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PubMed2003

Quantitative analyses of principal and secondary compound parieto-occipital feedback pathways in cat.

The purpose of our study was to quantify the magnitude of principal and secondary pathways emanating from the middle suprasylvian (MS) region of visuoparietal cortex and terminating in area 18 of primary visual cortex. These pathways transmit feedback signals from visuoparietal cortex to primary visual cortex. (1) WGA-HRP was injected into area 18 to identify inputs from visual structures. In terms of numbers of neurons, feedback projections to area 18 from MS sulcal cortex (areas PMLS, AMLS and PLLS) comprise 26% of inputs from all visual structures. Of these neurons, between 21% and 34.9% are located in upper layers 2-4 and the dominant numbers are located in deep layers 5 and 6. Areas 17 (11.8%) and 19 (11.2%) provide more modest cortical inputs, and another eight areas provide a combined total of 4.3% of inputs. The sum of neurons in all subcompartments of the lateral geniculate nucleus (LGN) accounts for another 34.8% of the input to area 18, whereas inputs from the lateral division of the lateral-posterior nucleus (LPl) account for the final 11.9%. (2) Injection of tritiated-((3)H)-amino acids into MS sulcal cortex revealed substantial direct projections from MS cortex that terminated in all layers of area 18, but with a markedly lower density in layer 4. Projections from MS cortex to both areas 17 and 19 are of similar density and characteristics, whereas those to other cortical targets have very low densities. Quantification also revealed minor-to-modest axon projections to all components of LGN and a massive projection throughout the LP-Pul complex. (3) Superposition of the labeled terminal and cell fields identified secondary, compound feedback pathways from MS cortex to area 18. The largest secondary pathway is massive and it includes the LPl nucleus. Much more modest secondary pathways include areas 17 and 19, and LGN. The relative magnitudes of the secondary pathways suggest that the one through LPl exerts a major influence on area 18, whereas the others exert more modest or minor influences. MS cortex in the contralateral hemisphere also innervates area 18 directly. These data are important for interpreting the impact of deactivating feedback projections from visuoparietal cortex on occipital cortex.

Animals↗
PubMed1997

Orientation selectivity in pinwheel centers in cat striate cortex.

In primary visual cortex of higher mammals neurons are grouped according to their orientation preference, forming "pinwheels" around "orientation centers." Although the general structure of orientation maps is largely resolved, the microscopic arrangement of neuronal response properties in the orientation centers has remained elusive. The tetrode technique, enabling multiple single-unit recordings, in combination with intrinsic signal imaging was used to reveal the fine-grain structure of orientation maps in these locations. The results show that orientation centers represent locations where orientation columns converge containing normal, sharply tuned neurons of different orientation preference lying in close proximity.

Action Potentials↗
PubMed2002

Anatomical evidence of multimodal integration in primate striate cortex.

The primary visual cortex (area 17 or V1) is not thought to receive input from nonvisual extrastriate cortical areas. However, this has yet to be shown to be the case using sensitive tracers in the part of area 17 subserving the peripheral visual field. Here we show using retrograde tracers that peripheral area 17 subserving the visual field at an eccentricity of 10-20 degrees receives projections from the core and parabelt areas of the auditory cortex as well as from the polysensory area of the temporal lobe (STP). The relative strength of these projections was calculated for each injection by computing the proportions of retrogradely labeled neurons located in the auditory and STP areas with respect to number of labeled neurons constituting the established projection from the superior temporal sulci (STS) motion complex (middle temporal area, medial superior temporal, fundus of the superior temporal area). In peripheral area V1 the projection from auditory cortex corresponds to 9.5% of that of the STS motion complex and STP to 35% of that from the STS motion complex. Compared to peripheral area 17, central and paracentral area 17 showed considerably weaker inputs from auditory cortex (0.2-0.8%) but slightly more from STP cortex (3.5-6.1%). The present results show that the connectivity of area 17 is eccentricity dependent. Direct projections from auditory and STP cortex to peripheral area 17 have important consequences for higher visual functions of area 17, including multimodal integration at early stages of the visual cortical pathway.

Animals↗
PubMed2003

Expression of occ1 mRNA in the visual cortex during postnatal development in macaques.

We previously reported that the occ1 gene is specifically expressed in the primary visual cortex of adult monkeys in an activity-dependent manner (Tochitani et al., Eur. J. Neurosci., 3, 297-307, 2001). In this report, we compared occ1 mRNA expression in the primary visual cortex during the development of newborn, 3-month-old and adult monkeys. occ1 mRNA was already expressed preferentially in the primary visual cortex of newborn monkeys, but the laminar pattern of occ1 expression in the visual cortex changed as development proceeded. This suggests the possible importance of experience-dependent developmental regulations of occ1 in the developing primary visual cortex.

Aging↗
PubMed2001

When the auditory cortex turns visual.

We studied visually guided behavior and the visual response properties of single auditory cortex (A1) neurons in neonatally operated hamsters with surgically induced, permanent, ectopic retinal projections to auditory thalamic nuclei and to visual thalamic nuclei which normally receive little direct retinal input. The surgically induced retino-thalamo-cortical pathways can mediate visual guided behaviors whose normal substrate, the pathway from the retina to the primary visual cortex via the primary thalamic visual nucleus, is missing. The visually evoked response properties of A1 neurons resemble in many respects those of neurons in V1 of normal hamsters: many A1 neurons have well-defined visual receptive fields and preferences for orientation or direction of movement. In addition, some visually responsive cells in A1 are bimodal--they also respond to auditory stimuli. The visually responsive neurons in A1 probably account for the capacity of the auditory cortex to mediate visual behavior in 'rewired hamsters'.

Animals↗
PubMed2003

Morphology and histology of chimpanzee primary visual striate cortex indicate that brain reorganization predated brain expansion in early hominid evolution.

Human brain evolution is characterized by an overall increase in brain size, cerebral reorganization, and cerebral lateralization. It is generally understood when brain enlargement occurred during human evolution. However, issues concerning cerebral reorganization and hemispheric lateralization are more difficult to determine from brain endocasts, and they are topics of considerable debate. One region of the cerebral cortex that may represent the earliest evidence for brain reorganization is the primary visual cortex (PVC), or area 17 of Brodmann. In nonhuman primates, this region is larger in volume (demarcated anteriorly by the lunate sulcus), and extends further rostrally than it does in modern humans. In early hominid fossil (Australopithecus) endocasts, this region appears to occupy a smaller area compared to that in nonhuman primates. Some have argued that the brain first underwent size expansion prior to reorganization, while others maintain that reorganization predated brain expansion. To help resolve this question, we provide a description of two male, common chimpanzee (Pan troglodytes) brains, YN77-111 and YN92-115, which clearly display a more posterior lunate sulcal morphology than seen in other chimpanzees. These data show that neurogenetic variability exists in chimpanzees, and that significant differences in organization (e.g., a reduced PVC) can predate brain enlargement. While the human brain has experienced numerous expansion and reorganization events throughout evolution, the data from these two chimpanzees offer significant support for the hypothesis that the neurogenetic basis for brain reorganization was present in our early fossil ancestors (i.e., the australopithecines) prior to brain enlargement.

Animals↗
PubMed1995

Development of a crossed corticotectal pathway following cerebral hemispherectomy in cats: a quantitative study of the projecting neurons.

A hypothetical mechanism for the partial sparing of visual function in the contralateral visual field following cerebral hemispherectomy early in life is the formation of a new corticotectal pathway arising from the remaining primary visual cortex (areas 17 and 18) that projects to the contralateral superior colliculus. To test this hypothesis, the left superior colliculus of intact adult and neonatal (5-15 days old) cats and of adult cats with a left cerebral hemispherectomy sustained neonatally (7-9 days old) or in adulthood, was injected with WGA-HRP and the brains were processed for combined TMB/DAB histochemistry. The primary visual cortex was examined, labelled neurons were counted and the cross sectional area of their somata was measured. The left primary visual cortex of intact adult animals exhibited a mean of 959.68 labelled cells +/- 406.5 (S.E.), with a mean soma size of 366.7 microns2 +/- 131.2. For the neonatal intact cats, there was a mean of 75.31 +/- 21.08 cells within the left primary visual cortex which exhibited a mean soma size of 249.56 microns2 +/- 68.18. The peak cell size distribution for both intact groups was similar at 300 microns2. Virtually no labelled neurons were detected in the right primary visual cortex of intact animals (neonatal or adult). For neonatal-hemispherectomized cats, the remaining right primary visual cortex exhibited a mean cell count of 351.09 +/- 126.3 cells, with a mean soma size of 436.1 microns2 +/- 131.5, and a peak cell size distribution of 400 microns2. Finally, for adult-hemispherectomized animals, the contralateral primary visual cortex exhibited 68.27 +/- 20.13 neurons having a mean soma size of 486.6 microns2 +/- 143.2 with a peak cell size distribution of 500 microns2. These results indicate that reorganization of the corticotectal pathway occurs in both adult- and neonatal-hemispherectomized cats but is more pronounced in neonatal-lesioned animals. In addition, the cells of origin of this reorganized pathway tended to be larger, perhaps in response to a greater axonal arborization.

Animals↗
PubMed1989

Corticopontine projection in the rat: the distribution of labelled cortical cells after large injections of horseradish peroxidase in the pontine nuclei.

The distribution of cortical cells projecting to the pontine nuclei in rats was studied by making large injections of horseradish peroxidase that filled the basilar pons and measuring the density of labelled cells in each cortical area. All retrogradely labelled cells were layer V pyramidal cells. The highest densities of labelled cells were observed in the motor areas. The lowest densities were in temporal association cortex and perirhinal cortex. Visual cortical areas, including the primary visual cortex, provided a major source of pontine projections. The distribution of corticopontine cells within the primary visual cortex was studied in more detail. In all cases the highest density of labelled cells was observed in the region of cortex that represents the nasal visual field. Control injections into brainstem regions adjacent to the pontine nuclei produced a much lower absolute density of retrogradely labelled cortical cells and the distribution of those cells was different from that observed following pontine injections. We conclude that every area of the rat's cerebral cortex projects to the pontine nuclei and that there are consistent variations in the density of the projections both between and within areas.

Animals↗
PubMed1991

Prosencephalic connections of striate and extrastriate areas of rat visual cortex.

Afferent connections of rat primary visual cortex (area 17 or V1 area) and the rostral and caudal parts of areas 18a and 18b were studied, by placing in each of the areas, small electrophoretic injections of enzyme horseradish peroxidase (HRP) or wheat germ agglutinated-HRP. The results indicate that: 1) each of the areas has a distinct pattern of distribution of afferent neurons in the ipsilateral visual thalamus - area 17 receives its principal thalamic input from the dorsal lateral geniculate nucleus, the caudal parts of areas 18a and 18b receive a major thalamic input from the lateral posterior nucleus and a minor input from the posterior nucleus, while the rostral parts of areas 18a and 18b receive a major input from the posterior nucleus, and a minor projection from the lateral posterior nucleus; 2) the rostral and caudal parts of areas 18a and 18b each receive an associational input from area 17; 3) the rostral parts of areas 18a and 18b each receive associational input from three different extrastriate regions, the caudal part of the same extrastriate area, and the rostral and caudal parts of the other extrastriate area, whereas the caudal parts of areas 18a and 18b receive associational inputs only from one or two extrastriate regions; 4) area 17, area 18b and rostral area 18a each receive a substantial associational input from lamina V of the caudal part of the frontal eye field (FEF) in the motor cortex; however the input from the FEF to caudal area 18a (if present) is very small; 5) The extrastriate areas studied receive associational input from the restrosplenial cingulate area 29d; however, the input from area 29d to area 17 appears to be very small. The distinct patterns of distribution of prosencephalic afferents suggest to us that multiple retinotopically organized areas described previously in the rat cortex (cf Montero 1981; Espinoza and Thomas 1983) represent functionally distinct areas.

Afferent Pathways↗
PubMed1989

Neuropeptide Y-containing neurons are situated predominantly outside cytochrome oxidase puffs in macaque visual cortex.

Layers II/III of the primary visual cortex contain a regular pattern of histochemically detectable cytochrome oxidase (CO)-rich "puffs," which differ from the interpuff regions in their thalamo-cortical and cortico-cortical connectivity, receptive-field properties, and the density of inhibitory GABA-containing synaptic terminals. We used an immunocytochemical method, in combination with cytochrome oxidase histochemistry, to analyze the spatial relationship between neurons that contain neuropeptide Y (NPY) and the CO puffs. Of a total of 606 neurons, only 2.6% of the NPY-containing cells are located in the puffs, whereas the rest are situated in the interpuffs, or at the interface between puffs and interpuffs. The number of NPY-containing neurons in the puffs is substantially less than that expected in an equal volume of the interpuffs (X2 = 13.86; df = 1; P less than 0.001). These observations indicate that columns containing the puffs may differ also from those in the interpuff regions in that they contain a unique array of chemically and morphologically distinct local circuit neurons.

Animals↗
PubMed1994

Interocular control of neuronal responsiveness in cat visual cortex.

Neurons in the cat primary visual cortex are selective for particular contour orientations but their responsiveness can vary under certain conditions. After prolonged stimulation (adaptation), the contrast sensitivity of cortical cells is reduced and the 'gain' (the strength of response as a function of contrast) falls. The response to an optimal contour is also reduced when a different stimulus is superimposed on the receptive field in the same eye. Here we report that the sudden appearance of an inappropriate stimulus in one eye can interocularly suppress the activity of cortical neurons if they are already responding to an optimally oriented stimulus in the other eye. In strabismic cats, whose cortical neurons lack binocular facilitation, even contours of similar orientation shown to the two eyes trigger such suppression. This interocular control of cortical responsiveness could serve to veto signals from one eye under conditions that would otherwise cause double vision and perceptual confusion.

Animals↗
PubMed1995

Motion and shape perception in cerebral akinetopsia.

Motion cues serve many purposes in primate vision. Consequently, akinetopsia, a defect of movement perception due to cerebral lesions, would be expected to comprise a range of motion-related defects. To address this issue we explored further the perceptual profiles in akinetopsic subject L.M. who has motion perception deficits due to extensive bilateral lesions of the dorsolateral visual association cortex that spare primary visual cortex, area V1. We used several different experimental procedures. Using random-dot motion stimuli, we showed that L.M. can still perceive global coherent motion and discriminate motion direction, yet these abilities fail even at moderate levels of background noise. L.M. also viewed a two-frame apparent (phi) motion stimulus known as a Ternus display. Her performance on this test suggests defective 'long-range' apparent motion mechanisms. These results are clearly in agreement with previous reports. Additional experiments showed that L.M. can still perceive 2-D shape and 3-D structure-from-motion (SFM). However, like motion direction discrimination, these abilities broke down at moderate levels of moving and stationary noise. Surprisingly, L.M. also had trouble perceiving 2-D shapes defined by non-motion signals including 'on' and 'off' transients, dynamic and static binocular disparity, and static texture cues. Our findings highlight the role of the visual association cortex in extracting salient information from noise.

Adult↗
PubMed1989

The Role of Horizontal Connections in Generating Long Receptive Fields in the Cat Visual Cortex.

The cells in the primary visual cortex possess numerous functional properties that are more complex and varied than those seen in the cortical input. These properties result from the network of intrinsic cortical connections running across the cortical layers and between cortical columns. In the current study we relate the long receptive fields that are characteristic of layer 6 cells to the input that these cells receive from layer 5. The axons of layer 5 pyramidal cells project over long distances within layer 6, enabling layer 6 cells to collect input from regions of cortex representing large parts of the visual field. When layer 5 was locally inactivated by injection of the inhibitory transmitter GABA, layer 6 cells lost sensitivity over the portion of their receptive fields corresponding to the inactivated region of layer 5. This suggests that the extensive convergence in the projection from layer 5 to layer 6 is responsible for generating the long receptive fields characteristic of the layer 6 cells.

Journal Article↗
PubMed2002

Acupuncture stimulation of the vision-related acupoint (Bl-67) increases c-Fos expression in the visual cortex of binocularly deprived rat pups.

Our previous study with functional magnetic resonance imaging (MRI) demonstrated that acupuncture stimulation of the vision-related acupoint, B1-67, activates the visual cortex of the human brain. As a further study on the effect of B1-67 acupuncture stimulation on the visual cortex, we examined c-Fos expression in binocularly deprived rat pups. Binocular deprivation significantly reduced the number of c-Fos-positive cells in the primary visual cortex, compared with that of normal control rat pups. Interestingly, acupuncture stimulation of B1-67 resulted in a significant increase in the number of c-Fos-positive cells in the primary visual cortex, while acupuncture stimulation of other acupoints less important for visual function had no significant effect on c-Fos expression in the primary visual cortex. The results suggest the possibility of vision-related acupoint (B1-67) having an influence over the activity of the primary visual cortex.

Acupuncture Points↗
PubMed2004

The role of spared calcarine cortex and lateral occipital cortex in the responses of human hemianopes to visual motion.

Some patients, who are rendered perimetrically blind in one hemifield by cortical lesions, nevertheless exhibit residual visual capacities within their field defects. The neural mechanism that mediates the residual visual responses has remained the topic of considerable debate. One explanation posits the subcortical visual pathways that bypass the primary visual cortex and innervate the extrastriate visual areas as the substrate that underlies the residual vision. The other explanation is that small islands of the primary visual cortex remain intact and provide the signals for residual vision. We have performed behavioral and functional magnetic resonance imaging experiments to investigate the validity of the two explanations of residual vision. Our behavioral experiments indicated that of the seven hemianopes tested, two had the ability to discriminate the direction of a drifting grating. This residual visual response was shown with fMRI to be the result of spared islands of calcarine cortical activity in one of the hemianopes, whereas only lateral occipital activity was documented in the other patient. These results indicate that the underlying neural correlates of residual vision can vary between patients. Moreover, our study emphasizes the necessity of ruling out the presence of islands of preserved function and primary visual cortex before assigning residual visual capacities to the properties of visual pathways that bypass the primary visual cortex.

Adult↗
PubMed1993

Development of blobs in the visual cortex of macaques.

We have examined the area of the primary visual cortex and the number and size of blobs within it in 10 neonatal and 11 adult rhesus monkeys. The average area of the primary visual cortex (V1) increases from 919 mm2 in newborns to 1,069 mm2 in adult animals (16%). The number of blobs decreases per unit area from an average of 5.2/mm2 at birth to 4.3/mm2 in maturity (18%). As a consequence, the number of blobs remains approximately the same during maturation, at about 4,800/hemisphere. These observations correct a preliminary report on a subset of the animals studied here (Purves and LaMantia: Proc Natl Acad Sci 87:5765, '90), in which it appeared that blob number might increase between birth and maturity. As in other regions of the developing postnatal brain, we found no net loss of modular circuitry.

Animals↗
PubMed1999

Gaze direction controls response gain in primary visual-cortex neurons.

To localize objects in space, the brain needs to combine information about the position of the stimulus on the retinae with information about the location of the eyes in their orbits. Interaction between these two types of information occurs in several cortical areas, but the role of the primary visual cortex (area V1) in this process has remained unclear. Here we show that, for half the cells recorded in area V1 of behaving monkeys, the classically described visual responses are strongly modulated by gaze direction. Specifically, we find that selectivity for horizontal retinal disparity-the difference in the position of a stimulus on each retina which relates to relative object distance-and for stimulus orientation may be present at a given gaze direction, but be absent or poorly expressed at another direction. Shifts in preferred disparity also occurred in several neurons. These neural changes were most often present at the beginning of the visual response, suggesting a feedforward gain control by eye position signals. Cortical neural processes for encoding information about the three-dimensional position of a stimulus in space therefore start as early as area V1.

Animals↗
PubMed2004

Thinning of the cerebral cortex in aging.

The thickness of the cerebral cortex was measured in 106 non-demented participants ranging in age from 18 to 93 years. For each participant, multiple acquisitions of structural T1-weighted magnetic resonance imaging (MRI) scans were averaged to yield high-resolution, high-contrast data sets. Cortical thickness was estimated as the distance between the gray/white boundary and the outer cortical surface, resulting in a continuous estimate across the cortical mantle. Global thinning was apparent by middle age. Men and women showed a similar degree of global thinning, and did not differ in mean thickness in the younger or older groups. Age-associated differences were widespread but demonstrated a patchwork of regional atrophy and sparing. Examination of subsets of the data from independent samples produced highly similar age-associated patterns of atrophy, suggesting that the specific anatomic patterns within the maps were reliable. Certain results, including prominent atrophy of prefrontal cortex and relative sparing of temporal and parahippocampal cortex, converged with previous findings. Other results were unexpected, such as the finding of prominent atrophy in frontal cortex near primary motor cortex and calcarine cortex near primary visual cortex. These findings demonstrate that cortical thinning occurs by middle age and spans widespread cortical regions that include primary as well as association cortex.

Adolescent↗