Biomedical subjects
Publications and source records attributed to B L Finlay.
During normal development of the hamster eye, there is a substantial loss of cells from the retinal ganglion cell layer in the first two postnatal weeks. If one eye is lost at birth, this cell death is reduced in the remaining eye. This may account for the increased ipsilateral projection from this eye to the thalamus and midbrain observed in these animals.
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The great diversity of the niches inhabited by rodents, and the variety of the visual demands of these niches, present an excellent prospect for a comprehensive neuroethological analysis of rodent visuomotor behavior. To this end, rodent taxonomy is reviewed, with special attention to the multiple independent invasions of arboreal, terrestrial, fossorial and aquatic niches by distantly related rodent species. Current work on rat, gerbil and hamster is reviewed with emphasis on visual contributions to naturalistic behaviors such as exploration, foraging, predator detection and conspecific recognition.
The contributions of the midbrain optic tectum to visuomotor behaviors likely to be important to hamsters in the wild were studied, including aperture detection, insect catching, and barrier avoidance. Following tectal undercuts, hamsters ceased to make direct approaches to apertures in the posterior 180 degrees of the visual field; this appeared to be mediated by a loss of exploratory or scanning head movements. Reorientation to and pursuit of crickets jumping out of grasp into the visual periphery was impaired, though initial approach to them was not. Barrier avoidance was unaffected by tectal undercuts. This pattern is similar to the contribution of the frog and toad optic tectum to analogous visuomotor tasks. The contribution of the tectum to searching and scanning in the hamster is an extension of the basic orienting capabilities dependent on optic tectum in anurans.
After neonatal damage to the midbrain tectum, hamsters exhibit as adults substantially reorganized retinal projections to the thalamus and midbrain. These experiments investigated the pattern of acquisition of a range of visuomotor capacities in hamsters with neonatal tectal damage, and the role of visual experience in this acquisition, by examining the time course of visuomotor development and the effects of dark-rearing on visuomotor behavior. These hamsters acquired visual orientation and pursuit at the same age as normal hamsters, but orientation and pursuit were impaired and exhibited no improvement over time. Photophobia and edge and depth sensitivity were relatively unaffected. Dark-rearing disrupted all visuomotor behavior in the midbrain-damage animals but had no effect on normal animals. These results are consistent with a dominant role of neocortex in the residual visuomotor capacities of animals with neonatal ablations of the midbrain tectum.
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Retinotectal topography, response properties of neurons in superior colliculus, and visual orienting behavior were studied in hamsters whose superior colliculi were innervated by one or the other of two types of anomalous ipsilateral projections. For the first type, an abnormally large uncrossed projection was created by monocular enucleation on the day of birth. This projection extended over the superficial part of the rostral half of the colliculus. The upper visual field was represented medially, and the lower visual field laterally, which corresponds to a normal projection. The rostrocaudal axis was disordered, but showed a slight tendency for nasal visual field to be represented rostrally and temporal field caudally; this tendency corresponds to an inversion of the normal ipsilateral projection, fitting instead the pattern of a contralateral projection. For the second type of anomalous ipsilateral projection, an abnormal intertectal decussation of optic tract fibers was created by neonatal ablation of the superficial layers of one superior colliculus and removal of the ipsilateral eye (Schneider, '73). Retinotectal topography observed in this recrossing projection was predominantly mirror-symmetric to the normal contralateral projection; however, some distortions in retinotopic order were observed, including misplaced fields and local inversions of the mirror-symmetric topography, and distortions of local magnification factor. Response properties of single units found medially in the left colliculus were similar to those found in normal colliculus. Units found more laterally were underresponsive, showing response decrements with repeated stimulation which is abnormal for units in the superficial gray, and many had abnormally large receptive fields. This physiological pattern was reflected in the pattern of errors made in visual orienting to small targets. It was concluded that polarity cues exist in the tectum sufficient to order the terminals of the retinotectal projection independent of the direction of fiber arrival or order in the optic tract as it enters the tectum. In addition, the functional competence of the abnormal recrossing retinotectal projection has been demonstrated by both electrophysiological and behavioral methods.
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The topography of visual and somatosensory projections to the superior colliculus in the Syrian hamster was studied using electrophysiological techniques. The visual projection to the superficial layers of the colliculus is similar in general topography to that described for other rodents. The magnification of the visual field on the colliculus surface was greatest for nasal visual field. The magnification factor paralleled retinal ganglion cell density for corresponding visual field sectors. In the deep layers of the colliculus, a somatosensory projection is found in register with the visual projection such that the anterior somatosensory field and nasalmost visual field are both represented in rostral colliculus; posterior somatosensory fields and temporal visual fields are found in caudal colliculus. Likewise, upper visual and somatosensory fields are found in medial colliculus, and lower visual and somatosensory fields are found in lateral colliculus. Large receptive fields make the somatosensory topography less precise than the visual topography, but this lack of precision could serve to keep the two maps generally in register during eye and body movements.
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1. The properties of single cells in striate cortex of the rhesus monkey, representing the visual field 2 degrees -5 degrees from the fovea, were examined quantitatively with stationary and moving stimuli. Three distinct classes of cells were identified: S type, CX type, and T type. 2. S-type cells were defined as those oriented cells which to the optimal direction of movement in their receptive fields exhibited one or more spatially separate subfields within each of which a response was obtained to either a light or dark edge, but not to both. Several different types of S-cells were distinguished: a) S1-type cells for which moving edges revealed a single excitatory area within which a response was elicited by either a light or a dark edge but not by both. Most of these cells were unidirectional. b) S2-type cells for which moving edges revealed two spatially separate response areas, one of which was excited by a light edge and the other by a dark edge. Both regions responded to the same direction of movement. c) S3-type cells which had two response areas, one of which was excited by a stimulus moving in one direction (at right angles to the axis of orientation) and the other, of opposite contrast, which responded in the opposite direction, d) S4-type cells which to one direction of movement showed two spatially separate regions sensitive to a light and dark edge and which in the other direction of movement had only one responsive area (either light or dark). e) Cells which had multiple spatially separate subfields (S5-7 types). 3. CX-type cells were defined as those oriented cells which in their receptive fields exhibited no spatial separation for light- and dark-edge responses; they discharged to both edges in the same direction of movement and in the same spatial area. Flashing stimuli elicited both on and off responses throughout the receptive field. CX-type cells were predominantly of two types: those which were selective for direction of stimulus movement and those which were not. 4. A third class of cells (T-type) were those which were excited by only one sign of contrast change and responded in a sustained fashion even when there was no contour within the receptive field. These cells were poorly or not at all oriented; some of them were selective to wavelength. 5. Quantitative comparisons showed the following differences between S-type and CX-type cells: a) S-type cells had smaller receptive fields than CX-type cells but the populations over-lapped considerably. Receptive-field size was smallest in layer 4c. In all other layers S-type cells had the same size fields. CX-type cells, by contrast, tended to have larger fields in layer 5-6 than 2-3. b) The spatial separation between light and dark response areas was the best criterion for distinguishing S-type and CX-type cells. The distribution of this measure disclosed two populations of cells with relatively limited overlap. c) In layers 2 and 3, both S-type and CX-type cells had low spontaneous activity...
1. Quantitative analyses of orientation specificity and ocular dominance were carried out in striate cortex of the rhesus monkey. 2. Sharpness of orientation selectivity was greater for simple (S type) than for complex (CX type) cells. CX-type cells became more broadly tuned in the deeper cortical layers: S-type cells were equally well tuned throughout the cortex. 3. Sharpness of orientation selectivity for S-type cells was similar at all retinal eccentricities studied (0 degrees - 20 degrees from the fovea):in CX-type cells orientation selectivity decreased slightly with increasing eccentricity. 4. The orientation tuning of binocular cells was similar when mapped separately through each eye. 5. Orientation selectivity and direction selectivity are independent of each other, suggesting that separate neural mechanisms give rise to them. 6. More CX-type cells can be binocularly activated than S-type cells (88% versus 49%). The ocular dominance of S-type cells is similar in all cortical layers: for CX-type cells there is an increase in the number of cells in ocular-dominance category 4 in layers 5 and 6.
1. The response properties of single cells in monkey striate cortex were examined using moving bars, square-wave gratings, and sine-wave gratings. 2. The moving of cells studied were not selective for bar width or for the spatial frequency of square-wave gratings. 3. Most cells responded selectively to the spatial frequency of the sine-wave gratings. 4. The spatial frequency of the sine-wave grating eliciting the optimal response could not be predicted from the organization of the receptive field of each cell as determined by stationary or moving stimuli. 5. The sharpness of spatial-frequency selectivity is only slightly more pronounced in S-type cells than in CX-type cells. 6. S-type and CX-type cells differ significantly in the temporal modulation of their discharges to gratings. S-type cells discharge in sharp bursts to each cycle which traverses the receptive field. CX-type cells discharge in a rather continuous fashion. This measure can be used reliably to classify cells as S or CS type.
1. The receptive-field properties of corticotectal cells in the monkey's striate cortex were studied using stationary and moving stimuli. These cells were identified by antidromic activation from the superior colliculus. 2. Corticotectal cells form a relatively homogeneous group. They are found primarily in layers 5 and 6. These cells can usually be classified as CX-type cells but show broader orientation tuning, larger receptive fields, higher spontaneous activity, and greater binocular activation than CX-type cells do in general. A third of the corticotectal cells were direction selective. 3. These results suggest that the cortical input to the superior colliculus is not directly responsible for the receptive-field properties of collicular cells. We propose that this input has a gating function in contributing to the control of the downflow of excitation from the superficial to the deep layers of the colliculus.
1. Several statistical analyses were performed on 205 S-type and CX-type cells which had been completely analyzed on 12 response variables: orientation tuning, end stopping, spontaneous activity, response variability, direction selectivity, contrast selectivity for flashed or moving stimuli, selectivity for interaction of contrast and direction of stimulus movement, spatial-frequency selectivity, spatial separation of subfields responding to light increment of light decrement, sustained/transient response to flash, receptive-field size, and ocular dominance. 2. Correlation of these variables showed that within any cell group, these response variables vary independently. 3. A multivariate discriminant analysis showed that orientation specificity, receptive-field size, interaction of direction and contrast specificity ocular dominance, and spontaneous activity, taken together can adequately assign cells into the S-type or CX-type subgroups. 4. Various models of visual cortex are examined in view of the findings reported here and in the previous papers of this series, which suggest that a) orientation and direction selectivities are produced by separate neural mechanisms, b) there may be hierarchy among simple (S type) cells, and c) complex (CX-type) cells appear to receive a prominent S-type cell input.
The effects of changing retinal ganglion cell (RGC) density and availability of presynaptic sites on the development of RGC dendritic arbor in the developing chick retina were contrasted. Visual form deprivation was used to induce ocular enlargement and expanded retinal area resulting in a 20-30% decrease in RGC density. In these retinas, RGC dendritic arbors increased in a compensatory manner to maintain the inner nuclear layer to RGC convergence ratio in a way that is consistent with simple stretching; RGC dendritic arbors become larger with increased branch lengths, but without change in the total number of branches. In the second manipulation, partial optic nerve section was used to produce areas of RGC depletion of approximately 60% in the central retina. This reduction in density is comparable to the density of locations in the normal peripheral retina. In RGC depleted retinas, dendritic arbor areas of RGCs in the central retina grow to match the size of normal peripheral arbors. In contrast to the expanded case, two measures of intrinsic arbor structure are changed in RGC-depleted retinas; the branch density of RGC dendrites is greater, and the relative areas of the two arbors of bistratified cells are altered. We discuss the potential roles of retinal growth, local RGC density, and availability of presynaptic terminals in the developmental control of RGC dendritic arbor.
The diversity of neocortical cytoarchitecture could arise from genetic prespecification of cell types and numbers in the ventricular zone, by interaction of cells with their immediate environment, efferent targets, or afferent inputs. Here, we examine the role of the thalamus as efferent target or afferent source in the early control of cell number and type in the developing neocortex. Electrolytic lesions of the thalamus were made in hamsters at birth prior to the thalamic innervation of layer IV. By postnatal day 7, when migration to the cortex is complete, there were no differences in cell number between the cortical plate contralateral and ipsilateral to the thalamic lesion, showing that the absence of thalamic input does not influence the last phases of neocortical cell generation or migration. However, the incidence of pyknotic cells was elevated in the upper half of the cortical plate at this time. By adulthood, the number of cells per unit column of cortex was reduced, due to the apparent absence of small, nonpyramidal cells of layer IV, as determined from Nissl-stained material. Therefore, some of the cytoarchitectonic variability of the neocortex could arise epigenetically by the interaction of neocortical cells with their afferent connections.