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G G Blasdel

Publications and source records attributed to G G Blasdel.

15 recordsLinked to original sources
PubMed1995

Neural pattern formation via a competitive Hebbian mechanism.

In this contribution we investigate a simple pattern formation process [9,10] based on Hebbian learning and competitive interactions within cortex. This process generates spatial representations of afferent (sensory) information which strongly resemble patterns of response properties of neurons commonly called brain maps. For one of the most thoroughly studied phenomena in cortical development, the formation of topographic maps, orientation and ocular dominance columns in macaque striate cortex, the process, for example, generates the observed patterns of receptive field properties including the recently described correlations between orientation preference and ocular dominance. Competitive Hebbian learning has not only proven to be a useful concept in the understanding of development and plasticity in several brain areas, but the underlying principles have have been successfully applied to problems in machine learning [22]. The model's universality, simplicity, predictive power, and usefulness warrants a closer investigation.

Animals↗
PubMed1993

Geometry of orientation and ocular dominance columns in monkey striate cortex.

In addition to showing that ocular dominance is organized in slabs and that orientation preferences are organized in linear sequences likely to reflect slabs, Hubel and Wiesel (1974a) discussed the intriguing possibility that slabs of orientation might intersect slabs of ocular dominance at some consistent angle. Advances in optical imaging now make it possible to test this possibility directly. When maps of orientation are analyzed quantitatively, they appear to arise from a combination of at least two competing themes: one where orientation preferences change linearly along straight axes, remaining constant along perpendicular axes and forming iso-orientation slabs along the way, and one where orientation preferences change continuously along circular axes, remaining constant along radial axes and forming singularities at the centers of the spaces enclosed. When orientation patterns are compared with ocular dominance patterns from the same cortical regions, quantitative measures reveal (1) that singularities tend to lie at the centers of ocular dominance columns, (2) that linear zones (arising where orientation preferences change along straight axes) tend to lie at the edges of ocular dominance columns, and (3) that the short iso-orientation bands within each linear zone tend to intersect the borders of ocular dominance slabs at angles of approximately 90 degrees.

Animals↗
PubMed1992

Orientation selectivity, preference, and continuity in monkey striate cortex.

Maps of orientation preference and selectivity, inferred from differential images of orientation (Blasdel, 1992), reveal linear organizations in patches, 0.5-1.0 mm across, where orientation selectivities are high, and where preferred orientations rotate linearly along one axis while remaining constant along the other. Most of these linear zones lie between the centers of adjacent ocular dominance columns, with their short iso-orientation slabs oriented perpendicular, in regions enjoying the greatest binocular overlap. These two-dimensional linear zones are segregated by one- and zero-dimensional discontinuities that are particularly abundant in the centers of ocular dominance columns, and that are also correlated with cytochrome oxidase-rich zones within them. Discontinuities smaller than 90 degrees extend in one dimension, as fractures, while discontinuities greater than 90 degrees are confined to points, in the form of singularities, that are generated when orientation preferences rotate continuously through +/- 180 degrees along circular paths. The continuous rotations through 180 degrees imply that direction preferences are not organized laterally in striate cortex. And they also ensure that preferences for all orientations converge at each singularity, with perpendicular orientations represented uniquely close together on opposite sides. The periodic interspersing of linear zones and singularities suggests that orientation preferences are organized by at least two competing schemes. They are optimized for linearity, along with selectivity and binocularity, in the linear zones, and they are optimized for density near singularities. Since upper-layer neurons are likely to have similarly sized dendritic fields in all regions (Lund and Yoshioka, 1991), those in the linear zones should receive precise information about narrowly constrained orientations, while those near singularities should receive coarse information about all orientations--very different inputs that suggest different perceptual functions.

Algorithms↗
PubMed1992

Differential imaging of ocular dominance and orientation selectivity in monkey striate cortex.

Differential images of ocular dominance, acquired by comparing responses to the two eyes, reveal dark and light bands where cortical cells are dominated by the right and left eyes. These include most (but not all) histochemically stained cytochrome oxidase blobs in their centers. Differential images of orientation, acquired by comparing responses to orthogonal orientations, reveal dark and light bands that are reminiscent of the "orientation columns" reported earlier, on the basis of 2-deoxyglucose (2DG) autoradiograms (Hubel et al., 1978). However, they are shorter and more fragmented because they do not include regions lacking selectivity for orientation. Even though these "bands" derive from orientation-selective areas, comparisons with differential images of other orientations reveal that regions along their centers prefer different orientations. Hence, the orientation preferences inferred from "bands" in single differential images, or single 2DG autoradiograms, are not necessarily incorrect. Interactions between ocular dominance and orientation were investigated by comparing differential images of orientation obtained with binocular and monocular stimulation, as well as by comparing differential images of ocular dominance obtained with different orientations. In both cases, the elicited interactions were minimal, indicating a remarkable and unexpected independence that subsequent experiments revealed arises, at least in part, from a lateral segregation of regions most selective for one eye and regions most selective for one orientation, in the centers and edges of ocular dominance columns. Since this can also be viewed as a lateral correlation between binocularity and orientation selectivity, it fits with the simultaneous emergence of these properties in layers receiving input from layer 4c, and suggests that each of these properties requires the other.

Animals↗
PubMed1985

Intrinsic connections of macaque striate cortex: afferent and efferent connections of lamina 4C.

We have studied the intrinsic organization of macaque striate cortex by tracing the pattern of horseradish peroxidase (HRP)-labeled axons and cell bodies produced by microinjections of HRP into single cortical laminae. Both anterograde and retrograde transport results were used to examine: (1) the pattern of projections from lamina 4C to the superficial layers; (2) the projection from lamina 4C to deeper cortical layers; and (3) the projections to lamina 4C from other cortical laminae. Laminae 4C alpha and 4C beta differ in their pattern of projections to the superficial layers of striate cortex. Axons from neurons in lamina 4C beta ascend through lamina 4B without giving off collaterals and terminate in lamina 4A and in the base of lamina 3. By contrast, axons from neurons in lamina 4C alpha terminate in lamina 4B and less densely in the 4A/3B region. The projection from lamina 4C beta to lamina 4A is particularly dense and is distributed in a patchy fashion immediately above each injection site. The projection from lamina 4C beta to lamina 3B appears less dense and more widespread; we estimate that individual 4C beta axons may spread laterally for more than 400 micron. Furthermore, the pattern of HRP-labeled cell bodies in lamina 4C beta following injections into laminae 4A and 3B provides evidence for a subdivision within 4C beta. These injections always produce a large number of labeled neurons in the upper part of lamina 4C beta, whereas the lower portion contains few labeled neurons that are located immediately under the center of the injection site. Both lamina 4C alpha and lamina 4C beta also contribute less dense projections to the deeper layers of cortex. Lamina 4C beta projects mainly to lamina 6, whereas lamina 4C alpha contributes axon terminals to both lamina 5A and lamina 6. Neurons in lamina 6 provide the bulk of the intracortical projections to lamina 4C. The axons of these neurons are fine in caliber and have a delicate side-spine morphology that is quite distinct from lateral geniculate axon arbors. Neurons in lamina 5A also project onto lamina 4C, but the projections of these neurons appear concentrated in lamina 4C alpha. These results confirm or refine many conclusions about intrinsic connections of striate cortex drawn from Golgi material and suggest new patterns of connections not suspected from previous work.

Afferent Pathways↗
PubMed1985

Intrinsic connections of macaque striate cortex: axonal projections of cells outside lamina 4C.

We have exploited a technique for making small injections of horseradish peroxidase into single cortical laminae in order to study axonal projections in macaque striate cortex. In the preceding paper (Fitzpatrick, D., J. S. Lund, and G. G. Blasdel (1985) J. Neurosci. 5: 3329-3349) we examined the projections of cells in lamina 4C--cells that receive most of their input from the lateral geniculate nucleus. The present paper deals with the projections of neurons that lie outside of lamina 4C. Among our findings are several projections that previously had not been described in the monkey. These include: a strong and precise (point-to-point) projection from lamina 4B to lamina 2/3A, a reciprocal projection from 2/3A back to 4B, a definite projection from lamina 4B to 5B, as well as a prominent input to lamina 6 from 5B. In many cases, we find it possible to trace the flow of visual information through several "circuits" in striate cortex that have, as their output, projections to extrastriate cortex or to the brainstem. Our results offer additional insights in this regard since we are able, in many cases, to compare the lateral spreads of particular projections. These vary and can be separated into at least three categories: those that terminate in a precise, point-to-point, fashion, those that spread widely, and those that terminate in a laterally periodic fashion. In several cases we find evidence for a correlation between specific patterns of projection and known physiological differences between the topographies of laminae that are connected. In cases where two laminae possess similar topographies (for example, where both contain orderly maps for orientation) their interconnections appear precise, with little diffuse spread. In cases where two laminae are characterized by strikingly different topographies (where, for example, one contains an orderly map for orientation and the other a precise map for retinotopic position, but no specificity for orientation), the connections appear more diffuse.

Animals↗
PubMed1984

Physiological organization of layer 4 in macaque striate cortex.

Numerous highly angled electrode penetrations through the opercular region of macaque striate cortex reveal that layers 4A, 4C alpha, and 4C beta--the primary input sublaminae for axons from the lateral geniculate nucleus (LGN)--are retinotopically organized on a fine scale and populated mostly by monocularly driven cells having small receptive fields and lacking orientation selectivity. Layer 4B, which does not receive a direct thalamic input, contains orientationally selective cells, and many of these are also direction selective. To a significant degree the response properties of cells in layers 4C alpha and 4C beta reflect the response properties of their respective afferent inputs, from the magno- and parvocellular laminae of the LGN. Accordingly, cells in layer 4C alpha have lower contrast thresholds and larger minimum response fields than do the cells in layer 4C beta. In contrast to this clear-cut separation, the cells of layer 4A (whose major source of direct LGN input arises from the parvocellular layers) exhibit both high and low contrast thresholds. With regard to the precision of retinotopic mapping that is seen in lamina 4C, it is noteworthy that there is substantial overlap among the minimum response fields of neighboring neurons. Due to a larger mean receptive field size, this overlap is greater in layer 4C alpha than it is in 4C beta. In either sublamina, however, the minimum cortical distance that separates different and nonoverlapping parts of the visual field corresponds closely--within a factor of 2--to the known arborizational spreads of single geniculate afferents.

Animals↗
PubMed1983

Termination of afferent axons in macaque striate cortex.

We used horseradish peroxidase (HRP) to orthogradely label afferent axons in macaque striate cortex. Of the 38 axons that we recovered, nine were recorded intracellularly before being filled with HRP. Light microscope and computer reconstructions of filled processes reveal highly stereotyped patterns of arborization and suggest that there are at least five discrete populations of lateral geniculate nucleus (LGN) afferent axon: (1) those to layer 4C beta, which have extremely circumscribed, dense terminal fields (small branches of which occasionally intrude into 4C alpha) but which have not been shown to project to other laminae; (2) afferents to layer 4A, which in some cases send fine ascending collaterals into layer 2-3 and which do not, apparently, send collaterals to other laminae; (3) afferents to layer 1, which are fine, extend over large distances horizontally, and send collaterals to layer 6A; (4) afferents to the lower two-thirds of layer 4C alpha, which have few or no collaterals in layer 6; and (5) afferents to the upper half of layer 4C alpha, which have arborizing collaterals in layer 6B. Of the nine axons that were recorded intracellularly, those with projections to layer 4C beta (two axons) and to layer 1 (one axon) had color-selective properties, whereas those (six axons) which arborized in 4C alpha all had transient, broad band and highly contrast-sensitive receptive fields. These properties are consistent with derivations from somata in the parvocellular and magnocellular divisions of the LGN, respectively. Afferents to 4C alpha were found to cover approximately 6 times as much surface area as afferents to 4C beta. The preterminal trunks of all axons were found to follow tortuous paths through the neuropil--paths that may derive from axon segregation during development. The wide ranging, patchy distributions of single afferents in 4C alpha suggest that individual 4C alpha axons supply more than one ocular dominance stripe. In one case where the terminal arborization of a 4C alpha axon was mapped against the transneuronally determined pattern of ocular dominance, three separate patches of terminal boutons were indeed found to coincide with the bands of one eye.

Afferent Pathways↗
PubMed1980

Regional variation in the representation of the visual field in the visual cortex of the Siamese cat.

In Siamese cats, many ganglion cell fibers from the temporal retina misproject to the contralateral hemisphere; as a result, each lateral geniculate nucleus contains an abnormally large representation of the ipsilateral visual field. The manner in which the visual cortex processes this aberrant visual information has been examined in several previous studies. In some Siamese cats, the region of the 17/18 border was found to contain an extensive, systematic map of the ipsilateral field, while in other animals no such map was found, and the 17/18 border appeared to represent the zero meridian of azimuth (as in normal cats). These results have led to the suggestion that there are two distinct types of Siamese cat ("Boston" and "Midwestern") which can be distinguished on the basis of cortical topography and the anatomical organization oif the geniculocortical pathway. In the present study, we have recorded from four Siamese cats in order to examine the visual field map in the region of the 17/18 border; in each cat we recorded at anterior coronal levels corresponding to the representation of the lower visual field, and also at more posterior levels near the horizontal meridian representation. In all of the animals we found that the anterior penetrations (corresponding to mean receptive field elevations inferior to -7 degrees) yielded 15-20 degrees of ipsilateral field representation at the 17/18 border; however, the posterior, horizontal meridian penetrations (with mean elevations from +1 degrees to -4 degrees) showed excursions of only about 5 degrees into the ipsilateral field. This large difference in the representation of azimuth was not due to rotation of the eyes during our recording sessions. The finding of appreciable differences in the amount of ipsilateral field represented at different anterior-posterior levels of the same animal might lead to the suggestion that there are not two distinct populations (or types) of Siamese cat with regard to the cortical map of the ipsilateral field. Rather, we raise the possibility that Siamese cats form one population in which there is a continuous variation in the extent of ipsilateral field represented in the cortex.

Animals↗
PubMed1979

Degree of interocular synchrony required for maintenance of binocularity in kitten's visual cortex.

1. The importance of synchronous activation in maintaining cortical binocularity was studied physiologically in kittens that had been reared under different regimens of alternating monocular deprivation. 2. Three different techniques were employed to provide alternate monocular stimulation: a) mechanical shutters placed before the animals' eyes; b) goggles fitted with complementary colored cutoff filters, which restricted visual input to one eye at a time; and c) two rotating gratings that were 90 degrees out of phase. In the third technique, the gratings were always orthogonal to one another and viewed separately through cutoff filters. This allowed us to exploit the orientation selectivity of cortical cells and thereby stimulate them alternately through each eye without simultaneously affecting activity in the dorsal lateral geniculate nucleus (dLGN). 3. We based our conclusions on a sample of 691 neurons, which we recorded in 21 animals. Results with all techniques were remarkably consistent. Binocular cortical inputs predominated at normal or nearly normal levels, even when a number of seconds elapsed between successive exposures of each eye. 4. An interonset interval of at least 10 s was required to make a substantial reduction in binocularity. This interval can be separated into two parts--the duration of exclusive monocular stimulation and the time when neither channel receives input. Of these, the latter appeared to be less important. Blanking times of 0.15--1.0 s did not affect binocularity if the interonset interval was 1 or 10 s; and in one experiment where the blanking time was 9 s, the resulting disruption in binocularity was less than that found with shorter blanking times and the same interonset interval. 5. Our results imply that mechanisms responsible for the disappearance of binocular cortical inputs require independent stimulation of each eye for periods of at least a few seconds; this stimulation must be of a kind that is known to excite cortical cells. Our results with the rotating grafting show, in addition, that the mechanisms whose timing we have measured are intrinsic to the cortex.

Animals↗
PubMed1979

Improved use of tapetal reflection for eye-position monitoring.

A new technique is described for eye-position monitoring in species with strong tapetal reflections. A fiber optic is used to introduce light into the eye, whose optics then produce an image of the fundus on a tangent in front of the animal. The technique simplifies heretofore tedious measurement of cylotorsional changes, as well as providing a very wide view of the fundus. It has been used successfully in conjunction with single-unit recording from the visual system.

Animals↗
PubMed1978

Effect of prior visual experience on cortical recovery from the effects of unilateral eyelid suture in kittens.

The reversibility of monocular deprivation was tested physiologically in twelve kittens which had received varying amounts of normal visual experience prior to unilateral eye closure. Recordings obtained from cells in cortical area 17 indicated that the effects of monocular deprivation are reversible in cats as old as 8 weeks if the animals have been allowed normal visual experience before eye closure.1. Four kittens had their right eyes closed at 3, 4, and 5 weeks of age. All animals were reverse sutured at about 8 weeks of age, and after 2 weeks forced usage of the deprived eye, each animal was tested physiologically for recovery. Single unit recordings revealed a consistent relation between the extent to which the effects of initial deprivation could reverse and the age of the animal at the time of this deprivation. Recordings from a fifth, 8 week old kitten, which had been monocularly deprived for 3 weeks, ensured that our shortest period of deprivation induces a severe deficit.2. Two kittens which were reared in the dark for 4 and 5 weeks before unilateral eye closure, failed to show substantial (greater than 20%) reversal when reverse lid-sutured at 8 weeks of age. This failure suggests that kittens having their eyes closed after normal experience show enhanced reversibility, at least in part, on account of their early experience and not necessarily because of shorter deprivation. A litter-mate of one of these kittens was allowed 9 days of normal vision after dark rearing and before lid suture. Recordings which were performed both before and after 6 days of reverse suture demonstrated a near total (85%) reversal in ocular dominance.3. Three animals, monocularly deprived after some normal experience, recovered substantial numbers of binocular cells after both eyes had been allowed to remain open. In one of these cats the effectiveness of monocular deprivation was confirmed physiologically before eye opening. These findings contrast with those of other workers who have failed to find many binocular cells in kittens that had been allowed to recover, with eye opening, from monocular deprivation which had lasted since birth.4. One kitten was reared for 2 weeks (3-5 weeks of age) with alternating monocular deprivation. Single unit recordings from this cat, which were made after four additional weeks of unilateral eye closure, revealed some cells which had resisted monocular deprivation and which responded strongly to stimulation of the deprived eye. Reverse suture in the same animal led to the recovery of many cells (36%), dominated by the initially deprived eye, but did not result in a reversal in ocular dominance.

Action Potentials↗
PubMed1977

A physiological and behavioural study in cats of the effect of early visual experience with contours of a single orientation.

1. Three kittens were reared in visual environments that consisted of stripes at one of three orientations - horizontal, right oblique, or left oblique. Two additional cats were reared as controls. One of these matured viewing right and left oblique stripes on alternate days. The other experienced a normal visual environment. 2. Following the completion of rearing, and after several weeks of normal visual experience, behavioural testing of the stripe-reared animals demonstrated a deficit in visual acuity for orientations which were not present in the early visual environment. No comparable deficit emerged for either of the control cats. 3. Following 1-3 years of further, normal, visual experience, each of the cats was shipped separately to California where single units were recorded from area 17 of the visual cortex and an effort made to guess the early visual history of each animal which was unknown to the experimenters. Cell samples from each experimental cat and the normal control cat allowed the physiologist to guess their early visual experience correctly. The control cat which matured viewing orthogonal sets of oblique stripes on alternate days demonstrated a bias for horizontal contours in his cell sample. In contrast to units recorded from normal cats, about 80% of which are binocular, only about 30% of the cells recorded from the stripe-reared animals could be influenced by both eyes.

Animals↗
PubMedDate not supplied

Voltage-sensitive dyes reveal a modular organization in monkey striate cortex.

Voltage-sensitive dyes allow neuronal activity to be studied by non-invasive optical techniques. They provide an attractive means of investigating striate cortex, where important response properties are organized in two dimensions. In the present study, patterns of ocular dominance and orientation selectivity were obtained repeatedly from the same patch of cortex using the dye merocyanine oxazolone, together with current image-processing techniques. The patterns observed agree with most established features of monkey striate cortex and suggest a new unit of cortical organization; one that is modular in structure and which appears to link the organization of orientation selectivity with that of ocular dominance.

Animals↗