Early development of visual cortex in human fetuses

Prenatal development of visual cortex (area 17) was studied in human fetuses of 8-9, 13-15 and 16-18 weeks of gestation, with a view to analyse the early critical events. Under light microscope, five zones of development were seen in all the age groups. The total thickness of cortex of area 17 as well as that of its cortical plate was measured with the help of camera lucida. It was observed that the total thickness of the cortex increased with increase in age. Diversity in the shape, size, staining intensity and arrangement of neurons was noted in the different zones. Most of the cells were found to have a thin rim of cytoplasm and a prominent nucleus with multiple nucleoli. The cells in subventricular zone and cortical plate were regularly arranged in vertical rows while in other zones, they were irregularly scattered. Several mitotic figures were seen in the ventricular zone at 8-9 weeks but later they were also noticed in subventricular and intermediate zones. In the later ages the mitotic figures were observed to be fewer in the ventricular zone. No mitosis was seen in cortical plate at any age period.     

Changes in choline acetyltransferase activity and high-affinity choline uptake, but not in acetylcholinesterase activity and muscarinic cholinergic receptors, in rat somatosensory cortex after sciatic nerve injury

Selected cholinergic markers (choline acetyltransferase, acetylcholinesterase, muscarinic acetylcholine receptor, high-affinity choline uptake) were studied in the hindlimb representation areas of the rat somatosensory cortex and within the visual cortex 1 to 63 days after unilateral transection of the sciatic nerve. In the contralateral somatosensory cortex, peripheral deafferentation resulted in a significant reduction of choline acetyltransferase activity (by 15%) 3 days after sciatic nerve injury, and in a significant reduction of high-affinity choline uptake (by 30%) 1 day after nerve transection, in comparison to untreated control rats. Investigations in individual cortical layers revealed that the decrease of both choline acetyltransferase activity and high-affinity choline uptake sites was mainly due to reductions in cortical layer V. Acetylcholinesterase activity and [3H]quinuclidinyl benzilate binding to muscarinic acetylcholine receptors were not affected by unilateral transection of the sciatic nerve. In the ipsilateral somatosensory cortex, as well as in the visual cortex at both cortical hemispheres, no significant changes in the cholinergic parameters studied could be detected. The data indicate that peripheral deafferentation of the somatosensory cortex results in a transient change of presynaptic cholinergic parameters within the affected somatosensory area as early as 1 to 3 days after the lesion; thus, they emphasize the involvement of cholinergic mechanisms in cortical reorganizational events.     

Behavioral detection of electrical microstimulation in different cortical visual areas

The extent to which areas in the visual cerebral cortex differ in their ability to support perceptions has been the subject of considerable speculation. Experiments examining the activity of individual neurons have suggested that activity in later stages of the visual cortex is more closely linked to perception than that in earlier stages [1-9]. In contrast, results from functional imaging, transcranial magnetic stimulation, and lesion studies have been interpreted as showing that earlier stages are more closely coupled to perception [10-15]. We examined whether neuronal activity in early and later stages differs in its ability to support detectable signals by measuring behavioral thresholds for detecting electrical microstimulation in different cortical areas in two monkeys. By training the animals to perform a two-alternative temporal forced-choice task, we obtained criterion-free thresholds from five visual areas--V1, V2, V3A, MT, and the inferotemporal cortex. Every site tested yielded a reliable threshold. Thresholds varied little within and between visual areas, rising gradually from early to later stages. We similarly found no systematic differences in the slopes of the psychometric detection functions from different areas. These results suggest that neuronal signals of similar magnitude evoked in any part of visual cortex can generate percepts.     

Retinotopic maps, spatial tuning, and locations of human visual areas in surface coordinates characterized with multifocal and blocked FMRI designs

The localization of visual areas in the human cortex is typically based on mapping the retinotopic organization with functional magnetic resonance imaging (fMRI). The most common approach is to encode the response phase for a slowly moving visual stimulus and to present the result on an individual's reconstructed cortical surface. The main aims of this study were to develop complementary general linear model (GLM)-based retinotopic mapping methods and to characterize the inter-individual variability of the visual area positions on the cortical surface. We studied 15 subjects with two methods: a 24-region multifocal checkerboard stimulus and a blocked presentation of object stimuli at different visual field locations. The retinotopic maps were based on weighted averaging of the GLM parameter estimates for the stimulus regions. In addition to localizing visual areas, both methods could be used to localize multiple retinotopic regions-of-interest. The two methods yielded consistent retinotopic maps in the visual areas V1, V2, V3, hV4, and V3AB. In the higher-level areas IPS0, VO1, LO1, LO2, TO1, and TO2, retinotopy could only be mapped with the blocked stimulus presentation. The gradual widening of spatial tuning and an increase in the responses to stimuli in the ipsilateral visual field along the hierarchy of visual areas likely reflected the increase in the average receptive field size. Finally, after registration to Freesurfer's surface-based atlas of the human cerebral cortex, we calculated the mean and variability of the visual area positions in the spherical surface-based coordinate system and generated probability maps of the visual areas on the average cortical surface. The inter-individual variability in the area locations decreased when the midpoints were calculated along the spherical cortical surface compared with volumetric coordinates. These results can facilitate both analysis of individual functional anatomy and comparisons of visual cortex topology across studies.     

BDNF regulates the maturation of inhibition and the critical period of plasticity in mouse visual cortex.

Maturation of the visual cortex is influenced by visual experience during an early postnatal period. The factors that regulate such a critical period remain unclear. We examined the maturation and plasticity of the visual cortex in transgenic mice in which the postnatal rise of brain-derived neurotrophic factor (BDNF) was accelerated. In these mice, the maturation of GABAergic innervation and inhibition was accelerated. Furthermore, the age-dependent decline of cortical long-term potentiation induced by white matter stimulation, a form of synaptic plasticity sensitive to cortical inhibition, occurred earlier. Finally, transgenic mice showed a precocious development of visual acuity and an earlier termination of the critical period for ocular dominance plasticity. We propose that BDNF promotes the maturation of cortical inhibition during early postnatal life, thereby regulating the critical period for visual cortical plasticity.     

The future of mapping sensory cortex in primates: three of many remaining issues

After 100 years of progress in understanding the organization of cerebral cortex, three issues have persisted over the last 35 years, which are revisited in this paper. First, is V3 an established or questionable area of visual cortex? Second, does taste cortex include part of area 3b (S1 proper) and other somatosensory areas? Third, is primary auditory cortex, A1, of primates the homologue of A1 in cats? The existence of such questions about even the early stages of cortical processing reflects the difficulties in mapping cerebral cortex, and reminds us that the era of basic discovery is far from over.     

Responses to lightness variations in early human visual cortex

Lightness is the apparent reflectance of a surface, and it depends not only on the actual luminance of the surface but also on the context in which the surface is viewed [1-10]. The cortical mechanisms of lightness processing are largely unknown, and the role of early cortical areas is still a matter of debate [11-17]. We studied the cortical responses to lightness variations in early stages of the human visual system with functional magnetic resonance imaging (fMRI) while observers were performing a demanding fixation task. The set of dynamically presented visual stimuli included the rectangular version of the classic Craik-O'Brien stimulus [3, 18, 19] and a variant that led to a weaker lightness effect, as well as a pattern with actual luminance variations. We found that the cortical activity in retinotopic areas, including the primary visual cortex (V1), is correlated with context-dependent lightness variations.     

Visual Function and Cortical Organization in Carriers of Blue Cone Monochromacy

Carriers of blue cone monochromacy have fewer cone photoreceptors than normal. Here we examine how this disruption at the level of the retina affects visual function and cortical organization in these individuals. Visual resolution and contrast sensitivity was measured at the preferred retinal locus of fixation and visual resolution was tested at two eccentric locations (2.5° and 8°) with spectacle correction only. Adaptive optics corrected resolution acuity and cone spacing were simultaneously measured at several locations within the central fovea with adaptive optics scanning laser ophthalmoscopy (AOSLO). Fixation stability was assessed by extracting eye motion data from AOSLO videos. Retinotopic mapping using fMRI was carried out to estimate the area of early cortical regions, including that of the foveal confluence. Without adaptive optics correction, BCM carriers appeared to have normal visual function, with normal contrast sensitivity and visual resolution, but with AO-correction, visual resolution was significantly worse than normal. This resolution deficit is not explained by cone loss alone and is suggestive of an associated loss of retinal ganglion cells. However, despite evidence suggesting a reduction in the number of retinal ganglion cells, retinotopic mapping showed no reduction in the cortical area of the foveal confluence. These results suggest that ganglion cell density may not govern the foveal overrepresentation in the cortex. We propose that it is not the number of afferents, but rather the content of the information relayed to the cortex from the retina across the visual field that governs cortical magnification, as under normal viewing conditions this information is similar in both BCM carriers and normal controls.     

Changes in Choline Acetyltransferase Activity and High-Affinity Choline Uptake,but Not in Acetylcholinesterase Activity and Muscarinic Cholinergic Receptors,in Rat Somatosensory Cortex after Sciatic Nerve Injury

Selected cholinergic markers (choline acetyltransferase, acetylcholinesterase, muscarinic acetylcholine receptor, high-affinity choline uptake) were studied in the hindlimb representation areas of the rat somatosensory cortex and within the visual cortex 1 to 63 days after unilateral transection of the sciatic nerve. In the contralateral somatosensory cortex, peripheral deafferentation resulted in a significant reduction of choline acetyltransferase activity (by 15%) 3 days after sciatic nerve injury, and in a significant reduction of high-affinity choline uptake (by 30%) 1 day after nerve transection, in comparison to untreated control rats. Investigations in individual cortical layers revealed that the decrease of both choline acetyltransferase activity and high-affinity choline uptake sites was mainly due to reductions in cortical layer V. Acetylcholinesterase activity and [³H]quinuclidinyl benzilate binding to muscarinic acetylcholine receptors were not affected by unilateral transection of the sciatic nerve. In the ipsilateral somatosensory cortex, as well as in the visual cortex at both cortical hemispheres, no significant changes in the cholinergic parameters studied could be detected. The data indicate that peripheral deafferentation of the somatosensory cortex results in a transient change of presynaptic cholinergic parameters within the affected somatosensory area as early as 1 to 3 days after the lesion; thus, they emphasize the involvement of cholinergic mechanisms in cortical reorganizational events.     

The organization of somatosensory cortex in the short-tailed opossum ( Monodelphis domestica )

The organization of neocortex in the short-tailed opossum ( Monodelphis domestica ) was explored with multiunit microelectrode recordings from middle layers of cortex. Microelectrode maps were subsequently related to the chemoarchitecture of flattened cortical preparations, sectioned parallel to the cortical surface and processed for either cytochrome oxidase (CO) or NADPH-diaphorase (NADPHd) histochemistry. The recordings revealed the presence of at least two systematic representations of the contralateral body surface located in a continuous strip of cortex running from the rhinal sulcus to the medial wall. The primary somatosensory area (S1) was located medially while secondary somatosensory cortex (S2) formed a laterally located mirror image of S1. Auditory cortex was located in lateral cortex at the caudal border of S2, and some electrode penetrations in this area responded to both auditory and somatosensory stimulation. Auditory cortex was outlined by a dark oval visible in flattened brain sections. A large primary visual cortex (V1) was located at the caudal pole of cortex, and also consistently corresponded to a large chemoarchitecturally visible oval. Cortex just rostral and lateral to V1 responded to visual stimulation, while bimodal auditory/visual responses were obtained in an area between V1 and somatosensory cortex. The results are compared with brain organization in other marsupials and with placentals and the evolution of cortical areas in mammals is discussed.     

The organization of somatosensory cortex in the short-tailed opossum (Monodelphis domestica)

The organization of neocortex in the short-tailed opossum (Monodelphis domestica) was explored with multiunit microelectrode recordings from middle layers of cortex. Microelectrode maps were subsequently related to the chemoarchitecture of flattened cortical preparations, sectioned parallel to the cortical surface and processed for either cytochrome oxidase (CO) or NADPH-diaphorase (NADPHd) histochemistry. The recordings revealed the presence of at least two systematic representations of the contralateral body surface located in a continuous strip of cortex running from the rhinal sulcus to the medial wall. The primary somatosensory area (S1) was located medially while secondary somatosensory cortex (S2) formed a laterally located mirror image of S1. Auditory cortex was located in lateral cortex at the caudal border of S2, and some electrode penetrations in this area responded to both auditory and somatosensory stimulation. Auditory cortex was outlined by a dark oval visible in flattened brain sections. A large primary visual cortex (V1) was located at the caudal pole of cortex, and also consistently corresponded to a large chemoarchitecturally visible oval. Cortex just rostral and lateral to V1 responded to visual stimulation, while bimodal auditory/visual responses were obtained in an area between V1 and somatosensory cortex. The results are compared with brain organization in other marsupials and with placentals and the evolution of cortical areas in mammals is discussed.     

Correction of Distortion in Flattened Representations of the Cortical Surface Allows Prediction of V1-V3 Functional Organization from Anatomy

Several domains of neuroscience offer map-like models that link location on the cortical surface to properties of sensory representation. Within cortical visual areas V1, V2, and V3, algebraic transformations can relate position in the visual field to the retinotopic representation on the flattened cortical sheet. A limit to the practical application of this structure-function model is that the cortex, while topologically a two-dimensional surface, is curved. Flattening of the curved surface to a plane unavoidably introduces local geometric distortions that are not accounted for in idealized models. Here, we show that this limitation is overcome by correcting the geometric distortion induced by cortical flattening. We use a mass-spring-damper simulation to create a registration between functional MRI retinotopic mapping data of visual areas V1, V2, and V3 and an algebraic model of retinotopy. This registration is then applied to the flattened cortical surface anatomy to create an anatomical template that is linked to the algebraic retinotopic model. This registered cortical template can be used to accurately predict the location and retinotopic organization of these early visual areas from cortical anatomy alone. Moreover, we show that prediction accuracy remains when extrapolating beyond the range of data used to inform the model, indicating that the registration reflects the retinotopic organization of visual cortex. We provide code for the mass-spring-damper technique, which has general utility for the registration of cortical structure and function beyond the visual cortex.     

Predicting functional properties of visual cortex from an evolutionary scaling law

The number of neurons in the primary visual cortex (V1) is, across primate species, related to the number of neurons in the visual thalamus (the lateral geniculate nucleus [LGN]) by a power law with an exponent of 3/2. This evolutionary scaling law is explained by a simple relation according to which the fineness of resolution in cortex is related to the number of neurons in the area of cortex used to process the information from a single point of light (the point-spread area). The same theory provides a link between two functional properties of the visual cortex, the areal cortical magnification factor (ACMF) and the receptive field (RF) area.     

Structural rearrangements of the cerebral cortex in children and adolescents

The cortical formations of the brain involved in visual functions (the occipital and temporo-parieto- occipital areas, the oculomotor area of the prefrontal cortex), as well as the motor cortex in the representation zone of the arm and the medial region of the frontal cortex adjacent to the limbic lobe, were studied in post-mortem material. The thickness of the cortex and cortical layer III, the sizes of pyramidal neurons, the specific volumes of neurons and intracortical vessels were studied in subjects of both sexes, from birth to the age of 20 years, at yearly intervals (103 observations) using histological techniques, computer morphometric and stereological analysis. The thickness of the cortex of the cerebral hemispheres was observed to intensively increase from birth to the age of 3 years in the occipital, temporo-parieto-occipital and prefrontal cortical areas involved in visual recognition processes. The increase in thickness of the cerebral cortex continues until the age of 6 in the occipital cortex and in the oculomotor area, until the age of 7 years in the temporo-parietooccipital area and the medial prefrontal area, and until the age of 8–9 years in the motor cortex. The sizes of pyramidal neurons increase until the age of 6 years in the motor cortex, until the age of 8 years on the medial surface of the frontal lobe, and until the age of 9–10 years in the temporo-parieto-occipital area and in the dorsolateral area of the prefrontal cortex. The specific volume of neurons and blood vessels in the cortex of the cerebral hemispheres decreases and the volume of intracortical fibers increases throughout the ascending ontogeny, which is manifested most intensively in the prefrontal cortex.     

Organization of the cortico-ponto-cerebellar pathway to the dorsal paraflocculus. An experimental study with anterograde and retrograde transport of WGA-HRP in the cat.

To reveal the organization and relative magnitude of connections from various parts of the cerebral cortex to the dorsal paraflocculus via the pontine nuclei, WGA-HRP was injected in the dorsal paraflocculus in conjunction with injection of the same tracer in various parts of the cerebral cortex in 17 cats. Termination areas of cortical fibres (anterogradely labelled) and pontine neurons projecting to the dorsal paraflocculus (retrogradely labelled) were carefully plotted in serial transverse sections. As an average of countings in ten cats, 90% of the labelled cells were found in the pontine nuclei contralateral to the injection, and the majority (70%) were located in the rostral half of the nuclei. The highest degree of overlap between anterograde and retrograde labelling was found after injections of the parietal association cortex (areas 5 and 7). In an experiment with double anterograde tracing, it was shown that both area 5 and 7 contribute substantially to the cerebral inputs to the dorsal paraflocculus. High degree of overlap also occurred after injections of several visual cortical areas (areas 17, 18, 19, 20 and the posteromedial lateral suprasylvian visual area, PMLS). Cases with injections restricted to individual visual areas indicate that they all contribute to the parafloccular input. Considerably less overlap occurred after injections of the primary sensorimotor region (SI, MI) and second somatosensory area (SII), while the supplementary motor area, the auditory cortex and gyrus cinguli probably have no or very restricted access to the dorsal paraflocculus. It is concluded that the dorsal paraflocculus has its major cortical input from the parietal association cortex and the visual cortical areas. Since all the various cortical regions studied project to largely different parts of the pontine nuclei, and overlap with neurons projecting to the dorsal paraflocculus takes place at numerous places, it follows that the pontine neurons projecting to the dorsal paraflocculus must consist of many subgroups differing with regard to their cortical input.     

用脑光学成像精确测定猫初级视皮层视野拓扑投射关系

利用基于脑内源信号的光学成像和二维互相关分析的方法,对猫初级视皮层17区的视野拓扑离心度(即视网膜-皮层拓扑关系)进行了精确测量。当采用在同一屏幕内处于上下视野的、方位互相垂直的两个相邻光栅刺激时,皮层中一部分区域的绝大部分细胞因同时兴奋而导致方位功能图模糊不清。将这种方位功能图和用单一方位(水平或垂直)全屏光栅刺激所得到的功能图进行比较,通过计算每一像素的互相关系数,从而获得皮层的精确视野拓扑离心度。同时用电生理的方法测量了同一视皮层内的单细胞的感受野位置,证明这种方法得到的视野离心度和光学记录方法得到的相同。因此,本研究为大面积地确定视皮层细胞感受野在视野中的位置提供了一种快速和较准确的方法。     

Retino-cortical information transmission achievable with a retina implant

Blind subjects with photoreceptor degeneration perceive phosphenes when their intact retinal ganglion cells are stimulated electrically. Is this approach suitable for transmitting enough information to the visual cortex for partially restoring vision? We stimulated the retina of anesthetized cats electrically and visually while recording the responses in the visual cortex. Transmission of retino-cortical information T was quantified by information theory. T was 20-160 bit/s (per stimulation and recording site) with random electrical or visual impulse stimulation at rates between 20 and 40 s-1. While increasing spatial density of independent electrical stimulation channels T did not saturate with 7 electrodes/mm2 retina. With seven electrodes up to 500 bit/s was transmitted to 15 cortical recording sites. Electrical stimulation basically employs temporal stimulus patterns. They are intimately linked with intensity/contrast information coded by the spike density of retinal ganglion cells. From the cortical information spread we estimated the spatial resolution as 0.5mm cortex corresponding to 0.5-1.0 degrees visual angle. If the human cortex can receive and decode the information transmitted by a retina implant, our quantitative results measured in cats suggest that visuo-motor coordination and object recognition in many in- and out-door situations will be possible.     

Dynamic shape integration in extrastriate cortex

BACKGROUND: In anorthoscopic viewing conditions, observers can perceive a moving object through a narrow slit even when only portions of its contour are visible at any time. We used fMRI to examine the contribution of early and later visual cortical areas to dynamic shape integration. Observers' success at integrating the shape of the slit-viewed object was manipulated by varying the degree to which the stimulus was dynamically distorted. Line drawings of common objects were either moderately distorted, strongly distorted, or shown undistorted. Phenomenologically, increasing the stimulus distortion made both object shape and motion more difficult to perceive.RESULTS: We found that bilateral cortical activity in portions of the ventral occipital cortex, corresponding to known object areas within the lateral occipital complex (LOC), was inversely correlated with the degree of stimulus distortion. We found that activity in left MT+, the human cortical area specialized for motion, showed a similar pattern as the ventral occipital region. The LOC also showed greater activity to a fully visible moving object than to the undistorted slit-viewed object. Area MT+, however, showed more equivalent activity to both the slit-viewed and fully visible moving objects.CONCLUSIONS: In early retinotopic cortex, the distorted and undistorted stimuli elicited the same amount of activity. Higher visual areas, however, were correlated with the percept of the coherent object, and this correlation suggests that the shape integration is mediated by later visual cortical areas. Motion information from the dorsal stream may project to the LOC to produce the shape percept.     

Visual thalamic afferents to area 29 of the rat limbic cortex

Afferent connections of the retrosplenial area of the rat limbic cortex were investigated by the retrograde horseradish peroxidase axon transport method. After injection of horseradish peroxidase (HRP) into area 29 of the cortex, HRP-labeled cells were found in the dorsal part of the lateral geniculate body and the posterolateral, pretectal, and anterior dorsal thalamic nuclei. Connections were found between cortical area 29 and visual projection areas (areas 17 and 18a) and with area 29 on the contralateral side of the brain. The results are evidence that all the principal visual structures of the thalamus and the visual cortical projection area form direct projections to the retrosplenial cortex.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 14, No. 2, pp. 135–139, March–April, 1982.     

The Development of Visual Areas Depends Differently on Visual Experience

Visual experience plays an important role in the development of the visual cortex; however, recent functional imaging studies have shown that the functional organization is preserved in several higher-tier visual areas in congenitally blind subjects, indicating that maturation of visual areas depend unequally on visual experience. In this study, we aim to validate this hypothesis using a multimodality MRI approach. We found increased cortical thickness in the congenitally blind was present in the early visual areas and absent in the higher-tier ones, suggesting that the structural development of the visual cortex depends hierarchically on visual experience. In congenitally blind subjects, the decreased resting-state functional connectivity with the primary somatosensory cortex was more prominent in the early visual areas than in the higher-tier ones and were more pronounced in the ventral stream than in the dorsal one, suggesting that the development of functional organization of the visual cortex also depends differently on visual experience. Moreover, congenitally blind subjects showed normal or increased functional connectivity between ipsilateral higher-tier and early visual areas, suggesting an indirect corticocortical pathway through which somatosenroy information can reach the early visual areas. These findings support our hypothesis that the development of visual areas depends differently on visual experience.