their eyes on target. Eye movements were measured by the electromagnetic search coil technique. 2. Cat saccades are qualitatively similar to primate saccades, but exhibit more variability in their parameters. However, they have longer durations and lower maximum velocities than primate saccades. As in the monkey, the duration of the horizontal or vertical component of an oblique saccade is lengthened when the orthogonal component has a larger amplitude. Cat saccades can be modified in midflight like human saccades. Opening the visual feed-back loop by controlling target position with eye position causes the cat to execute a staircase of equal amplitude saccades if a retinal error is present. Increasing the amount of visual feed-back induces saccadic oscillations. 3. Horizontal smooth pursuit of a 0 . 5 deg visual target is limited to velocities of less than 1 deg/sec. However, moving an optokinetic background with the 0 . 5 deg target enables the cat to achieve higher horizontal smooth eye velocities of up to 8 . 5 deg/sec. Prolonged (10-20 sec) constant velocity rotation of an optokinetic drum evokes horizontal slow-phase velocities of up to 28 deg/sec. In response to vertical movements of the target and optokinetic background, smooth eye movements reached 6 deg/sec maximum upward velocities but only 2 . 5 deg/sec maximum downward velocities. Opening the feed back loop with no retinal error present causes the eye to exhibit a growing smooth trajectory. The response to a Rashbass step-ramp target suggests that the feline smooth response is a function of target movement rather than displacement. 4. These data suggest that cat saccadic eye movements resemble those of primates while the cat smooth pursuit and optokinetically induced eye movements are more similar to those of the rabbit.

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Role of cerebellum in adaptive modification of reflex blinks.

We investigated the involvement of the cerebellar cortex in the adaptive modification of corneal reflex blinks and the regulation of normal trigeminal reflex blinks in rats. The ansiform Crus I region contained blink-related Purkinje cells that exhibited a complex spike 20.4 msec after a corneal stimulus and a burst of simple spike activity correlated with the termination of orbicularis oculi activity. This occurrence of the complex spike correlated with trigeminal sensory information associated with the blink-evoking stimulus, and the burst of simple spike activity correlated with sensory feedback about the occurrence of a blink. Inactivation of the inferior olive with lidocaine prevented all complex and significantly reduced simple spike modulation of blink-related Purkinje cells, but did not alter orbicularis oculi activity evoked by corneal stimulation. In contrast, both acute and chronic lesions of the cerebellar cortex containing blink-related Purkinje cells blocked adaptive increases in orbicularis oculi activity of the lid ipsilateral but not contralateral to the lesion. These data are consistent with the hypothesis that the cerebellum is part of a trigeminal reflex blink circuit. Changes in trigeminal signals produce modifications of the cerebellar cortex, which in turn, reinforce or stabilize modifications of brainstem blink circuits. When the trigeminal system does not attempt to alter the magnitude of trigeminal reflex blinks, cerebellar input has little or no effect on reflex blinks.

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