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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

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

Physiological studies in deafferented visual cortex cells of cats following transplantation of fetal xenografts from the rat's cortex.

We have studied the physiological properties of cells (N = 822) in visual cortex area 17 of seven adult cats transplanted with visual cortex xenografts from fetal (E15-E17) rats. The transplants were assumed to induce recovery of adjacent neurons partially deafferented from visual input. The control group (eight cats, 564 cells) had just analogous sectioning in the cortex. The level of activity found, following visual stimulation, in the deafferented cortical region (medially to the graft) was 48.6% compared to the result (34.5%) obtained in the control cats, indicating the preservation of visual responsiveness. Furthermore, no disturbance could be found in the ocular dominance distribution or binocularity (64.4%) of the cells in the grafted region compared to those in the control cats, indicating preservation of the columnar organization. The deafferented cells in the grafted cortex thus demonstrated the absence of adverse immunological reaction there due to the presence of the xenogeneic tissue, indicating that the visual cortex is immunologically privileged.

Afferent Pathways

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

Nerve growth factor-induced ocular dominance plasticity in adult cat visual cortex.

Activity-dependent modifiability of cortical ocular dominance occurs only during early postnatal life, within the so-called "critical period," but not thereafter in adult visual cortex. To examine the role of neurotrophins in the activity- and age-dependent stimulation-induced modifiability of visual cortex, we tested whether intracortical infusion of nerve growth factor could induce ocular dominance plasticity in adult visual cortex. Nerve growth factor was continuously infused, by means of osmotic minipumps, into striate cortex of adult cats for 2 weeks. At the time of minipump implantation, one eyelid of the experimental animals was sutured closed. After 3 weeks of monocular deprivation, the ocular dominance distribution of neurons in the striate cortex was assessed using single unit recording. We found that monocular deprivation imposed on adult animals in conjunction with nerve growth factor infusion causes an ocular dominance shift toward the deprived eye. Although the underlying mechanisms remain uncertain, the results indicate that nerve growth factor can enhance activity-dependent synaptic modification and remodeling in adult visual cortex.

Age Factors

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

Receptive field expansion in adult visual cortex is linked to dynamic changes in strength of cortical connections.

1. Receptive field (RF) sizes of neurons in adult primary visual cortex are dynamic, expanding and contracting in response to alternate stimulation outside and within the RF over periods ranging from seconds to minutes. The substrate for this dynamic expansion was shown to lie in cortex, as opposed to subcortical parts of the visual pathway. The present study was designed to examine changes in cortical connection strengths that could underlie this observed plasticity by measuring the changes in cross-correlation histograms between pairs of primary visual cortex neurons that are induced to dynamically change their RF sizes. 2. Visually driven neural activity was recorded from single units in the superficial layers of primary visual cortex in adult cats, with two independent electrodes separated by 0.1-5 mm at their tips, and cross-correlated on-line. The neurons were then conditioned by stimulation with an "artificial scotoma," a field of flashing random dots filling the region of visual space around a blank rectangle enclosing the RFs of the recorded neurons. The neuronal RFs were tested for expansion and their visually driven output again cross-correlated. After this, the neurons were stimulated vigorously through their RF centers to induce the field to collapse, and the visually driven output from the collapsed RFs was again cross-correlated. Cross-correlograms obtained before and after conditioning, and after RF collapse, were normalized by their flanks to control for changes in peak size due solely to fluctuations in spike rate. 3. A total of 37 pairs of neurons that showed distinct cross-correlogram peaks, and whose RF borders were clearly discernible both before and after conditioning, were used in the final analysis. Of these neuron pairs, conditioning led to a clear expansion of RF boundaries in 28 pairs, whereas in 9 pairs the RFs did not expand. RFs that did expand showed no significant shifts in their orientation preference, orientation selectivity, or ocularity. 4. When the RFs of a pair of neurons expanded with conditioning, the area of the associated flank-normalized cross-correlogram peaks also increased (by a factor ranging from 0.84 up to 3.5). Correlograms returned to their preconditioning values when RFs collapsed.(ABSTRACT TRUNCATED AT 400 WORDS)

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

Biochemical evidence for glutamate and/or aspartate as neurotransmitters in fibers from the visual cortex to the lateral posterior thalamic nucleus (pulvinar) in rats.

The effects of visual cortex ablation on several neurotransmitter parameters in the lateral thalamic nucleus (pulvinar) in rats have been investigated. We found a 57% decrease in high affinity uptake of D-[3H]aspartate in the pulvinar after ablation of the ipsilateral visual cortex. The KCl-evoked release of exogenous D-[3H]aspartate and endogenous glutamate were decreased by 33 and 37%, respectively. Moreover, the contents of endogenous glutamate and aspartate were decreased by 35%, each. The glutamate decarboxylase and choline acetyltransferase activities and the contents of other amino acids were not affected by the lesion. Our biochemical data indicate that glutamate and/or aspartate may be transmitters in the fibers from visual cortex to pulvinar in rats.

Animals

Responses of single cells in cat visual cortex to prolonged stimulus movement: neural correlates of visual aftereffects.

1. The activity of single cortical cells in area 17 of anesthetized and unanesthetized cats was recorded in response to prolonged stimulation with moving stimuli. 2. Under the appropriate conditions, all cells observed showed a progressive response decrement during the stimulation period, regardless of cell classification, i.e., simple, complex, or hypercomplex. 3. The observed response decrement was shown to be largely cortical in origin and could be adequately described with an exponential function of the form R = Rf +(R1-Rf)e-t/T. Time constants derived from such calculations yielded values ranging from 1.92 to 12.45 s under conditions of optimal-stimulation. 4. Most cells showed poststimulation effects, usually a brief period of reduced responsiveness that recovered exponentially. Recovery was essentially complete in about 5-35 s. 5. The degree to which stimuli were effective at inducing response was shown to have significant effects on the magnitude of the response decrement. 6. Several cells showed neural patterns of response and recovery that suggested the operation of intracortical inhibitory mechanisms. 7. A simple two-process model that adequately describes the behavior of all the studied cells is presented. 8. Because the properties of the cells studied correlate well with human psychophysical measures of contour and movement adaptation and recovery, a causal relationship to similar neural mechanisms in humans is suggested.

Anesthesia, General