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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

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

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

Low-threshold Ca2+ channels mediate induction of long-term potentiation in kitten visual cortex.

1. The induction mechanism of long-term potentiation (LTP) in developing visual cortex was studied by recording intracellular responses from layer III-IV cells in slice preparations of kitten visual cortex at 30-40 days after birth. 2. Strong stimulation of white matter produced a late depolarizing response after an orthodromic action potential. This depolarizing response was abolished by membrane depolarization or hyperpolarization caused by current injection through the recording electrode. In addition, this response was reduced by bath application of a low concentration (100 microM) of Ni2+ without any changes in the rising slope of the excitatory postsynaptic potential (EPSP) or orthodromic action potential. This suggests that this response is mediated by low-threshold Ca2+ channels (LTCs). 3. The involvement of LTCs in the induction of LTP was tested. White matter was stimulated at 2 Hz for 15 min as a conditioning stimulus to induce LTP, and the resultant changes in EPSPs were tested by low-frequency (0.1 Hz) stimulation of white matter. Conditioning stimulation produced a large N-methyl-D-aspartate (NMDA) receptor-mediated depolarizing response in these cells, which obscured the presence of the late depoliarzation. Therefore the test was conducted in a solution containing an NMDA antagonist 2-amino-5-phosphonovalerate (APV). 4. Weak conditioning stimulation, which evoked no LTC responses, never induced LTP; whereas strong conditioning stimulation, which evoked LTC responses, always induced LTP. Strong conditioning stimulation failed to induce LTP when LTC responses were prevented either by membrane depolarization or hyperpolarization or by a bath application of 100 microM Ni2+. 5. In a solution without APV, the application of Ni2+ also prevented the induction of LTP. 6. When cells were impaled by an electrode containing a Ca2+ chelator 1,2-bis-(o-aminophenoxy)ethane-N,N,N',N'-tetraacetic acid (BAPTA), LTP was never induced, even though LTC responses were evoked by conditioning stimulation. These results indicate that Ca2+ influx into postsynaptic cells through LTCs induces the LTP. 7. The responses mediated by LTCs, which were evoked by the injection of current pulses into the cells, were maximum at the critical period of visual cortical plasticity, suggesting that LTCs in postsynaptic cells regulate the plastic changes in developing visual cortex.

2-Amino-5-phosphonovalerate

Interhemispheric synchronization of oscillatory neuronal responses in cat visual cortex.

Neurons in area 17 of cat visual cortex display oscillatory responses that can synchronize across spatially separate columns in a stimulus-specific way. Response synchronization has now been shown to occur also between neurons in area 17 of the right and left cerebral hemispheres. This synchronization was abolished by section of the corpus callosum. Thus, the response synchronization is mediated by corticocortical connections. These data are compatible with the hypothesis that temporal synchrony of neuronal discharges serves to bind features within and between the visual hemifields.

Animals

The rat claustrum: afferent and efferent connections with visual cortex.

We have examined the afferent and efferent projections between the claustrum and visual cortex in the Long-Evans rat using anterograde and retrograde axonal transport techniques. Injections of either wheat germ agglutinin/horseradish peroxidase (WGA/HRP) or Fast Blue were made into each of the main visual regions (17, 18a or 18b) as well as directly into the claustrum. The cortical injections were placed in either the upper, middle or deep layers so as to assist in determining the laminar organization of these connections. Of the 3 visual areas, only area 18b appears to have extensive and reciprocal connections with the claustrum. After a WGA/HRP injection of this area, dense labeled terminals and numerous labeled cells were found intermixed throughout the full extent of the claustrum. The density of this labeled activity was found to vary directly with the amount of the infragranular layers involved by the injections. Injections in the other visual areas did produce labeled cells in the claustrum, but their number was always small or even negligible. There was never any evidence of anterograde labeled terminals in the claustrum from any injection of areas 17 or 18a. Tracer injections directly in the claustrum confirmed and extended these findings by showing that the labeled terminals and/or labeled cells were localized predominantly in layer VI of area 18b of visual cortex. On the basis of these injections, two major conclusions are reached. First, the pattern of connections between the claustrum and visual cortex in the rat differs fundamentally with that found in other species.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals

Functional organization in the visual cortex of the golden hamster.

The visual cortex of the golden hamster was studied by means of multi-unit and single unit recording, which revealed three separate retinotopic maps of the visual field in the posterior cortex. V1, corresponding to cyto-architectonic area 17, has the contralateral temporal field represented medially, the central visual field (extending about 10 deg ipsilateral) represented laterally and the lower field anteriorly. The borders of the map, especially for the upper field, seem to be more restricted than the whole visual field available to the contralateral hemiretina: V1 probably does not represent the extreme periphery of the field. A large fraction of V1 has binocular input, for up to about 50 deg lateral to the vertical midline. There is a retinotopic reversal near the representation of the vertical midline where V1 meets V2 (corresponding to the more lateral "area 18a"). There is another retinotopic reversal at the extremity of the contralateral field representation, where V1 meets Vm (the medial visual area, corresponding to "area 18"). V2 and Vm each contain a reduced mirror image version of the map in V1. Almost all isolated single units in V1 have receptive fields that can be classified as radially symmetrical (60%) or asymmetrical (35%). Symmetrical fields have ON (13%), OFF (4%), ON-OFF (30%) or "SILENT" (12%) central areas when plotted with flashing spots. There are minor but not striking differences between these groups in their field sizes, velocity preferences and so on. They almost invariably prefer moving to stationary stimuli but are not selective for orientation or direction of movement. Asymmetrical fields are of four types, three of which (type 1, 11%; type 2, 17%; and type 3, 2%) are orientation selective and resemble simple, complex and hypercomplex cells in the cat cortex. Some of these have direction as well as orientation preference. Axial movement detectors (5%) have a selectivity for one axis of motion, and thus prefer one orientation of edge, but respond equally well to movement of a spot. Vertical and horizontal orientation preferences, especially the latter, are much the most common. There is some evidence of clustering of cells according to receptive field type and, possibly, preferred orientation. Asymmetrical cells are, relatively somewhat rarer in the deeper cortical layers. Within the binocular segment, fully 89% of cells are binocularly driven and the receptive fields are similar in the two eyes. Receptive fields tend to increase in size away from the area centralis representation and, in a complementary fashion, the magnification factor decreases from up to 0.1 mm/deg at the area centralis representation to about 0.02 mm/deg for the peripheral field.

Animals

Disparity tuning and binocularity of single neurons in cat visual cortex.

Activity from single neurons in the visual cortex of anaesthetized and paralyzed cats were recorded in response to monocular and binocular stimuli of different retinal disparities. Three different types of disparity sensitive neurons were found, characterized by (1) a response maximum near zero disparity, (2) a response minimum near zero disparity, and (3) an asymmetric disparity sensitivity. Usually, neurons of the first type receive about the same monocular input from the two eyes; the other disparity sensitive neurons respond better to monocular stimulation of one eye as compared to the other. The results closely resemble those obtained recently in the rhesus monkey.

Animals

Age dependence of the effect of squint on cells in kittens' visual cortex.

The period of susceptibility of the visual cortex of kittens to the effect of squint is limited to the first three postnatal months. The reduction of binocularity found in these kittens as reflected by the distribution of neurons according to their ocular dominance is especially emphasized in animals operated on between the ages of 4-7 weeks in comparison to animals operated on between the ages of 8-11 weeks. The proportion of monocularly dominated neurons within the first three postnatal months is significantly (0.05 greater than p greater than 0.025) reduced with age. Similar effect on cortical neurons was found for animals who were under the influence of squint for 6 weeks or 15 months and for animals with wide range of deviation angles.

Age Factors

[Effect of visual deafferentiation on the ultrastructure of synapses of the rat visual cortex].

Electron microscopic study and quantitative analysis of the visual cortex synapses in 14, 30 and 60-day-old rats were performed after bilateral enucleation of newly-forn rats. A great amount of synapses of other functional systems was shown to be functioning in the area striata in addition to the synapses formed by specific visual afferents. Alterations in the synapses of the area striata of blind rats are developing gradually, achieving the greatest pronouncement in 60-day-old rats. These changes develop according to the type of atrophic process in connection with dysfunction. The atrophic alterations of the synapses were found both in axo-somatic and axo-dendritic synapses on the dendrite trunks and on the thorns. The alterations of synapses being concentrated in layer IV. The quantitative ratio of different kinds of atrophied synapses in the cross-section of the visual cortex was different suggesting the following conclusion about the distribution of the visual afferents. In layers I and III the visual afferents formed mostly axon-thorn contacts and less amount of axo-somatic and axo-dendritic synapses on the dendrite trunks. In layer IV they mainly formed axo-somatic and axo-thorn synapses and less amount of axo-dendritic ones on the dendrite trunks. In layers V and VI they mainly contact with the dendrite trunks and with the nervous cell bodies and more rarely with thorns.

Afferent Pathways

The development of ventral tegmental area (VTA) projections to the visual cortex of the rat.

The development of the ventral tegmental area (VTA) projections to the rat visual cortex was studied with the wheat germ agglutinin-horseradish peroxidase (WGA-HRP) retrograde tracing technique. Large injections of WGA-HRP in the visual cortex of newborn, early postnatal, and adult rats resulted in a substantial number of retrogradely labelled neurons in the VTA showing the same distribution pattern at all ages examined. Contrary to other reports, labelled cells were never found in the pars compacta of the substantia nigra but occasionally were seen in the contralateral VTA near the midline. These neurons showed a continuous growth from the day of birth to the end of the second postnatal week, when they acquired morphological features comparable to the adult; they subsequently showed a substantial decrease in soma size. The present results clearly demonstrate that there exists a substantial mesocortical projection to the rat visual cortex which arises exclusively from the VTA. This projection is already established at birth, but the neurons which give rise to it appear morphologically immature during the first two postnatal weeks.

Animals

Elaborate organization of visual cortex in the hamster.

We have studied the organization of visual cortex in the hamster by analyzing and correlating the distributions of callosal and striate-extrastriate connections. Our results show that, in lateral extrastriate cortex, callosal connections form an elaborate pattern which delineates at least 3 regions poor in callosal connections. These regions receive projections from striate cortex which are distributed into multiple separate fields. Two additional striate-projection fields were observed in medial extrastriate cortex, and another field was often observed near the posterolateral border of primary somatosensory cortex. All striate recipient fields are reciprocally connected with striate cortex. The patterns of callosal and striate-extrastriate connections in the hamster are similar, although not identical, to the corresponding patterns in rats and mice. Our results provide evidence that the organization of visual cortex in the hamster is highly elaborate, and suggest that extrastriate cortex is subdivided into multiple visual areas.

Animals

Direct evidence for recurrent inhibition in sliced brain preparation of the cat's visual cortex.

To reveal the recurrent inhibitory circuit in the visual cortex, a depolarizing current was applied through a glass microelectrode to an impaled cell in an in vitro slice of the visual cortex obtained from a cat anesthetized with pentobarbital. The cell reported here produced inhibitory postsynaptic potentials (IPSPs) following single spikes or bursts of spikes which were elicited by intracellularly applied current. This observation indicates that IPSPs have been mediated by the recurrent inhibitory circuit via axon collaterals of the impaled cell.

Action Potentials

[Non-specific facilitating influences on the responses of rabbit visual cortex neurons].

A study was made of neuronal responses in the rabbit visual cortex to patterned visual stimuli and their change during non-specific activation reaction. Non-specific activation while only slightly affecting the background neuronal activity, enhances the responses of most units to patterned visual stimuli. A comparison of responses with a different degree of inhibition participation shows that the depression of inhibitory processes may act as a mechanism of facilitation of the responses during non-specific activation. After facilitated responses evoked by the action of the stimulus during non-specific activation, the neurones retain a state of enhanced excitability. Repetition of stimuli not attended with non-specific activation leads to the diminution of excitability. The possible connection of the indicated effects with extinction processes is discussed.

Animals

Beta-adrenergic receptors: astrocytic localization in the adult visual cortex and their relation to catecholamine axon terminals as revealed by electron microscopic immunocytochemistry.

It has long been recognized that noradrenaline, the most abundant catecholamine within the visual cortex, plays important roles in modulating the sensitivity of cortical neurons to visual stimuli. However, whether or not these noradrenaline effects are confined to a discrete synaptic specialization or mediated by diffuse modulation of a group of synapses has remained an issue open for debate. The aim of this study was to examine the cellular basis for noradrenaline action within the visual cortex of adult rats and cats. To this end, I used electron microscopic immunocytochemistry to examine the relationship between (1) catecholamine axon terminals and beta-adrenergic receptors (beta AR), which, together, may define the effective sphere of noradrenaline modulation; and then (2) these putative sites for catecholamine modulation and axospinous asymmetric junctions where excitatory neurotransmission is likely to dominate. Antibodies against beta AR were used at light and electron microscopic levels on the visual cortex of rat and cat. Rat visual cortex was also labeled simultaneously for beta AR and the catecholamine-synthesizing enzyme, tyrosine hydroxylase (TH), to determine the ultrastructural relationships between catecholamine terminals and beta AR. Immunoperoxidase labeling revealed that beta AR404, a polyclonal antibody directed against the C-terminal tail of hamster lung beta AR (beta 2-type), recognized astrocytic processes predominantly. In contrast, beta AR248, a polyclonal antibody directed against the third cytoplasmic loop, recognized neuronal perikarya as observed in previous studies. Dual labeling for beta AR404 and TH revealed that catecholamine axon terminals that contained numerous vesicles formed direct contacts with astrocytic processes exhibiting beta AR404 immunoreactivity. However, some catecholamine axon terminals that lacked dense clusters of vesicles were positioned away from beta AR404-immunoreactive astrocytes. Frequently, beta AR-immunoreactive astrocytic processes surrounded asymmetric axospinous junctions while also contacting catecholamine axon terminals. These observations support the possibility that, through activation of astrocytic beta AR, noradrenaline modulates astrocytic uptake mechanism for excitatory amino acids, such as L-glutamate. Astrocytic beta AR might also define the effective sphere of catecholamine modulation through alterations in the morphology of distal astrocytic processes and the permeability of gap junctions formed between astrocytes.

Animals