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R W Rhoades

Publications and source records attributed to R W Rhoades.

At least 181 records · Page 10Linked to original sources

Expansion of the ipsilateral visual corticotectal projection in hamsters subjected to partial lesions of the visual cortex during infancy: anatomical experiments.

Electrophysiological methods were employed to determine whether or not partial visual cortical lesions in neonatal (7--11-day) hamster produced large scotomas in the cortical visual representation. In cases where such scotomas were present electrophoretic deposits of radioactive amino acids in the visually responsive "cortical remnant" of the damaged hemisphere resulted in labelling throughout the lower portion of the stratum griseum superficiale and the stratum opticum of the ipsilateral superior colliculus. No differential labeling of the part of the colliculus which was topographically matched with the remaining visual representation in the cortical remnant was observed. In normal hamsters relatively localized, visual cortical deposits of radioactive amino acids resulted in superficial layer labeling only in portions of the colliculus which corresponded to the locus of the cortical deposit. In a similar fashion, small lesions at physiologically defined loci in the cortical remnant produced degeneration throughout most of the superficial tectal laminae, but a more restricted "focus" of denser degeneration was also visible in these cases. The position of this focus in the colliculus for a given cortical lesion varied with the nature of the visual map in the cortical remnant. In several additional neonatally brain-damaged hamsters large lesions of the visual cortex in the intact hemisphere were combined with radioactive amino acid deposits in the cortical remnant to determine whether or not axons from the crossed corticocollicular pathway previously demonstrated in such hamsters were intermingled with fibers from the ipsilateral corticotectal projection. In alternate sections processed for autoradiography or by the Fink-Heimer ('67) method autoradiographic label and degeneration argyrophilia were both observed in the medical part of the colliculus ipsilateral to the neonatal cortical lesion.

Aging↗

Expansion of the ipsilateral visual corticotectal projection in hamster subjected to partial lesions of the visual cortex during infancy: electrophysiological experiments.

Single unit recording from cells in the superior colliculus ipsilateral to the damaged hemisphere in hamsters subjected to unilateral removal of a part of the posterior neocortex during infancy was combined with electrical stimulation of the cortical remnant and the visual cortex in the undamaged hemisphere. Cells activated by stimulation of the cortical remnant were recorded in all portions of the colliculus. No differences in percentages of driven cells or threshold current intensities were noted between electrode penetrations in which collicular neurons having receptive fields within the remaining visual cortical representation were recorded and tracks where units with receptive fields outside this region were isolated. In the medical part of the tectum ipsilateral to the damaged hemisphere cells driven by stimulation of either cortex were encountered. It was also demonstrated that stimulation of the ipsilateral cortical remnant and/or the contralateral cortex was capable of suppressing discharges normally elicited by optic chiasm or visual stimulation in a manner qualitatively similar to that observed for collicular cells in normal hamsters. The response properties of cells functionally influenced by the ipsilateral and/or contralateral corticles were not different from those of neurons which received no demonstrable cortical input. The receptive field characteristics of the sample of neurons recorded were, on the whole, quite similar to those of collicular neurons in hamsters subjected to lesions of the visual cortex as adults.

Animals↗

Anatomical and electrophysiological demonstration of tectotectal pathway in the golden hamster.

Both the retrograde transport of horseradish peroxidase (HRP) and extracellular single unit recording have been employed to demonstrate the existence of an intertectal pathway in the golden hamster. The cells of origin of this projection are located primarily in the stratum griseum intermediate and stratum griseum profundum of the anterior one-half of the colliculus and respond, in most cases, only to somatosensory stimuli. These findings cast doubt on the recently proposed generalization [13] that this pathway may be rudimentary or even absent in animals with limited collicular binocularity or a small ipsilateral hemifield representation in the colliculus.

Animals↗

Cortical and spinal somatosensory input to the superior colliculus in the golden hamster: an anatomical and electrophysiological study.

The horseradish peroxidase technique was used to identify the sources of somatosensory afferent fibers to the hamster superior colliculus. These experiments demonstrated that the tectum receives axons from pyramidal cells in layer V of the ipsilateral sensorimotor cortex, contralateral lamina IV of all levels of the spinal cord, the contralateral dorsal column nuclei, lateral cervical nucleus, internal basilar nucleus, and nucleus of the spinal trigeminal tract. Electrical stimulation of the spinal cord coupled with extracellular single unit recordings concentrated, for the most part, in the posterior portion of the tectum revealed that such stimuli activated approximately 40% of the cells tested. Almost all of these units were isolated ventral to the stratum opticum and 86% were responsive only to somatosensory stimulation. Analysis of the latencies of collicular responses obtained with two point spinal stimulation in intact hamsters and in animals subjected to somatosensory cortical and/or spinal damage indicated that the initial impulse elicited from most collicular cells was mediated by a polysynaptic pathway(s) which probably synapses in the dorsal column, lateral cervical, and/or internal basilar nuclei. Damage to the dorsal spinal cord and/or somatosensory cortex altered neither the incidence nor the response characteristics of spinally driven collicular neurons. This indicated that most somatosensory collicular cells also received input from the spinotectal fibers which travel in the ventrolateral quadrant. Electrical stimulation of somatosensory cortex activated about 20% of the cells tested in the ipsilateral superior colliculus. If cortical and spinal stimulation were delivered with an interstimulus interval ranging between 50 and 80 msec the response of the tectal neuron to the latter stimulus was suppressed in most cases. This was true regardless of the order of the stimulus pairing. Concurrent somatosensory cortical shocks also suppressed responses to tactile stimuli for 21% of the cells tested.

Afferent Pathways↗

Organization of somatosensory input to the deep collicular laminae in hamster.

Retrograde transport of horseradish peroxidase (HRP) was used to delineate the sources of somatosensory input to the hamster's superior colliculus. Cells in the ipsilateral somatosensory cortex and contralateral dorsal horn of the spinal cord, dorsal column nuclei, lateral cervical nucleus, internal basilar nucleus, nucleus of the spinal trigeminal tract and deep layers of the superior colliculus were labeled following HRP injections centered in the deep tectal laminae. The response characteristics of somatosensory corticotectal, spinotectal and intertectal neurons were investigated with extracellular single unit recording methods and, with the exception of the fact that the receptive fields of corticotectal and spinotectal neurons were consistently smaller than those of cells recorded in the colliculus, the response characteristics of these neurons were quite similar to those of somatosensory neurons in the deep layers of the tectum. Lesions of the somatosensory cortex or dorsal half of the spinal cord were also combined with single unit recording in the colliculus to determine whether or not such damage altered the incidence and/or response characteristics of deep layer somatosensory cells. These lesions had no appreciable effect upon the functional organization of the deep tectal laminae. The implications of these results with regard to the convergence of visual and somatosensory information in the tectum are discussed.

Animals↗

Neonatal enucleation induces an asymmetric pattern of visual callosal connections in hamsters.

Autoradiography and the horseradish peroxidase (HRP) technique were used to investigate visual callosal connections in normal hamsters and in animals which had one eye removed at birth. In normals and in the cortex contralateral to the remaining eye in the neonatal enucleates, callosal cells and terminal labeling were restricted to the region around the area 17-18a border. In the cortex ipsilateral to the remaining eye in the neonatal enucleates labeled cells and terminal labeling were observed throughout areas 17 and 18a.

Animals↗

Compatibility of DAPl and silver staining for combined anterograde and retrograde tracing of neural connections.

The geniculocortical and corticogeniculate pathways in hamster were used to test the compatibility of 4'6 diamidino-2 phenylindole 2HCl (DAPl)6,10 and anterograde degeneration techniques for tracing reciprocal connections in the brain. The two methods were compatible within a single brain and, with some loss of sensitivity in the retrograde labeling, within a single section.

Animals↗

Response suppression induced by afferent stimulation in the superficial and deep layers of the hamster's superior colliculus.

Stimulation of the optic chiasm (OX) or visual cortex (VC) elicited a burst of impulses from visual cells in the superficial layers of the hamster's superior colliculus which was followed by a period of response suppression which lasted from 50-200 ms. During this period responses to normally suprathreshold OX and VC shocks, spontaneous activity and even injury discharges were markedly attenuated. For approximately 50% of the visual cells tested VC stimulation also reduced responses to visual stimuli. No correlations between receptive field properties and whether or not VC shocks diminished a given cell's visual responses were noted. Stimulation of either the cervical spinal cord (SC) or somatic sensory cortex (SMCTX) evoked action potentials from somatosensory neurons in the deep tectal laminae. These responses were followed by a period of suppression identical to that seen in the superficial layers after OX or VC shocks. SMCTX stimulation attenuated responses to tactile stimuli for 30% of the cells tested in the deep layers. Again, no correlation was observed between somatosensory response characteristics and whether or not a given cell exhibited response suppression.

Afferent Pathways↗

Effects of neonatal enucleation on the functional organization of the superior colliculus in the golden hamster.

1. The responses of visual, auditory and somatosensory superior collicular neurones were investigated using extracellular single unit recording techniques in hamsters which were subjected to the removal of one eye on the day of birth. 2. Neonatal enucleation resulted in a marked increase in the region of the colliculus from which visual neurones activated by stimulation of the ipsilateral eye could be recorded. In most cases the visuotopic representation in the colliculus ipsilateral to the remaining eye mirrored that observed in the contralateral tectum along both the rostrocaudal and mediolateral axes: in both colliculi temporal retina projected rostrally and inferior retina medially. In some animals, however, there appeared to be a dual mapping of the remaining eye onto the ipsilateral tectum. In these hamsters the central portion of the visual field was represented twice along the rostrocaudal axis of colliculus. 3. No changes in the topography of the somatosensory and auditory representations in the tectum were observed following neonatal enucleation. 4. The laminar distribution of visual neurones in the ipsilateral colliculus was markedly altered in the neonatally enucleated hamsters. Very few exclusively visual units were encountered in the layers ventral to the stratum opticum and almost all of the visual cells recorded in the ipsilateral colliculus were isolated within 150 microM of the tectal surface. 5. In the posterior half of the ipsilateral tectum a large number of extravisually responsive cells were encountered in the stratum griseum superficiale and stratum opticum. This was not the case in the colliculus contralateral to the remaining eye, nor has it ever been observed in normal hamsters. 6. Recordings from animals subjected to both neonatal enucleation and acute bilateral removal of somatosensory and auditory cortex indicated that the projections from these areas to the colliculus were not essential to the observed changes in laminar organization. 7. Recordings from normally reared hamsters which were subjected to removal of one eye at the time of the recording experiment suggested further that the isolation of extravisual cells in the superficial tectal aminae of the neonatal enucleates was probably not the result of the 'unmasking' of extravisual influences in the superficial layers which are present, but ineffective, in the normal case.

Action Potentials↗

Effects of neonatal enucleation on receptive-field properties of visual neurons in superior colliculus of the golden hamster.

1. Monocular enucleation in infant hamsters results in a marked expansion of the normally very limited ipsilateral retinotectal projection (13). In 34 hamsters subjected to removal of one eye within 12 h of birth, the receptive-field characteristics of superior collicular neurons ipsilateral and contralateral to the remaining eye were investigated quantitatively and compared to those of normal animals. In six additional neonatal enucleates, the density of the expanded retinotectal projection was studied with the autoradiographic method and an attempt was made to relate the anatomical reorganization with the electrophysiological findings, 2. The response characteristics of visual cells in the colliculus contralateral to the remaining eye were not significantly different from those observed in normal animals. In the ipsilateral tectum, however, numerous changes were observed. Visual receptive fields were abnormally large. The incidence of directional selectivity was markedly reduced, as were the magnitudes of the discharges elicited by either flashed or moving stimuli. Fewer cells were activated by small flashed spots and most of the units that were responsive to such stimulation failed to exhibit the surround suppression typical for the majority of tectal neurons in normal hamsters. Most cells in the ipsilateral colliculus responded only to relatively low (less than 50 degrees/s) stimulus velocities and response decrements resulting from repeated stimulation also occurred much more readily for the neurons tested on this side. 3. The results of additional experiments in neonatal enucleates (n = 8), which were also subjected to acute bilateral removal of the visual cortex, demonstrated that such damage resulted in a marked reduction in the incidence of directional selectivity in the colliculus contralateral to the remaining eye but had no effect on the responses of cells innervated by the aberrant ipsilateral pathway. 4. A correlation between the relative density of the ipsilateral retinal projection at different points in the colliculus, as demonstrated by the autroradiography and the nature of the visual responses obtained in different portions of the structure, indicated that receptive-field size was negatively correlated with the density of the aberrant retinotectal projection and that absolute responsivity (number of impulses elicited by an optimal stimulus) was positively correlated with autoradiographic grain density. 5. These findings demonstrate that while the aberrant retinocollicular projection can, along with the other visual inputs to the tectum, result in the organization of normal response properties for a small number of tectal neurons, the majority of the visual cells innervated by this pathway have responses that are appreciably different from normal.

Animals↗

An autoradiographic study of the retinotectal projection in the golden hamster.

Intraocular injections of tritiated leucine and proline were used to examine the retinotectal projection of the golden hamster. In the contralateral superior colliculus intense and complete label was seen in the stratum zonale, stratum griseum superficiale and the upper portion of the stratum opticum, with relatively less dense label in the lower part of the optic layer. On the ipsilateral side no label was found in the most rostral portion of the tectum. This area comprised about 10% of the rostro-caudal extent of the colliculus, and most likely, it receives a crossed input from the temporal retina (as demonstrated in the cat by Harting and Guillery, '76). Very sparse label was observed in an anterior segment of the ipsilateral colliculus. In coronal sections it appeared as discrete clumps or patches which were confined to the stratum opticum. Within this layer there was a tendency for the clumps to be located more dorsally with increasing laterality. There was considerable variability between and within animals in the size of the clumps as well as the distance between clumps. Reconstruction of coronal sections showed that the ipsilateral label forms discontinuous ribbons which extend up to 180 microns in the rostro-caudal dimension. No label was seen on the ipsilateral side in the remaining tectum (caudal 60%).

Animals↗

An electronmicroscopic analysis of the optic nerve in the golden hamster.

The electronmicroscopic examination of sections taken from the hamster's optic nerve 5 mm behind the globe indicated that the nerve contains 110,165 +/- 4,177 (p less than 0.05) fibres of which 96.4% are myelinated. The fibre diameter distribution is unimodal with a peak at 1.2 micrometer and axon diameters ranging from 0.20 micrometer to 3.93 micrometer. Fibres of all sizes are distributed uniformly throughout the cross section of the nerve. The thickness of the myelin sheath surrounding a given axon is highly (0.80) correlated with axonal diameter and the degree of myelination for a fibre of a given size is nearly constant throughout the nerve's cross section. In nerve sections taken just posterior to the globe most (64%) of the fibres counted are unmyelinated and the percentage of unmyelinated axons is highest near the peripheral boundary of the nerve. The process of myelination is essentially complete in sections taken 3.5 mm behind the eye. These differences in the myelination of the proximal and distal nerve most probably account for the discrepancy between the results reported here and those provided by a previous study (Tiao and Blakemore, '76) concerned with the structure of the optic nerve in this species.

Animals↗

Conduction velocity distribution of the retinal input to the hamster's superior colliculus and a correlation with receptive field characteristics.

Cells driven reliably by shocks delivered to the optic nerve or optic chiasm were encountered throughout the depth of the colliculus. The incidence of such cells, however, decreased markedly in the laminae ventral to the stratum opticum. The distribution of conduction velocities for the retinal afferents to the tectum was quite broad (range: 1.7-25.5 m/sec) and clearly biomodal with peaks at about 6 and 12 m/sec. A small number of cells were innervated by rapidly (greater than 15 m/sec) conducting axons. No evidence of an indirect-fast pathway from the retina to the colliculus via the lateral geniculate nucleus and visual cortex was obtained. Afferent conduction velocity was not correlated with retinal eccentricity, collicular depth or speed selectivity. It was, however, clearly related to directional selectivity. Ninety percent of the tectal neurons receiving inputs from axons having conduction velocities of less than 5 m/sec were directionally selective while only 41% of those neurons innervated by more rapidly conducting fibers (greater than 5 m/sec) exhibited selectivity. One hundred and sixteen cells in the anterior portion of the colliculus were tested with shocks delivered to the ipsilateral optic nerve and photic stimulation of the ipsilateral eye. Of these, 11% exhibited some degree of binocularity and only 6% were responsive to optic nerve shocks. These electrophysiological findings were correalted with the limited nature of the retinal input to the ipsilateral superior colliculus.

Animals↗