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R E Soodak

Publications and source records attributed to R E Soodak.

13 recordsLinked to original sources

Simulation of visual cortex development under lid-suture conditions: enhancement of response specificity by a reverse-Hebb rule in the absence of spatially patterned input.

In this report, I show that a reverse-Hebb synaptic modification rule leads to the enhancement of response specificity of simulated visual cortex neurons in the absence of spatial patterning of the afferent activity. Although it is clear that receptive fields in the visual cortex can be modified by experience, many studies have shown a substantial increase of response specificity in cats deprived of pattern vision by lid suture, leading some to conclude that receptive field properties are essentially hard-wired. The hard-wired vs. experience-dependent controversy can be resolved by assuming that while Hebb-type plasticity is responsible for developmental synaptic changes, the organization of presynaptic activity which exists under conditions of visual deprivation is sufficient to drive the neurons towards greater specificity (Linsker 1986a-c; Miller 1989, 1992; Miller et al. 1989). As a reverse-Hebb rule enhances response specificity by balancing the push-pull system of ON- and OFF-center afferents, the sufficient condition is that the activity of ON- and OFF-center retinal ganglion cells be negatively correlated, a condition which will be met by diffuse illumination as seen through sutured eyelids. Unlike the models of Linsker and Miller and colleagues, which are based on a standard-Hebb rule, the model presented here does not require the presence of a "Mexican hat" spatial patterning of the afferent correlations, which has not been observed experimentally.

Animals↗

Reverse-Hebb plasticity leads to optimization and association in a simulated visual cortex.

The effects of a variable-phase synaptic modification rule on the experience-dependent development of a simulated visual cortex were investigated. Of interest were the process of optimization of the internal representation with respect to orientation, through which the weakly tuned neurons of visually inexperienced animals attain their tightly tuned adult characteristics, and the process of association by which identical stimuli presented to either eye come to evoke identical cortical representations. In its general form, the synaptic modification rule was Hebbian. However, it was not assumed that positive correlation of presynaptic and postsynaptic activity would lead to an increase in synaptic weight. The relative phase of presynaptic vs. postsynapatic activity that would effect an increase in synaptic weight was a parameter of the modification rule. When this parameter was zero, synaptic modification conformed to the standard Hebbian type. With a value of 180 deg, or the reverse-Hebb condition, negative correlation between presynaptic and postsynaptic activity led to increased synaptic weight. It was found that a synaptic modification rule of the reverse-Hebb type not only optimized the cortical representation, and associated the representations from the two eyes, but was quite stable with respect to retaining the optimized state for long periods of learning.

Animals↗

Fine structure of receptive-field centers of X and Y cells of the cat.

We investigated the fine structure of receptive field centers of X and Y cells of the retina and lateral geniculate nucleus of the cat using sinusoidal grating stimuli of high spatial frequency. By measuring orientation tuning and spatial-frequency tuning at multiple orientations, the two-dimensional sensitivity distribution was examined. We found that receptive-field centers typically have multiple sensitivity peaks that can be modeled as several spatially offset subunits. A subunit structure was found in both X and Y cells, with an average number of subunits per receptive-field center of approximately 2.9 in X cells and approximately 4.6 in Y cells. In X cells these subunits may correspond to individual cone bipolar inputs. In Y cells, the subunits may reflect the structure of the dendritic tree. The observation of the subunit structure of the receptive-field center, in conjunction with manipulation of the retinal wiring through pharmacological intervention, may provide a new tool for probing the circuitry of the retina.

Animals↗

Directional selectivity and spatiotemporal structure of receptive fields of simple cells in cat striate cortex.

1. Simple cells in cat striate cortex were studied with a number of stimulation paradigms to explore the extent to which linear mechanisms determine direction selectivity. For each paradigm, our aim was to predict the selectivity for the direction of moving stimuli given only the responses to stationary stimuli. We have found that the prediction robustly determines the direction and magnitude of the preferred response but overestimates the nonpreferred response. 2. The main paradigm consisted of comparing the responses of simple cells to contrast reversal sinusoidal gratings with their responses to drifting gratings (of the same orientation, contrast, and spatial and temporal frequencies) in both directions of motion. Although it is known that simple cells display spatiotemporally inseparable responses to contrast reversal gratings, this spatiotemporal inseparability is demonstrated here to predict a certain amount of direction selectivity under the assumption that simple cells sum their inputs linearly. 3. The linear prediction of the directional index (DI), a quantitative measure of the degree of direction selectivity, was compared with the measured DI obtained from the responses to drifting gratings. The median value of the ratio of the two was 0.30, indicating that there is a significant nonlinear component to direction selectivity. 4. The absolute magnitudes of the responses to gratings moving in both directions of motion were compared with the linear predictions as well. Whereas the preferred direction response showed only a slight amount of facilitation compared with the linear prediction, there was a significant amount of nonlinear suppression in the nonpreferred direction. 5. Spatiotemporal inseparability was demonstrated also with stationary temporally modulated bars. The time course of response to these bars was different for different positions in the receptive field. The degree of spatiotemporal inseparability measured with sinusoidally modulated bars agreed quantitatively with that measured in experiments with stationary gratings. 6. A linear prediction of the responses to drifting luminance borders was compared with the actual responses. As with the grating experiments, the prediction was qualitatively accurate, giving the correct preferred direction but underestimating the magnitude of direction selectivity observed.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

The accessory optic system of rabbit. I. Basic visual response properties.

1. The response properties of accessory optic system (AOS) neurons were assessed using single-unit extracellular recording from each of the three AOS terminal nuclei [medial, lateral, and dorsal terminal nuclei (MTN, LTN, and DTN)] in the anesthetized rabbit. 2. AOS neurons had large, monocular (contralateral) receptive fields (tens of degrees on a side) and exhibited a pronounced selectivity to the speed and direction of movement of large, textured patterns. The greatest responses occurred at slow speeds on the order of 0.5 degrees/s. 3. MTN and LTN neurons responded best to movement in near vertical directions. However, the stimulus directions corresponding to the greatest excitation and the greatest inhibition both had a posterior component and, thus, the preferred excitatory and inhibitory directions were not opposite each other. DTN neurons responded most strongly to horizontal movement and were excited by temporal to nasal movement. 4. AOS neurons were unresponsive to natural vestibular stimulation presented as sinusoidal oscillations of the rabbit about the yaw, pitch, and roll axes. 5. The response properties of AOS neurons are remarkably similar to those of the ON, direction-selective ganglion cells of the rabbit retina, and therefore this class of ganglion cell is most likely the predominant, if not the only, direct retinal input to the AOS. The local direction-selective properties of AOS neurons can be accounted for by combining the tuning curves of ON, direction-selective ganglion cells in a simple manner. 6. The low speed preference of AOS neurons, along with their large receptive fields suggests that they are suited to complement the vestibular system in detecting self-motion.

Animals↗

The accessory optic system of rabbit. II. Spatial organization of direction selectivity.

1. To compare the spatial organization of the direction selectivity of neurons in the medial terminal nucleus (MTN) of the accessory optic system with that of neurons in the adjacent ventral tegmentum, extracellular single-unit recordings were made in the anesthetized rabbit. The ventral tegmental neurons were located in a region called the visual tegmental relay zone (VTRZ), which is defined by the ventral tegmental terminal field of contralaterally projecting MTN neurons. 2. Some of the present sample of MTN neurons (5 of 34) had monocular receptive fields composed of two parts distinguished by a marked difference in the orientation of their respective direction-selective tuning curves. For one part of the receptive field the preferred excitatory direction was "up," while for the other part it was "down." Such receptive fields for one eye were called bipartite, whereas the more usually encountered MTN receptive fields, which could be characterized by a single direction-selective tuning curve, were called uniform. 3. Of the 16 neurons recorded from the VTRZ, all but one were binocular. For these neurons, both uniform and bipartite receptive fields were found for each eye alone. The only monocular neuron encountered in the VTRZ had a contralateral, bipartite receptive field. 4. The spatial organization of the direction selectivity of bipartite receptive fields strongly suggests that they are suited to represent rotation of the visual field about a horizontal axis located in the vertical plane that divides the receptive field into two parts. 5. The boundary between the two parts of the bipartite receptive fields was found using handheld visual stimuli at one of two azimuthal locations, either close to 45 degrees or between 95 and 125 degrees (the 0 degree reference was rostral in the midsagittal plane). This particular structure of the bipartite receptive fields suggests that their preferred rotation axes have a close spatial relation to the best-response axes of the semicircular canals. 6. Seven VTRZ neurons were antidromically activated by electrical stimulation of the ipsilateral dorsal cap of the inferior olive. Since the receptive fields of VTRZ neurons have many of the structural features characteristic of the receptive fields of rostral dorsal cap neurons we conclude that the spatial organization of the receptive fields of dorsal cap neurons is, for the most part, synthesized prior to the inferior olive.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

The retinal ganglion cell mosaic defines orientation columns in striate cortex.

A computer simulation was used to demonstrate that the tangential organization of orientation columns is a natural consequence of the orderly projection of the mosaic of retinal ganglion cells onto the visual cortex. Parameters of the simulation were taken from published anatomical and electrophysiological data, and the resulting columnar organization of the simulated visual cortex shows many similarities with observations from animals. The model is able to account for a variety of experimental observations, including the presence of orientation columns in visually inexperienced animals.

Animals↗

Linear mechanisms of directional selectivity in simple cells of cat striate cortex.

The role of linear spatial summation in the directional selectivity of simple cells in cat striate cortex was investigated. The experimental paradigm consisted of comparing the response to drifting grating stimuli with linear predictions based on the response to stationary contrast-reversing gratings. The spatial phase dependence of the response to contrast-reversing gratings was consistent with a high degree of linearity of spatial summation within the receptive fields. Furthermore, the preferred direction predicted from the response to stationary gratings generally agreed with the measurements made with drifting gratings. The amount of directional selectivity predicted was, on average, about half the measured value, indicating that nonlinear mechanisms act in concert with linear mechanisms in determining the overall directional selectivity.

Action Potentials↗

Linear mechanism of orientation tuning in the retina and lateral geniculate nucleus of the cat.

1. The orientation tuning of lateral geniculate nucleus (LGN) neurons and retinal ganglion cells (recorded as S potentials in the LGN) was investigated with drifting grating stimuli. 2. Results were compared with a quantitative model, in which receptive fields were constructed from linear, elliptical Gaussian center and surround subunits, and responses could be predicted to gratings of any spatial frequency at any orientation. 3. The orientation tuning of X and Y retinal ganglion cells and LGN neurons was shown to result from the linear mechanism of receptive-field elongation, as data from these cells could be well fit with this model. 4. The responses of LGN neurons and their input retinal ganglion cells were compared. The orientation tuning of LGN neurons was found to be a reflection of the tuning of their retinal inputs, showing that neither intrageniculate neural interactions nor the corticogeniculate projection play any role in LGN orientation selectivity.

Action Potentials↗

Two-dimensional modeling of visual receptive fields using Gaussian subunits.

Retinal ganglion cell receptive fields have been successfully described using the difference of Gaussians model introduced by Rodieck. As the basic elements of retinal receptive fields are well described by the Gaussian function, it is natural to model receptive fields beyond this level as a convergence of Gaussian subunits. In this paper the full two-dimensional solution to the problem of calculating the response to drifting gratings of a model receptive field composed of Gaussian subunits is presented. The subunits are not required to be radially symmetric, any number is allowed, with any temporal phase delays; and responses are predicted to gratings of any spatial frequency at any orientation. This solution will greatly extend the range of receptive fields that can be modeled as a convergence of Gaussian subunits, including those with orientational and directional selectivities.

Geniculate Bodies↗

Retinal ganglion cells projecting to the rabbit accessory optic system.

Recent evidence from extracellular recording studies indicates that the medial terminal nucleus (MTN) of the rabbit accessory optic system receives inputs from a particular functional class of retinal ganglion cells--specifically, the on-type direction-selective cells. These ganglion cells have been selectively labeled by the retrograde transport of horseradish peroxidase (HRP) injected into the MTN. The number of labeled cells, their distribution over the retina, and their soma areas were determined. In one animal in which the HRP injection completely filled the nucleus, two thousand ganglion cells were labeled. This number agrees with previous estimates of the number of retinal axons terminating in the MTN. Unlike results in avians, none of the ganglion cells was displaced--i.e., they were not Dogiel cells. The density of labeled cells was highest in the visual streak and, overall, the distribution of labeled cells corresponded to the physiologically determined distribution of on-type direction-selective cells. Cells labeled by the HRP injection were among the 20% largest cells in the retina. This result, in conjunction with conclusions from other studies, leads to the prediction that on-type direction-selective cells can be characterized morphologically as cells with large cell bodies and a very extensive dendritic spread in which the dendrites ramify in the vitreal sublamina of the inner plexiform layer.

Animals↗