14C-Deoxyglucose mapping of orientation subunits in the cats visual cortical areas

Summary The orientation domain in the cortical visual areas of anesthetized cats has been investigated by employing the ¹⁴C-Deoxyglucose technique (Sokoloff et al., 1977).Orientation subunits (OS) are seen in the first (V1), the second (V2) and the third visual area (V3) as well as in the visual areas of the suprasylvian sulcus. In the latter regions OS are less elaborated than in V1, V2, and V3. The OS are continuous through all cortical layers; in V1 however, only weak label is detected in layer 4C. In V1, V2, and V3 the width of the OS is about 0.4 mm and the average distance between two OS centers is 0.9 mm. The spatial pattern of the OS seems to be more regular in the visual field periphery than in regions representing the vertical meridian.     

A developmental study of retinal afferents and visual responses in the cat pretectum

Summary Neuronal responses in the pretectum (PT) were analyzed in 4–16 week old kittens after visual and electrical stimulation and compared with adult responses from a previous study. All three retinal fiber types projecting to the adult PT could be electrically activated in kittens from 4 weeks on. There was a dramatic reduction of response latencies to electric shocks to retinal afferents applied at the optic chiasm (OX) and optic tract (OT) in postsynaptic cells as a function of age, involving X-, Y-, and Wfibers. At four through six weeks postnatally the reduction in latency was found to be due to enhanced signal transmission at the axonal terminal region. Latency reduction continued after six weeks of life due to sharp increases in conduction velocity of the afferent fibers. Different steps in the maturation of visual response specifity were found for neurons of different functional types. Possible relationships are discussed between the development of neuronal responses of pretectal cells and the maturation of oculomotor behavior.     

Bilateral projections from the visual cortex to the striatum in the cat

Summary Direct projections from visual areas 17, 18, 19, and lateral suprasylvian visual area (LS) to the striatum were searched for in 12 adult cats using the autoradiographic technique to detect neuronal pathways. Striatal labels were found only after injections in areas 19 and LS. Projections homolateral to the injection sites were observed from both areas to the head and body of the caudate nucleus and to the putamen. Contralateral projections were found from both areas 19 and LS: however, area 19 did not project to the contralateral putamen. The extent of contralateral projections was smaller and they were confined within the same regions as the homolateral ones. Silver grains were often arranged in cluster-like patches, which were more evident ipsilaterally, in the head of the caudate nucleus and after injections in area LS.The present data support the view of a not strictly topographical segregation of striatal projections from the cat visual cortex.Supported by a grant from the CNR, Rome, Italy     

Failure to find luxotonic responses for single units in visual cortex of the rabbit

Summary The firing frequency of a population of 213 units in striate and circumstriate cortex of the moderately restrained rabbit was studied under the influence of alternating 1-min periods of darkness versus steady, diffuse, featureless illumination. The intent was to determine whether luxotonic responses, so prominent in striate cortex of primates, are indeed absent in rabbits. Such was the case, there being only transient occurrences in three units where the continuing rate of discharge in darkness was double that in the light. There were, however, much more modest differences in rate of continuing discharge in light versus darkness, and for 46% of the units discharging > 1/s this difference exceeded 10% and/or 1/s. The rate of discharge in any case did not provide a reliable index as to the characteristics of a unit's receptive field in response to patterned visual stimuli. The nature and function of luxotonic activity in primates still not being understood, it cannot be decided whether its absence in rabbits represents a true qualitative or merely a quantitative difference between species.Supported by Grant NS 03606 and Contract 70-2279 from the National Institute of Neurological and Communicative Disorders and Stroke, National Institutes of HealthDeceased November 5, 1978     

Topographic organization of orientation columns in the cat visual cortex

Summary Three-dimensional reconstructions of the orientation column system were obtained from the visual cortex of four cats using the deoxyglucose technique. One cat had normal visual experience, one was monocularly deprived and two had selective experience with vertical and horizontal contours, respectively. In areas 17 and 18 orientation columns form a remarkably regular system of equally spaced parallel bands whose trajectory is orthogonal to the borderline between areas 17 and 18. This topographic organization is resistant to manipulations of early visual experience.     

Identification of the projection from the visual cortex to the claustrum by anterograde axonal transport in the cat

Summary Visual cortex, including areas 17, 18, and sometimes 19, was injected with tritiated leucine. Terminal labelling could be detected by autoradiography in the dorsocaudal part of the ipsilateral claustrum in all cases.     

An “oblique effect” in the visual evoked potential of the cat

Summary An oblique effect was observed in the amplitude of the VEP recorded from area 17 of the cat. The ratio of the responses to oblique gratings compared with responses to horizontal and vertical gratings averaged 0.77. Orientation dependence was strongest at low spatial frequencies, unlike the effect found in primates.This work was supported by EY 01175 and EY 03260 from the National Eye Institute     

Retinotopic organization of extra-retinal saccade-related input to the visual cortex in the cat

Summary Single unit activity of 842 cells has been recorded in cat visual cortex and analyzed with respect to vestibular induced, and spontaneous saccadic eye movements performed in the dark. This study has been done in awake, chronically implanted cats, subsequently placed in acute conditions to achieve the precise retinotopic mapping of the cortical areas previously investigated.In areas 17 and 18, respectively, 27% and 24% of the cells tested were influenced by horizontal saccadic eye movements in the dark (E. M. cells). In the Clare-Bishop area, the proportion of E. M. cells was 12%, while only 2% of such cells were found in areas 19 and 21.The distribution of E.M. cells in areas 17 and 18 with respect to retinotopy showed that E.M. cells were more numerous in the cortical zones devoted to the representation of the area centralis (38% in area 17, 27% in area 18) than in the zones subserving the periphery of the visual field (17% and 12%, respectively).Two of the characteristics of E. M. cell activations appear dependant on the retinotopic organization. First, a larger number of E.M. cells presenting an asymmetry in their responses to horizontal saccadic eye movements in opposite directions (directional E.M. cells) were encountered in the cortical representation of the peripheral visual field. 53% of E. M. cells recorded in area 17 and 71% in area 18 were directional in the cortex corresponding to the peripheral visual field. This percentage was of 23% and 25% respectively in the cortex devoted to area centralis. Second, E.M. cells were found to have a latency from the onset of the saccade systematically larger than 100 ms (i.e, they discharged at, or after the end of the eye movement) if they were located in the cortical representation of the area centralis, while E.M. cells related to the peripheral visual field displayed a wider range of latencies (0–240 ms).Results obtained in Clare Bishop area, although limited to the representation of the peripheral visual field, were quantitatively and qualitatively similar to those observed in the homologous retinotopic zones of areas 17 and 18.It is concluded that an extra-retinal input related to oculomotor activity is sent to the cat visual cortex and is organized, at least in areas 17 and 18, with respect to the retinotopic representation of the visual field. These data support the hypothesis of a functional duality between central and peripheral vision and are discussed in the context of visual-oculomotor integration.Supported by INSERM (C.R.L. 79-53336)     

The retinotopic distribution of visual callosal projections in the suprasylvian visual areas compared to the classical visual areas (17, 18, 19) in the cat

Summary The distribution of the interhemispheric projection from area 17 and 18 was studied using the anterograde degeneration technique. Besides the classical visual areas (17, 18, 19), area 21 and several visual areas in the middle suprasylvian sulcus also received visual callosal input. In the four terminal areas of the middle suprasylvian sulcus the projection was found to be focused on representations of the vertical meridian including the area centralis, as in the classical visual areas. An increase of the width of visual field represented in the zone of callosal terminations can be seen from area 17 through area 18 to area 19 and possibly this trend continues in the suprasylvian visual areas.     

A study of binocular convergence in cat visual cortex neurons

Summary The average latency of cortical neuronal responses to electrical optic nerve (ON) stimulation was 3.0±0.7 s.d. msec. No significant difference between latencies to ipsi- and contralateral ON stimulation was found. Binocularly excitable cells showed almost equal response latencies to stimulation of both nerves. The average latency of subcortically recorded geniculo-cortical fibers was 0.3 msec less, but showed the same variance as that of cortical cells, suggesting that in all cases direct monosynaptic excitation of cortical cells by fibers of either ocularity is possible. Classes of ocular dominance based on electrical stimulation were positively, but not 100% correlated with classes of ocular dominance to visual stimulation. An anatomical study revealed that in cat terminals of geniculo-cortical projection are segregated to a lesser degree into ocularity stripes than in monkey. Direct monosynaptic excitation of cells by fibers of either ocularity which was found physiologically would also on these grounds appear possible for all cells.A preliminary report has been presented at the 46th German Physiological Society Meeting in Spring 1976, Pflügers Archiv, Vol. 362, Abstract No. 155, 1976     

The pattern of ocular dominance columns in flat-mounts of the cat visual cortex

Summary Ocular dominance (OD) columns in the cat visual cortex were visualized with autoradiography after intravitreal injection of (³H)proline. Extending previous studies, a flat-mount technique was applied that enabled the analysis of the distribution of label throughout extensive regions of the visual cortex without requiring reconstructions from serial sections. OD-columns were confined to layer IV and consisted of isolated patches and short bands. The latter were parallel to each other and regularly spaced, the main trajectory being orthogonal to the 17/18 border. This pattern of the geniculo-cortical terminals was similar in the hemispheres ipsi- and contralateral to the injected eye. The mean periodicities of the OD-bands were virtually identical in the two hemispheres of the same animal: 850 m and 830 m in cat D1 and 770 m and 800 m in cat D2. However, the ipsilateral OD-columns appeared smaller, more heavily labeled and more sharply delineated than the contralateral columns.     

Response covariance in cat visual cortex

Summary The activity of pairs of neurons in the visual cortex (area 17) of anaesthetized, paralysed cats was recorded using two independently manipulated micropipettes. The number of spikes in the evoked responses of pairs of single neurons were analyzed for response covariance. Responses of the majority of cell pairs (83%) did not covary. Covariance was restricted to closeby neurons with distances of less than 150 m and with identical orientation and ocular dominance preference.This work was partially supported by grants NS10332 and EY03796     

Effects of subcortical ablations on cortical association responses in the cat

The effects of subcortical lesions on cortical polysensory association area responses to peripheral sensory and direct reticular stimulation were investigated in acute chloralosed cats. Large lesions of the mesencephalic reticular formation were followed by a reduction in the amplitude of polysensory association responses to approximately 0–20% of control levels, with little or no reduction of evoked potentials in the primary sensory cortical areas. Unilateral lesions of posterior medial thalamus or the rostral pole of the thalamus had similar effects of polysensory cortical association responses, and also abolished cortical association area responses to direct electrical stimulation of the mesencephalic reticular formation, but only in the ipsilateral cortex. More ventral lesions, in the subthalamic region, reduced nonspecific evoked responses of orbital cortex to both peripheral and reticular stimulation without impairing responses in dorsal cortical association areas. The results are discussed with regard to the pathways of sensory input to the cortical association responses areas.     

The distribution of degenerating axons after small lesions in the intact and isolated visual cortex of the cat

Summary The extent of the spread of axonal degeneration was investigated in the visual cortex of the cat after making small lesions restricted to the grey matter. Two series of experiments were undertaken. In the first, normal adult cats were used, and in the second, the cortex of the postlateral gyrus was isolated from its extrinsic afferents by surgical undercutting 3 months before making the lesions. The results were similar in the two series in most respects. 1. Horizontal fibres extended in considerable numbers for some 500 m from the lesion, mainly in layers I, III/IV and V, a few reaching 2–3 mm. These fibres were better seen in the intact than in the isolated cortex. Their spread was usually asymmetrical, being greater posteromedially than anterolaterally. 2. Oblique axons ran downwards from the middle layers into layers V and VI, or upwards into layers I and II. 3. Axons arising from layers II to VI descended vertically into the white matter. Degeneration patterns after lesions in areas 17 and 18 were compared.     

Postnatal development of thalamocortical projections upon striate and extrastriate visual cortical areas in the cat

The development of visual thalamocortical projections was analyzed quantitatively by comparing, in cresyl violet-stained brain sections of early postnatal (10–17 days) and adult cats, the cell body dimensions and total cell packing density (CPD) of neuronal populations in different laminae (A, A1 and C) of the dorsal lateral geniculate (dLGN), medial interlaminar nucleus (MIN), and in lateral (LPl), intermediate (LPi) and medial (LPm) subdivisions of the lateral posterior complex. Following injections of different fluorescent tracers (FB, NY, EB, RITC) into cortical visual areas 17/18, posterior medial (PMLS) and posterior lateral (PLLS) lateral suprasylvian and anterior ectosylvian (AEV), the thalamic distribution and densities of retrogradely labeled neurons were analyzed. Projection CPDs and ratios of projection/total CPDs were determined and compared within the different thalamic components in the kitten and adult cat. A significant decrease in total cell packing density was observed in the various thalamic components of the adult cat, varying between 43% and 65%, and a marked increase in mean cell body diameter in the A, A1 and C laminae and MIN from kitten to adult (8.4±1.8 and 11.8±2.8 μm respectively) compared to the LP subnuclei (9.0±1.3 and 9.1±1.5 μm). The ratios of projection/total CPDs decreased significantly for projections upon areas 17/18 stemming from layers A and A1 (20 and 25%, respectively) and from LPi upon both PMLS (34%) and AEV (16%). Thalamocortical projections observed in the kitten from LPi upon areas 17/18 and from the A-laminae upon PMLS were absent in the adult cat. The data indicate that, in comparison to the lateral posterior nucleus, the maturation of neurons within the dLGN and MIN is incomplete with respect to cell body size during the early postnatal period. In addition, the developmental changes observed involve both reductions in the total number of thalamic neurons and a differential loss of cortical projections. The selective elimination of early cortical connections stemming from dorsal lateral geniculate laminae A and A1 and from the intermediate division of the lateral posterior nucleus may occur through a process of axon collateral withdrawal from the expanded cortical sites, thereby giving rise to the adult pattern. Accepted: 15 June 2000     

Contralateral corticofugal projections to cat visual thalamus

Summary Contralateral corticofugal projections from visual cortical areas to thalamic nuclei were demonstrated in the cat using anterograde transport of tritiated proline. Thalamic nuclei receiving projections from contralateral visual cortex include both subdivisions of the lateral-posterior nucleus, the posterior nucleus of Rioch, and the posterior nuclear complex.Abbreviations BIC brachium of the inferior colliculus - BN nucleus of the brachium of the inferior colliculus - BSC brachium of the superior colliculus - C dorsal lateral geniculate nucleus, C laminae - CG central gray matter - D nucleus of Darkschewitz - FR fasciculus retroflexus - FTC central tegmental field - H habenula - IPN interpeduncular nucleus - LGNd dorsal lateral geniculate nucleus, A laminae - LGNv ventral lateral geniculate nucleus - LP lateral posterior complex - LPi interjacent division of lateral posterior complex - LPl lateral division of lateral posterior complex - LPm medial division of lateral posterior complex - M mammillary body - MGM magnocellular division of medial geniculate nucleus - MGN medial geniculate nucleus - MGP parvocellular division of medial geniculate nucleus - MIN medial interlaminar division of lateral geniculate nucleus - MML medial medullary lamina - NOT nucleus of the optic tract - OT optic tract - P cerebral peduncle - PA anterior pretectal nucleus - PC nucleus of the posterior commissure - PM medial pretectal nucleus - PO posterior nuclear group - PoC posterior commissure - POi intermediate division of posterior nuclear complex - POL pretectal olivary nucleus - POm medial division of posterior nuclear complex - PPT posterior pretectal nucleus - PUL pulvinar - RN red nucleus - RNR posterior nucleus of Rioch - SG suprageniculate nucleus - SGI stratum griseum intermedium of superior colliculus - SGP stratum griseum profundum of superior colliculus - SCSl lower division of stratum griseum superficiale of superior colliculus - SGSu upper division of stratum griseum superficiale of superior colliculus - SN substantia nigra - SO stratum opticum of superior colliculus - TC tectal commissure - III III nerve - IIIN nucleus of III nerve     

Velocity sensitivity mechanisms in cat visual cortex

Summary To understand why some cells in the visual cortex respond to high stimulus velocities while others fail to do so, a sample of 71 of such cells were examined for their responses to stationary presented stimuli as well as to moving edges or slits of different widths. When presented with stationary stimuli it was found that cells which respond best to slowly moving stimuli generally have tonic discharges, long time to peak latencies and often long minimal durations of stimulation. In contrast, cells which respond preferentially to fast stimuli have phasic discharges, short latencies and short critical durations of stimulation when presented with stationary flashed slits. In the latter type of cells the responses to very fast stimulus movement were abolished selectively when contrast and width of the stimulus were not optimal. A few cells exhitited a velocity-response (VR) curve with a central dip indicating good responsiveness to either slow or fast movement but little to medium velocities. These cells responded both phasically and tonically to stationary slits and the latency of the tonic and phasic responses corresponded well to the latency of the responses at low and high velocities, respectively. It is suggested that the ability of phasic cells to respond to high velocities is linked to their limited need for temporal summation.Supported by an NFWO grant     

Stimulus contrast and visual cortical lesions

Summary Intact animals can make fine orientation discriminations over a wide range of contrasts. After ablation of area 17 deficits in orientation discrimination are observed only at low contrast. The relevance of this finding for the design of sensitive ablation experiments is discussed.     

Evidence for a nicotinic component to the actions of acetylcholine in cat visual cortex

Summary Radioligand binding assays, receptor autoradiography and iontophoresis have been used to look for evidence of a nicotinic component to the actions of acetylcholine in cat visual cortex. [³H]Nicotine bound to a uniform population of high affinity binding sites in cat primary visual cortex. This binding was inhibited by nicotine agonists and antagonists but not muscarinic antagonists. The concentration of nicotinic binding sites was about 10% of that of muscarinic binding sites measured with [³H]N-methylscopolamine. The muscarinic sites were resolved into M1 and M2 subtypes. Quantitative receptor autoradiography showed that there were muscarinic sites in all layers, although they were least abundant in layer IV of area 17. In contrast, the nicotinic sites were most concentrated in layer IV in area 17. The concentration of this labelling was reduced at the 17/18 border and also at the 18/19 border. Layer I of the cingulate and suprasylvian gyri were also labelled. Electrolytic lesions of the lateral geniculate nucleus (LGN) led to a loss of nicotinic binding sites in layer IV of area 17, indicating that these sites are most likely located on the LGN terminals. Iontophoresis of mecamylamine, a nicotinic antagonist, decreased evoked responses in visual cortex, providing evidence that the [³H]nicotine binding sites are functional receptors and suggesting that the release of acetylcholine onto these receptors on the LGN terminals facilitates the input of visual information into visual cortex.     

A comparison of magnification functions in area 19 and the lateral suprasylvian visual area in the cat

A retinotopic map can be described by a magnification function that relates magnification factor to visual field eccentricity. Magnification factor for primary visual cortex (VI) in both the cat and the macaque monkey is directly proportional to retinal ganglion cell density. However, among those extrastriate areas for which a magnification function has been described, this is often not the case. Deviations from the pattern established in V1 are of considerable interest because they may provide insight into an extrastriate area's role in visual processing. The present study explored the magnification function for the lateral suprasylvian area (LS) in the cat. Because of its complex retinotopic organization, magnification was calculated indirectly using the known magnification function for area 19. Small tracer injections were made in area 17, and the extent of anterograde label in LS and in area 19 was measured. Using the ratio of cortical area labeled in LS to that in area 19, and the known magnification factor for area 19 at the corresponding retinotopic location, we were able to calculate magnification factor for LS. We found that the magnification function for LS differed substantially from that for area 19: central visual field was expanded, and peripheral field compressed in LS compared with area 19. Additionally, we found that the lower vertical meridian's representation was compressed relative to that of the horizontal meridian. We also examined receptive field size in areas 17, 19, and LS and found that, for all three areas, receptive field size was inversely proportional to magnification factor.