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Sex differences in blood-oxygenation-level-dependent functional MRI with primary visual stimulation.

OBJECTIVE: The authors evaluated the effect of sex on data derived from activation studies using blood-oxygenation-level-dependent (BOLD) functional magnetic resonance imaging (MRI). METHOD: Gradient echo-echo planar imaging was used to measure BOLD signal response in the primary visual cortex in response to binocular photic stimulation in 16 healthy, young subjects (eight women and eight men). RESULTS: BOLD signal response was 38% lower in women than in men, and much of the difference was lateralized to the right hemisphere. CONCLUSIONS: Lower BOLD signal response in women may reflect a sex difference in the brain's response to a primary visual stimulation or in the physiology underlying BOLD functional MRI signal changes.

Adult↗

Regional and laminar distribution of the dopamine and serotonin innervation in the macaque cerebral cortex: a radioautographic study.

The regional density and laminar distribution of dopamine (DA) and serotonin (5-HT) afferents were investigated in the cerebral cortex of cynomolgus monkeys using a radioautographic technique that is based on the high affinity uptake capacity of these aminergic neurons. Large vibratome sections, 50 micron thick, were incubated with [3H] DA (0.2 microM) and desipramine (5 microM) or with unlabeled norepinephrine (5 microM) and [3H] 5-HT (0.6 microM), which allowed for the specific labeling of the DA and 5-HT innervations, respectively. After fixation, these sections were dried, defatted, and radioautographed by dipping. Semiquantitative data on the DA innervation also were provided by counting [3H] DA-labeled axonal varicosities in radioautographs from 4-micron-thick sections of the slices obtained after epon embedding. The DA innervation was widespread and differed in density and laminar distribution in the agranular and granular cortices. DA afferents were densest in the anterior cingulate (area 24) and the motor areas (areas 4, 6, and supplementary motor area [SMA]). In the latter they displayed a trilaminar pattern of distribution, predominating in layers I, IIIa, and V-VI, with characteristic cluster-like formations in layer IIIa, especially in the medial part of motor areas. In the granular prefrontal (areas 46, 9, 10, 11, 12), parietal (areas 1, 2, 3, 5, 7), temporal (areas 21, 22), and posterior cingulate (area 23) cortices, DA afferents were less dense and showed a bilaminar pattern of distribution, predominating in the depth of layer I and in layers V-VI; density in layers II, III, and IV was only 20% of that in layer I. The lowest density was in the visual cortex, particularly in area 17, where the DA afferents were almost restricted to layer I. The density of 5-HT innervation was generally greater than that of DA except in the motor areas and in the anterior cingulate cortex. Region-specific laminar patterns characterized (1) motor areas where a lower density in layer III contrasted with the clusters of DA axons in the same layer; (2) the primary visual cortex (area 17), where two bands of higher density in layers III-IV and layer V outlined a poorly innervated zone in layer IVc-beta; (3) the peristriate area 18, where the 5-HT network was relatively loose but with a denser band in layer III. Thus, DA innervation of the cerebral cortex displays major differences between rodents and primates, characterized by expanded cortical targets and by a highly differentiated laminar distribution.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Transient patterns of serotonergic innervation in the rat visual cortex: normal development and effects of neonatal enucleation.

The transient aggregation of serotonin (5-HT)-containing fibers in the early development of rat visual cortex was examined immunohistochemically. The aggregation of 5-HT immunoreactive (IR) fibers consisted of three stages which were classified according to the course of time and degree of space occupied. The primary aggregation appeared in the subplate and moved upward along the development of the cortex. The aggregation proceeded to the secondary stage in presumptive layer IV. The fibers extended in a column-like structure following the secondary aggregation and formed the tertiary aggregation. The upper edge of the tertiary aggregation formed a lattice-like pattern in layer I and its structure was recognized to be similar to the structure of a 'blob' which characterizes the primary visual cortex in monkey. This transient aggregation of 5-HT-IR fibers began in the subplate of the anterior visual cortex on postnatal day 2 (PND 2) and progressed towards the posterior. On PND 11, the secondary and tertiary aggregations were completed in the entire region. No further aggregation of 5-HT-IR fibers was observed on PND 15. The anterior-to-posterior axis in the aggregation process corresponds to the direction of differentiation in the layer structure of cortex. In order to investigate the relationship between the transient aggregation of 5-HT-IR fibers and the development of the visual pathway, the secondary and tertiary aggregation on PND 11 were observed after postnatal monocular or binocular enucleation. Enucleation of eye balls did not affect either the area occupied by the 5-HT-IR fibers in the secondary aggregation or the number of column structures in the tertiary aggregation. However, the contralateral and ipsilateral cortices of monocularly enucleated cases were irregularly shaped in the secondary aggregation. The distribution of 5-HT-IR fiber terminals in the binocular area (Oc1B) increased in density on the contralateral side in the monocular enucleation, while that of both sides in the binocular enucleation was of non-homogeneous density and were shaped irregularly. The above results suggest that the transient aggregation of 5-HT-IR fibers observed in the early stage of development of visual cortex is regulated primarily by the intrinsic factors, and that extrinsic factors, such as visual pathway input, affect the aggregation within the boundary of such intrinsic factors. That is, the visual pathway input and the input balance from both eyes affect the distribution density of 5-HT-IR fibers and the shape of the visual cortex, respectively.

Animals↗

Correlation between cytochrome oxidase staining and the uptake and laminar distribution of tritiated aspartate, glutamate, gamma-aminobutyrate and glycine in the striate cortex of the squirrel monkey.

The cellular uptake and laminar distribution of tritium-labeled gamma-aminobutyrate, aspartate, glutamate and glycine were examined in the primary visual cortex of squirrel monkeys. The purpose was to correlate the distribution of these labeled neurons with their level of cytochrome oxidase activity, particularly in laminae II-III (puffs) and adjacent non-puff regions. In general, tritium-labeled neurons that had either high or low levels of cytochrome oxidase activity were present in all laminae with each amino acid tested; however, their density varied between laminae and with the amino acid injected. Specifically, in laminae II-III, very few neurons were labelled with either of the putative excitatory amino acids (aspartate and glutamate). An increased uptake for both was observed in lamina IVC, with the greatest increase for each occurring in laminae V and VI. Significantly more neurons in each lamina were labeled with the putative inhibitory transmitters (gamma-aminobutyrate and glycine) than with either aspartate or glutamate. gamma-Aminobutyrate-labeled neurons were more prevalent in lamina II than III, and an increase in labeling was observed in laminae IV-VI, with the most prominent increase found in laminae V and VI. Glycine-labeled neurons were larger, more uniformly distributed and more abundant throughout all cortical laminae than those labeled with the other amino acids. Significantly more gamma-aminobutyrate- and glycine-labeled neurons were found in the puff regions than in the non-puff areas. No difference was found between puff and non-puff regions for the tritium-labeled leucine controls. Labeled neurons included stellate, fusiform and pyramidal-shaped cells of varying sizes; however, gamma-aminobutyrate-labeled pyramidal cells were not observed outside of the intense injection site. Large glycine-labeled cytochrome-oxidase-reactive pyramidal cells (24-32 micron in diameter) were present at the boundary between laminae V and VI. In addition, a row of large glycine-labeled, fusiform neurons were present in lamina IVB. With each amino acid injected, the tritium-labeled neurons that were darkly reactive for cytochrome oxidase were, on average, larger than the tritium-labeled neurons that were only lightly reactive for cytochrome oxidase. Thus, each of the four amino acids tested had its unique pattern of distribution in the primate striate cortex. Whether one or all of them served as neurotransmitter(s) for distinct neuronal groups is beyond the scope of this study. Glycine, in particular, might be used in part or in whole for metabolic purposes.(ABSTRACT TRUNCATED AT 400 WORDS)

Amino Acids↗

Horizontal Interactions in Cat Striate Cortex: II. A Current Source-Density Analysis.

The current source-density (CSD) analysis was used to investigate the organization of tangential synaptic connections in primary visual cortex of normally reared (NR) kittens and of NR, binocularly deprived (BD) and dark-reared (DR) adult cats. Laminar profiles of field potentials, elicited by intracortical microstimulation were measured at various distances from the stimulating electrodes. To exclude contamination by axon collaterals of antidromically stimulated thalamo-cortical fibres, these were destroyed by injecting the cytotoxin N-methyl-D-aspartate (NMDA) into the lateral geniculate nucleus 13 - 27 days before recording. The CSD profiles revealed distinct layer-specific patterns of lateral spread of activity. Invariably, the most prominent, long-lasting and far reaching responses were recorded in supragranular layers. Responses in layer IV were brief and confined to the vicinity of the stimulation site. Responses in infragranular layers spread as far as those in supragranular layers, but were of smaller amplitude. Latency considerations, the results of double shock stimulation, and the effects of translaminar cuts, suggest that these responses were monosynaptic and mediated by intracortical pathways with a conduction velocity of 0.3-5 m/s. The spatial spread of these responses changed substantially with age but was not influenced by visual deprivation. In NR adults, supra- and infragranular responses were recordable up to 2.5 mm from the stimulation site and layer IV responses up to 1 mm from the stimulation site. In kittens, the former responses spread up to 5 mm and the latter up to 2 mm from the site of stimulation. The amplitude of the responses decreased with distance from the stimulation site. This decrease was not always monotonic suggesting inhomogeneities in the tangential projections. The laminar distribution of current sinks and sources indicates that the pathways mediating tangential interactions form excitatory synapses on apical dendrites of pyramidal cells. It is concluded that the spatial spread of tangential excitatory interactions decreases with age, but that neither the laminar pattern nor the age-dependent reduction in the strength of tangential interactions are influenced by visual deprivation.

Journal Article↗

A case of multiple sclerosis with homonymous hemianopia examined by positron emission tomography.

BACKGROUND: To demonstrate the efficacy of positron emission tomography (PET) for examining multiple sclerosis (MS) patients with hemianopia. CASE: A 20-year-old man visited us with a complaint of left homonymous hemianopia and headache. OBSERVATIONS: The patient's visual acuity was 1.2 (n.c.) OD and 0.9 (1.0) OS. Magnetic resonance imaging (MRI) showed a mass in the temporoparietal lobe. A pathological diagnosis of MS was made by brain biopsy. Low glucose metabolism in the lesion and visual cortex was observed by PET with (18)F-fluorodeoxy glucose. PET with (11)C-flumazenyl revealed a reduction of (11)C-uptake in the demyelinated optic radiation, and only a slight reduction of (11)C-uptake in the primary visual cortex. The results of (11)C-flumazenyl PET suggested a slight reduction of neuronal density. In 2 years, the visual field recovered to the normal state. CONCLUSION: PET can be a useful tool for estimating the visual outcome of patients with hemianopia in MS.

Adult↗

Reciprocal connections between the claustrum and visual thalamus in the tree shrew (Tupaia glis).

We previously described the existence of reciprocal connections between the dorsal claustrum and striate cortex in the tree shrew, Tupaia glis. These projections were found to originate and terminate in a distinct topographic manner within the mid region of the upper portion of the dorsal claustrum. In this investigation, we examined the afferent and efferent projections between the claustrum and the lateral intermediate nucleus (Li) of the visual thalamus using small electrophoretic injections of anterograde and/or retrograde tracers (horseradish peroxidase, wheat germ agglutinin/horseradish peroxidase, or tritiated amino acids) into the claustrum, as well as the Li. Following tracer injections in the dorsal claustrum, labeled cells and/or terminal grains were found throughout the Li, except at the more caudal levels where the activity was confined to the lateral and medial borders. Tracer injections within the same nuclear region of the dorsal thalamus confirmed the existence of reciprocal projections between the Li and the claustrum. Following anterograde tracer injections, labeled terminals were found only within the most ventral zone of the dorsal claustrum--the 'hilum'; while retrograde tracer injections, produced labeled cells principally along the outer margins of the claustrum, including the hilum and tended to encapsulate the nucleus at all levels. Both sets of labeled activity were found to extend in this specific fashion over the majority of the dorsal claustrum, but appeared not to overlap with regions interconnected with striate cortex. These results thus suggest that the claustrum is capable of exerting a neural influence on cortex directly as well as indirectly via the visual thalamus. However, since the thalamic projection terminates in a claustral region not known to project to visual cortex, it is uncertain what function such projections have within the claustrum. It is possible that since Li receives ascending projections from the pretectum and superior colliculus, that it provides a multisensory input to the claustrum for relay onto areas outside of primary visual cortex.

Animals↗

Reorganization of visual cortical maps after focal ischemic lesions.

Plasticity after central lesions may result in the reorganization of cortical representations of the sensory input. Visual cortex reorganization has been extensively studied after peripheral (retinal) lesions, but focal cortical lesions have received less attention. In this study, we investigated the organization of retinotopic and orientation preference maps at different time points after a focal ischemic lesion in the primary visual cortex (V1). We induced a focal photochemical lesion in V1 of kittens and assessed, through optical imaging of intrinsic signals, the functional cortical layout immediately afterwards and at 4, 13, 33, and 40 days after lesion. We analyzed histologic sections and evaluated temporal changes of functional maps. Histological analysis showed a clear lesion at all time points, which shrank over time. Imaging results showed that the retinotopic and orientation preference maps reorganize to some extent after the lesion. Near the lesion, the cortical retinotopic representation of one degree of visual space expands over time, while at the same time the area of some orientation domains also increases. These results show that different cortical representations can reorganize after a lesion process and suggest a mechanism through which filling-in of a cortical scotoma can occur in cortically damaged patients.

Animals↗

Nitric oxide synthase inhibition prevents neuronal death in the developing visual cortex.

During postnatal development of the visual cortex of golden hamster, there is a transient increase in both the expression and the activity of nitric oxide synthase (NOS), which coincides temporally with the formation of ipsilateral retino-collicular and retino-geniculate projections and the functional differentiation of primary visual cortex, suggesting the involvement of NO in the maturation of the visual cortex. In the present study, an inhibitor of NOS, N-nitro-L-arginine (L-NNA) was used to block the NOS activity of newborn golden hamster, and effects on development were examined. L-NNA treatment caused an increase in mortality, and suppression of both body weight gain and nicotinamide adenine dinucleotide phosphate-diaphorase (NADPH-d) activity in the early phase of treatment (before postnatal day 14, PD14). The growth of NADPH-d-positive neurons in the visual cortex was also suppressed by the treatment. In control animals, significant numbers of apoptotic neurons were detected by terminal deoxynucleotidyl transferase-mediated dUTP nick end labelling assay on PD14, and this apoptosis mainly affected cells in cortical layers II and III. NOS inhibition largely rescued neurons from undergoing apoptosis, indicating that NO may serve as a signal triggering apoptosis and play a role in the maturation of the visual cortex.

Animals↗

The processing of stereoscopic information in human visual cortex: psychophysical and electrophysiological evidence.

Three-dimensional depth perception relies in part on the binocular fusion of horizontally disparate stimuli presented to the left and right eye. The mammalian visual system offers a unique possibility to study electrophysiologically cortical neuronal mechanisms: since the input of the two eyes remains separated up to the level of the visual cortex, evoked potential components that are generated exclusively by cortical structures may be explored when dynamic random-dot stereograms (dRDS) are presented. In a series of independent studies, we determined the scalp topography of dRDS evoked brain activity in different groups of healthy subjects, and we found consistent results. Major differences between stereoscopic and contrast evoked brain activity are seen in the strength of the potential fields as well as in their topography. Our findings suggest that there are fewer neurons in the human visual cortex that are responsive to horizontal disparity, and that higher visual areas like V2 are more engaged with stereoscopic processing than the primary visual cortex. On the other hand, component latencies of evoked brain activity show no effect signifying that the binocular information flow to the visual cortex has a similar time course for both the processing of contrast information and of dRDS stimuli. We could also verify that healthy subjects can learn to perceive 3D structure contained in dRDS. Changes in perceptual ability as measured with psychophysical tests are paralleled by systematic alterations in the topography of stereoscopically evoked potential fields. Stereoscopic VEP recordings may also be of clinical use: in patients with selectively disturbed depth perception but normal visual acuity there is a high correlation between clinical symptoms, perceptual deficiency, and altered VEP amplitudes and latencies.

Brain Mapping↗

Cortical blindness and visual imagery.

Controversy exists concerning the neural basis underlying visual imagery. Some propose that visual images evoked from memory are mediated by primary visual cortices. Others argue that these primary visual areas perform computations on elementary visual features when constructing visual representations from retinal input but that they are not activated during recall of these representations. The visual imagery abilities of patients with cortical blindness may resolve this controversy. The proposition that primary visual cortex is necessary for visual imagery predicts a cortically blind subject's inability to perceive visual stimuli would be accompanied by an inability to image visually. Our investigations of three patients with cortical blindness provide strong evidence that primary visual cortices are not essential for the mediation of visual images recalled from memory.

Adult↗

Investigations of origins of serotonergic projection to developing rat visual cortex: a combined retrograde tracing and immunohistochemical study.

The present study investigated whether the raphe neurons which give rise to the transient serotonergic fibers in the visual cortex of neonatal rats persist or disappear as the rats mature. Three experiments were performed employing the WGA-apoHRP-Au retrograde transport technique in conjunction with 5-HT or WGA-HRP immunohistochemical staining. WGA-apoHRP-Au was injected into the primary visual cortex of all rats 9 days postnatally. In the first experiment, the animals were examined after 2 days; retrogradely labeled cells were observed in the dorsal raphe nucleus (DR), the median raphe nucleus (MR), and in the B9 and B6 cell groups; the majority (82.5%) of the cells was serotonergic. In the second experiment, the examinations took place following a survival time of 8 weeks: virtually all of the original raphe-visual cortical serotonergic neurons were found to the present. In the third experiment, also performed after 8 weeks relabeling the raphe-visual cortical neurons by WGA-HRP, it was found that 37.2% of the raphe neurons which had projected to the neonatal visual cortex no longer possessed such projections.

Aging↗

Infusion of nerve growth factor (NGF) into kitten visual cortex increases immunoreactivity for NGF, NGF receptors, and choline acetyltransferase in basal forebrain without affecting ocular dominance plasticity or column development.

Intracerebroventricular or intracortical administration of nerve growth factor (NGF) has been shown to block or attenuate visual cortical plasticity in the rat. In cats and ferrets, the effects of exogenous NGF on development and plasticity of visual cortex have been reported to be small or nonexistent. To determine whether locally delivered NGF affects ocular dominance column formation or the plasticity produced by monocular deprivation in cats at the height of the critical period, we infused recombinant human NGF into the primary visual cortex of kittens using an implanted cannula minipump. NGF had no effect on the normal developmental segregation of geniculocortical afferents into ocular dominance columns as determined both physiologically and anatomically. The plasticity of binocular visual cortical responses induced by monocular deprivation was also normal in regions of immunohistochemically detectable NGF infusion, as measured using intrinsic signal optical imaging and single-unit electrophysiology. Immunohistochemical analysis of the basal forebrain regions of the same animals demonstrated that the NGF infused into cortex was biologically active, producing an increase in the number of NGF-, TrkA-, p75(NTR)-, and choline acetyltransferase-positive neurons in basal forebrain nuclei in the hemisphere ipsilateral to the NGF minipump compared to the contralateral basal forebrain neurons. We conclude that NGF delivered locally to axon terminals of cholinergic basal forebrain neurons resulted in increases in protein expression at the cell body through retrograde signaling.

Animals↗

Effects on binocular activation of cells in visual cortex of the cat following the transection of the optic tract.

Cells in area 17 of the cortex are generally activated either directly through a retino-thalamic pathway or indirectly via a contralateral hemisphere-callosal pathway. The aim of the present experiment was to evaluate the effects of eliminating this second pathway on the binocular activation of cells in the primary visual cortex. The optic tract was sectioned on one side in 18 cats and unit activity was recorded in the contralateral hemisphere. This hemisphere should receive normal thalamo-cortical inputs but no visual callosal input. These animals were compared to 21 normal cats. Extracellular electrophysiological recordings were carried out in the conventional way using tungsten microelectrodes and N2O anaesthesia. Results indicated that the proportion of binocular cells found in the cortex of tract sectioned animals was lower than that found in normal animals. However, this decrease in binocularity could be essentially attributed to cells having receptive fields situated to within 4 degrees of the vertical meridian of the visual field. These results are interpreted as being congruent with the demonstrated anatomo-physiological projections of the callosal system.

Animals↗

Facilitation of probabilistic classification learning by transcranial direct current stimulation of the prefrontal cortex in the human.

The aim of our study was to test if the electrical stimulation of the prefrontal cortex (PFC) could modify probabilistic classification learning (PCL). Transcranial direct current stimulation (tDCS) was administered to the left prefrontal and to the primary visual cortex of 22 healthy subjects while they performed a PCL task. In this task subjects learned which of two outcomes would occur on each trial after presentation of a particular combination of cues. Ten minutes of anodal, but not cathodal, stimulation improved implicit learning only when the left PFC was stimulated. Our results show that implicit PLC can be modified by weak anodal tDCS, which probably increases neural excitability, as has been shown in the motor and visual cortices previously. Our results suggest that further studies on the facilitation of learning and memory processes by tDCS are warranted.

Adult↗

Analysis of pathways mediating preserved vision after striate cortex lesions.

This study investigated the neural substrates of preserved visual functioning in a patient with homonymous hemianopsia and Riddoch syndrome after a posterior cerebral artery stroke affecting the primary visual cortex (area V1). The limited visual abilities of this patient included above-chance verbal reports of movement and color change as well as discrimination of movement direction in the hemianopic field. Functional magnetic resonance imaging showed that motion and color-change stimuli presented to the hemianopic field produced activation in several extrastriate areas of the lesioned hemisphere that were defined using retinotopic mapping. Magnetoencephalographic recordings indicated that evoked activity occurred earlier in the higher-tier visual areas V4/V8 and V5 than in the lower-tier areas V2/V3 adjacent to the lesion. In addition, the functional magnetic resonance imaging analysis showed an increased functional connectivity between areas V4/V8 and V5 of the lesioned hemisphere in comparison with the same areas in the intact hemisphere during the presentation of color changes. These results suggest that visual perception after the V1 lesion in Riddoch syndrome is mediated by subcortical pathways that bypass V1 and project first to higher-tier visual areas V5 and V4/V8 and subsequently to lower-tier areas V2/V3.

Adult↗

Reduced minicolumns in the frontal cortex of patients with autism.

Cell minicolumns were shown to be narrower in frontal regions in brains of autistic patients compared with controls. This was not found in primary visual cortex. Within the frontal cortex, dorsal and orbital regions displayed the greatest differences while the mesial region showed the least change. We also found that minicolumns in the brain of a 3-year-old autistic child were indistinguishable from those of the autistic adult in two of three frontal regions, in contrast to the control brains. This may have been due to the small size of the columns in the adult autistic brain rather than to an accelerated development. The presence of narrower minicolumns supports the theory that there is an abnormal increase in the number of ontogenetic column units produced in some regions of the autistic brain during corticoneurogenesis.

Adult↗

Effect of electrical stimulation of locus coeruleus on the activity of neurons in the cat visual cortex.

1. We studied the effect of electrically stimulating the locus coeruleus (LC) and iontophoresing noradrenergic antagonists on visual responses and spontaneous activity of individual cells in the cat primary visual cortex. 2. A bilateral projection from LC to visual cortex was demonstrated anatomically, by retrograde labeling using horseradish peroxidase. Where electrical stimulation of both ipsilateral and contralateral LC affected a cortical neuron, the effect induced by stimulating each side was similar. 3. One hundred and two cells were recorded in area 17: 52% of them had their activity suppressed and 36% had their activity facilitated by LC stimulation. The suppressive effect was predominant in cortical layers II + III and IV, whereas most cells in layer V and one-half of the cells in layer VI were facilitated by LC stimulation. This suggests that LC neurons innervate each cortical layer in a different manner. 4. Simple and complex cells were equally sensitive to LC stimulation. For simple cells, the suppressive effect of LC stimulation was dominant throughout all layers. For complex cells, the suppressive effect was dominant in layers II + III and IV, whereas the facilitatory effect was dominant in layers V and VI. 5. The suppressive effect of LC stimulation was blocked by iontophoretic application of beta-adrenergic receptor antagonists and the facilitatory effect was blocked by either alpha- or beta-adrenergic receptor antagonists. 6. Nonselective alpha-, and selective alpha 1- and alpha 2-receptor antagonists suppressed visual and spontaneous activity in almost all neurons tested, suggesting that these receptors are either facilitatory at a postsynaptic site or inhibitory at a site presynaptic to an inhibitory synapse in the visual cortex. 7. beta-Receptor antagonists facilitated activity in 45% and suppressed activity in 36% of the cells tested, suggesting there are both suppressive and facilitatory types of beta-receptors. 8. The effectiveness of alpha- and beta-antagonists on the activity of neurons without LC stimulation also suggested that spontaneously released noradrenaline activated noradrenergic receptors in the visual cortex even in the anesthetized and paralyzed cat. 9. In most cells tested, both alpha- and beta-receptor antagonists exerted effects on single neurons suggesting that endogenous noradrenaline acts on both alpha- and beta-receptors on the same cell. 10. The activation of LC did not improve the signal- (visual response)to-noise (spontaneous discharge) ratio of neurons in the visual cortex. 11. LC seemed to control the activity of each cortical layer differently, by activating different kinds of noradrenergic receptors in different layers.

Adrenergic alpha-Antagonists↗