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Blindsight: the role of feedforward and feedback corticocortical connections.

When human subjects suffer from a lesion to the primary visual cortex, they lose all visual percepts in the region of space that corresponds to the site of the lesion. However, they are still capable of responding to stimuli in this region when asked to 'guess' or to execute forced-choice motor commands related to these stimuli. This phenomenon, termed blindsight, is still only partly understood. Here, the possible roles of feedforward and feedback corticocortical connections in the visual brain in the understanding of blindsight are reviewed. What emerges is substantial evidence in favor of the theory that unconscious visuo-motor transformations, as in blindsight, may be executed in an entirely feedforward processing cycle, while visual awareness is critically dependent on feedback connections to the primary visual cortex.

Awareness↗

Optical imaging of the retinotopic organization of V1 in the common marmoset.

We examined the retinotopic mapping of the visual world in the primary visual cortex of the marmoset monkey using differential optical imaging. Two sets of complementary stripe-like locations were visually stimulated in turn. Their difference depicts the cortical representations of continuous bands of visual space. By rotating the sets of stripe-like locations it is possible to map different spatial axes. Analogous to the macaque we found that the V1/V2 border represented the vertical meridian, while horizontal, 45-, and 135-degree angled stripes of space were also represented in a continuous manner. We developed a new automatic method of calculating local measures of cortical magnification from our optical retinotopic maps. Using this method we found no evidence of any local anisotropies in cortical representation. Overall our results indicate that space is mapped isotropically in the primary visual cortex of the common marmoset.

Algorithms↗

How are complex cell properties adapted to the statistics of natural stimuli?

Sensory areas should be adapted to the properties of their natural stimuli. What are the underlying rules that match the properties of complex cells in primary visual cortex to their natural stimuli? To address this issue, we sampled movies from a camera carried by a freely moving cat, capturing the dynamics of image motion as the animal explores an outdoor environment. We use these movie sequences as input to simulated neurons. Following the intuition that many meaningful high-level variables, e.g., identities of visible objects, do not change rapidly in natural visual stimuli, we adapt the neurons to exhibit firing rates that are stable over time. We find that simulated neurons, which have optimally stable activity, display many properties that are observed for cortical complex cells. Their response is invariant with respect to stimulus translation and reversal of contrast polarity. Furthermore, spatial frequency selectivity and the aspect ratio of the receptive field quantitatively match the experimentally observed characteristics of complex cells. Hence, the population of complex cells in the primary visual cortex can be described as forming an optimally stable representation of natural stimuli.

Animals↗

Emergence of simple-cell receptive field properties by learning a sparse code for natural images.

The receptive fields of simple cells in mammalian primary visual cortex can be characterized as being spatially localized, oriented and bandpass (selective to structure at different spatial scales), comparable to the basis functions of wavelet transforms. One approach to understanding such response properties of visual neurons has been to consider their relationship to the statistical structure of natural images in terms of efficient coding. Along these lines, a number of studies have attempted to train unsupervised learning algorithms on natural images in the hope of developing receptive fields with similar properties, but none has succeeded in producing a full set that spans the image space and contains all three of the above properties. Here we investigate the proposal that a coding strategy that maximizes sparseness is sufficient to account for these properties. We show that a learning algorithm that attempts to find sparse linear codes for natural scenes will develop a complete family of localized, oriented, bandpass receptive fields, similar to those found in the primary visual cortex. The resulting sparse image code provides a more efficient representation for later stages of processing because it possesses a higher degree of statistical independence among its outputs.

Algorithms↗

Effect of monocular deprivation on NMDAR1 immunostaining in ocular dominance columns of the marmoset Callithrix jacchus.

We previously showed that immunoreactivity to N-Methyl-D-aspartate (NMDA) receptors in primary visual cortex of Callithrix jacchus is regulated by visual activity during the second and third postnatal months (Fonta et al., 1997). The purpose of the present study was to show that the columnar pattern of high and low NMDAR1 immunoreactivity observed in monocularly deprived animals corresponds to ocular dominance columns linked to the nondeprived and deprived eye, respectively. We compared cortical distribution of NMDAR1 receptors and the projection zones of thalamic afferents, revealed by transneuronal transport of tritiated proline, in 2-month-old, either monocularly deprived or control, marmosets. The data show that ocular dominance columns exist in 2-month-old marmosets and that a 2-week monocular deprivation by means of eyelid suture leads to a modification of the thalamo-cortical afferents organization. Experiments of neuronal tracing and immunohistochemistry performed on the same animals demonstrated that cortical domains with decreased NMDAR1 level correspond to the deprived eye columns. These investigations, coupled to the previous results, strongly suggest that the NMDA receptors, regulated by visual activity, are involved in the refining of ocular dominance columns in the primary visual cortex of juvenile marmoset.

Animals↗

Visual system: how is the retina wired up to the cortex?

A single retinal output neuron transmits to primary visual cortex through multiple pathways with different strengths. A new study in which activity was simultaneously recorded in pairs of retinal and cortical neurons provides evidence that these pathways converge on a single cortical neuron.

Animals↗

Effects of early monocular deprivation on response properties and afferents of nucleus of the optic tract in the ferret.

Effects of early monocular deprivation on visual response properties of neurons in the nucleus of the optic tract (NOT) were studied in six adult ferrets. Retinal input to NOT was investigated by orthodromic electrical stimulation of optic chiasm and optic nerves. Electrical stimulation of the ipsilateral primary visual cortex was applied to reveal the presence of a cortical pathway to NOT. All 75 neurons studied in the NOT displayed the typical strongly direction-specific response to horizontal stimulus motion; they were activated by ipsiversively directed motion (i.e. motion towards the recorded hemisphere) similar to NOT-cells in animals with normal visual experience. When tested binocularly most of the NOT-cells preferred velocities of 10 or 20 deg/s, revealing no significant difference from animals reared with normal binocular experience. The most pronounced effect of monocular deprivation was observed on ocular dominance: In the hemisphere contralateral to the non-deprived eye, NOT-cells were almost exclusively driven through the contralateral eye. In the hemisphere contralateral to the deprived eye, three of the six animals studied showed a marked dominance of the ipsilateral, non-deprived eye. In the other three animals, most neurons were binocularly activated, but over all they were significantly more strongly activated by the ipsilateral eye than found in normal animals. In four animals, dependence of ocular dominance on stimulus velocity was tested in the NOT contralateral to the deprived eye. In one of them, neurons were almost exclusively driven by the ipsilateral, non-deprived eye, irrespective of stimulus velocity. In the other animals, ocular dominance shifted from contralateral to ipsilateral with increasing stimulus velocities. Electrical stimulation of the optic chiasm revealed a mean latency of 5.53 +/- 0.48 ms. In both hemispheres, NOT-units could only be activated by stimulation of the contralateral optic nerve. Thus, no significant difference in the retinofugal conduction velocities from the deprived and the normal nerve could be detected. Of 52 cells studied, 28 (= 54%) could be activated by stimulation of primary visual cortex, mean latency being 3.9 +/- 1.7 ms. No significant difference in the percentage of cortically excitable cells between the two hemispheres as well as compared to normal animals was found (contralateral to the deprived eye: 67%, contralateral to the non-deprived eye: 53%). Therefore, cortical projections to NOT seem not to be affected by monocular deprivation. The effects of monocular deprivation in the ferret NOT, especially on ocular dominance and cortical input, are compared to the results previously described for the cat.

Afferent Pathways↗

Morphological study of neocortical areas in Rett syndrome.

Various neocortical areas from four females aged 16-24 years with Rett syndrome (RS) were investigated and compared with brains of therapy-resistant partial epilepsy (TRPE) patients (18-25 years), infantile autism (IA), and control brains (24 and 58 years). The cytoarchitecture of area 10 (frontal), area 21 (temporal), area 4 (primary motor cortex), and area 17 (primary visual cortex) was studied by the combined Klüver-Barrera (luxol fast blue and cresyl violet) standard procedure. Autofluorescence of lipofuscin, immunofluorescence of synaptic vesicle proteins [synaptophysin (p38)] and lectin-stained (Wisteria floribunda agglutinin) perineuronal nets (PNs) were studied in the cortices using dual-channel confocal laser scanning microscopy. The brains of RS females show various types of morphological/cytoarchitectonical abnormalities of single pyramidal neurons in layers II-III, and V-VII of different cortical areas. The abnormalities include mild losses of pyramidal neurons, more pronounced in layers II and III than in layers V and VII, and more evident in frontal and temporal areas than in the visual cortex. Microdysgenesis, including abnormalities due to neuronal migration disorders, was not found in RS, in contrast to the observations in TRPE patients, strongly indicating that RS is not a neuronal migration disorder. Lipofuscin distribution was normal but amounts were lower in RS cases than in control and TRPE brains. PNs were less expressed in cortices of the IA case but were clearly overexpressed in the motor cortex of RS. Quantitative analysis of p38 showed a decrease in the area occupied by p38 immunoreactivity by 20-40% in RS compared with controls. It is concluded that RS could best be explained by a postnatal synaptogenic developmental deficiency; the basic defect, however, is still completely unknown.

Adolescent↗

The effects of aging on the frequency of nerve fibers in rhesus monkey striate cortex.

In the rhesus monkey primary visual cortex, there are bundles of vertically oriented myelinated axons, which mainly contain efferent fibers originating from pyramidal cells. At the level of layer 4Cbeta, the bundles are regularly arranged and the nerve fibers in them are closely packed. In order to determine if a significant loss of intracortical nerve fibers occurs as the primate cerebral cortex ages, the frequency of vertically oriented myelinated fibers was examined at the level of layer 4Cbeta in 1 microm-thick, tangential sections. The results show no statistically significant differences in the numbers of vertically oriented fibers beneath 1 mm(2) of cortical surface between young, middle-aged, and old monkeys, and electron microscopic examination reveals few signs of degenerating axons. There is, however, an age-related breakdown of the myelin in sheaths that surround some axons. Thus, the data indicate that there is not a loss of vertically oriented myelinated fibers from the cortical gray matter during aging, although their sheaths may be altered.

Aging↗

Neurochemical compartmentation of monkey and human visual cortex: similarities and variations in calbindin immunoreactivity across species.

The compartmental organization of visual cortical neurons was examined across species of primates by directly comparing the pattern of immunoreactivity for the 28-kD vitamin D-dependent calcium-binding protein (calbindin) in area 17 of squirrel monkeys, macaques, and neurologically normal adult humans. Area 17 of macaques and squirrel monkeys was similar in that somata and processes intensely immunoreactive for calbindin were present in the same layers (II-III, IVB, and V) and in both species formed a well-stained matrix that surrounded the CO-rich puffs in layer III. These intensely calbindin-immunoreactive neurons were identified as subpopulations of GABA-immunoreactive neurons. Among the most obvious differences in the two monkey species was the distribution of calbindin-positive elements outside of layer III: a dense immunostained matrix surrounded the puffs in layers II, IVB, V, and VI of squirrel monkeys but the immunostained neurons adopted no regular pattern outside layer III in macaques. In addition, although somata lightly immunoreactive for calbindin were present in both species, they were much more abundant in squirrel monkeys than macaques. The pattern of calbindin immunostaining in human area 17 resembled that of macaques in forming an intense matrix that surrounded puffs only in layer III, yet also resembled that of squirrel monkeys by including large numbers of light immunoreactive somata. These lightly immunostained somata included a very dense population forming a prominent band in layer IVA of human visual cortex. We conclude that for layer III of primary visual cortex, a similar pattern of neuronal chemistry exists across species of primates which is related to this layer's compartmental organization. Yet for other layers, the expression of calbindin immunoreactivity varies from one species to the next, perhaps reflecting variations in other neuronal properties.

Aged↗

Localization of hemiachromatopsia.

Impaired color perception with relative preservation of form vision (achromatopsia) caused by cerebral lesions was first described over a century ago. However, for many years some researchers questioned the existence of an area of cerebral cortex apart from the primary visual cortex specialized in color processing. The development of sophisticated structural and functional neuroimaging techniques has allowed verification of the cortical structures important in color perception. We describe a case of a patient with impaired color perception in one hemifield of vision (hemiachromatopsia) and compare the localization of the lesion with previous cases described in the literature. These cases show that lesions of the ventromedial occipital cortex can impair color perception and leave form vision intact.

Aged↗

Effects of cholinergic depletion on neuron activities in the cat visual cortex.

1. Unilateral lesions of the nucleus basalis magnocellularis (nBM), a source of cholinergic projection to the cerebral cortex, were produced by injection of kainic acid in the cat. The lesions caused a significant reduction in density of choline acetyltransferase-immunoreactive terminals in the visual cortex ipsilateral to the lesions. 2. In the primary visual cortex ipsilateral to the lesions [acetylcholine (ACh)-depleted cortex], about half of the cells had weak or undetectable visual responses, whereas in the contralateral visual cortex almost all the cells had normal responsivity. The response selectivity, such as orientation and direction selectivities, of cortical cells was not affected by the depletion of ACh. 3. The microionophoretic application of ACh to cells under observation facilitated visual responses in 83% of the cells recorded from the ACh-depleted cortex, whereas it suppressed the responses in only 9%. The application of a muscarinic antagonist, atropine, to cells in the ACh-depleted cortex was ineffective, suggesting no residual ACh activity. 4. The mean current required to induce facilitation in the cortex ipsilateral to the lesion was significantly smaller than that required in the contralateral cortex and the visual cortex of the normal cat, suggesting a supersensitivity of receptors mediating the effect or a reduction in catabolism of exogenous ACh in the ACh-depleted cortex. 5. More than half of the cells that had been unresponsive to visual stimuli became clearly responsive during the ACh application. The response magnitude of cortical cells, as a whole, increased to the same degree as that observed during the ACh application in the normal cat. 6. In addition to the decrease in the average response magnitude, there was a remarkable variability in responses of cells to motion of the slit from sweep to sweep in the ACh-depleted cortex. The application of ACh to cortical cells decreased the variability of responses and consequently made the responses much more consistent. 7. These results suggest that without ACh supplied from the nBM, most of the cortical neurons could not respond briskly and consistently to excitatory inputs and that exogenously applied ACh could reverse such an impairment of cortical neurons through intact or even supersensitive postsynaptic receptors.

Acetylcholine↗

Tomographic mapping of human cerebral metabolism visual stimulation and deprivation.

Positron computed tomography was used to investigate changes in the local cerebral metabolic rate for glucose (LCMRGlc) of the visual cortex. Progressive increases in LCMRGlc were found from eyes-closed control to stimulation with white light, alternating black/white checkerboard pattern, and a complex visual scene of a park, with the associative visual cortex increasing at a faster rate than the primary visual cortex as the visual scene complexity increased. A graded decrease in LCMRGlc of the visual cortex was found with a stepwise deletion of spontaneous cell firing at the retinal, geniculate and cortical level due to lesions. Left/right metabolic symmetry of the visual cortex during monocular stimulation confirms 50% crossing of the human visual system. Neonatal blindness showed no apparent degeneration of the visual cortex and was equivalent to eyes-closed controls. The interictal state of a patient with visual seizures demonstrated a hypometabolic visual cortex with a 2.5-fold increase in metabolism during an ictal visual hallucination.

Adult↗

Immunohistochemical localization of calcium-binding proteins, parvalbumin and calbindin-D 28k, in the adult and developing visual cortex of cats: a light and electron microscopic study.

In the cat primary visual cortex, we investigated with immunohistochemical techniques the developmental changes in the cellular and subcellular localization of the Ca2+-binding proteins parvalbumin (PV) and calbindin-D 28K (CBP), in order to determine whether there is a correlation between the expression of Ca2+-dependent processes and the time course of the critical period for use-dependent plasticity. On the 54th day of gestation and at 1 week postnatally, both calcium-binding proteins were present only in a subpopulation of neurons in layers V and VI. During subsequent maturation, the number of PV(+) and CBP(+) neurons increased significantly and labeled cells were detected in more superficial layers. Moreover, the homogeneous labeling of some CBP(+) neurons in layers IV to VI decreased and changed to a punctate pattern. In adult cats PV(+) neurons were evenly distributed throughout layers II to VI, whereas CBP(+) neurons were concentrated in layers II/III. Only a few immunoreactive cells had morphological features characteristic of pyramidal cells; the large majority were nonpyramidal. Electron microscopy confirmed the presence of PV- and CBP-reaction product within the perikarya, axons, and dendrites of labeled cells. It was associated preferentially with microtubules, postsynaptic densities, and intracellular membranes. Immunoreactive neurons received immunonegative asymmetric synapses on their dendritic shafts and made symmetric synaptic contacts with labeled and unlabeled somata and with unlabeled dendritic shafts. The large number and widespread distribution of immunoreactive neurons implies that PV and CBP play an important role in the regulation of calcium-dependent processes in the visual cortex. Furthermore, the developmental redistribution of PV and CBP points to changes in the organization of Ca2+-dependent processes during maturation.

Aging↗

Spatial summation in lateral geniculate nucleus and visual cortex.

We have compared the spatial summation characteristics of cells in the primary visual cortex with those of cells in the dorsal lateral geniculate nucleus (LGN) that provide the input to the cortex. We explored the influence of varying the diameter of a patch of grating centred over the receptive field and quantitatively determined the optimal summation diameter and the degree of surround suppression for cells at both levels of the visual system using the same stimulus parameters. The mean optimal summation size for LGN cells (0.90 degrees) was much smaller than that of cortical cells (3.58 degrees). Virtually all LGN cells exhibited strong surround suppression with a mean value of 74%+/-1.61% SEM for the population as a whole. This potent surround suppression in the cells providing the input to the cortex suggests that cortical cells must integrate their much larger summation fields from the low firing rates associated with the suppression plateau of the LGN cell responses. Our data suggest that the strongest input to cortical cells will arise from geniculate cells representing areas of visual space located at the borders of a visual stimulus. We suggest that analysis of response properties by patterns centred over the receptive fields of cells may give a misleading impression of the process of the representation. Analysis of pattern terminations or salient borders over the receptive field may provide much more insight into the processing algorithms involved in stimulus representation.

Animals↗

Cognitive control signals in visual cortex: flashes meet spotlights.

At the intersection of two intensely belabored fields, primary visual cortex (V1) function and neural mechanisms of cognitive control, Jack et al. (in this issue of Neuron) report a neural signal that is neither related to stimulus representation nor spatial attention. Instead, this endogenous signal correlates with task structure and raises new questions.

Afferent Pathways↗

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.

Animals↗

Synchronous activity in cat visual cortex encodes collinear and cocircular contours.

We explored how contour information in primary visual cortex might be embedded in the simultaneous activity of multiple cells recorded with a 100-electrode array. Synchronous activity in cat visual cortex was more selective and predictable in discriminating between drifting grating and concentric ring stimuli than changes in firing rate. Synchrony was found even between cells with wholly different orientation preferences when their receptive fields were circularly aligned, and membership in synchronous groups was orientation and curvature dependent. The existence of synchrony between cocircular cells reinforces its role as a general mechanism for contour integration and shape detection as predicted by association field concepts. Our data suggest that cortical synchrony results from common and synchronous input from earlier visual areas and that it could serve to shape extrastriate response selectivity.

Animals↗