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Magnetic stimuli applied over motor and visual cortex: influence of coil position and field polarity on motor responses, phosphenes, and eye movements.

Transcranial magnetic stimulation was performed over the motor and visual cortex with the Novametrix 12-cm diameter stimulation coil. The influence of coil position on the size of electromyographic responses and on the intensity and position of phosphenes in the binocular visual field was studied. Furthermore, it was investigated whether stimulation over the visual cortex or over the frontal eye field evoked or disturbed eye movements. Coil position was recorded by constructing grids on the skull surface using extracranial bony landmarks, and was then related individually to underlying cerebral sulci by analysis of magnetic resonance images of the brain. Motor responses. Excitatory effects were maximal when the coil windings in the middle of the coil ring lay over the particular motor representation field of area 4. The response size depended on the direction of the coil currents passing over the motor cortex. For example, coil currents passing over the lateral part of the precentral gyrus from the front and transversely to the central sulcus elicited maximal responses in hand muscles. With the currents passing over the same area in the opposite direction, response amplitudes were much smaller, suggesting activation of different inputs to cortical motoneurons. Phosphenes. Fields of fixed, white and unstructured phosphens occurred in the lower half of the visual field when the coil center was placed about 7 cm anterior to the inion on the inion-nasion line. Counterclockwise or clockwise coil currents elicited phosphenes within the left or right lower quadrant of the binocular visual field, respectively, which could be attributed to an activation of the right or left primary visual cortex (area 17). The 'cortical' phosphenes moved with voluntary eye movements, but not during caloric and optokinetic nystagmus. Phosphenes resulting from an excitation of the optic nerve rather than the retina could be evoked by stimulation over frontal parts of the skull. Eye movements. The application of single magnetic field pulses over the frontal eye field or over the visual cortex did not elicit eye movements except for small vertical eye movements as part of a magnetically elicited blink.

Adolescent↗

Synaptic characteristics of identified pyramidal and multipolar non-pyramidal neurons in the visual cortex of young and adult rabbits. A quantitative Golgi-electron microscope study.

The visual cortex of 20 day old rats and rabbits has been considered as mature on the basis of the observations that the dendritic arborization and the overall synaptic population have almost reached their adult stage in these animals. In the present study we have investigated the visual cortex of 20 day and 7 month old (adult) rabbits in order to determine whether this apparent adult appearance also holds for the synaptic organization of individual neurons. Neurons mainly located in layers III and IV of the primary visual cortex (area 17) were Golgi-impregnated, gold toned and deimpregnated and were then, after embedding in plastic, sectioned serially. The number and length of synaptic profiles, and the length of the neuronal boundaries were analysed in every tenth section. From these counts and measurements the size distribution of the synaptic discs, the number of synapses per 100 micron2 neuronal surface and the receptive surface expressed as the percentage of the total neuronal surface covered with synaptic contacts were estimated using stereological methods. At both ages studied, the density of synapses was significantly higher for the non-pyramidal neurons than for the pyramidal neurons. Differences in the amount of receptive surface were parallel to the differences observed for the number of synapses per 100 micron2. At day 20 the receptive surface of the non-pyramidal neurons was significantly larger than that of the pyramidal neurons. The receptive surface of the non-pyramidal neurons in the adult stage was not only larger than that of the pyramidal neurons in the adults, but also larger than that of the day 20 non-pyramidal neurons. From our results the following conclusions can be drawn: (1) The synaptic input received by the pyramidal neurons is mainly established at day 20 of postnatal life, i.e. prior to the establishment of adult visual behaviour. (2) The non-pyramidal neurons complete their maturation in a later stage than the pyramidal neurons. (3) Medium to large sized synaptic contacts are newly formed after day 20 and are mainly added to the synaptic population on dendrites of non-pyramidal neurons. (4) The specific increase in the number of synapses on non-pyramidal neurons is discussed in relation to intracortical inhibition which is thought to be important for the fine regulation of visual function during development.

Animals↗

Selective magnetization transfer ratio decrease in the visual cortex following optic neuritis.

Patients with clinically isolated syndromes suggestive of multiple sclerosis have evidence for abnormality in normal appearing grey matter detected using the magnetization transfer ratio (MTR), a quantitative MRI measure. One potential mechanism for the decreased grey matter MTR (GM MTR) observed is trans-synaptic morphological abnormality secondary to demyelinating lesions that are in an anatomically linked pathway but remote location. We investigated this potential association by studying the location of abnormalities using voxel-based analysis of GM MTR maps in a group of 80 patients studied within 6 months of presenting with isolated optic neuritis and compared the findings with those seen in 50 age- and sex-matched healthy controls. Occipital cortex and whole brain analysis comparing all optic neuritis patients and controls revealed a selective decrease of MTR bilaterally in the visual cortex in patients [Brodmann area (BA) 17]. Whole brain analysis of patients fulfilling the McDonald criteria for multiple sclerosis (n = 20) showed a lower MTR compared to controls bilaterally in the visual cortex (BA 17/18), left hippocampus, bilateral superior temporal gyrus, bilateral lenticular nuclei and the right cerebellum. There was no significant difference in the percentage of grey matter between patients and controls in the regions of abnormal MTR detected in the visual cortex. The intrinsic MTR decrease seen in patients suggests that there are structural changes in the visual cortex following an attack of optic neuritis. Potential mechanisms for this include trans-synaptic neuronal degeneration and cortical synaptic morphological changes; such abnormalities may also contribute to MTR abnormalities observed in the normal appearing grey matter in multiple sclerosis.

Adult↗

Cross-modal innervation of primary visual cortex by auditory fibers in congenitally anophthalmic mice.

Auditory-visual cross-modal innervation was examined in control (sighted, ZRDCT-N) and congenitally anophthalmic (eyeless, ZRDCT-AN) mice using electrophysiological recording and pathway tracing with carbocyanine dyes. Electrophysiological data demonstrate that the primary visual cortex of congenitally eyeless, blind, mice receives auditory stimuli. Neuroanatomical data demonstrate a direct connection between the inferior colliculus (IC) and visual cortex. Our experiments provide new information about how the brain adapts to the loss of sight.

Adaptation, Physiological↗

[Contemporary knowledge on the subject of visual cortex structure and function].

The present state of knowledge concerning the structure and the function of the visual cortex was discussed. Special attention was focused on the: (1) retino-geniculo-cortical pathway (2) representation and topography of the visual cortex (3) anatomy, physiology, and functional architecture of the primary visual cortex (4) organization and function of the parvocellular, magnocellular, and koniocellular pathway (5) functional specialization and connections between the visual areas (6) neurotransmitters' presence and function in the Brodmann's 17th area.

Humans↗

Involvement of muscarinic acetylcholine receptors in regulation of kitten visual cortex plasticity.

Receptor autoradiographic methods specific for M3 and M1 muscarinic acetylcholine receptors were used to investigate the development and input-dependent laminar redistribution of these receptor populations during the critical period for kitten visual cortex plasticity. Analysis of the binding curves of [3H]4-diphenylacetyl-N-methyl-piperidine (4-DAMP) and [3H]pirenzepine (PZ) indicated that these two ligands bound heterogeneously to muscarinic acetylcholine receptors with different affinities. While [3H]4-DAMP showed a high affinity for M3 receptors and much lower affinities for M1 and M2 receptors, [3H]PZ displayed higher affinity for M1 receptors. By carefully choosing concentrations of labelled and unlabeled ligands, the patterns of laminar distribution for both receptor subtypes within visual cortex were obtained. Both receptors were most concentrated in cortical layer IV immediately after birth and during the most sensitive period of visual cortex plasticity. The binding density for both receptor subtypes thinned out progressively in this layer to concentrate in more superficial layers as plasticity waned with age. Moreover, interruption of visual or spontaneous input to visual cortex induced either by lesion or by tetrodotoxin infusion into lateral geniculate nucleus prevented the developmental redistribution of these receptors from layer IV to superficial layers, that is, the pattern of laminar distribution remained that of the age at which the lesion or tetrodotoxin infusion into the lateral geniculate nucleus was performed. The results indicate that the developmental expression of M3 and M1 muscarinic acetylcholine receptors in kitten visual cortex depends on cortical inputs.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Spatially asymmetric response to moving patterns in the visual cortex: re-examining the local sign hypothesis.

One of the most fundamental functions of the visual system is to code the positions of objects. Most studies, especially those using fMRI, widely assume that the location of the peak retinotopic activity generated in the visual cortex by an object is the position assigned to that object-this is a simplified version of the local sign hypothesis. Here, we employed a novel technique to compare the pattern of responses to moving and stationary objects and found that the local sign hypothesis is false. By spatially correlating populations of voxel responses to different moving and stationary stimuli in different positions, we recovered the modulation transfer function for moving patterns. The results show that the pattern of responses to a moving object is best correlated with the response to a static object that is located behind the moving one. The pattern of responses across the visual cortex was able to distinguish object positions separated by about 0.25 deg visual angle, equivalent to approximately 0.25 mm cortical distance. We also found that the position assigned to a pattern is not simply dictated by the peak activity-the shape of the luminance envelope and the resulting shape of the population response, including the shape and skew in the response at the edges of the pattern, influences where the visual cortex assigns the object's position. Therefore, visually coded position is not conveyed by the peak but by the overall profile of activity.

Attention↗

Response properties in the pulvinar complex after neonatal ablation of the primary visual cortex.

Injuries to specific areas of the brain (such as cerebrovascular accidents or surgical procedures) and particularly to the primary visual cortex, yield profound visual defects. The level of spared visual functions or residual vision depends on the extent and location of the lesion as well as the age at which the trauma occurs. For instance, in primate as well as non-primate species, it is well established that lesions in adulthood have a more profound effect than those occurring in young animals. The recovery of visually guided behavior observed after massive destruction of the occipital cortex in young animals across many species has been generally associated with the reorganization of the pathways from the extrageniculate thalamus to the spared visual cortex, i.e. the extrastriate areas. In this chapter, we present some evidence that the lateral posterior-pulvinar (LP-pulvinar) complex may contribute to maintaining visual capacities in brain-damaged cats. Our data indicate that the overall visual responsiveness of the lateral part of the LP (LPl) cells is not altered by the early removal of the visual cortex. However, some specific properties differ from those of intact animals: on average, LPl neurons in brain-damaged animals are more broadly tuned for orientation than that in intact cats. Spatial frequency tuning functions are also affected since most units in lesioned animals are of the low-pass type. Moreover, most LPl cells of lesioned cats responded to drifting gratings with modulated discharges and a linear spatial summation within their receptive field, a characteristic that is infrequently observed in intact animals. The change in LPl response properties observed in the present study is likely to come from the reorganization of cortical and retinal fibers reaching this extrageniculate nucleus.

Animals↗

Cortisol reduces plasticity in the kitten visual cortex.

We investigated the effect of elevated levels of cortisol on plasticity in the visual cortex of the cat. Animals were given daily injections of cortisol i.m. for 20 days starting around 35 days of age. After 10 days they were monocularly deprived, and after an additional 10 days recordings were made from the visual cortex to construct an ocular dominance histogram. The results were compared with those from normal animals of the same age, and with animals monocularly deprived for the same period but not treated with cortisol. Cortisol reduced the ocular dominance shift in a dose-dependent manner, but did not totally abolish it even at the highest doses used. Two other series of animals were recorded, one slightly later in the critical period and one slightly earlier, with care taken to give cortisol before the animals were exposed to light in the morning. In both cases, cortisol reduced the ocular dominance shift but did not abolish it. To interpret these results, we measured levels of plasma cortisol in normal cats of various ages. Average levels were fairly constant between birth and 12 months of age (0.5-1 microgram/dl), and increased slightly after that, but there was a large variation between animals. Thus elevated levels of cortisol can have a substantial effect on plasticity in the visual cortex of the cat, but the decline of the critical period for plasticity between 6 weeks and 3-5 months of age does not seem to be due to a rise in cortisol levels during this time.

2-Amino-5-phosphonovalerate↗

Influence of pulse stimulation of the visual cortex on the function of the superior colliculus of the awake rabbit.

It has been demonstrated in awake rabbits that stimulation of the visual cortex by a solitary pulse of electrical current leads to the formation of a short-latency response in the superior colliculus. The formation of this response is suppressed when a light stimulus precedes it. At the same time, a conditioning solitary electrostimulation of the visual cortex induces a short inhibition of the formation of the response to the test light stimulus. This fact suggests that the influences of the visual cortex on the functioning of the superior colliculus may be biphasic in character. When the adrenergic apparatus of the reticular formation is blocked this inhibitory influence bears a more pronounced and prolonged character. The stimulation of the reticular formation, on the other hand, by means of anodic polarization leads to the diametrically opposite effect: the inhibitory character of the influence of the cortex is replaced by a facilitatory one. The inference is drawn that the character and the directionality of the influence of the visual cortex on the functioning of the superior colliculus is determined to a significant degree by the initial functional state of nonspecific brain systems.

Animals↗

Enigmatic bipolar cell of rat visual cortex.

Our earlier Golgi-electron microscopic study of bipolar cells in the rat visual cortex showed the axons of these neurons as forming asymmetric synapses (Peters and Kimerer; J. Neurocytol, 10:921-946, '81) in which the most common postsynaptic elements were dendritic spines. This result was unexpected, since Parnavelas (Parnavelas, Sullivan, Lieberman, and Webster: Cell Tissue Res. 183:499-517, '77) had earlier shown a bipolar cell from the same cortex to have an axon forming symmetric synapses with dendritic shafts. Here then was an enigma, strengthened by examination of neuronal components labelled by antibodies to two compounds in particular--namely, vasoactive intestinal polypeptide (VIP) and choline acetyltransferase (ChAT). Antibodies to these compounds preferentially label bipolar cells in the rat cerebral cortex, and the labelled axon terminals form symmetric synapses. Against this background the present study was performed, and it has been shown that the resolution to the enigma is that there are two different populations of bipolar cells in the rat visual cortex. Thus some Golgi-impregnated bipolar cells examined by electron microscopy after gold toning have been found to possess axons forming asymmetric synapses, and others have been found to have axons forming symmetric synapses. The axons of the bipolar cells forming asymmetric synapses most commonly synapse with dendritic spines (67%), although other terminals synapse with dendritic shafts (33%). In contrast, the bipolar cells with axons forming symmetric synapses preferentially synapse with dendritic shafts (100%). The population of bipolar cells that form symmetric synapses includes the ones that label with antibodies to vasoactive intestinal polypeptide (VIP), for the axons of VIP-labelled bipolar cells have been traced to labelled terminals forming symmetric synapses. However, examination of the population of VIP-labelled axon terminals shows that in addition to dendritic shafts, some of the labelled terminals synapse with the cell bodies of pyramidal and nonpyramidal cells. This includes bipolar cells, some of which receive large numbers of VIP-labelled axon terminals. It is also shown that some VIP-positive bipolar cells have myelinated axons. Analysis of tissue labelled with VIP antibody reveals that about 50% of the total population of bipolar cells in the rat visual cortex is VIP positive. These results are discussed in the light of information about labelling of bipolar cells with antibodies to gamma-aminobutyric acid (GABA) and to other peptides, and it is suggested that most VIP-positive bipolar cells also contain GABA.

Animals↗

Age-related changes in the visual cortex.

The ability to accurately perceive the speed of moving objects is one of many visual functions that decline with age. One factor that may contribute to this is a deterioration in temporal processing speed. At present, there is a dearth of information concerning how this may occur in the central nervous system, particularly in the visual cortex. Thus, in the present study, we investigated the neural basis of speed and temporal processing in areas 17 and 18 of visual cortex in young and aged rats using either a moving bar of light or a series of flashing lights. Our results showed that the mean preferred speed of a moving bar of light was significantly reduced in aged as compared to young animals. We also found that cells recorded from young animals were able to entrain to a higher frequency of flashing light stimuli than those recorded from aged animals. In addition, we found no age-related differences between cortical fields. These results suggest an age-related difference in temporal processing speed at the level of visual cortex.

Aging↗

Evidence for a nicotinic component to the actions of acetylcholine in cat visual cortex.

Radioligand binding assays, receptor autoradiography and iontophoresis have been used to look for evidence of a nicotinic component to the actions of acetylcholine in cat visual cortex. [3H]Nicotine bound to a uniform population of high affinity binding sites in cat primary visual cortex. This binding was inhibited by nicotine agonists and antagonists but not muscarinic antagonists. The concentration of nicotinic binding sites was about 10% of that of muscarinic binding sites measured with [3H]N-methylscopolamine. The muscarinic sites were resolved into M1 and M2 subtypes. Quantitative receptor autoradiography showed that there were muscarinic sites in all layers, although they were least abundant in layer IV of area 17. In contrast, the nicotinic sites were most concentrated in layer IV in area 17. The concentration of this labelling was reduced at the 17/18 border and also at the 18/19 border. Layer I of the cingulate and suprasylvian gyri were also labelled. Electrolytic lesions of the lateral geniculate nucleus (LGN) led to a loss of nicotinic binding sites in layer IV of area 17, indicating that these sites are most likely located on the LGN terminals. Iontophoresis of mecamylamine, a nicotinic antagonist, decreased evoked responses in visual cortex, providing evidence that the [3H]nicotine binding sites are functional receptors and suggesting that the release of acetylcholine onto these receptors on the LGN terminals facilitates the input of visual information into visual cortex.

Animals↗

The effects of dark rearing on the development of the visual cortex of the rat.

The effects of dark rearing on the development of the visual cortex has been studied in Wistar rats, as have the effects of subsequent light exposure on recovery. Five groups of animals were used: (1) light exposed until 30-40 days post partum (dpp) (2) dark reared until 30-40 ddp (3) dark reared until 80-120 dpp (4) dark reared to 21 dpp, then light exposed until 40 dpp (5) light exposed to 21 dpp and then dark reared until 40 dpp. Golgi-Cox impregnations of layer IV stellate cell dendritic fields were analysed and total neuronal and glial counts were also done within layer IV of the primary visual cortex. Normal visual stellate cell dendritic fields were radially organised, with the highest dendritic density being recorded below the soma. In short term visually deprived animals and in the exposed only for 21 dpp and then reared in light until 40 dpp the radial distribution of dendrites was maintained but the peak density shifted to above the soma. In all other experimental groups this abnormal polarisation was still present but not as marked. Measurement of branching indices suggested that these field changes resulted from increased branching and growth in the superficial domain and not from the reorientation of dendrites. Differential glial counts revealed a significantly higher number of microglia in dark reared animals than in controls. Neuronal numbers were not affected.

Animals↗

Distinct contrast response functions in striate and extra-striate regions of visual cortex revealed with magnetoencephalography (MEG).

OBJECTIVE: To spatially and temporally characterise the cortical contrast response function to pattern onset stimuli in humans. METHODS: Magnetoencephalography (MEG) was used to investigate the human cortical contrast response function to pattern onset stimuli with high temporal and spatial resolution. A beamformer source reconstruction approach was used to spatially localise and identify the time courses of activity at various visual cortical loci. RESULTS: Consistent with the findings of previous studies, MEG beamformer analysis revealed two simultaneous generators of the pattern onset evoked response. These generators arose from anatomically discrete locations in striate and extra-striate visual cortex. Furthermore, these loci demonstrated notably distinct contrast response functions, with striate cortex increasing approximately linearly with contrast, whilst extra-striate visual cortex followed a saturating function. CONCLUSIONS: The generators that underlie the pattern onset visual evoked response arise from two distinct regions in striate and extra-striate visual cortex. SIGNIFICANCE: The spatially, temporally and functionally distinct mechanisms of contrast processing within the visual cortex may account for the disparate results observed across earlier studies and assist in elucidating causal mechanisms of aberrant contrast processing in neurological disorders.

Adult↗

Hydrocephalus in developing cats: physiological properties of visual cortex cells.

We have studied electrophysiologically by single cell recording in the visual cortex, whether modification of the visual system in developing and in adult cats by hydrocephalus has an effect on processing of visual information. One of our cats (H1) had developed a complete hydrocephalus and the others partial, as proved by either complete or partial dilation of the lateral ventricles, respectively and by the thinning of the cortex. Despite this, the horizontal lamination and the vertical organization of the cortex were fully preserved. Except for the optic radiation and the corpus callosum which was remarkably modified, the optic tract, chiasm, nerve and retina were morphologically and histologically normal. The visual behavior of the hydrocephalic cats was normal. This was also reflected, by and large, in the physiological properties of the visual cortex. However, in cat H1 there were many more visually unresponsive cortical cells in comparison to its matched controls (C1) and the normal cats. A reduced responsiveness was also found in cat H2 with partial hydrocephalus but not in the other partial hydrocephalic cats. Similarly, the ocular dominance distribution of the cells was affected in cat H1 in comparison to the control cats as indicated by the changes found in the relative proportions of contralaterally and ipsilaterally driven cells in the two hemispheres. No change was, however, found in the partially hydrocephalic cats. Most of the cells in the hydrocephalic cats were orientation specific, similarly to the result of their matched controls. Direction specific cells were much smaller in proportion in cat H1 but not in the other cats, in comparison with their matched controls. In keeping with this, a large increase was found in the receptive field area of cat H1, a smaller one in cat H2 and none in the other hydrocephalic cats in comparison to the matched controls. The eccentricity distribution of the receptive fields in the hydrocephalic cats was the same as expected under normal conditions. It was concluded that in the way hydrocephalus had modified the brain of several of our cats, a quantitative effect was induced in visual cortex cells leading to some degradation of function; this change, however, did not interfere with their basic visual properties.

Animals↗

Recovery of tactile placing after visual cortex ablation in cat: a behavioral and metabolic study of diaschisis.

After bilateral visual cortex ablation, cats show a transient deficit in tactile placing and a permanent deficit in visual placing of both forelimbs. Amphetamine administration (four doses, 5 mg/kg, i.p., spaced at 4-day intervals beginning 10 days after surgery) accelerated the rate of recovery of tactile placing compared with saline controls whereas visual placing was not affected. The catecholamine antagonist, haloperidol (0.4 mg/kg, i.p.), blocked the amphetamine-enhanced recovery of tactile placing. Additionally, the visual cortex lesions produced a depression of oxidative metabolism, measured by cytochrome oxidase histochemistry, in subcortical regions remote from the injury. Animals treated with amphetamine exhibited an alleviation of this metabolic depression in the superior colliculus but not in other regions.

Amphetamine↗

The neuroanatomical organization of pathways between the dorsal lateral geniculate nucleus and visual cortex in Old World and New World primates.

Pathways between the dorsal lateral geniculate nucleus (dLGN) and visual cortex in Old World (Macaca, Papio, Erythrocebus, Cercopithecus) and New World (Saimiri, Cebus) primates were studied after injections of horseradish peroxidase and H3 or S35 amino acids into the dLGN or visual cortex. Trans-synaptic autoradiography was also used to study these pathways after an injection of H3 proline-fucose into one eye. The subsequent autoradiographs of visual cortex showed that Old World primates have separate eye inputs (ocular dominance columns) in the striate cortex, whereas New World monkeys have overlapping or non-separated eye inputs. In both primate groups the geniculocortical input to layer IVA formed a pattern which resembled a honeycomb in tangential sections, unlike the solidly labeled layer IVC. Also common to the two primate groups was a projection from dLGN to layer VI. There was no dLGN projection to any prestriate area in any of the primates. However, after an injection limited to the prestriate cortex of Macaca, light autoradiographic labeling was seen in the interlaminar zones and the magnocellular and S laminae, demonstrating a prestriate-dLGN pathway. Our results indicate that the primate visual system differs significantly from the cat in having no dLGN projection to area 18. There are also signficant differences between primates in the level at which the possibility of binocularity (of an excitatory nature) first occurs in the striate cortex because in the species studied thus far with neuroanatomical methods, Old World primates have ocular dominance columns in layer IV but most New World monkeys lack them.

Animals↗