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R D Freeman

Publications and source records attributed to R D Freeman.

At least 19 recordsLinked to original sources

Direction selectivity of cells in the cat's striate cortex: differences between bar and grating stimuli.

We have investigated the notion that directional responses of cells in the visual cortex depend on the type of stimulus used to drive the cell. Specifically, we have asked if sinusoidal gratings provide a different estimate of direction selectivity than bars that are brighter or darker than the background. Using standard techniques, we recorded from 176 cells in the visual cortex of nine cats. For each cell, bright bars, dark bars, and sinusoidal gratings were presented in a randomly interleaved fashion. Complex cells exhibited around twice as many direction-selective as nondirection-selective responses. Estimates of direction selectivity were nearly identical for bright and dark bars and for gratings. For simple cells, a similar ratio of direction-selective to nondirection-selective responses was observed for gratings. However, a larger proportion of simple cells were classified as direction selective when bars were used for stimulation. A simple cell that exhibited direction selectivity to a grating behaved in a similar manner when stimulated with bright or dark bars. However, in contrast to complex cells, some simple cells classed as directionally nonselective on the basis of their responses to gratings, displayed directionally selective behavior to bars. In addition, the preferred directions for dark and bright bars sometimes differed. These results demonstrate that the classification of a simple cell as directionally selective or nonselective can depend critically on the visual stimulus used.

Animals

Organization of suppression in receptive fields of neurons in cat visual cortex.

1. The response to an optimally oriented stimulus of both simple and complex cells in the cat's striate visual cortex (area 17) can be suppressed by the superposition of an orthogonally oriented drifting grating. This effect is referred to as cross-orientation suppression. We have examined the spatial organization and tuning characteristics of this suppressive effect with the use of extracellular recording techniques. 2. For a total of 75 neurons, we have measured the size of each cell's excitatory receptive field by use of rectangular patches of drifting sinusoidal gratings presented at the optimal orientation and spatial frequency. The length and width of these grating patches are varied independently. Receptive-field length and width are determined from the dimensions of the smallest grating patch required to elicit a maximal response. 3. The extent of the area from which cross-orientation suppression originates has been measured in an analogous manner. Each neuron is excited by a patch of drifting grating the same size as the receptive field. The response to this stimulus is modulated by a superimposed patch of grating having an orthogonal orientation. After selecting the spatial frequency that produces maximal suppression, the response of each cell is examined as a function of the length and width of the orthogonal (suppressive) grating patch. Results from 29 cells show that the dimensions of the orthogonal grating patch required to elicit maximal suppression are similar to, or smaller than, the dimensions of the excitatory receptive field. Thus cross-orientation suppression originates from within the receptive field. 4. For some cells the spatial frequency tuning of the suppressive effect is much broader than the spatial frequency tuning for excitation. In these cases it is possible to find a spatial frequency that produces suppression but not excitation. With the use of a suppressive stimulus having this spatial frequency, we examined the strength of suppression as a function of orientation for 11 cells. These tests show that suppression occurs at all orientations, including the preferred orientation for excitation. In some cases, suppression is somewhat stronger at the preferred orientation for excitation than at any other orientation. 5. For 12 cells we varied the relative spatial phase between the optimally oriented and orthogonal gratings. In all cases the magnitude of suppression is largely independent of the relative spatial phase. 6. For three binocular cells we examined whether the suppressive effect of a grating oriented orthogonal to the optimum could be mediated dichoptically.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals

Oscillatory discharge in the visual system: does it have a functional role?

1. The discharge of individual neurons in the visual cortex and lateral geniculate nucleus (LGN) of anesthetized and paralyzed cats and kittens was examined for the presence of oscillatory activity. Neural firing was evoked through the monoptic or dichoptic presentation of drifting gratings and random sequences of flashed bars. The degree to which different oscillatory frequencies were present in neural discharge was quantified by computation of the power spectra of impulse train responses. 2. Action potentials from single cells were recorded extracellularly and isolated on the basis of amplitude. Receptive-field properties of the neurons under study were characterized initially by their discharge in response to gratings of sinusoidal luminance. By varying orientation and spatial frequency, optimal stimulus characteristics were determined. Oscillation analysis was performed on spike trains acquired during repeated presentations of the optimal stimulus by identification of power spectra peaks in the frequency range of rhythmic potentials observed in electroencephalograph studies (30-80 Hz). The amplitude and frequency of the largest peak in this range was used to characterize oscillatory strength and frequency. All discharge in which the peak amplitude exceeded the high-frequency noise by a factor > 1.5 was classified as oscillatory. 3. Of the 342 cortical cells examined, 147 cells displayed oscillatory activity in the 30 to 80-Hz range during portions of their visual response. Sixty out of 169 simple cells, 82 out of 166 complex cells, and 5 out of 7 special complex cells exhibited oscillations. There was no laminar bias in the distribution of oscillatory cells; the proportions of oscillatory cells were similar in all layers. All oscillatory discharge was variable with respect to frequency and strength between successive presentations of the same optimal stimulus. In as little as 10 s, for example, peak frequencies shifted by a factor of two. For many cells, these trial-to-trial variations obscured detectable oscillations when all trials were averaged together. 4. The potential role of neuronal maturation in the generation of oscillatory activity was investigated by studying neuronal responses from kittens at 4 wk postnatal. Of the 80 kitten cells studied, 27 exhibited oscillatory discharge. Although oscillations in the kitten visual cortex spanned the same frequency range as that seen in the adult, oscillations in the midfrequency range (36-44 Hz) are more common in the adult cortex. 5. To explore the possibility that oscillations might play a functional role in vision, we investigated the dependence of oscillations on different stimulus parameters.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals

Development of binocular vision in the kitten's striate cortex.

Studies of the development and plasticity of the visual pathway are well documented, but a basic question remains open: what is the physiological status of the system prior to extensive visual experience? Somewhat conflicting answers have been put forward, and in a major area, binocular vision, reports have ranged from severe immaturity to well-developed maturity. This is an important question to resolve since binocular cells in the visual cortex are thought to be the neural substrate for stereoscopic depth perception. We have addressed this question by recording from single cells in the striate cortex of kittens at postnatal ages 2, 3, and 4 weeks and from adults for comparison. Gratings with sinusoidal luminance distribution are presented to left, right, or both eyes. For each cell, we determine optimal values for orientation and spatial frequency. Relative phase (retinal disparity) is then varied in a dichoptic sequence so that binocular interaction may be studied. Results are as follows. In the normal adult, we have shown in previous work that most binocular interaction in the visual cortex can be accounted for on the basis of linear summation. Results from 3 and 4 week postnatal kittens are closely similar to those from adults. All types of binocular interaction found in adults are present in kittens. This includes phase-specific and non-phase-specific suppression or facilitation. Furthermore, monocular and binocular tuning characteristics are comparable in kittens and adults. The clear changes that occur with age are optimal spatial frequencies and peak responses. In addition, at 2 weeks, there is a substantially higher proportion of monocular cells compared to other ages and correspondingly, lower relative numbers of cells that exhibit phase-specific or suppressive binocular interactions. From increases in optimal spatial frequency and interpupillary distance with age, we calculated predicted changes in binocular disparity thresholds (stereo acuity) with age. Although there are methodological limits with respect to the behavioral testing of young kittens, the predicted results are comparable to some of the values obtained. Considered together, our results show that the physiological apparatus for binocular vision is functional at an early stage in postnatal development. It is possible that the connections that underlie this function are developed rapidly during early postnatal experience. An alternative possibility is that there is an elaborate genetic organization of binocular vision, but our study does not address this issue directly. A combination of these factors may be applicable.(ABSTRACT TRUNCATED AT 400 WORDS)

Aging

Contrast sensitivity in amblyopia: masking effects of noise.

Contrast sensitivities were determined for sinusoidal gratings of varying spatial frequencies with and without the presence of a random noise pattern superimposed on the gratings. Control subjects with normal binocular vision and observers with amblyopia were tested to determine the relative effects of noise on contrast sensitivity. For both amblyopes and normal subjects, contrast sensitivities are reduced by the presence of noise. Effects are maximal at 4 cycles/degree and are minimal at low and high spatial frequencies. Dichoptic presentation of noise and gratings to opposite eyes is equivalent to monoptic results for both amblyopes and normal subjects. Masking effects are eliminated if gratings are drifted while noise patterns are static. The contrast sensitivity of amblyopes is reduced by relatively similar amounts to that of normal subjects when noise is added to the stimulus. Overall, masking effects are virtually identical for amblyopes and for subjects with normal binocular vision.

Adult

Depth is encoded in the visual cortex by a specialized receptive field structure.

Binocular neurons in the visual cortex are thought to perform the first stage of processing for the fine stereoscopic depth discrimination exhibited by animals with frontally located eyes. Because lateral separation of the eyes gives a slightly different view to each eye, there are small variations in position (disparities), mainly along the horizontal dimension, between corresponding features in the two retinal images. The visual system uses these disparities to gauge depth. We studied neurons in the cat's visual cortex to determine whether the visual system uses the anisotropy in the range of horizontal and vertical disparities. We report here that there is a corresponding anisotropy in the cortical representation of binocular information: receptive-field profiles for left and right eyes are matched for cells that are tuned to horizontal orientations of image contours. For neurons tuned to vertical orientations, left and right receptive fields are predominantly dissimilar. Therefore, a major modification is required of the conventional notion of disparity processing. The modified scheme allows a unified encoding of monocular form and binocular disparity information.

Action Potentials

Stereoscopic depth discrimination in the visual cortex: neurons ideally suited as disparity detectors.

The possibility has been explored that a subset of physiologically identifiable cells in the visual cortex is especially suited for the processing of stereoscopic depth information. First, characteristics of a disparity detector that would be useful for such processing were outlined. Then, a method was devised by which detailed binocular response data were obtained from cortical cells. In addition, a model of the disparity detector was developed that includes a plausible hierarchical arrangement of cortical cells. Data from the cells compare well with the requirements for the archetypal disparity detector and are in excellent agreement with the predictions of the model. These results demonstrate that a specific type of cortical neuron exhibits the desired characteristics of a disparity detector.

Animals

On the neurophysiological organization of binocular vision.

The considerable mixing in the visual cortex, of signals from left and right eyes, provides an abundant population of binocularly activated neurons. Based on this and on the fact that cortical cells respond best to different ranges of retinal disparities, it has been proposed that these neurons form the physiological substrate of stereoscopic depth discrimination. We outline reasons here for addressing first the more fundamental issue of the rules of convergence in the visual cortex, for input from the two eyes. We show that most of this convergence may be described by a linear summation process. However, there is a nonlinear mechanism that maintains binocular interaction regardless of large differences in stimulus strength between the eyes. This finding suggests that a cell which appears to be dominated by one eye, when monocular tests are conducted, may respond equally under binocular conditions. In this case, binocular processing for all cortical cells could be uniform and independent of the ocular dominance values determined monocularly. With respect to a neural mechanism for the processing of information concerning different depths in space, we propose an alternative to the conventional notion. First, we identify fundamental problems with the current view. Second, we describe a procedure which allows us to distinguish between the conventional view and our alternative proposal. Standard receptive field mapping techniques are not adequate for determining phase-disparity relationships of the type we require. Therefore, we have employed a reverse correlation procedure which enables efficient and detailed mapping of receptive field structure. Third, we describe preliminary data concerning the physiological mechanism of stereoscopic depth discrimination.

Action Potentials

Cortical plasticity in monocularly deprived immobilized kittens depends on eye movement.

A marked reduction of binocular cells in striate cortex is found if 4-week-old kittens are visually stimulated monocularly while anesthetized and held in a stereotaxic apparatus. If the kittens are paralyzed and artificially respirated, changes are not found unless an eye is moved mechanically. It appears that eye movement and visual stimulation are necessary conditions for deactivation of binocular connections, but neither is sufficient to induce such changes alone.

Animals

Effects of brief uniocular 'patching' on kitten visual cortex.

Studies of the striate cortex of the kitten are reviewed with reference to the effects of very brief periods of uniocular occlusion. It is shown that changes occur in binocular connections within 4 hours after the vision of an eye is blocked. Effects are also found if the occlusion is instituted while an animal is anaesthetized but not if it is paralysed in addition. If 4-hour unilateral 'patching' sessions are given daily, a cumulative uniocular deprivation effect is found if the animals are kept in darkness between the patching sessions. However, the visual cortex is nearly normal if the kittens are allowed binocular vision between patching periods.

Anesthesia

Rotating grating treatment for amblyopia.

The recently published method of treatment of amblyopia by the use of rotating gratings is very similar to an older technique. Used apparently extensively during the 1930s in the United States, the method, called 'visual gyration' was based on use of a rotating disk pattern like a grating. Differences between the old and new procedures are described in this paper.

Amblyopia

Visuomotor restriction of one eye in kittens reared with alternate monocular deprivation.

It is known that kittens reared in ways that restrict movement while visual stimulation is received exhibit deficits in visually guided behavior. Presumably, the behavioral dysfunction is due to a lack of sensorimotor coordination during visual exposure. The current investigation was undertaken to study this effect both physiologically and behaviorally. Two groups of kittens were normally reared until they were nearly 4 weeks old. They were then placed in a darkroom and exposed daily for 1--2 hours while one eye was occluded. On alternate days, alternate eyes were covered. One group was restrained in a body cast while a given eye was exposed, but the kittens were free to move about while the other eye received visual experience. A second control group was alternately occluded, but freely mobile during all exposures. An additional cat was also unrestrained during daily rearing periods and neither eye was ever occluded. Behavioral tests showed clear deficits when the "passive" eye of the restrained-unrestrained group was used. Deficiencies were found in visually guided paw placing, pursuit eye movements, and jumping behavior to a platform. Responses of single cells were studied in area 17 of the visual cortex. Ocular dominance distributions showed marked reductions in binocularity for alternated occluder cats. The eye that had received exposure while animals were active in the restrained-unrestrained group also tended to predominate but the difference was not statistically significant.

Animals