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P D Gamlin

Publications and source records attributed to P D Gamlin.

29 records · Page 2Linked to original sources

Antidromic identification of midbrain near response cells projecting to the oculomotor nucleus.

Medial rectus motoneurons carry both conjugate and vergence eye position signals. Abducens internuclear neurons, whose axons travel in the medial longitudinal fasciculus, provide these motoneurons with the major signal for conjugate eye movements but not for vergence eye movements. A vergence signal appropriate for these motoneurons is seen on the near response cells that are found in the mesencephalic reticular formation within 2 mm of the oculomotor nucleus. The goal of the present study was to determine if midbrain near response cells project to the medial rectus subdivision of the oculomotor nucleus. Near response cells were recorded in two trained rhesus monkeys with ocular search coils. A stimulating electrode was positioned within the medial rectus subdivision of the oculomotor nucleus. Twenty-eight near response cells were found that could be driven by single pulse microstimulation of the ipsilateral medial rectus subdivision. In all cases, antidromic activation was confirmed by collision testing. Attempts to antidromically activate midbrain near response cells from the contralateral medial rectus subdivision were unsuccessful. Most antidromically activated cells had a steady state firing rate proportional to vergence angle. One cell also showed burst activity during the vergence eye movements. Divergence cells were not antidromically activated.

Animals↗

Trochlear unit activity during ocular convergence.

1. Ocular convergence is usually accompanied by excyclotorsion of the eyes. Furthermore, the magnitude of cyclotorsion is dependent on the elevation of the eyes. The reason for this excyclotorsion during convergence is not understood. 2. Excyclotorsion could be produced by either increased activity in the inferior oblique muscle or decreased activity in the superior oblique muscle. An earlier study indicated that convergence may also be accompanied by a temporal (lateral) translation of the eye. This observation is more consistent with a relaxation of the superior oblique than contraction of the inferior oblique. 3. This hypothesis was tested by recording the activity of 31 neurons in the trochlear nucleus, which contains the superior oblique motoneurons. This was done in alert monkeys that were trained to make both versional and vergence eye movements. In addition, the cyclotorsion associated with convergence was measured in one of these monkeys. 4. A consistent excyclotorsion associated with convergence was observed. Trochlear unit activity decreased during convergence in all cells tested. The magnitude of this decrease was significantly greater than that seen with conjugate adduction. Furthermore, the size of the decrease varied systematically with ocular elevation in a manner that was consistent with earlier measures of cyclotorsion during convergence. 5. These results suggest that the excyclotorsion seen during convergence, and perhaps the lateral translation of the eye, are due to a relaxation of the superior oblique muscle. This relaxation during convergence is greater than that which accompanies similar conjugate movements of the eyes. We hypothesize that this peculiar pattern of muscle innervation has a motor rather than sensory function.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Abducens internuclear neurons carry an inappropriate signal for ocular convergence.

1. Single-unit recording studies in alert Rhesus monkeys characterized the vergence signal carried by abducens internuclear neurons. These cells were identified by antidromic activation and the collision of spontaneous with antidromic action potentials. The behavior of abducens internuclear neurons during vergence was compared with that of horizontal burst-tonic fibers in the medial longitudinal fasciculus (MLF) and to that of a large sample of unidentified abducens cells (presumably both motoneurons and internuclear neurons). 2. The results indicate that abducens internuclear neurons and lateral rectus motoneurons behave similarly during vergence eye movements: the majority of both groups of cells decrease their firing rate for convergence eye movements: a minority show no change for vergence. This finding is strongly supported by recordings of horizontal burst-tonic fibers in the MLF, the majority of which decrease their activity significantly for convergence eye movements. 3. These findings indicate that a net inappropriate vergence signal is sent to medial rectus motoneurons via the abducens internuclear pathway. Because medial rectus motoneurons increase their activity appropriately during symmetrical convergence, this inappropriate MLF signal must be overcome by a more potent direct vergence input. 4. Overall, both abducens internuclear neurons and lateral rectus motoneurons decrease their activity for convergence less than would be expected based on their conjugate gain. This implies that some degree of co-contraction of the lateral and medial rectus muscles occurs during convergence eye movements. 5. Some horizontal burst-tonic MLF fibers decrease their activity more for convergence than any recorded abducens neuron. These fibers may arise from cells in the nucleus prepositus hypoglossi or vestibular nuclei.

Abducens Nerve↗

Lidocaine-induced unilateral internuclear ophthalmoplegia: effects on convergence and conjugate eye movements.

1. To characterize the vergence signal carried by the medial longitudinal fasciculus (MLF), it was subjected to reversible blockade by small injections of 10% lidocaine hydrochloride. The effects of these blockades on both conjugate and vergence eye movements were studied. 2. With this procedure, experimentally induced internuclear ophthalmoplegia (INO) and its effects on conjugate eye movements could be studied acutely, without possible contamination from long-term oculomotor adaptation. In the eye contralateral to the MLF blockade, saccadic and horizontal smooth-pursuit eye movements were normal. Horizontal abducting nystagmus, often seen in patients with INO, was not observed in this eye. 3. As previously reported for INO, profound oculomotor deficits were seen in the eye ipsilateral to the MLF blockade. During maximal blockade, adducting saccades and horizontal smooth-pursuit movements in this eye did not cross the midline. Adducting saccades were reduced in amplitude and peak velocity and showed significantly increased durations. Abducting saccades, which were slightly hypometric, displayed a marked postsaccadic centripetal drift. 4. The eye ipsilateral to the blockade displayed a pronounced, upward, slow drift, whereas the eye contralateral to the blockade showed virtually no drift. Furthermore, although vertical saccades to visual targets remained essentially conjugate, the size of the resetting quick phases in each eye was related to the amplitude of the slow phase movement in that eye. Thus the eye on the affected side displayed large quick phases, whereas the eye on the unaffected side showed only slight movements. On occasion, unilateral downbeating nystagmus was seen. This strongly suggests that the vertical saccade generators for the two eyes can act independently. 5. The effect of MLF blockade on the vergence gain of the eye on the affected side was investigated. As a measure of open-loop vergence gain, the relationship of accommodative convergence to accommodation (AC/A) was measured before, during, and after reversible lidocaine block of the MLF. After taking conjugate deficits into account, the net vergence signal to the eye ipsilateral to the injection was found to increase significantly during the reversible blockade. 6. The most parsimonious explanation for this increased vergence signal is suggested by the accompanying single-unit study. This study showed that abducens internuclear neurons, whose axons course in the MLF, provide medial rectus motoneurons with an appropriate horizontal conjugate eye position signal but an inappropriate vergence signal. Ordinarily, this incorrect vergence signal is overcome by another, more potent, v

Animals↗

Retinal projections to the pretectum in the pigeon (Columba livia).

The retinal projection to the pretectum in the pigeon has previously been described in detail only by means of anterograde degeneration techniques (Repérant, '73). The present study reinvestigated these retinal projections by using the more sensitive anterograde autoradiographic technique. In general, our results confirm and extend those of Repérant ('73). We have found that three pretectal nuclei--the nucleus lentiformis mesencephali, tectal gray, and the area pretectalis--received heavy retinal input. A fourth pretectal nucleus, pretectalis diffusus, receives a slight retinal input. The nucleus lentiformis mesencephali can be divided into two closely apposed subnuclei that are cytoarchitecturally similar. We have termed them "lentiformis mesencephali, pars medialis" and "lentiformis mesencephali, pars lateralis." The tectal gray can be divided into a rostral, retinorecipient region and a narrow, caudal, nonretinorecipient region. The cytoarchitecture and retinal terminal field in area pretectalis have been described previously by us (Gamlin et al., '84). Pretectalis diffusus is located caudal to the retinorecipient dorsal thalamus and rostral to area pretectalis. Localized retinal injections of 3H proline delineated the number and extent of the retinal representations in the pretectum. Separate retinal representations were present in lentiformis mesencephali, pars medialis, lentiformis mesencephali, pars lateralis, the tectal gray, area pretectalis, and pretectalis diffusus. Only in the lentiformis mesencephali, pars medialis, lentiformis mesencephali, pars lateralis, and the tectal gray could the retinal representations be analyzed. Whereas the retinal representations in the lentiformis mesencephali, pars medialis and the tectal gray are comparable, the retinal representation in the lentiformis mesencephali, pars lateralis is different, being a mirror-image mediolaterally. In this study we introduce a conservative nomenclature for the retino-recipient pretectal nuclei that is consistent with earlier studies, in particular, those of Kuhlenbeck ('39), but has been modified in the light of our findings. We believe that this nomenclature, combined with the detailed cytoarchitectural descriptions provided, should facilitate future studies of the avian pretectum.

Animals↗

Projections of the retinorecipient pretectal nuclei in the pigeon (Columba livia).

We have used anterograde autoradiographic and retrograde HRP techniques to investigate the efferent connections of the retinorecipient pretectal nuclei in the pigeon. In the accompanying paper we identified these nuclei in the pigeon as the nucleus lentiformis mesencephali--pars lateralis and pars medialis, the tectal gray, the area pretectalis, and pretectalis diffusus. Although there are reports of a few of the projections of these nuclei, they had not previously been the subject of a detailed study. We found that different cell types in the lentiformis mesencephali, pars medialis and the lentiformis mesencephali, pars lateralis have descending projections to different targets. These targets include the inferior olive, the cerebellum, the lateral pontine nucleus, the nucleus papillioformis, the nucleus of the basal optic root, the nucleus mesencephalicus profundus, pars ventralis, the nucleus principalis precommissuralis, and the stratum cellulare externum. We found that a few cells in the lentiformis mesencephali project to the medial pontine nucleus, but that a much heavier projection arises from the nucleus laminaris precommissuralis, which is medial to the nucleus lentiformis mesencephali, pars medialis. The tectal gray has predominantly ascending projections to the diencephalon. The nuclei that it projects to are the nucleus intercalatus thalami, the nucleus of the ventral supraoptic decussation, the nucleus posteroventralis, the ventral lateral geniculate nucleus, the nucleus dorsolateralis medialis, and the nucleus dorsolateralis anterior. The tectal gray also projects topographically to layers 4 and 8-13 of the optic tectum. Area pretectalis has both ascending and descending projections. It has ipsilateral ascending projections to the nucleus dorsolateralis anterior, pars magnocellularis, the nucleus lateralis anterior, and the nucleus ventrolateralis thalami. It has ipsilateral descending projections to the central gray, the nucleus of the basal optic root, pars dorsalis, the lateral pontine nucleus, and the deep layers of the optic tectum. It has contralateral projections to the area pretectalis, the nucleus Campi Foreli, the interstitial nucleus of Cajal, the nucleus of Darkschewitsch, the cerebellum, and the Edinger-Westphal nucleus. The efferent projections of pretectalis diffusus are limited. It projects contralaterally to the pretectalis diffusus, and ipsilaterally to the nucleus of the ventral supraoptic decussation, the lateral pons, and the cerebellum.4

Animals↗

A second ascending visual pathway from the optic tectum to the telencephalon in the pigeon (Columba livia).

Previous studies in the pigeon (Karten and Revzin: Brain Res. 2:368-377, '66; Karten and Hodos: J. Comp. Neurol. 140:35-52, '70) have described an ascending tectofugal visual pathway from the optic tectum to the ectostriatum by way of the nucleus rotundus of the thalamus. This present study used anterograde autoradiographic and retrograde horseradish peroxidase pathway-tracing techniques to investigate another ascending tectofugal pathway in the pigeon. Injections of 3H-proline/leucine confirmed a previous report that the optic tectum projects to the nucleus dorsolateralis posterior of the thalamus (DLP). This projection is predominantly ipsilateral and is confined to a large-celled caudal region of the nucleus (DLPc); the rostral region of the nucleus (DLPr) is not tectorecipient. Injections of horseradish peroxidase in DLPc labeled cells predominantly ipsilaterally in layers 8-15 of the optic tectum. Injections of 3H-proline/leucine placed in the DLPc labeled a discrete region of the ipsilateral telencephalon. Similar injections of DLPr labeled a contiguous, but more rostral, region of the neostriatum intermedium. Nissl- and silver-stained material indicated that the region in which DLP terminates is cytoarchitecturally distinct from ventromedial ectostriatal core and belt. Injections of horseradish peroxidase at various locations in the neostriatal DLP terminal field demonstrated a rostrocaudal ordering of the DLP projection upon the neostriatum intermedium. Single-unit recording demonstrated that cells in DLPc respond to whole-field illumination at the same latency as cells in the nucleus rotundus, indicating that the tecto-DLPc-neostriatal pathway transmits visual information to the telencephalon. We suggest that comparable pathways may exist in both reptiles and mammals.

Animals↗

Neural control of vergence eye movements: neurons encoding vergence velocity.

Single-unit recordings were made from midbrain areas in monkeys trained to make both conjugate and disjunctive (vergence) eye movements. Previous work had identified cells with a firing rate proportional to the vergence angle, without regard to the direction of conjugate gaze. The present study describes the activity of neurons that burst for disjunctive eye movements. Convergence burst cells display a discrete burst of activity just before and during convergence eye movements. For most of these cells, the profile of the burst is correlated with instantaneous vergence velocity and the number of spikes in the burst is correlated with the size of the vergence movement. Some of these cells also have a tonic firing rate that is positively correlated with vergence angle (convergence burst-tonic cells). Divergence burst cells have similar properties, except that they fire for divergent and not convergent movements. Divergence burst cells are encountered far less often than convergence burst cells. Both convergence and divergence burst cells were found in an area of the mesencephalic reticular formation just dorsal and lateral to the oculomotor nucleus. Convergence burst cells were also recorded in another more dorsal mesencephalic region, rostral to the superior colliculus. Both of the areas also contain cells that encode vergence angle. Models of the vergence system derived from psychophysical data imply the existence of a vergence integrator, the output of which is vergence angle. Some models also suggest the presence of a parallel element that improves the frequency response of the vergence system, but has no effect on the steady-state behavior of the system. Vergence burst cells would be suitable inputs to a vergence integrator. By providing a vergence velocity signal to motoneurons, they may improve the dynamic response of the vergence system. The behavior of vergence burst cells during vergence movements is similar to that of the medium-lead burst cells during saccades. The proposed roles for vergence velocity cells are analogous to those of the saccadic burst cells. In this respect, the neural organization of the vergence system resembles that of the saccadic system, despite the distinct difference in the kinematics of these two types of eye movements.

Action Potentials↗

The neural substrate for the pupillary light reflex in the pigeon (Columba livia).

The neural substrate of the pupillary light reflex in the pigeon was investigated using anatomical, stimulation, and lesion techniques. In birds, as in mammals, the sphincter pupillae muscle (which constricts the iris) is innervated by cells in the ciliary ganglion (Pilar and Tuttle, '82). These cells are in turn innervated by cells in the Edinger-Westphal nucleus (EW) (Cowan and Wenger, '68; Narayanan and Narayanan, '76; Lyman and Mugnaini, '80). The efferent link of the pupillary light reflex must therefore involve cells in EW. To study the central course of this reflex pathway, injections of horseradish peroxidase (HRP) were placed in EW. These injections labeled cells in a number of regions including a contralateral pretectal nucleus, area pretectalis (AP). Only a limited number of cells in AP project to EW. Injections of tritiated amino acids into AP labeled a discrete region of the contralateral EW. This projection is confined to a dorsolateral region of caudal EW and overlies the somata of approximately 100 cells. Tritiated proline was injected into the eye, and the results confirmed an earlier report (Reperant, '73) that AP receives retinal input from the contralateral eye. Immunohistochemical studies demonstrated fibers in AP that stained positively for substance-P-like, enkephalin-like and tyrosine-hydroxylase-like immunoreactivity. Injections of HRP were placed in AP to examine the retinal ganglion cells mediating the reflex. Cells with an average diameter of approximately 14 microns (5-25 microns range) were labeled and averaged approximately 6 microns greater in diameter than the retinal ganglion cells (mean = 7.3 microns) labeled by an optic chiasm injection. The cells labeled by AP injections were distributed unevenly throughout the retina with a higher concentration in the central and temporal retina and a paucity in the red field and fovea. Our results demonstrate that AP receives input from a distinct subpopulation of large retinal ganglion cells that comprises a very small percentage of the total population of retinal ganglion cells. Unilateral lesions of AP abolished the pupillary light reflex in the eye contralateral to the lesion; stimulation of AP elicited pupilloconstriction in the eye contralateral to the stimulation site. These results delineate the central course of the pupillary light reflex pathway in the pigeon and identify the retinal ganglion cells that subserve this reflex. They show that, at every point in the pathway, only a few cells mediate this simple reflex.(ABSTRACT TRUNCATED AT 400 WORDS)

Afferent Pathways↗

Substance P-containing neurons of the avian suprachiasmatic nucleus project directly to the nucleus of Edinger-Westphal.

Retrograde and anterograde pathway tracing techniques were used in the pigeon to study afferent visual input from the suprachiasmatic nucleus (SCN) of the hypothalamus to the nucleus of Edinger-Westphal (EW), the parasympathetic visceral efferent component of the oculomotor complex. Horseradish peroxidase injected into the EW retrogradely labeled numerous neurons in the contralateral SCN, a retinorecipient hypothalamic nucleus, as well as a few neurons in the ipsilateral SCN. Autoradiographic orthograde pathway tracing experiments confirmed that the SCN projects heavily upon the medial subdivision of the contralateral EW and lightly upon the medial subdivision of the ipsilateral EW. Some neurons of the SCN contain substance P and a substance P-positive plexus of fibers was seen in precisely that medial portion of the EW to which the SCN was found to project in the autoradiographic experiments. This substance P-positive fiber plexus in the medial EW was eliminated by bilateral electrolytic lesions of the SCN. These results show that the avian suprachiasmatic nucleus has a heavy contralateral and a much lighter ipsilateral projection to the medial subdivision of the EW and that this projection may be largely substance P positive. Previous studies have suggested that neurons of the medial EW specifically project to the neurons of the ciliary ganglion that control the choriocapillary blood flow of the eye. Since the SCN has been implicated in the control of circadian rhythms in vertebrates, the projection of the SCN to the EW may represent a pathway by which a circadian rhythmicity is imposed on choriocapillary blood flow. Alternatively or in addition, the SCN-EW pathway described in this paper may provide the central neural substrate for a homeostatic regulatory mechanism by which choriocapillary blood flow is controlled by the intensity of retinal illumination.

Animals↗

Central neural circuits for the light-mediated reflexive control of choroidal blood flow in the pigeon eye: a laser Doppler study.

Electrical stimulation in pigeons of the input from the medial subdivision of the nucleus of Edinger-Westphal (EWM) to the choroidal neurons of the ipsilateral ciliary ganglion, which themselves have input to the choroidal blood vessels of the ipsilateral eye, increases choroidal blood flow (ChBF). Since the EWM receives input from the contralateral suprachiasmatic nucleus (SCN), which in turn receives contralateral retinal input, the present study sought to determine if activation of the SCN by microstimulation or by retinal illumination of the contralateral eye would also yield increases in ChBF in that same eye. Using laser Doppler flowmetry (LDF) to measure ChBF, we found that electrical activation of the contralateral SCN by 100-Hz anodal pulse trains yielded increases in ChBF that were stimulus related and proportional to the stimulating current. These increases in ChBF elicited by the SCN stimulation were accompanied by increases in choroidal volume (vasodilation), but not by increases in systemic blood pressure. Furthermore, the increases could be blocked reversibly by lidocaine injection into the EWM. These results suggest that the increases in ChBF in the eye contralateral to the SCN stimulation were specifically mediated by the SCN-EWM pathway. Retinal illumination with a fiber optic light source was also found to increase ChBF in the illuminated eye, and these effects too could be blocked reversibly with lidocaine injection into the EWM or permanently by the EWM lesion. Control studies confirmed that the light-elicited increases were mediated by increases in choroidal volume (i.e. vasodilation), were not accompanied by systemic blood pressure increases, and were not artifactually generated by transocular illumination of the LDF probe. Thus, the SCN-EWM circuit may be involved in regulating ChBF in response to the level of retinal illumination and/or the visual patterns falling on the retina.

Analysis of Variance↗