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Blindsight, conscious vision, and the role of primary visual cortex.

What is the role the primary visual cortex (V1) in vision? Is it necessary for conscious sight, as indicated by the cortical blindness that results from V1 destruction? Is it even necessary for blindsight, the nonreflexive visual functions that can be evoked with stimuli presented to cortically blind fields? In the context of this controversial issue, I present evidence indicating that not only is blindsight possible, but that conscious vision may, to a varying degree, return to formerly blind fields with time and practice even in cases where functional neuroimaging reveals no V1 activation.

Animals↗

Endstopped neurons in the visual cortex as a substrate for calculating curvature.

Neurons in the visual cortex typically respond selectively to the orientation, and velocity and direction of movement, of moving-bar stimuli. These responses are generally thought to provide information about the orientation and position of lines and edges in the visual field. Some cells are also endstopped, that is selective for bars of specific lengths. Hubel and Wiesel first observed that endstopped hypercomplex cells could respond to curved stimuli and suggested they might be involved in detection of curvature, but the exact relationship between endstopping and curvature has never been determined. We present here a mathematical model relating endstopping to curvature in which the difference in response of two simple cells gives rise to endstopping and varies in proportion to curvature. We also provide physiological evidence that endstopped cells in area 17 of the cat visual cortex are selective for curvature, whereas non-endstopped cells are not, and that some are selective for the sign of curvature. The prevailing view of edge and curve determination is that orientations are selected locally by the class of simple cortical cells and then integrated to form global curves. We have developed a computational theory of orientation selection which shows that measurements of orientation obtained by simple cells are not sufficient because there will be strong, incorrect responses from cells whose receptive fields (RFs) span distinct curves (Fig. 1). If estimates of curvature are available, however, these inappropriate responses can be eliminated. Curvature provides the key to structuring the network that underlies our theory and distinguishes it from previous lateral inhibition schemes.

Animals↗

Functional organization of lateral geniculate cells following removal of visual cortex in the newborn kitten.

When the visual cortex of a newborn kitten is removed, most neurons in the dorsal lateral geniculate nucleus degenerate, but a small population of large cells is spared. Electrophysiological recording revealed that detailed visual topography in the nucleus is abnormal and that single cells have unusually large receptive fields. These results suggest that optic axons deprived of their normal synaptic targets rearrange their connections to converge on local surviving neurons.

Animals↗

Roles of protein kinase A and protein kinase G in synaptic plasticity in the visual cortex.

Monocular deprivation leads to clear physiological and anatomical changes in the visual cortex known as ocular dominance plasticity. Protein kinase A (PKA) is necessary for ocular dominance plasticity, while protein kinase G (PKG) is not. We have now tested the role of PKA and PKG in long-term potentiation (LTP) and long-term depression (LTD). We have shown that PKA inhibitors have a major effect on both LTP and LTD in the visual cortical slices, whereas a PKG inhibitor affects LTP but not LTD. The PKA activator, 8-chloroadenosine-3',5'-monophosphorothioate, Sp-isomer (Sp-8-Cl-cAMPS), by itself induces a slowly rising form of LTP, which is occluded by theta-burst stimulation (TBS)-induced LTP. These results support the point that the PKA signaling pathway is crucial for neuronal plasticity in visual cortex, and the dissociation of the role of PKA and PKG in long-term synaptic plasticity in the visual cortex suggests that LTP alone is not sufficient to support ocular dominance plasticity, or LTD plays a more fundamental role than LTP in ocular dominance plasticity.

Animals↗

Distribution and morphology of functionally identified neurons in the visual cortex of the rat.

The distribution and morphology of functionally identified neurons were examined in the visual cortex of Long Evans pigmented rats. The results, based on qualitative and quantitative analysis of single cell spike activity, have shown that neurons in the rat visual cortex have well-defined receptive field properties and are similar to those reported for animals with more highly developed visual systems. Unlike the cat and monkey, the distribution of receptive field types appeared even throughout the visual cortex. Exception was provided by layer IV which, similar to the more 'visual' animals, contained the largest percentage of simple cells. Horseradish peroxidase injected into single, physiologically identified neurons allowed for detailed morphological characterization of functional cell types. Of the cells successfully filled with horseradish peroxidase, complex cells were pyramidal in morphology and located in layers II through VI. Simple cells were both pyramidal and non-pyramidal in appearance and were located in layers II + III and IV. Finally, hypercomplex cells were pyramidal in appearance and their perikarya were situated in layers II + III and V.

Animals↗

Amphetamine-induced recovery of visual cliff performance after bilateral visual cortex ablation in cats: measurements of depth perception thresholds.

After bilateral visual cortex ablation, cats exhibit a loss of depth perception as measured on a visual cliff, which recovers following administration of d-amphetamine. In this Study, 3 amphetamine-treated cats with visual cortex ablations showed a rapid and enduring recovery, with 2 of these animals obtaining levels of performance seen only with binocular vision, suggesting a restoration of binocular depth perception. Cats with asymmetrical lesions showed only a transient improvement during amphetamine treatment, and some animals not displaying autonomic signs of amphetamine intoxication did not improve. Saline-treated cats showed no signs of improvement, and the effect of amphetamine was blocked by the catecholaminergic antagonist haloperidol. These results indicate that amphetamine can induce an enduring recovery from a behavioral deficit after brain injury, which if left untreated would not spontaneously recover.

Amphetamine↗

Computational design and nonlinear dynamics of a recurrent network model of the primary visual cortex.

Recurrent interactions in the primary visual cortex make its output a complex nonlinear transform of its input. This transform serves preattentive visual segmentation, that is, autonomously processing visual inputs to give outputs that selectively emphasize certain features for segmentation. An analytical understanding of the nonlinear dynamics of the recurrent neural circuit is essential to harness its computational power. We derive requirements on the neural architecture, components, and connection weights of a biologically plausible model of the cortex such that region segmentation, figure-ground segregation, and contour enhancement can be achieved simultaneously. In addition, we analyze the conditions governing neural oscillations, illusory contours, and the absence of visual hallucinations. Many of our analytical techniques can be applied to other recurrent networks with translation-invariant neural and connection structures.

Animals↗

Early extrastriate activity without primary visual cortex in humans.

Damage to the primary visual cortex (V1) destroys the major source of anatomical input to extrastriate cortical areas (V2, V3, V4 and V5) and produces cortical blindness--an absence of any sensation of light and colour--in the visual field contralateral to the side of the lesion. Neuroimaging studies, nevertheless, have recently demonstrated dorsal and ventral extrastriate activation for stationary stimuli presented to the blind visual field in the absence of V1 activity in human subjects. To clarify the moment in time that visual information reaches extrastriate areas, by means of event-related potentials (ERPs) we tracked the temporal course of responses to complex visual stimuli (faces) presented in the blind field of a hemianopic patient. Stimulation of the normal visual field elicited a positive occipital deflection (P1) at 140 ms. A P1 response was also observed with stimulation of the blind field, although slightly delayed (20 ms) and reduced. Its topography and timing demonstrate that early neural activity for stationary stimuli takes place within extrastriate regions despite V1 denervation.

Adult↗

Differential effects of quinolinic acid lesions on muscarinic acetylcholine receptors in cat visual cortex during postnatal development.

Quinolinic acid (QA) lesions of neurons in cat visual cortex were combined with conventional in vitro autoradiographic methods in order to define the cellular locus of the muscarinic acetylcholine receptor (mAChR). Animals of various postnatal ages had QA unilaterally injected into the visual cortex. Four to fourteen days later they were sacrificed and processed for electron microscopy (EM) or in vitro autoradiography. QA lesions at the various postnatal ages were found to eliminate intrinsic cortical neurons and their processes while leaving intact glia, fibers of passage and axon terminals from outside the lesion zone. Autoradiograms of visual cortex labelled with [3H]QNB (which labels M1 and M2 subtypes) showed an age-dependent loss of binding sites, with the greatest decreases occurring after 65 days postnatal. Examined separately, only the M1 mAChRs labelled with [3H]pirenzepine exhibited these age-dependent alterations. The results indicate a differential distribution of the M1 mAChRs during postnatal development. The loss of receptors late in postnatal life following QA suggests a dominantly neuronal locus; the relatively small loss early in postnatal life suggests a locus on other cellular elements.

Aging↗

[Differences in the properties of 2 groups of orientation detectors of the visual cortex in the cat].

Functional properties of 149 neurons in field 17 of the visual cortex were studied in immobilized cats. They were divided into two groups. In 37% of neurons the initial and peak latencies of responses changed not more than by 10 ms in the orientation range. They were named "timer"-neurons. 63% of neurons were called "scanners". "Timers" reliably differed from "scanners" in shorter latencies, time of the discharge frequency increase, duration of the response and higher discharge frequency in all stimuli orientations. Scanners had a higher orientation selectivity, they considerably more often scanned the range of orientations in time. Neurons of the two groups had contraphase dynamics of the width of tuning orientation during response while distribution of their preferential orientations supplemented each other. In orientation columns of the visual cortex, both timers and scanners were most often found with a predominance of the latter. The columns consisting of only timers or only scanners were more seldom. The role of differences in properties of two neuronal groups in the cat visual cortex for orientation detection is discussed.

Animals↗

Functional organization of the cat visual cortex in relation to the representation of a uniform surface.

Neuronal activity in the early visual cortex has been extensively studied from the standpoint of contour representation. On the other hand, representation of the interior of a surface surrounded by a contour is much less well understood. Several studies have identified neurons activated by a uniform surface covering their receptive fields, but their distribution within the cortex is not yet known. The aim of the present study was to obtain a better understanding of the distribution of such neurons in the visual cortex. Using optical imaging of intrinsic signals, we found that there are a group of surface-responsive regions located in area 18, along the area 17/18 border, that tend to overlap the singular points of the orientation-preference map. Extracellular recordings confirmed that neurons responsive to uniform plane stimuli are accumulated in these regions. Such neurons also existed outside the surface-responsive regions around the singular points. These results suggest that there exists a functional organization related to the representation of a uniform surface in the early visual cortex.

Animals↗

How does the cerebral cortex work? Learning, attention, and grouping by the laminar circuits of visual cortex.

The organization of neocortex into layers is one of its most salient anatomical features. These layers include circuits that form functional columns in cortical maps. A major unsolved problem concerns how bottom-up, top-down, and horizontal interactions are organized within cortical layers to generate adaptive behaviors. This article models how these interactions help visual cortex to realize: (i) the binding process whereby cortex groups distributed data into coherent object representations; (ii) the attentional process whereby cortex selectively processes important events; and (iii) the developmental and learning processes whereby cortex shapes its circuits to match environmental constraints. New computational ideas about feedback systems suggest how neocortex develops and learns in a stable way, and why top-down attention requires converging bottom-up inputs to fully activate cortical cells, whereas perceptual groupings do not.

Attention↗

Experience-dependent plasticity of mouse visual cortex in the absence of the neuronal activity-dependent marker egr1/zif268.

Neuronal activity elicits a rapid increase in the expression of several immediate early genes (IEGs). To clarify a role for IEG response in activity-dependent development, we examined the contribution of the egr1/zif268 gene during visual cortical processing and plasticity in mice. We first analyzed the expression of egr1 mRNA in wild-type (WT) mice using Northern blot hybridization. In the visual cortex, expression of egr1 mRNA increased dramatically after eye opening, systemic injection of kainate, or 30 min of photostimulation after a brief (5 d) period of dark adaptation. Thus, the expression of egr1 is regulated by synaptic activity in the mouse visual cortex, as it is in other species (e.g., monkeys, cats, and rats). To evaluate whether this transcription factor is directly involved in activity-dependent plasticity, mice lacking Egr1 were deprived of the use of one eye during the developmental critical period [postnatal day 24 (P24)-P34]. Extracellular in vivo single-unit recordings from the binocular zone of the visual cortex revealed that visual responses developed normally in egr1 knock-out (KO) mice. Moreover, a similarly significant shift of responsiveness in favor of the open eye was produced in both KO and WT mice by either brief (4 d) or long-term (>2 weeks) occlusion of one eye. There was no apparent compensation among egr2, egr3, or c-fos mRNA and protein expression in the visual cortex of egr1 KO mice. Taken together, these results indicate that egr1 is a useful marker of sensory input in mice but is not intrinsically necessary for the experience-dependent plasticity of the visual cortex. Our findings underscore a mechanistic distinction between sensory plasticity and long-lasting forms of synaptic potentiation in the hippocampus, for which egr1/zif268 was recently found to be essential.

Aging↗

Muscarinic acetylcholine receptor subtypes in rat visual cortex--a comparative study using quantitative receptor autoradiography and in situ hybridization.

The laminar pattern of M1- and M2-muscarinic acetylcholine receptors (mAChR) in rat visual cortex has been compared with the distribution of the corresponding m1, m2, m3 and m4 receptor genes using both quantitative receptor autoradiography and in situ hybridization histochemistry. The laminar distribution of 3H-pirenzepine binding to M1-mAChRs in rat visual cortex shows a bimodal pattern with higher binding levels in upper layer III and deeper layer VI. In contrast, highest binding of 3H-AF-DX384 to M2-mAChRs was observed in upper layer IV (100%) and upper layer VI (about 80% of highest binding). The m1 receptor mRNA is almost homogeneously distributed throughout the visual cortex, whereas the m2mAChr mRNA predominates in layer IV with lower levels in layers I and V. The highest amounts of m3mAChR mRNA in rat visual cortex were observed in layer II, while the distribution of m4mAChR transcripts shows a bimodal pattern with peaks in layers III and upper layer VI. The distinct laminar pattern of mRNA muscarinic receptor subtypes in rat visual cortex suggest specific roles of the muscarinic receptor in visual function.

Animals↗

Evidence for a laminar organization of basal forebrain afferents to the visual cortex.

The present study shows that restriction of HRP injections to layer I within the visual cortex results in negligible retrograde labeling within the nuclei of the basal forebrain. In contrast, when the injections of either HRP or WGA-HRP are restricted to the granular and infragranular layers of visual cortex, extensive retrograde labeling occurs within the basal forebrain. Based upon these findings, we argue that the projection from the basal nucleus terminates preferentially within the deep layers of the visual cortex, and thus contributes minimally to the supragranular layers, including layer I.

Acetylcholinesterase↗

Domain specificity in visual cortex.

We investigated the prevalence and specificity of category-selective regions in human visual cortex. In the broadest survey to date of category selectivity in visual cortex, 12 participants were scanned with functional magnetic resonance imaging while viewing scenes and 19 different object categories in a blocked-design experiment. As expected, we found selectivity for faces in the fusiform face area (FFA), for scenes in the parahippocampal place area (PPA), and for bodies in the extrastriate body area (EBA). In addition, we describe 3 main new findings. First, evidence for the selectivity of the FFA, PPA, and EBA was strengthened by the finding that each area responded significantly more strongly to its preferred category than to the next most effective of the remaining 19 stimulus categories tested. Second, a region in the middle temporal gyrus that has been reported to respond significantly more strongly to tools than to animals did not respond significantly more strongly to tools than to other nontool categories (such as fruits and vegetables), casting doubt on the characterization of this region as tool selective. Finally, we did not find any new regions in the occipitotemporal pathway that were strongly selective for other categories. Taken together, these results demonstrate both the strong selectivity of a small number of regions and the scarcity of such regions in visual cortex.

Adult↗

The representation of perceived angular size in human primary visual cortex.

Two objects that project the same visual angle on the retina can appear to occupy very different proportions of the visual field if they are perceived to be at different distances. What happens to the retinotopic map in primary visual cortex (V1) during the perception of these size illusions? Here we show, using functional magnetic resonance imaging (fMRI), that the retinotopic representation of an object changes in accordance with its perceived angular size. A distant object that appears to occupy a larger portion of the visual field activates a larger area in V1 than an object of equal angular size that is perceived to be closer and smaller. These results demonstrate that the retinal size of an object and the depth information in a scene are combined early in the human visual system.

Brain Mapping↗

Image features selected by neurons of the cat primary visual cortex.

The sensitivity of neurons in field 17 of the visual cortex in cats to cross-shaped, Y-shaped, and star-shaped figures flashing in the receptive field was studied. About 40% of the neurons studied (114 of 289) were found to generate large responses (with an average response factor of 3.06 +/- 0.32) to one of the figures flashing in the center of the receptive field, as compared with the responses produced to a single bar in the optimal orientation. Most of these neurons (72%) were selectively sensitive to the shape and orientation of figures; the remainder demonstrated some degree of tuning invariance to these properties. The latent periods of responses to figures were usually shorter than those of responses to bars. Tuning parameters for bars and figures were generally related: neurons with acute orientational tuning to a bar were usually highly selective to both the configuration and the orientation people figures. Separate or combined stimulation with crosses in the center and near periphery of the receptive fields demonstrated summation, antagonism, or the lack of any interaction between these zones in producing sensitivity to crosses. Local blockade of intracortical GABAergic inhibition by microiontophoretic application of bicuculline showed that in one third of the neurons studied, sensitivity to figures was generated or enhanced by inhibition in normal conditions, while one third of cells showed suppression by inhibition, and sensitivity in the remainder was independent of inhibition. These data show that reconsideration of existing concepts of the role of field 17 in selecting only first-order shape features of images (i.e., the orientations of single lines) is needed, since almost half the neurons in the cat primary visual cortex can efficiently detect second-order features (angles and line intersections).

Action Potentials↗