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Critical period-dependent reduction of the permissiveness of cat visual cortex tissue for neuronal adhesion and neurite growth.

During postnatal development, the visual cortex undergoes an experience-dependent refinement of its circuitry. This process includes synapse formation, as well as synapse elimination. Both mechanisms appear to be restricted to a limited 'critical period' which lasts for approximately 2 months in cats. We tested whether the termination of the critical period for cortical malleability is paralleled by changes in the growth permissiveness of the tissue. These changes may inhibit progressive reorganization of functional circuitries mediated by axon growth. Embryonic cortical neurons were cultured on unfixed cryostat sections of the visual cortex obtained from cats aged 2-50 weeks. After 2-3 days in vitro the distribution of viable cells and the percentage of neurite-bearing cells were determined and analysed with respect to the developmental age and subdivisions of the underlying tissue substrate. It was shown that cell adhesion and neurite formation are correlated with the developmental age of the substrate tissue and the time period of myelination. While embryonic neurons adhered and survived on grey and white matter tissue from 2- and 4-week-old kittens, there was a significant reduction in cell adhesion on the myelinated white matter regions of the tissue sections of older animals. Quantitative analyses showed that neurite formation by cultured neurons also became successively impaired on grey and white matter areas of tissue substrates, corresponding to the time course of the critical period for cortical malleability. On grey matter tissue this effect was most pronounced between the second and sixth postnatal weeks. The effects were not antagonized by coating the substrate sections with the growth-promoting molecule laminin. It is therefore proposed that neurite growth-inhibiting factors, most probably associated with central nervous system myelin, are gradually expressed postnatally and may contribute to the termination of the critical period in the visual cortex of cats.

Animals↗

Object-based attention in the primary visual cortex of the macaque monkey.

Typical natural visual scenes contain many objects, which need to be segregated from each other and from the background. Present theories subdivide the processes responsible for this segregation into a pre-attentive and attentive system. The pre-attentive system segregates image regions that 'pop out' rapidly and in parallel across the visual field. In the primary visual cortex, responses to pre-attentively selected image regions are enhanced. When objects do not segregate automatically from the rest of the image, the time-consuming attentive system is recruited. Here we investigate whether attentive selection is also associated with a modulation of firing rates in area V1 of the brain in monkeys trained to perform a curve-tracing task. Neuronal responses to the various segments of a target curve were simultaneously enhanced relative to responses evoked by a distractor curve, even if the two curves crossed each other. This indicates that object-based attention is associated with a response enhancement at the earliest level of the visual cortical processing hierarchy.

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↗

Orientation formed by a spot's trajectory: a two-dimensional population approach in primary visual cortex.

There exist a large number of visual illusions indicating that perception differs from pure representation of physical input. For example, a spot of light can be characterized by its position, but it does not contribute any information about orientation. However, when moved fast enough, a continuous streak along its trajectory is perceived that helps to determine the orientation of the movement path. The question arises whether the processing of the trajectory and its orientation are simultaneously represented in the primary visual cortex. Here I show that decoding neural population activity within a two-dimensional parameter space represents both (1) physical input given by the actual position of the moving spot and (2) orientation. This latter parameter has no physical counterpart in the stimulus but must be actively formed by spatiotemporal integration of the spot's trajectory.

Action Potentials↗

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↗

Recurrent inhibition and clustered connectivity as a basis for Gabor-like receptive fields in the visual cortex.

A continuous-space model of visual cortex interactions which, starting from elongate Gaussian functions, leads to high-selective Gabor-like receptive fields, is proposed. The model is based on intracortical inhibition mechanisms occurring through medium-range clustered connections. The analysis, carried out under the assumption of a linear superposition of geniculate and intracortical contributions, shows how inhibition enhances both spatial and spatial-frequency selectivity. The effects of inhibition strength and of its spatial organization on the properties of the resulting receptive field are investigated. Specifically, the relationships between receptive fields and inhibition schemes are investigated by deriving analytical expressions for their dependence and through a systematic numerical parametric study. The emergence of periodic receptive fields, like the ones observed in neurophysiological measurements, is also pointed out in relation to the clustered nature of the inhibitory scheme.

Animals↗

Deactivation of human visual cortex during involuntary ocular oscillations. A PET activation study.

Prompted by the observation of decreased glucose metabolism in the striate and extrastriate visual cortex in a patient with opsoclonus, we studied the influence of involuntary eye movements on visual cortex activity. Repeated measurements of cerebral blood flow (CBF) by PET were performed in 12 healthy volunteers using H2(15)O-bolus technique after ear canal irrigation with ice cold or warm (44 degrees C) water with the subjects eyes closed. In addition to blood flow increases in areas involved in central vestibular processing, statistical subtraction analysis revealed a nearly symmetrical, bilateral, highly significant decrease in the occipital cortex covering Brodmann areas 17, 18, and 19 after ice water stimulation of either ears. Region of interest analysis revealed in all subjects a mean decrease in regional CBF (rCBF) of 12.8% (range 4.6-21.0%) in these areas. A similar but less pronounced effect (mean rCBF decrease in visual cortex 4.8%, range 1.1-11.5%) was observed after warm water irrigation. The observations suggest that deactivation of the visual cortex is induced by involuntary ocular oscillations. This deactivation is not dependent on changes of the retinal input (eyes closed). The physiological significance of this hitherto unknown phenomenon may be the protection from inadequate visual input (oscillopsia) during involuntary ocular oscillations.

Adult↗

Differential contributions of magnocellular and parvocellular pathways to the contrast response of neurons in bush baby primary visual cortex (V1).

How neurons in the primary visual cortex (V1) of primates process parallel inputs from the magnocellular (M) and parvocellular (P) layers of the lateral geniculate nucleus (LGN) is not completely understood. To investigate whether signals from the two pathways are integrated in the cortex, we recorded contrast-response functions (CRFs) from 20 bush baby V1 neurons before, during, and after pharmacologically inactivating neural activity in either the contralateral LGN M or P layers. Inactivating the M layer reduced the responses of V1 neurons (n = 10) to all stimulus contrasts and significantly elevated (t = 8.15, P < 0.01) their average contrast threshold from 8.04 (+/- 4.1)% contrast to 22.46 (+/- 6.28)% contrast. M layer inactivation also significantly reduced (t = 4.06, P < 0.01) the average peak response amplitude. Inactivating the P layer did not elevate the average contrast threshold of V1 neurons (n = 10), but significantly reduced (t = 4.34, P < 0.01) their average peak response amplitude. These data demonstrate that input from the M pathway can account for the responses of V1 neurons to low stimulus contrasts and also contributes to responses to high stimulus contrasts. The P pathway appears to influence mainly the responses of V1 neurons to high stimulus contrasts. None of the cells in our sample, which included cells in all output layers of V1, appeared to receive input from only one pathway. These findings support the view that many V1 neurons integrate information about stimulus contrast carried by the LGN M and P pathways.

Animals↗

Cone inputs in macaque primary visual cortex.

To understand the role of primary visual cortex (V1) in color vision, we measured directly the input from the 3 cone types in macaque V1 neurons. Cells were classified as luminance-preferring, color-luminance, or color-preferring from the ratio of the peak amplitudes of spatial frequency responses to red/green equiluminant and to black/white (luminance) grating patterns, respectively. In this study we used L-, M-, and S-cone-isolating gratings to measure spatial frequency response functions for each cone type separately. From peak responses to cone-isolating stimuli we estimated relative cone weights and whether cone inputs were the same or opposite sign. For most V1 cells the relative S-cone weight was <0.1. All color-preferring cells were cone opponent and their L/M cone weight ratio was clustered around a value of -1, which is roughly equal and opposite L and M cone signals. Almost all cells (88%) classified as luminance cells were cone nonopponent, with a broad distribution of cone weights. Most cells (73%) classified as color-luminance cells were cone opponent. This result supports our conclusion that V1 color-luminance cells are double-opponent. Such neurons are more sensitive to color boundaries than to areas of color and thereby could play an important role in color perception. The color-luminance population had a broad distribution of L/M cone weight ratios, implying a broad distribution of preferred colors for the double-opponent cells.

Action Potentials↗

[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↗

Immediate early gene expression in cat visual cortex during and after the critical period: differences between EGR-1 and Fos proteins.

Immediate early gene (IEG) expression in the cat visual cortex is highly responsive to visual input and may initiate genetic mechanisms responsible for neuronal plasticity. The present study used immunohistochemical methods to address two issues regarding IEG expression in response to visual input. One was to define the differential response of distinct IEG families by comparing EGR-1 (also termed zif-268, NGFI-A, and Krox-24) and Fos proteins. The second was to determine whether IEG expression, in addition to reflecting neural activity, is related to the state of plasticity by comparing young and adult visual cortex. Immunoreactivity of the two IEG proteins was compared between 5-week-old and adult cats under three conditions of visual input: ambient light to assess basal levels of expression, 1 week of darkness to assess the effect of reduced activity, and exposure to light after 1 week of darkness to determine rapid changes in expression as a result of visual input. At both ages, there were marked differences in the expression of the two IEG proteins. EGR-1 responded to visual input with sustained changes in its level of expression. It showed high basal levels, reduced expression in darkness, and a rapid return to high constitutive levels with the introduction of light. Fos showed a markedly different profile. It had very low basal expression which was not demonstrably affected by darkness and its principal response was a marked transient induction upon exposure to light after darkness. These unique changes in expression highlight the complex response across IEGs to environmental input and suggest a genetic "on/off' signaling mechanism. There were marked differences in the laminar distribution of EGR-1 and Fos proteins between young and adult cats. In young animals, cells in all visual cortical layers showed high levels of EGR-1 and Fos proteins. In adults, immunostaining was largely specific to cells located above and below layer IV and only very faint labeling occurred within layer IV. These differences in laminar distribution between ages are inconsistent with a simple explanation of IEG expression in terms of neural activity level; rather, they suggest a relation between IEG expression and the state of plasticity in visual cortex.

Animals↗

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↗

The role of functional MR imaging in patients with ischemia in the visual cortex.

BACKGROUND AND PURPOSE: Functional MR (fMR) imaging is based on changes in regional blood flow. The purpose of this study was to evaluate the role of fMR imaging for detection of a vascular compromised status in the occipital lobe in patients with ischemia in the visual cortex. METHODS: We performed fMR imaging in seven control subjects and seven patients with symptoms and signs of visual cortical transient ischemia and/or infarct. fMR imaging was performed using a gradient-echo sequence with the 2D fast low-angle shot technique. An axial slice including both visual cortices was selected, and stimulation of the visual cortex was performed using a red photostimulator. The number of activated pixels in each primary visual cortex area were counted and an asymmetry ratio [AR (%) = 100 x (R-L)/(R+L)/2] was calculated. Patients and control subjects underwent visual field examination, conventional MR imaging, and vascular imaging (MR angiography in all patients and control subjects, conventional catheter angiography in two patients). fMR imaging results were compared with the results of a visual field examination, conventional MR imaging, and vascular imaging. RESULTS: fMR imaging of the patients showed significant activation asymmetry (P <.05) compared with that of control subjects. Vascular abnormalities in the posterior circulation were found in all seven patients. By conventional MR imaging, five patients were found to have infarction in the occipital lobe and the remaining two patients showed no abnormality. In visual field examination, six of the seven patients showed homonymous hemi- or quadrantanopsia suggesting postchiasmic abnormalities, and the remaining patient had normal findings. fMR imaging showed decreased activity in the visual cortices corresponding to vascular abnormalities (seven of seven patients), permanent infarction (five of seven patients), or visual field defect (six of seven patients). Two patients with normal conventional MR imaging had vascular lesions in the posterior circulation, and fMR imaging showed decreased activity in the corresponding visual cortices. One patient with normal visual field examination had multifocal stenosis in the posterior cerebral artery without infarction, and fMR imaging showed decreased activity in the corresponding visual cortex. CONCLUSION: fMR imaging of the visual cortex may be a sensitive method for the detection of vascular-compromised status in the occipital lobe.

Adult↗