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E J Tehovnik

Publications and source records attributed to E J Tehovnik.

12 recordsLinked to original sources

Compensatory saccades made to remembered targets following orbital displacement by electrically stimulating the dorsomedial frontal cortex or frontal eye fields of primates.

If the eye-position signal during visually-evoked saccades is dependent on the dorsomedial frontal cortex (DMFC), one would expect that saccades generated to briefly presented visual targets would be disrupted after displacement of the eyes via electrical stimulation of this cortical area. Compared are compensatory saccades evoked to brief targets following stimulation of the DMFC and frontal eye fields (FEF). Compensatory saccades produced to brief targets following perturbation via the DMFC were not affected. Accordingly, electrical stimulation of the DMFC does not disrupt the eye-position signal during the execution of visually-evoked saccades.

Animals

Electrical stimulation of neural tissue to evoke behavioral responses.

This review yields numerous conclusions. (1) Both unit recording and behavioral studies find that current activates neurons (i.e., cell bodies and axons) directly according to the square of the distance between the electrode and the neuron, and that the excitability of neurons can vary between 100 and 4000 microA/mm2 using a 0.2-ms cathodal pulse duration. (2) Currents as low as 10 microA, which is considered within the range of currents typically used during micro-stimulation, activate from a few tenths to several thousands of cell bodies in the cat motor cortex directly depending on their excitability; this indicates that even low currents activate more than a few neurons. (3) Electrode tip size has no effect on the current density--or effect current spread--at far field, but tip size limits the current-density generated at near field. (4) To minimize neuronal damage, the electrode should be discharged after each pulse and the pulse duration should not exceed the chronaxie of the stimulated tissue. (5) The amount of current needed to evoke behavioral responses depends not only on the excitability of the stimulated substrate but also on the type of behavior being studied.

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Topographic distribution of fixation-related units in the dorsomedial frontal cortex of the rhesus monkey.

Most cells in the dorsomedial frontal cortex of the rhesus monkey had activity related to saccadic eye movement and/or visual fixation. This activity changed depending upon the position of a fixation target, which suggested coding for the target location in spatial coordinates. Further analysis of such activity revealed a topographical distribution of neurons: neurons in the rostral part of the area were more active with eyes to the contralateral position, while those in the caudal part were more active with eyes to the ipsilateral position; also, cells in the medial part of the area had higher activity with a downward fixation position, whereas those in the lateral part had higher activity with an upward fixation position. This distribution of units was in agreement with the map of termination zones of saccadic eye movements evoked by electrical stimulation of the same area. These observations provide evidence for the hypothesis that the dorsomedial frontal cortex is organized in spatial coordinates and is involved in specifying the position of visual fixation.

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The dorsomedial frontal cortex: eye and forelimb fields.

This review yields three conclusions: first, the eye field as described using unit recording and electrical stimulation on behaving monkeys trained to fixate visual targets is much larger than the 4 mm2 area originally described. Second, the eye field and forelimb field share a similar neural space within the dorsomedial frontal cortex (DMFC); thus the electrophysiogical studies that have been conducted on visually guided and sensory-triggered forelimb movements must be re-evaluated, since none of these studies controlled eye movement and eye position independently. Third, a topographic map representing eye position in orbit has been discovered in the DMFC; it is proposed that this topographic map records the order of positions of the eyes and forelimbs during the acquisition of visually guided movement sequences.

Animals

Stimulation-evoked saccades from the dorsomedial frontal cortex of the rhesus monkey following lesions of the frontal eye fields and superior colliculus.

This study examined whether signals for the generation of eye movements from the dorsomedial frontal cortex (DMFC) reach brainstem oculomotor centers either through the frontal eye fields (FEF) or through the superior colliculi (SC). The DMFC was stimulated when the monkeys studied were intact and after either one FEF or one SC was ablated. Following lesions of either the FEF or SC, the topographic order of the DMFC was largely preserved. After either lesions, stimulation of anterior DMFC sites still evoked saccades that terminated in contralateral space, and stimulation of posterior DMFC sites still evoked saccades that terminated in central space. The probability of evoking saccades decreased and the latency to evoke saccades increased as fixation neared the termination zone (a restricted region within craniotopic space) both before and after either lesion. Ablation of the SC, but not of the FEF, eliminated the saccadic inhibition to visual targets which resulted when the DMFC was stimulated in the intact animal. The findings suggest that additional channels besides those coursing through the FEF and SC are utilized by the DMFC to access the saccade generator in the brainstem.

Animals

The dorsomedial frontal cortex of the rhesus monkey: topographic representation of saccades evoked by electrical stimulation.

The dorsomedial frontal cortex (DMFC) of monkeys has been implicated in mediating visually guided saccadic eye movements. The purpose of this study was to determine whether the DMFC has a topographic map coding final eye position, and to ascertain whether this region subserves the maintenance of eye position. The DMFC was stimulated electrically while monkeys fixated a target presented somewhere in visual space. A series of parametric tests was conducted to ascertain the best stimulation parameters to evoke saccades. Electrical stimulation typically produced contraversive saccades that converged onto a region of space, the termination zone. For some stimulation sites, however, stimulation produced ipsiversive saccades. This occurred when the termination zone was located straight ahead of the monkey. Convergence onto an orbital position was never observed during stimulation of the frontal eye fields (FEF), stimulation of which evoked fixed-vector saccades. The latency to evoke a saccade from the DMFC varied with fixation position, such that it increased monotonically the closer the fix spot was to the termination zone. Moreover, the probability of evoking a saccade from the DMFC decreased the closer the fix spot was to the termination zone. The latency for evoking a saccade and the probability of evoking a saccade from the FEF did not vary with fixation position. Horizontal head movements were not evoked from the DMFC while a monkey fixated targets presented in different positions of visual space. Moveover, changing the position of the head with respect to the body did not change the location of a termination zone with respect to the head. The DMFC was found to contain a topographic coding of termination zones, with rostral sites representing zones in extreme contralateral visual space, and caudal sites representing zones straight ahead or ipsilaterally. Furthermore, lateral sites represented zones in upper visual space, whereas medial sites represented zones in lower visual space. Once the eyes were positioned within a termination zone, further stimulation fixed the gaze and inhibited visually evoked saccades. Following release from inhibition, which occurred shortly after the end of stimulation, the saccades reached the visual target accurately. This shows that the stimulation delayed the execution of the saccades without actually aborting their execution. We conclude that the DMFC contains a map representing eye position in craniotopic coordinates, and we argue that this map is utilized to maintain eye position.

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Efferent projections of the anteromedial cortex of the rat as described by Phaseolus vulgaris leucoagglutinin immunohistochemistry.

Phaseolus vulgaris leucoagglutinin (PHA-L) immunohistochemistry was used to describe the corticofugal projections of the anteromedial cortex (AMC) of rats. PHA-L was injected iontophoretically into an area of the AMC which, when stimulated electrically, is known to induce contraversive head and body movements. It was found that the AMC innervates the midbrain via three separate pathways: a dorsal transthalamic pathway terminating in the pretectum, superior colliculus, and central grey area; and a ventral transthalamic pathway and a ventral capsular-peduncular pathway projecting to the central grey and mesencephalic and pontine reticular formation. The strongest terminations were found bilaterally in the mediodorsal thalamic nucleus and nucleus caudato-putamen. The functional significance of the pathways and terminations is discussed.

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Head and body movements evoked electrically from the caudal superior colliculus of rats: pulse frequency effects.

The effects of pulse frequency and current intensity on circling elicited from the caudal superior colliculus (SC) of rats were studied. The displacement of the head with respect to the body were measured for different levels of frequency (20, 29, and 50 Hz) and current (200 or 500 microA) at a pulse duration of 0.1 ms. The rate of circling increased monotonically with frequency and current. The rate at which the head was displaced laterally varied as a function of frequency. It is postulated that lateral head and body movements are affected by the firing frequency of SC output neurons.

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Contraversive circling elicited from the internal capsule and substantia nigra: evidence for a continuous axon bundle mediating circling.

Electrical stimulation of many brain sites (e.g., anteromedial cortex, internal capsule, substantia nigra, superior colliculus, rostro-medial tegmentum, and medial pons) evokes circling. The collision method of Shizgal et al. (J. Comp. Physiol. Psychol., 94 (1980) 227-237) was used to determine whether these sites are functionally connected for the production of circling in rats. If connectivity was evidenced, then refractory period and conduction velocity distributions were determined for axons passing through the connected stimulation sites. Collision of up to 90% was found between electrodes placed in internal capsule and substantia nigra, suggesting that these sites are connected by continuous axons that mediate circling. The refractory periods of these axons ranged from 0.5 to 4.5 ms, and the conduction velocities of these axons ranged from 0.9 to 4.4 ms. These velocities are similar to those of striatonigral axons. No collision was found between anteromedial cortex and any other sites tested, nor between pontine sites and internal capsule or substantia nigra.

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Turning responses evoked by stimulation of visuomotor pathways.

Lateral eye, head, and body movements are produced by electrical stimulation of many brain regions from frontal cortex to pons. A new collision method shows that at least 5 separate axon bundles mediate stimulation-elicited lateral head and body movements in rats. One bundle passes between the rostromedial tegmentum and medial pons, with conduction velocities of 0.8-18 m/s. A second bundle passes between the superior colliculus and contralateral medial pons, with conduction velocities of 1.7-13 m/s. A third bundle passes between the superior colliculus and ventrolateral pons, with conduction velocities of 1.3-20 m/s. A fourth bundle passes between the internal capsule and medial substantia nigra, with conduction velocities of 0.9-4.4 m/s. A fifth bundle passes between the anteromedial cortex and rostral striatum, with conduction velocities of 2.4-36 m/s. Collision effects have not been observed between the anteromedial cortex and the internal capsule, medial substantia nigra, superior colliculus, rostromedial tegmentum, or medial pons, which suggests that these sites are not connected by axons mediating turning. Possible synaptic linkages between the 5 bundles and possible transmitters are discussed.

Animals

Circling elicited from the anteromedial cortex and medial pons: refractory periods and summation.

Contraversive circling is evoked by stimulating the anteromedial cortex (AMC) of rats, and ipsiversive circling is evoked by stimulating the medial pons (PONS). During AMC circling, lateral and vertical head movements and vibrissae movements were exhibited. During PONS circling, although lateral head movements were exhibited, vertical head movements and vibrissae movements were not exhibited. Refractory periods were estimated by delivering trains of paired pulses and measuring the frequency thresholds for circling at various intrapair intervals. Refractory periods at AMC circling sites were much longer (range 1.4-3.3 ms) than at PONS circling sites (range 0.5-1.0 ms). To determine the degree of summation between the AMC and contralateral PONS, the two sites were stimulated concurrently. Summation of 95-100% was observed for AMC and PONS circling. No collision was observed at short intrapair intervals of paired pulses. Thus, the AMC and PONS are not connected axonally but are related, perhaps serially, for the production of circling.

Animals

Two converging brainstem pathways mediating circling behavior.

Ipsiversive circling results from stimulation of the rostromedial tegmentum (RMT) or medial pons (PONS), and contraversive circling results from stimulation of the superior colliculus (SC). To determine whether these sites are functionally connected, the collision method of Shizgal, Bielajew, Corbett, Skelton and Yeomans (1980) was used in rats. Pairs of stimulation pulses were presented to two sites, and the degree of collision between stimulation-evoked action potentials was assessed by measuring the frequency required for circling at short and long intrapair conditioning-testing (C-T) intervals. Collision was evidenced when the required frequencies were higher at short C-T intervals than at long C-T intervals. Collision of 46-62% was observed between RMT and PONS, and collision of 15-29% was observed between SC and PONS. Sites from which collision was obtained were located along the trajectories of the medial tegmental tract and the crossed tectospinal pathway. Refractory periods in all sites were similar, ranging from 0.3 to 1.7 ms. Conduction velocities of axons connecting RMT and PONS and SC and PONS were comparable, ranging from 0.8 to 13.3 m/s and 1.7 to 13.8 m/s, respectively, with lower conduction velocities associated with more ventral pontine sites. Thus, RMT and PONS, and SC and PONS are connected by myelinated axons that mediate circling.

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