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Regional distribution of binding sites for neuropeptide Y in cat and monkey visual cortex determined by in vitro receptor autoradiography.

The goal of this study was to elucidate the precise regional and laminar distribution of neuropeptide Y (NPY) binding sites in feline and primate visual cortex. By means of in vitro receptor autoradiography, NPY binding sites in primate and feline visual cortex were specifically labeled with 3H-NPY. In cat area 17, the highest density of NPY-binding sites was present in lamina I and the upper half of lamina II. The density then gradually decreased towards lamina VI. Areas 18 and 19 exhibited a similar binding site-density profile. The decrease in density from superficial to deep layers was more gradual in area 18 than in areas 17 and 19. In monkey primary visual cortex (V1), layer IVc presented a high concentration of NPY binding sites, in addition to a dense zone of binding sites in layer I. Monkey secondary visual cortex (V2) displays a similar dense zone in layer I, but lacks such high density of NPY binding sites in layer IV. Therefore, the border between primary and secondary visual cortex coincides with the abrupt disappearance of this latter high density in layer IV. In cat as well as in monkey visual cortex, no significant differences were found between regions representing central vision and those representing the peripheral parts of the visual field. Comparison of our results for NPY binding sites with the distribution of alpha 1-adrenergic receptors, as recently described by Rakic et al. (J. Neurosci. 8(10):3670-3690, 1988) for primate and Parkinson et al. (Brain Res. 457:70-78, 1988) for feline visual cortex, revealed that those two patterns are very similar.

Animals↗

Retinotopy within rat primary visual cortex using optical imaging.

The purpose of this study was to determine the retinotopic organization of rat primary visual cortex (area 17) using optical imaging technology. Stimulating discrete regions of visual space resulted in localised changes in the remitted light during optical imaging of visual cortex in rat. From these localised changes, our results confirm previous electrophysiological studies on the location, size and organization of rat primary visual cortex. Small differences in the cortical magnification factor (CMF) were found between visual field areas with the highest CMF confined to the upper nasal region. No significant CMF differences were found within the horizontal and vertical visual field axes. No secondary visual areas were activated either anterior or medial to area 17 with the pattern stimuli used in the current study. However, there was evidence of activity to upper nasal stimulation on the posterior lateral extrastriate area. The location of area 17 from optical imaging activity was confirmed anatomically using conventional immunohistochemical techniques. This study shows the retinotopic organization of rat primary visual cortex and serves as a precursor before examining animal models of retinal degeneration and the effectiveness of potential therapies to stem retinal disease.

Animals↗

Repetitive optical stimulation elicits fast receptive field changes in mature visual cortex.

Fast associative cellular plasticity of visual cortical cells has been shown in vitro with electrical stimulation as well as in vivo by pairing of natural stimuli with artificial depolarization. Here we experimentally changed receptive field properties of single cells in the mature cat visual cortex with purely natural associative stimulation in vivo. Single cortical cells expanded their receptive fields within minutes into unresponsive regions and changed their functional receptive field structure for hours after associative co-stimulation of active and primarily unresponsive regions. The effects are interpreted as strengthening of subthreshold synaptic inputs. Repetitive stimulation with the same visual pattern can modify single cell properties in vivo within minutes and may be related to preattentive fast perceptual learning.

Animals↗

[Electron microscopic analysis of expression of NMDA-R1 in the developmental process of visual cortex in strabismic amblyopic cat].

OBJECTIVE: To investigate the expression and distribution of N-methyl-D-aspartate receptor subunit 1 (NMDA-R1) in neuronal ultrastructure in visual cortex of strabismic amblyopic cat during development. METHODS: Eleven kittens were used for this study. Esotropia in six kittens had been made monocularly by tenotomy at two weeks of age. Two pairs of normal and strabismic kittens were sacrificed in three weeks of age, one week after tenotomy. Another two pairs of normal and strabismic kittens were sacrificed in five weeks of age, three weeks after tenotomy. One normal and two strabismic amblyopic cats were sacrificed after 6 months of age. Animals were deeply anaesthetized and perfused transcardially with 4% paraformaldehyde. Cryostat sections of frontal central area P5-P0 were cut to 25 micro m thickness. The mouse anti-NMDA-R1 monoclonal antibody (mAb54.1, PharMingen) was used. After stained, a light microscope was used to select regions of layer II-III, layer IV and layer V-VI of visual cortex area 17 for re-embedding. HITACHI H-7000 transmission electron microscope at magnifications ranging from 30 000 - 300 000 X was used for observation. RESULTS: Three hundred and twenty-eight neurons of strait cortex were observed. NMDA-R1 receptor was located at the nuclei, Nissl body, cytoplasm, plasma membrane and the postsynaptic element of axons and dendrites. The ultrastructural morphology, including the mitochondrion, smooth endoplasmic reticulum, rough endoplasmic reticulum, and the Golgi apparatus, was not significantly different in the comparison between the cells in visual cortex of normal and strabismic groups. In the entire normal group, the percentage density of NMDA-R1 labeled cells was higher than that of strabismic groups (chi(2) = 4.280, 4.41, 4.89; P < 0.05). One thousand and three hundred and twenty NMDA-R1 immunopositive synapses were counted. The NMDA-R1 immunopositive synapses were dominated in layer II-III of visual cortex and increased during the development of normal kittens (F = 3.28, P < 0.05). There was no significant difference of NMDA-R1 immunopositive synapse distribution between the normal and strabismic kitten at 3 weeks (one week after operation) of age (F = 0.17, P > 0.05). The reduction of NMDA-R1 immunopositive synapse of plasma membrane in visual cortex of strabismic kitten was started at 5 weeks (threes weeks after surgery) of age. It was decreased significantly in strabismic amblyopic cat compared with that of the normal cat (F = 26.94, 47.01; P < 0.001). The ratios of nuclear membrane invagination of cells in visual cortex of normal and strabismic cat were higher than those of normal and squint kittens (chi(2) = 36.24, P < 0.01), but the ratio was not significantly different between the normal and strabismic group. CONCLUSION: (1) In the normal developmental process of cat, the plasticity of the neuronal synapsis in II and III layer of visual cortex is relatively great. (2) In the strabismus amblyopia occurring in the plastic critical period of visual development, no pathological changes of neuronal organelle in the visual cortex are found, but there are changes at molecular level in the neuronal synapsis.

Amblyopia↗

Effects of serial unilateral and serial bilateral visual cortex lesions on brightness discrimination relearning in rats.

In Experiment 1, rats with serial unilateral (SU), serial bilateral (SB), or one-stage bilateral (B) visual cortex lesions (areas 17, 18 and 18a) were compared in their retention of a preoperatively learned brightness discrimination. Both the first-stage and second-stage lesions for the SU and SB groups involved the same amount of cortical tissue and the same composite areas of visual field projection in primary and extraprimary visual cortex. Groups SU and SB showed substantial savings of the brightness discrimination after the completed bilateral visual cortex removals, while Group B showed a complete postoperative loss. In Experiment 2, the effects of two additional types of serial bilateral visual cortex lesions were investigated. These involved the medial and then lateral aspects of visual cortex in two stages (ML), or vice versa (LM). Once again, the serial bilateral lesion groups (SB, ML, and LM) showed substantial savings of the discrimination after the completed lesion, while Group B showed a complete post-operative loss. These results demonstrate a general advantage of serial damage over one-stage damage to visual cortex in recovery of a preoperatively learned brightness discrimination and indicate that the serial lesion effect is not specific to interhemispheric relationships.

Animals↗

A linear model fails to predict orientation selectivity of cells in the cat visual cortex.

1. Postsynaptic potentials (PSPs) evoked by visual stimulation in simple cells in the cat visual cortex were recorded using in vivo whole-cell technique. Responses to small spots of light presented at different positions over the receptive field and responses to elongated bars of different orientations centred on the receptive field were recorded. 2. To test whether a linear model can account for orientation selectivity of cortical neurones, responses to elongated bars were compared with responses predicted by a linear model from the receptive field map obtained from flashing spots. 3. The linear model faithfully predicted the preferred orientation, but not the degree of orientation selectivity or the sharpness of orientation tuning. The ratio of optimal to non-optimal responses was always underestimated by the model. 4. Thus non-linear mechanisms, which can include suppression of non-optimal responses and/or amplification of optimal responses, are involved in the generation of orientation selectivity in the primary visual cortex.

Animals↗

Development of neuronal responses in cat posteromedial lateral suprasylvian visual cortex.

We studied the normal development of responses to visual stimulation among neurons in the posteromedial lateral suprasylvian (PMLS) visual cortex, an extrastriate visual cortical area in cats. Recordings were made from 495 single neurons in 19 kittens that were 2, 3, 4, 8, or 12 weeks of age, and the results were compared with those from normal adult cats. The percentage of neurons that respond to light increased from 57% in 2-week-old kittens to approximately adult values in 8-week-old kittens (81%). The strength and consistency of neuronal responses also increased with age. Nearly all of the responsive cells had well-defined excitatory receptive-field centers, and the receptive-field center sizes were similar to adults at all ages studied. However, few cells (5%) had inhibitory receptive-field surrounds in 2-week-old kittens. The incidence of surround inhibition increased to adult levels (about 40% of the cells) by 8 weeks of age, and the strength of surround inhibition also increased with age. Most cells responded best to moving stimuli in 2-week-old kittens, just as in adults. However, only about 20% of the responsive cells were direction sensitive at 2 weeks of age. The percentage of direction-sensitive cells increased gradually with age and reached approximately adult values by 8 weeks of age (74%). Once cells developed complete direction selectivity, with no response in the null direction, directional tuning width was similar to that in adults. When tested with slits of light flashed at various orientations or with spots and slits moving in various directions, few cells (8% or less) showed orientation selectivity at any age, just as in adults. Most of the cells were binocularly driven, and the ocular dominance distribution was similar to adults at all ages studied. These results indicate that many response properties of PMLS neurons are similar to those of adults as early as 2 weeks of age, soon after the time of eye opening. However, some properties show marked developmental changes. The mechanisms and sources of these changes are considered. In addition, the relevance of these results to mechanisms of compensation following early damage to visual cortical areas 17, 18 and 19 is discussed.

Aging↗

Intracortical microstimulation of neurons in the visual cortex of the cat.

The response of visual cortex neurons to local intracortical microstimulation was measured in the anesthetized cat. When the recording microelectrode was very close (about 20 micrometers) to the tip of the stimulating electrode, threshold currents as low as 10 micro A were capable of firing neurons. Over a 20-fold range in distance from the site of stimulation, an 80-fold increase in threshold current was observed. The mean latency of activation for 30 neurons tested with intracortical stimulation was 2.88 +/- 0.45 msec. The majority of these cells were probably synaptically activated. The mean threshold current for these neurons was 0.55 +/- 0.12 mA (N = 30). These values were significantly smaller than the thresholds found previously when stimulating electrodes were located on the pia-arachnoid surface of the visual cortex.

Animals↗

Diversity of relative intensities of acetylcholinesterase staining within the laminae of the visual cortex of four mammals.

The histological staining patterns of acetylcholinesterase fibers in the visual cortex of the hamster, bank vole, yellow necked wood mouse, and rabbit are described. Between the species there are variations in staining intensities of the laminae in the visual cortex. In the hamster and bank vole layers IV, V, and VI are intensely stained. The yellow necked wood mouse demonstrates little difference in staining intensity between the layers of the visual cortex. The rabbit evidences a dense precipitate in the lower portion of layer IV and also in layer V. Staining intensity patterns suggest that the cholinergic afferents to the visual cortex may develop preferential innervation to laminae which receive low sensory stimulation during the formative period.

Acetylcholinesterase↗

Correspondence of presaccadic activity in the monkey primary visual cortex with saccadic eye movements.

We continuously scan the visual world via rapid or saccadic eye movements. Such eye movements are guided by visual information, and thus the oculomotor structures that determine when and where to look need visual information to control the eye movements. To know whether visual areas contain activity that may contribute to the control of eye movements, we recorded neural responses in the visual cortex of monkeys engaged in a delayed figure-ground detection task and analyzed the activity during the period of oculomotor preparation. We show that approximately 100 ms before the onset of visually and memory-guided saccades neural activity in V1 becomes stronger where the strongest presaccadic responses are found at the location of the saccade target. In addition, in memory-guided saccades the strength of presaccadic activity shows a correlation with the onset of the saccade. These findings indicate that the primary visual cortex contains saccade-related responses and participates in visually guided oculomotor behavior.

Animals↗

Receptive fields in human visual cortex mapped with surface electrodes.

Most of our understanding of the functional organization of human visual cortex comes from lesion and functional imaging studies and by extrapolation from results obtained by neuroanatomical and neurophysiological studies in nonhuman primates. Although some single-unit and field potential recordings have been made in human visual cortex, none has provided quantitative characterization of spatial receptive fields (RFs) of individual sites. Here we use subdural electrodes implanted for clinical purposes to quantitatively measure response properties in different regions of human visual cortex. We find significant differences in RF size, response latency, and response magnitude for sites in early visual areas, versus sites in later stages of both the dorsal and ventral streams. In addition, we use this technique to estimate the cortical magnification factor in early human visual cortex. The spatial and temporal resolution of cortical surface recordings suggest that this technique is well suited to examine further issues in visual processing in humans.

Brain Mapping↗

Primary visual cortex neurons that contribute to resolve the aperture problem.

It is traditional to believe that neurons in primary visual cortex are sensitive only or principally to stimulation within a spatially restricted receptive field (classical receptive field). It follows from this that they should only be capable of encoding the direction of stimulus movement orthogonal to the local contour, since this is the only information available in their classical receptive field "aperture." This direction is not necessarily the same as the motion of the entire object, as the direction cue within an aperture is ambiguous to the global direction of motion, which can only be derived by integrating with unambiguous components of the object. Recent results, however, show that primary visual cortex neurons can integrate spatially and temporally distributed cues outside the classical receptive field, and so we reexamined whether primary visual cortex neurons suffer the "aperture problem." With the stimulation of an optimally oriented bar drifting across the classical receptive field in different global directions, here we show that a subpopulation of primary visual cortex neurons (25/81) recorded from anesthetized and paralyzed marmosets is capable of integrating informative unambiguous direction cues presented by the bar ends, well outside their classical receptive fields, to encode global motion direction. Although the stimuli within the classical receptive field were identical, their directional responses were significantly modulated according to the global direction of stimulus movement. Hence, some primary visual cortex neurons are not local motion energy filters, but may encode signals that contribute directly to global motion processing.

Action Potentials↗

Temporal characteristics of visual receptive fields in primary visual cortex and medial superior temporal cortex areas.

We mapped the receptive fields of 49 cells from primary visual cortex and 19 cells from medial superior temporal cortex in two awake monkeys. The receptive field structures we obtained lasted a mean time of 32.7 ms in primary visual cortex and 38.4 ms in medial superior temporal cortex, showing no statistical difference. This result suggests that both areas have the same time requirements for processing visual information. In primary visual cortex, 100% of cells had conformed the receptive field structure at 65 ms pre-spike, whereas in medial superior temporal cortex it occurred at 150 ms. In both areas, cells with shorter response latencies had receptive field structures with longer durations. This may indicate that cells tend to synchronize their output to other areas.

Action Potentials↗

Contrast constancy in natural scenes in shadow or direct light: A proposed role for contrast-normalisation (non-specific suppression) in visual cortex.

The range of contrasts in natural scenes is generally thought to far exceed the limited dynamic ranges of individual contrast-encoding neurons in the primary visual cortex. The visual system may employ gain-control mechanisms (Ohzawa et al. 1985) to compensate for the mismatch between the range of natural contrast energies and the limited dynamic range of visual neurons; one proposed mechanism is contrast normalisation or non-specific suppression (Heeger 1992a). This paper aims to evaluate the role of contrast normalisation in human contrast perception, using a computer model of primary visual cortex. The model uses orthogonal pairs of Gabor patches to simulate simple-cell receptive-fields to calculate local, band-limited contrast in a series of 50 digitised photographs of natural scenes. The average range of contrast energies in each image was 2.29 log units, while the "lifetime range" each model simple cell would see across all images was 2.98 log units. These ranges are greater than the dynamic range of real mammalian simple cells. Contrast normalisation (dividing contrast responses by the summed responses of all nearby neurons) reduces contrast ranges, perhaps sufficiently to match them to neurons' limited dynamic ranges. Comparison of images taken under diffuse and direct lighting conditions showed that contrast normalisation can sometimes match these conditions effectively. This may lead to perceptual contrast constancy in the face of spurious changes in contrast caused by natural environmental conditions.

Computer Simulation↗

Unconscious processing of orientation and color without primary visual cortex.

In humans, the primary visual cortex (V1) is essential for conscious vision. However, even without V1 and in the absence of awareness, some preserved ability to accurately respond to visual inputs has been demonstrated, a phenomenon referred to as blindsight. We used transcranial magnetic stimulation (TMS) to deactivate V1, producing transient blindness for visual targets presented in a foveal, TMS-induced scotoma. Despite unawareness of these targets, performance on forced choice discrimination tasks for orientation (experiment 1) and color (experiment 2) were both significantly above chance. In addition to demonstrating that TMS can be successfully used to induce blindsight within a normal population, these results suggest a functioning geniculoextrastriate visual pathway that bypasses V1 and can process orientation and color in the absence of conscious awareness.

Color Perception↗

The pharmacology of synapses formed by identified corticocollicular neurons in primary cultures of rat visual cortex.

Primary cultures of neurons from the visual cortex of 7-10-d-old Long Evans rats were used to study the pharmacology of synaptic transmission. Dissociated cells were grown either in mass cultures, which contained 8000-10,000 neurons, or in miniature island cultures of 50-100 cells. Prior to dissociation, cells in layer V of visual cortex that project to the superior colliculus were labeled in vivo by retrograde transport of fluorescent latex microspheres-a permanent fluorescent marker. After 2 d to 8 weeks in culture, labeled neurons were identified by epifluorescent illumination, and electrophysiological recordings were obtained from a labeled cell and, simultaneously, from a nearby unlabeled neuron in the same field of view. The 2 neurons were stimulated sequentially by current injection and the pharmacology of evoked postsynaptic potentials (PSPs) was investigated. In mass cultures, relatively few pairs of neurons from which we recorded were synaptically connected, although nearly every cell exhibited abundant spontaneous EPSPs and IPSPs. Neurons grown on island cultures generally did not exhibit spontaneous synaptic activity; however, stimulation of one of the cells in a pair frequently elicited a short-latency PSP in the follower neuron. Retrogradely labeled corticocollicular neurons produced only excitatory PSPs in follower cells, while unlabeled neurons were either excitatory or inhibitory. Three antagonists of excitatory amino acid receptors, kynurenic acid, piperidine dicarboxylic acid, and gamma-D-glutamylglycine, completely blocked EPSPs produced by labeled corticocollicular neurons, as well as EPSPs produced by nearly all of the unlabeled excitatory cells. We have previously shown that these compounds block both N-methyl-D-aspartate (NMDA)-type and non-NMDA receptors on cultured cortical neurons (Huettner and Baughman, 1986). The specific NMDA receptor antagonist 2-amino-5-phosphonovaleric acid (APV) did not alter short-latency EPSPs recorded in 1 mM Mg2+, but did reduce longer-latency EPSPs polysynaptic activity. Since responses mediated by the NMDA receptor are known to be antagonized by Mg2+ (Mayer and Westbrook, 1985), we perfused cultures with Mg2+-free medium and found that the falling phase of some monosynaptic EPSPs was prolonged. Addition of APV to Mg2+-free medium reduced the duration of the falling phase of EPSPs such that they returned to the time course obtained in 1 mM Mg2+.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗

Immunohistochemical study of the pattern of rapid expression of C-Fos protein in the visual cortex of dark-reared kittens following initial exposure to light.

Brief alterations to the nature of the visual input during critical periods in the early life of cats and monkeys can result in rapid anatomical and physiological changes in the central visual pathways. The immediate early genes (IEGs) represent a possible way in which these changes could be mediated since the protein products of a number of these genes have been shown to be induced rapidly in neurons in response to a variety of transsynaptic stimuli. Immunohistochemical methods were employed to examine the tempo and pattern of expression of Fos, the protein product of the c-fos gene, induced in the visual cortex of kittens dark-reared from birth to 30 days of age by brief periods of binocular visual exposure. In visual cortical area 17, the number of Fos immunoreactive cells increased rapidly from virtually zero in control kittens that received no visual exposure, to reach high levels in animals that received between 1 and 2 hours of visual experience. Immunoreactive cells were absent in the dorsal lateral geniculate nucleus, but were numerous in the ventral lateral geniculate nucleus, and in area 17, were most numerous in the extragranular layers (2, 3 and 6) but sparse in lower layer 4 and layer 5, and virtually absent in layer 1. Substantial constitutive Fos immunoreactivity was observed in area 17 of normal 30-day-old kittens but very few immunopositive cells were evident in adult animals. However, Fos immunoreactivity was observed in the visual cortex of a dark-reared (for 30 days) adult animal following a brief period of visual exposure, a finding that suggests that Fos might serve other roles in the visual cortex in addition to those it might play uniquely during development. It is suggested that Fos, in combination with the protein products of a select number of other IEGs, may mediate a variety of rapid changes in the visual cortex including those that underlie visual system plasticity during early postnatal life.

Animals↗

Calretinin-immunoreactive neurons in the primary visual cortex of dolphin and human brains.

A new class of gamma-aminobutyric acid (GABA)ergic neurons immunoreactive to the calcium-binding protein calretinin (CR) was demonstrated in primary visual cortices of the bottlenose dolphin (Tursiops truncatus) and humans (Homo sapiens). Comparative analysis revealed several differences between dolphin and human visual cortex in the laminar distribution of CR-positive perikarya, although general typology of the immunoreactive CR-positive neurons was similar in both species. Thus, in both human and dolphin primary visual cortex almost all CR-positive neurons are non-pyramidal, either fusiform or bipolar cells, oriented with their long axis along the radial axis of the cortex. Large multipolar stellate cells were also observed in layers I and VI. The CR-positive neurons in the dolphin visual cortex are concentrated almost exclusively in layer I and, to a lesser extent, in layer II. In all other layers (IIIa, b, IIIc/V and VI) of the dolphin visual cortex CR-positive neurons were only rarely seen. In the human primary visual cortex CR-positive neurons are located mainly in layers II, III and IVa, b, c, with considerably lower densities of these cells observed in layers V and VI. CR-positive neurons in layer I of the human visual cortex are represented by Cajal-Retzius horizontal cells, whereas no such cells were seen in layer I of the dolphin neocortex. The numerical density of CR-positive neurons in the dolphin primary visual cortex is significantly lower than in the area of cortex in humans.(ABSTRACT TRUNCATED AT 250 WORDS)

Aged↗