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Transcranial magnetic stimulation of the occipital pole interferes with verbal processing in blind subjects.

Recent neuroimaging studies in blind persons show that the occipital cortex, including the primary visual cortex (V1), is active during language-related and verbal-memory tasks. No studies, however, have identified a causal link between early visual cortex activity and successful performance on such tasks. We show here that repetitive transcranial magnetic stimulation (rTMS) of the occipital pole reduces accuracy on a verb-generation task in blind subjects, but not in sighted controls. An analysis of error types revealed that the most common error produced by rTMS was semantic; phonological errors and interference with motor execution or articulation were rare. Thus, in blind persons, a transient 'virtual lesion' of the left occipital cortex interferes with high-level verbal processing

Analysis of Variance↗

Two inhibitory postsynaptic potentials, and GABAA and GABAB receptor-mediated responses in neocortex of rat and cat.

1. Pyramidal neurones from layers II and III of the rat primary somatosensory cortex and cat primary visual cortex were studied in vitro. Inhibitory postsynaptic potentials (IPSPs) and responses to exogenously applied gamma-aminobutyric acid (GABA) and its analogue baclofen were characterized. The results from rats and cats were very similar. 2. Single electrical stimuli to deep cortical layers evoked a sequence of PSPs in the resting neurone: (a) an initial, brief excitation (EPSP), (b) a short-latency, fast inhibition (the f-IPSP) and (c) a long-latency, more prolonged inhibition (the l-IPSP). The f-IPSP was accompanied by a large conductance increase (about 70-90 nS) and reversed polarity at -75 mV; the l-IPSP displayed a relatively small conductance increase (about 10-20 nS) and reversed at greater than -90 mV. 3. Focal application of GABA near the soma evoked a triphasic response when measured near the threshold voltage for action potentials: (a) the GABAhf (hyperpolarizing, fast) phase was very brief and was generated by a large conductance increase with a reversal potential of -78 mV, (b) the GABAd (depolarizing) phase also had a high conductance but reversed at -51 mV, (c) the GABAhl (hyperpolarizing, long-lasting) phase had a relatively low conductance and reversed at -70 mV. The GABAhf response was specifically localized to the soma, whereas the apical or basilar dendrites generated predominantly GABAd responses. 4. Baclofen, a selective GABAB receptor agonist, caused a small (about 2 mV), slow hyperpolarization of the resting potential, which reversed at -90 mV. Saturating baclofen doses increased membrane conductance by a maximum of about 12 nS. Baclofen depressed the amplitude and conductance of PSPs; when baclofen was focally applied near the soma. IPSPs were selectively depressed. 5. The GABAA receptor antagonists bicuculline methiodide or picrotoxin (10 microM) greatly depressed f-IPSPs, but either enhanced or did not affect l-IPSPs. Concomitantly, GABAhf and GABAd responses were antagonized, leaving a more prominent GABAhl response that reversed polarity at a more negative level of -87 mV. Baclofen responses were unaffected by bicuculline and picrotoxin. Extracellular barium abolished the baclofen response, and shifted the reversal potentials of the GABAd and GABAhl responses in the positive direction; the GABAhf response was unaffected. 6. Both focal GABA and f-IPSPs strongly depressed the intrinsic excitability of pyramidal neurones. Each greatly increased spike threshold and abolished or vastly reduced the capacity of the cells to fire repetitively during intense stimuli.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗

[Expansion and thickness of the visual cortex of microphthalmic mice and healthy Wistar animals].

In connection with our studies about the development and the structure of the visual system of the brain of normal and genetically microphthalmic mice in this paper we have investigated the extend of the visual cortex and the thickness of the primary visual cortex (area Oc1). The most important results are: The size of the surface of the neocortex and of the areas Oc1 and Oc2m shows no significant differences between the normal and the microphthalmic mice. The whole thickness of area Oc1 is at day 20 in the microphthalmic mice significantly greater than in normal animals. In the microphthalmic mice lamina I, III and V are thicker than in normal mice. Lamina IV is in microphthalmic mice significantly thinner than in normal animals. The interpretation of these results is difficult because at PD 20 the cortex and his laminae are not full developed. But there are in the microphthalmic mice not such clear changes (reduction) on the cortical level as in the subcortical structures of the visual system. The reduction of the thickness of lamina IV maybe caused by a reduced projection from the lateral geniculate nucleus.

Animals↗

Single unit studies in the visual cortex of rodents.

This paper deals with the visual topography in the rat and hamster visual cortex, and with receptive field maps and trigger features of visual units in rat visual cortex. The visuotopic organization of the primary visual cortex (area 17) and the extrastriate visual regions surrounding it (areas 18a and 18) was studied in gray rats and hamsters using standard microelectrode techniques. The results confirm and extend previous observations in the rat and reveal in hamster an organization more complex than previously reported. In the rat, apart from the representation of the contralateral visual field (VF) in area 17, in which the upper VF is represented caudally and the nasal VF laterally, there are additional representations of the VF in the extrastriate cortex. In lateral extrastriate cortex (area 18a) there are at least four such representations, namely latero-medial (LM), antero lateral (AL), latero intermediate (LI) and latero-lateral (LL). In LM (second visual area) the upper VF is represented caudally at the nasal FV, medially, being thus a mirror image of V1. In AL (third visual area) the upper VF is represented rostrally and the nasal VF, medially, being thus a mirror image of LM. In LI the upper VF is medial, and the nasal VF, lateral, being thus a mirror image of LM, or a reduced copy of V1. In LL, the upper VF is caudal and the nasal VF, medial, being thus a mirror image of LI, or a reduced copy of LM. In medial extrastriate cortex (area 18) there are two representations of the temporal VF, labelled antero-medial (AM) and postero medial (PM). In AM, the upper temporal VF is medial and the lower temporal VF, lateral, the extreme temporal field being rostral. The 30 degree azimuth provides the boundary between AM and PM. Thus, AM is organized as a counter clock wise rotation by 90 degrees of the V1 representation. In PM, the upper lower VF topography is like in AM, but the extreme temporal VF is caudal, being thus a mirror image of AM. Our preliminary study of visual topography is hamster shows an organization at least as elaborate as that of rat. Finally, we have compared RFs of striate vs extrastriate units. Our extrastriate units showed similar trigger features and field maps as in V1, but with larger RF sizes and a proportion of rapidly habituating cells. The percent distribution of field types, at least in LM and AM was similar to that in V1.

Animals↗

Increased oxidative metabolism in middle suprasylvian cortex following removal of areas 17 and 18 from newborn cats.

We measured changes in metabolic activity in middle suprasylvian (MS) cortex of cats subjected to early or late removal of areas 17 and 18 to localize shifts in activity possibly indicative of regions within MS cortex that may receive expanded inputs and be involved in the sparing of some visual behaviors following early primary visual cortex damage. Cytochrome oxidase (CO) activity was measured in MS cortex of mature, intact cats and of others with areas 17 and 18 removed in adulthood (P180), or on postnatal day 28 (P28) or postnatal day 1 (P1). Not less than 9 months after the ablation, brain sections were prepared and reacted for the presence of CO. The density of CO reactivity in each of the six cortical layers in MS cortex was measured and standardized against densities from ventral periaqueductal gray or hypothalamus on the same section. Following lesions on P1, significant increases in CO activity occurred in deep layer III and in layer IV of the medial bank of the MS sulcus, including all of area PMLS and the posterior portion of AMLS. In contrast, there were no significant differences in the level of CO activity among P28, P180, or intact cats for any of the cortical layers, and all had lower levels than the P1 cats. This metabolic change provides an anatomical marker for localizing adjustments in MS cortex and can be linked to amplified projections into MS cortex from the thalamus (LPm and A and C laminae of the dorsal lateral geniculate nucleus) and ventral posterior suprasylvian cortex following P1 ablations. Furthermore, this neurochemical analysis implicates a distinct region of MS cortex as the cortical locus of some spared visual functions following early primary visual cortex damage.

Analysis of Variance↗

Delayed striate cortical activation during spatial attention.

Recordings of event-related potentials (ERPs) and event-related magnetic fields (ERMFs) were combined with functional magnetic resonance imaging (fMRI) to study visual cortical activity in humans during spatial attention. While subjects attended selectively to stimulus arrays in one visual field, fMRI revealed stimulus-related activations in the contralateral primary visual cortex and in multiple extrastriate areas. ERP and ERMF recordings showed that attention did not affect the initial evoked response at 60-90 ms poststimulus that was localized to primary cortex, but a similarly localized late response at 140-250 ms was enhanced to attended stimuli. These findings provide evidence that the primary visual cortex participates in the selective processing of attended stimuli by means of delayed feedback from higher visual-cortical areas.

Adult↗

Parvalbumin in the monkey striate cortex: a quantitative immunoelectron-microscopy study.

Parvalbumin (PV) is present in a subpopulation of interneurons in the visual cortex, and also in thalamic afferents to the neocortex of primates. The object of this study is to confirm by immunoelectron-microscopy the presence of intrinsic and extrinsic connections containing parvalbumin in the monkey visual cortex, by the demonstration of parvalbumin-immunoreactivity in symmetric and asymmetric synapses. We analyzed the distribution of parvalbumin-immunoreactive profiles at the ultrastructural level in the primary visual cortex of old world monkeys (Macaca fascicularis). It has been shown by others that parvalbumin-immunoreactive cells resemble non-spiny stellate cells, double-bouquet cells, chandelier and basket cells. These neurons are known to be inhibitory and to form symmetric synapses. In fact, we observed that the vast majority of parvalbumin-immunoreactive synaptic contacts in the primary visual cortex of Macaca fascicularis are of the symmetric type (81.7%). Since parvalbumin-positive asymmetric contacts are also present (18.3%) and occur mostly in the thalamic recipient layers, 4C and 4A (9.9%), these afferents probably derive from parvalbumin-immunoreactive neurons located in the dorsal lateral geniculate nucleus of the thalamus.

Animals↗

Separate neural pathways for the visual analysis of object shape in perception and prehension.

BACKGROUND: Earlier work with neurological patients has shown that the visual perception of object size and orientation depends on visual pathways in the cerebral cortex that are separate from those mediating the use of these same object properties in the control of goal-directed grasping. We present evidence suggesting that the same dissociation between perception and action is evident in the visual processing of object shape. In other words, discrimination between objects on the basis of their shape appears to be mediated by visual mechanisms that are functionally and neurally distinct from those controlling the pre-shaping of the hand during grasping movements directed at those same objects. RESULTS: We studied two patients with lesions in different parts of the cerebral visual pathways. One patient (RV), who had sustained bilateral lesions of the occipitoparietal cortex, was unable to use visual information to place her fingers correctly on the circumference of irregularly shaped objects when asked to pick them up, even though she had no difficulty in visually discriminating one such object from another. Conversely, a second patient (DF), who had bilateral damage in the ventrolateral occipital region, had no difficulty in placing her fingers on appropriate opposition points during grasping, even though she was unable to discriminate visually amongst such objects. CONCLUSIONS: This double dissociation lends strong support to the idea that the visual mechanisms mediating the perception of objects are functionally and neurally distinct from those mediating the control of skilled actions directed at those objects. It also supports the recent proposal of Goodale and Milner that visual perception depends on a ventral stream of projections from the primary visual cortex to the inferotemporal cortex, whereas the visual control of skilled actions depends on a dorsal stream from the primary visual cortex to the posterior parietal cortex.

Adult↗

The visual cortical association field: a Gestalt concept or a psychophysiological entity?

The receptive field of a visual neurone is classically defined as the region of space (or retina) where a visual stimulus evokes a change in its firing activity. Intracellular recordings in cat area 17 show that the visually evoked synaptic integration field extends over a much larger area than that established on the basis of spike activity. Synaptic depolarizing (dominant excitation) responses decrease in strength for stimuli that are flashed at increasing distances away from the centre of the discharge field, while their onset latency increases. A detailed spatio-temporal analysis of these electrophysiological data shows that subthreshold synaptic responses observed in the 'silent' surround of cortical receptive fields result from the intracortical spread of activation waves carried by slowly conducting horizontal axons within primary visual cortex. They also predict that a perceptual facilitation may occur when feedforward activation produced by the motion signal in the retina travels in phase in the primary visual cortex with the visually induced spread of horizontal activation. A psychophysical correlate has been obtained in humans, showing that apparent motion produced by a sequence of co-linear Gabor patches, known to preferentially activate V1 orientation selective cells, are perceived by human observers as much faster than non co-linear sequences of the same physical speed.

Animals↗

An approach to probe some neural systems interaction by functional MRI at neural time scale down to milliseconds.

In this paper, we demonstrate an approach by which some evoked neuronal events can be probed by functional MRI (fMRI) signal with temporal resolution at the time scale of tens of milliseconds. The approach is based on the close relationship between neuronal electrical events and fMRI signal that is experimentally demonstrated in concurrent fMRI and electroencephalographic (EEG) studies conducted in a rat model with forepaw electrical stimulation. We observed a refractory period of neuronal origin in a two-stimuli paradigm: the first stimulation pulse suppressed the evoked activity in both EEG and fMRI signal responding to the subsequent stimulus for a period of several hundred milliseconds. When there was an apparent site-site interaction detected in the evoked EEG signal induced by two stimuli that were primarily targeted to activate two different sites in the brain, fMRI also displayed signal amplitude modulation because of the interactive event. With visual stimulation using two short pulses in the human brain, a similar refractory phenomenon was observed in activated fMRI signals in the primary visual cortex. In addition, for interstimulus intervals shorter than the known latency time of the evoked potential induced by the first stimulus ( approximately 100 ms) in the primary visual cortex of the human brain, the suppression was not present. Thus, by controlling the temporal relation of input tasks, it is possible to study temporal evolution of certain neural events at the time scale of their evoked electrical activity by noninvasive fMRI methodology.

Animals↗

Mexican hats and pinwheels in visual cortex.

Many models of cortical function assume that local lateral connections are specific with respect to the preferred features of the interacting cells and that they are organized in a Mexican-hat pattern with strong "center" excitation flanked by strong "surround" inhibition. However, anatomical data on primary visual cortex indicate that the local connections are isotropic and that inhibition has a shorter range than excitation. We address this issue in an analytical study of a neuronal network model of the local cortical circuit in primary visual cortex. In the model, the orientation columns specified by the convergent lateral geniculate nucleus inputs are arranged in a pinwheel architecture, whereas cortical connections are isotropic. We obtain a trade-off between the spatial range of inhibition and its time constant. If inhibition is fast, the network can operate in a Mexican-hat pattern with isotropic connections even with a spatially narrow inhibition. If inhibition is not fast, Mexican-hat operation requires a spatially broad inhibition. The Mexican-hat operation can generate a sharp orientation tuning, which is largely independent of the distance of the cell from the pinwheel center.

Animals↗

Dynamics of orientation tuning in the cat striate cortex neurons.

Dynamics of orientation tuning was studied by a flashing light bar in 233 single units of cat primary visual cortex. Orientation tuning was plotted by the criterion of spike number in successive fragments of responses with a step of 10 or 20 ms. Preferred orientation and the width of orientation tuning were measured for each of such curves. It was found that successive dynamic change of preferred orientation during response was typical for 63% of units under study, while such change of orientation tuning width was typical for 93%. Neurons with or without shift in preferred orientation differed in their response characteristics: latency and duration of discharge, its frequency and response rise time, as well as quality of orientation tuning. Possible functional significance of two groups of cortical units, "scanners" (with systematic preferred orientation shift) and "timers" (without it), is discussed in accordance with the idea of spatiotemporal orientation coding in the primary visual cortex.

Animals↗

The projection from V1 to extrastriate area 21a: a second patchy efferent pathway colocalizes with the CO blob columns in cat visual cortex.

The different patchy organizations of neurons projecting from primary visual cortex (area 17) to the various extrastriate areas may contribute to functional differences in the output to each of these areas. The pattern of neurons projecting to extrastriate area 21a was examined using large injections of retrograde tracers and compared to the pattern shown by neurons projecting to the lateral suprasylvian area (LS). Patches of neurons projecting to 21a showed a bimodal laminar distribution, with numerous labeled cells in the upper and lower third of layer 3 bracketing a sparsely labeled central third; LS-projecting neurons were confined to the lower and middle thirds of layer 3. The 21a projecting cells were relatively tighter in their clustering pattern than the LS projecting cells, i. e. the difference in labeling density between patch and interpatch zones was greater for 21a-projecting cells than for LS-projecting cells. As previously shown for the LS-projecting cells, patches of 21a-projecting cells colocalized with CO blob columns in area 17. Combined with our earlier results, this study shows that the CO blob compartments in area 17 give rise to at least two distinct efferent pathways, one projecting to LS and the other to 21a, and furthermore suggest that each pathway may carry unique information to its extrastriate target.

Animals↗

A tonic hyperpolarization underlying contrast adaptation in cat visual cortex.

The firing rate responses of neurons in the primary visual cortex grow with stimulus contrast, the variation in the luminance of an image relative to the mean luminance. These responses, however, are reduced after a cell is exposed for prolonged periods to high-contrast visual stimuli. This phenomenon, known as contrast adaptation, occurs in the cortex and is not present at earlier stages of visual processing. To investigate the cellular mechanisms underlying cortical adaptation, intracellular recordings were performed in the visual cortex of cats, and the effects of prolonged visual stimulation were studied. Surprisingly, contrast adaptation barely affected the stimulus-driven modulations in the membrane potential of cortical cells. Moreover, it did not produce sizable changes in membrane resistance. The major effect of adaptation, evident both in the presence and in the absence of a visual stimulus, was a tonic hyperpolarization. Adaptation affects a class of synaptic inputs, most likely excitatory in nature, that exert a tonic influence on cortical cells.

Adaptation, Physiological↗

5-HT1 receptors in the structures of visual pathway of normal and monocularly deprived kittens.

The density and pattern of distribution of 5-HT1 receptor sites was examined using quantitative in vitro autoradiography with [3H]5-HT as a ligand in the visual structures of 5 weeks old kittens. One group of animals had normal binocular vision, the other was monocularly deprived during the last three days of life. The density of 5-HT1 receptor sites and the pattern of their distribution in the visual structures showed distinct regional, areal and laminar differences. In the primary visual cortex (area 17) a high labelling density was found as compared with other cortical areas investigated. Three bands of high binding density were observed corresponding to cortical layers II-III, IV c and VI. This pattern distinguished area 17 from other cortical areas investigated. In subcortical visual structures very high labelling was present in the superficial visual layers of superior colliculus, but LGN showed rather weak labelling although the lamination of LGN was also seen in the pattern of distribution of 5-HT1 sites. Neither density nor pattern of 5-HT1 sites in the visual cortex and superior colliculus were affected by 3 days of monocular deprivation. High density of labelling and the distinct pattern of 5-HT1 receptor sites in the primary visual cortex suggest the important role of serotonergic transmission in the modulation of visual afferent input activity.

Animals↗

Immunoreactivity for a calmodulin-dependent protein kinase is selectively increased in macaque striate cortex after monocular deprivation.

Immunocytochemical methods were used to localize type II Ca2+/calmodulin-dependent protein kinase in the macaque primary visual cortex. Neurons that stain for the kinase include both pyramidal and nonpyramidal cells and they appear to form a subset of cortical neurons. They are densely packed in layers II and IVB, somewhat more sparse in layers III, IVC beta, and VI, and nearly absent in layer V. In normal animals the distribution of kinase-positive cells within each layer is relatively uniform. However, in animals in which one eye is removed 7-14 days before sacrifice or sutured shut for 9 or 11 weeks, the cells in layer IVC beta are divided into alternating lightly and darkly stained bands. Comparison of immunocytochemically stained sections with adjacent sections stained for the mitochondrial enzyme, cytochrome oxidase, reveals that the kinase staining increases in ocular dominance columns originally driven by the removed or closed eye. These findings suggest that either the concentration of type II Ca2+/calmodulin-dependent protein kinase or its accessibility to the antibody probe increases dramatically and selectively in neurons of macaque primary visual cortex that have been deprived of their normal visual input. This may indicate that changing levels of activity in cortical neurons can alter their regulatory machinery.

Animals↗

Topography of cortical activation differs for fundamental and harmonic frequencies of the steady-state visual-evoked responses. An EEG and PET H215O study.

In humans, visual flicker stimuli of graded frequency (2-90 Hz) elicit an electroencephalographic (EEG) steady-state visual-evoked response (SSVER) with the same fundamental frequency as the stimulus and, in addition, a series of harmonic responses. The fundamental component of the SSVER is generated by increased synaptic activity in primary visual cortex (V1). We set out to determine the cortical origin of the harmonic responses in humans. For this purpose, we recorded the SSVERs at 5 different frequencies (5, 10, 15, 25, and 40 Hz) and measured regional cerebral blood flow (rCBF) with positron emission tomography-H(2)(15)O at rest and during visual stimulation at the same frequencies. The rCBF contrast weighted by the amplitude of the SSVERs first harmonics showed activation of a swath of cortex perpendicular to V1, including mostly the inferior half of the parieto-occipital sulcus. This area overlapped minimally with the primary visual cortex activated by the fundamental frequency. A different method, estimating EEG cortical source current density with low-resolution brain electromagnetic tomography, gave the same results. Our finding suggests that the inferior portion of the banks of the parieto-occipital sulci contains association visual cortex involved in the processing of stimuli that can be as simple as a flickering light source.

Adult↗

[New results in the experimental psychopathology of visual dysfunctions in schizophrenia].

New theories of schizophrenia put a special emphasis on cognitive deficits, including executive functions, attention, verbal memory, and psychomotor speed. Dysfunctions of low level visual processing are less known. In this paper, the author summarizes recent results from two new tasks and integrates them with the relevant literature of schizophrenia. First, the relationship between the precortical magno (M) and parvocellular (P) pathways was investigated with a new contrast categorization task assessing low level perception, memory, and abstraction. Patients with schizophrenia showed a generalized dysfunction, which suggests contrast-independent memory and abstraction problems. In contrast, siblings were selectively impaired on the task variant that included the 10-15% contrast range, the transition zone between M and P pathway contrast sensitivity. In a second paradigm, the author investigated intrinsic lateral connectivity in the primary visual cortex, the main target of M and P pathways. Using Gabor patches, a reduced facilitation effect of collinear flankers was demonstrated in schizophrenia, which suggests impaired lateral connections in the primary visual cortex. These results are closely related to backward masking and smooth pursuit eye movement abnormalities. As a neurobiological basis of these phenomena, N-methyl-D-aspartate (NMDA) glutamate receptors and integrated synaptic networks may be of special relevance.

Adult↗