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Single thalamic neurons project to both lateral suprasylvian visual cortex and area 17: a retrograde fluorescent double-labeling study.

Area 17 and the posteromedial lateral suprasylvian (PMLS) visual cortex receive inputs from three thalamic nuclei in common: the lateral division of the lateral posterior nucleus (LPl), the C-laminae of the lateral geniculate nucleus (LGNd), and the medial interlaminar nucleus (MIN). The present study determined whether these projections originate from the same cells via bifurcating axons or from separate populations of cells. Double-label retrograde transport techniques were used to label cells projecting to area 17 with one fluorescent dye and to label cells projecting to PMLS cortex with a different dye. The two dyes used were fast blue and Evans blue. Following injections into the two cortical areas, some cells were double labeled and some were single labeled in all three thalamic nuclei studied. However, the relative number of double- and single-labeled cells, as well as the relative number of cells single-labeled following injections into each cortical area, differed among the three thalamic nuclei. In both MIN and the C-laminae of the LGNd, the number of double-labeled cells was small. Similarly, the number of cells single labeled with the dye placed in PMLS cortex was small in these two nuclei. In contrast, a relatively large number of cells were single labeled with the dye placed in area 17, especially in the C-laminae of the LGNd. These results suggest that in both MIN and the C-laminae of the LGNd, few cells project to both area 17 and the PMLS cortex, few cells project only to PMLS cortex, and a relatively greater number of cells project only to area 17. In LPl, many cells were labeled after the cortical injections. In fact, when the areas of densest labeling for both dyes overlapped, almost every labeled cell in LPl was double labeled. This indicates that almost all LPl cells that project to one cortical area also project to the other via a bifurcating axon.

Animals↗

Inhibition of axoplasmic transport in the developing visual system of the rat: IV. Quantitative Golgi, electron microscopic, and histochemical analyses of the maturation of the visual cortex.

Intraocularly injected colchicine suppresses axonal transport within the developing rat's optic nerve throughout the critical period of visual system development. This results in a stunting of retinofugal terminals and relay neurons in the lateral geniculate nucleus. The present study focuses upon the effects of this unique form of developmental deprivation on the maturation of the visual cortex. Colchicine, in concentrations of from 10(-5) to 10(-2) M, was injected into the eyes of albino rats at birth or at 5, 10, or 15 days of age. Litters were killed at 5 to 50 days after this single injection, and the brains were processed for Nissl, rapid Golgi, histochemical, or electron microscopic analysis. The following results were obtained: Planimetry of coronal sections of the striate cortex revealed a reduction in the thickness of the cortex and in the ratio of neuropil area to neuronal soma area contralateral to the injected eye which was confined principally to layer IV, lower layer III, and upper layer V. This effect was inversely related to postnatal age at injection and directly proportional to colchicine concentration. A rapid Golgi analysis of 51 pairs of layer V pyramidal neurons in control and experimental cortex demonstrated a reduction in the number and size of spines along the portion of the apical dendrite passing through lower layer III and IV following colchicine administration at birth or 5 or 10 days of age but no significant change in the branching pattern of the entire dendritic arbor. Electron microscopy revealed a reduction in the number of small, asymmetric synaptic complexes with the result that the average size of remaining profiles was increased in layers III and IV. Histochemical analysis of cortical succinic dehydrogenase and cytochrome oxidase revealed a distinct band of intense enzyme activity in lower layers III and IV in normal cortex at 20-30 days of age. This band was significantly reduced in intensity after neonatal injection of colchicine as shown by densitometric measurements and comparison of experimental and control cortex. It is concluded that the geniculocortical projection, while not affected directly by colchicine administration, is altered by the secondary effects of axonal transport suppression, leading to an alteration in the establishment of cortical synaptic patterns and arborizations of their postsynaptic neurons whose dendrites are located in those layers recipient to this projection.

Animals↗

Reduction of transiently expressed acetylcholinesterase activity in developing thalamocortical projections does not affect the mature pattern of basal forebrain projections to visual cortex.

Experiments tested the hypothesis that acetylcholinesterase (AChE) activity, expressed transiently in developing thalamocortical projections, serves to limit the growth of basal forebrain cholinergic projections into thalamocortical recipient zones. Newborn rats were subjected to enucleation, a procedure that eliminates transient AChE activity in developing visual cortex. After 3-8 weeks survival, AChE histochemical techniques revealed no alteration in the pattern of AChE positive basal forebrain axons in visual cortex. These data indicate that transient AChE activity in developing sensory cortex does not limit ingrowth of basal forebrain cholinergic axons.

Acetylcholinesterase↗

Inhibition of nitric oxide synthase does not alter ocular dominance shifts in kitten visual cortex.

1. Since nitric oxide has been proposed as a feedback factor in plasticity in the hippocampus, we tested whether it might also be a feedback factor in sensory-dependent plasticity in the cat visual cortex. 2. The effects of monocular deprivation were compared between eight hemispheres with infusion of a nitric oxide synthase inhibitor, and eight control hemispheres with either infusion of the inactive isomer, or no infusion. Although nitric oxide synthase activity was reduced significantly, the ocular dominance histograms were not substantially different in the two groups of animals. We conclude that the feedback factor for sensory-dependent plasticity in the visual cortex is likely to be some factor other than nitric oxide.

Animals↗

Long-term depression is induced in Ca2+/calmodulin kinase-inhibited visual cortex neurons.

To elucidate a role of Ca2+/calmodulin-dependent protein kinase II (CaMKII) in induction of long-term potentiation (LTP), KN-62, a selective inhibitor for CaMKII, was injected into layer 2/3 neurons of sliced visual cortex obtained from young rats. Tetanic stimulation (5 Hz, 1 min) applied to the white matter after the KN-62 injection induced long-term depression (LTD) of excitatory postsynaptic potentials (EPSPs) evoked by test stimulation of the white matter in 9 of the 14 cells tested. However, EPSPs evoked by test stimulation of the non-tetanized site were not changed, indicating that the induction of LTD was input-specific. Simultaneously, recorded field potentials which were derived from neurons with intact CaMKII showed LTP. These results suggest that postsynaptic CaMKII plays a role in the induction of LTP/LTD in visual cortex.

Animals↗

[Spatial-frequency characteristics of the receptive fields of cat visual cortex with different rates of stimulus movement].

Velocity ranges exist for complex and simple receptive fields of cat's visual cortex. In these ranges there is a narrow band frequency tuning. In these velocity ranges the location of the maximum on frequency characteristics is constant and does not depend on the velocity of stimulating lattice movement. An increase of the movement velocity brings about a reduction of the reaction value of the receptive fields and a disappearance of the narrow band tuning expressed in the optimal velocity range.

Animals↗

Ontogeny of neurotensin receptor binding sites in the rat visual cortex.

The ontogeny of neurotensin receptor binding sites in the rat visual cortex was examined by in situ receptor autoradiography. Binding sites were present in the embryonic cortex and showed extremely high densities at birth and in early postnatal life. Their densities began a gradual decline in the second postnatal week to reach very low levels in adult animals. These results suggest that neurotensin receptor binding sites may play a role in cortical development.

Animals↗

Sleep enhances plasticity in the developing visual cortex.

During a critical period of brain development, occluding the vision of one eye causes a rapid remodeling of the visual cortex and its inputs. Sleep has been linked to other processes thought to depend on synaptic remodeling, but a role for sleep in this form of cortical plasticity has not been demonstrated. We found that sleep enhanced the effects of a preceding period of monocular deprivation on visual cortical responses, but wakefulness in complete darkness did not do so. The enhancement of plasticity by sleep was at least as great as that produced by an equal amount of additional deprivation. These findings demonstrate that sleep and sleep loss modify experience-dependent cortical plasticity in vivo. They suggest that sleep in early life may play a crucial role in brain development.

Action Potentials↗

Temporal integration in cat visual cortex: a test of Bloch's law.

Some units in the cat visual cortex fail to respond to a briefly flashed bar and it has been suggested that such neurons function as visual integrators with a long time constant. To test this integrator hypothesis, a study was made using presentations of a bar, flashed over the receptive field for various durations and at different luminances. Some cortical cells indeed showed an increase in the time to peak latency and in the response amplitude when stimulus duration was prolonged up to 320 msec. Such units obeyed Bloch's law for durations over 100 msec.

Animals↗

A model for the estimate of local image velocity by cells in the visual cortex.

Some computational theories of motion perception assume that the first stage en route to this perception is the local estimate of image velocity. However, this assumption is not supported by data from the primary visual cortex. Its motion sensitive cells are not selective to velocity, but rather are directionally selective and tuned to spatio-temporal frequencies. Accordingly, physiologically based theories start with filters selective to oriented spatio-temporal frequencies. This paper shows that computational and physiological theories do not necessarily conflict, because such filters may, as a population, compute velocity locally. To prove this point, we show how to combine the outputs of a class of frequency tuned filters to detect local image velocity. Furthermore, we show that the combination of filters may simulate 'Pattern' cells in the middle temporal area (MT), whereas each filter simulates primary visual cortex cells. These simulations include three properties of the primary cortex. First, the spatio-temporal frequency tuning curves of the individual filters display approximate space-time separability. Secondly, their direction-of-motion tuning curves depend on the distribution of orientations of the components of the Fourier decomposition and speed of the stimulus. Thirdly, the filters show facilitation and suppression for responses to apparent motions in the preferred and null directions, respectively. It is suggested that the MT's role is not to solve the aperture problem, but to estimate velocities from primary cortex information. The spatial integration that accounts for motion coherence may be postponed to a later cortical stage.

Animals↗

Images of illusory motion in primary visual cortex.

Illusory motion can be generated by successively flashing a stationary visual stimulus in two spatial locations separated by several degrees of visual angle. In appropriate conditions, the apparent motion is indistinguishable from real motion: The observer experiences a luminous object traversing a continuous path from one stimulus location to the other through intervening positions where no physical stimuli exist. The phenomenon has been extensively investigated for nearly a century but little is known about its neurophysiological foundation. Here we present images of activations in the primary visual cortex in response to real and apparent motion. The images show that during apparent motion, a path connecting the cortical representations of the stimulus locations is filled in by activation. The activation along the path of apparent motion is similar to the activation found when a stimulus is presented in real motion between the two locations.

Adult↗

An incremental Hebbian learning model of the primary visual cortex with lateral plasticity and real input patterns.

We present a simplified binocular neural network model of the primary visual cortex with separate ON/OFF-pathways and modifiable afferent as well as intracortical synaptic couplings. Random as well as natural image stimuli drive the weight adaptation which follows Hebbian learning rules stabilized with constant norm and constant sum constraints. The simulations consider the development of orientation and ocular dominance maps under different conditions concerning stimulus patterns and lateral couplings. With random input patterns realistic orientation maps with +/- 1/2-vortices mostly develop and plastic lateral couplings self-organize into mexican hat type structures on average. Using natural greyscale images as input patterns, realistic orientation maps develop as well and the lateral coupling profiles of the cortical neurons represent the two point correlations of the input image used.

Algorithms↗

[Binocular interaction in the visual cortex during changes in the intensity of monocular stimuli in different directions].

The characteristics of binocular interaction in the visual cortex were studied in rats and cats in conditions of dichotic presentation of monocular stimuli changing in intensity in opposite directions. Different combinations of stimuli intensities produced in intact animals a predominance of facilitation of binocular EPs. Depression of EPs was seen in rats subjected to weak contralateral and strong ipsilateral stimuli. After callosotomy EP facilitation in cats was less pronounced. In rats both effects of binocular interaction were enhanced and were recorded in conditions of photostimulation opposite to preoperational ones. It was found that the differences in the effects of binocular interaction before and after callosotomy resulted from changes in occlusional interrelations and elimination of transcallosal influences. An assumption is made that the effects of binocular interaction are largely responsible for specific characteristics of the formation of interhemispheric asymmetry in binocular vision.

Animals↗

Intracortical connectivity revealed by spike-triggered averaging in slice preparations of cat visual cortex.

Intracortical connectivity was studied in slice preparations of cat's visual cortex by spike-triggered averaging. The experiments documented the unitary postsynaptic potentials underlying the inhibitory and excitatory connections from layer III-IV border cells to supragranular cells, as demonstrated previously by cross-correlation studies. In addition the analysis demonstrated the existence of two excitatory connections, between supragranular and layer V cells, that were not detectable in previous cross-correlation studies.

Animals↗

Functional morphology of the feedback pathway from area 17 of the cat visual cortex to the lateral geniculate nucleus.

Two approaches were adopted to study the pattern of connectivity between the cat visual cortex and lateral geniculate nucleus. Fourteen individual cortico-geniculate axons were labeled and reconstructed after intracellular or extracellular injection of biocytin into regions of known receptive-field position and ocular dominance preference, and the distribution of boutons from multi-axon clusters was mapped in three dimensions and compared with the locations of strategically placed geniculate recordings made in the same tissue. The results show that the feedback has an accurate retinotopic component but that individual axons are both more extensive and more selective than described previously. Area 17 feedback axons terminate primarily in layers A and A1, but the distribution of terminal boutons is strongly biased (3:1 ratio) toward the layer that matches their eye preference. Thus, those driven by the contralateral eye preferentially target layer A, and those driven by the ipsilateral eye target layer A1. Each axon also innervates the perigeniculate nucleus (PGN), but the pattern is otherwise variable, suggesting that there are different axonal classes. The terminal fields of individual axons are much larger than described previously, with a maximum spread of 500-1500 microns. Nevertheless, the projection from a given location in area 17 has a center of maximum terminal density 400-500 microns across, which is in retinotopic correspondence with the aggregate receptive field of the cortical cells of origin. The surrounding zone of relatively sparse boutons, however, must permit corticofugal cells to influence visual processing well beyond the regions over which their own responses summate. It follows that any geniculate cell receives corticofugal input covering an equally extensive area of visual space.

Animals↗

Neural network model of visual cortex for determining surface curvature from images of shaded surfaces.

The visual system can extract information about shape from the pattern of light and dark surface shading on an object. Very little is known about how this is accomplished. We have used a learning algorithm to construct a neural network model that computes the principal curvatures and orientation of elliptic paraboloids independently of the illumination direction. Our chief finding is that receptive fields developed by units of such model network are surprisingly similar to some found in the visual cortex. It appears that neurons that can make use of the continuous gradations of shading have receptive fields similar to those previously interpreted as dealing with contours (i.e. 'bar' detectors or 'edge' detectors). This study illustrates the difficulty of deducing neuronal function within a network solely from receptive fields. It is also important to consider the pattern of connections a neuron makes with subsequent stages, which we call the 'projective field'.

Algorithms↗

Neural connections between the lateral geniculate nucleus and visual cortex in vitro.

Neural connections were established in cocultures of rat visual cortex (VC) and lateral geniculate nucleus (LGN), which were isolated in early infancy. Morphological and electrophysiological studies showed that the cortical laminar organization of afferent and efferent connections in the coculture preparations was similar to that in the adult VC. The results indicate the existence of intrinsic mechanisms in VC and LGN that guide the formation of synaptic connections with the appropriate targets.

Afferent Pathways↗