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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

A physiological correlate of the Pulfrich effect in cortical neurons of the cat.

When a swinging pendulum is viewed with a light-attenuating filter before one eye, the pendulum bob is perceived to move in an elliptical path in depth. It is believed that the filter causes this illusion, the Pulfrich effect, by delaying processing of the image in the filtered eye relative to that of the unfiltered eye. We sought a physiological correlate of this effect by studying binocular integration in cortical neurons of cats while they viewed moving stimuli. Special attention was focused on single unit disparity tuning because it is widely believed that depth perception is related to the responses of disparity selective neurons in visual cortex. We found that placing a filter before one of the cat's eyes produced a temporal delay in the cortical response. The temporal delay was always associated with a shift in the neuron's spatial disparity tuning. The observed temporal delays and disparity shifts are comparable with the magnitude of the Pulfrich effect in humans.

Action Potentials

Cortical processing of hyperacuity tasks.

Spatial discrimination thresholds were determined by having subjects make comparisons between stimuli presented successively at the same or at different locations in the visual field. Two tasks were employed, discrimination of line orientation and discrimination of the distance between two parallel lines (spatial interval discrimination). We find that discrimination thresholds based on the comparison of features in spatially-separated stimuli are comparable to those based on the comparison of features in two stimuli presented successively at the same location. This holds even when the stimuli are presented in nonhomologous positions in the visual field or are presented in a manner such that they activate cells in opposite cortical hemispheres in the early visual areas. This shows that discriminability is not determined solely by differences in the firing rates of striate or pre-striate neurons. Rather, it appears that the attributes of visual stimuli are precisely coded and available for comparison at higher levels of visual processing. Implications of this finding for models of hyperacuity are discussed.

Discrimination, Psychological

Monocular and binocular response properties of cells in the striate-recipient zone of the cat's lateral posterior-pulvinar complex.

1. We have studied response properties of single cells in the striate-recipient zone of the cat's lateral posterior-pulvinar (LP-P) complex. This zone is in the lateral section of the lateral posterior nucleus (LP1). Our purpose was to determine basic response characteristics of these cells and to investigate the possibility that the LP-P complex is a center of integration that is dominated by input from visual cortex. 2. The majority (72%) of cells in the striate-recipient zone respond to drifting sinusoidal gratings with unmodulated discharge. 3. Cells in the LP1 are selective to the orientation of gratings, and tuning functions have a mean bandwidth of 31 degrees. More than one-half of these units are direction-selective. The preferred orientation and the tuning widths for the two eyes are generally well matched. However, a few cells exhibited the interesting property of opposite preferred directions for the two eyes. Orientation tuning for a small group of cells was different for the mean discharge and first harmonic components, suggesting a convergence from different inputs to these cells. 4. Two-thirds of LP1 cells are tuned to low spatial frequencies (less than 0.5 c/deg). The tuning is broad with a mean bandwidth of 2.2 octaves. The remaining one-third of the units are low-pass because they show no attenuation of their responses to low spatial frequencies. Both eyes exhibit the same spatial frequency preference and the same spatial frequency tuning. There is a high correlation between spatial frequency and orientation selectivities. 5. All cells tested are tuned for temporal frequency with a sharp attenuation for low frequencies. The optimal values range between 4 and 8 Hz, and the mean bandwidth is 2.2 octaves. 6. Cells in LP1 are mostly binocular. When monocular, cells are almost always contralaterally driven. Dichoptic presentation of gratings reveals the presence of strong binocular interaction. In almost all cases, these interactions are phase specific. The cell's discharge is facilitated at particular phases and inhibited at phases 180 degrees away. These binocular interactions are orientation dependent. 7. Twenty-five percent of the cells with phase-specific binocular facilitation appear to be monocular when each eye is tested separately. For three cells, we observed a non-phase-specific inhibitory effect of the silent eye. 8. Our findings indicate that LP1 cells form a relatively homogeneous group, suggesting a high degree of integration of multiple cortical inputs.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals

Tourette syndrome and otolaryngology.

Tourette syndrome (TS) is the most complex tic disorder and presents primarily in the pediatric population between the ages of two and 15. The otolaryngologist may be consulted to see such a patient because of head and neck or facial tics or more often because of phonic or vocal tics such as throat-clearing, sniffing, coughing or abnormal noises. As this disorder has not appeared in the otolaryngologic literature we undertook a retrospective chart review (N = 72) and follow-up questionnaire related to the otolaryngologic aspects of this disorder.

Child

Binocular interaction in the perigeniculate nucleus of the cat.

We have recorded from single cells in the perigeniculate nucleus (PGN) of the cat to determine their response properties. Quantitative tests have been conducted with sinusoidal gratings. Using optimal stimulus parameters, determined monocularly, we explored binocular interaction by varying the relative phase between dichoptically presented gratings. Monocularly, cells exhibit varying degrees of response specificities with respect to stimulus orientation and spatial frequency. Binocularly, we have identified six types of response. The most prominent, type 1, found for half the cells, is phase-specific binocular interaction at the fundamental frequency component of the drifting grating. For these cells, mean response rate is independent of interocular phase. The remaining types of binocular responses involve varying degrees of interaction at different harmonic components. For a quarter of the sample, no binocular interaction was observed. To investigate the role of cortical input to PGN, visual cortex was removed from some cats. Subsequent study of PGN cells indicated that response properties were generally similar to those found in intact animals. We conclude that PGN response properties are determined primarily by subcortical inputs.

Afferent Pathways

Blockade of intracortical inhibition in kitten striate cortex: effects on receptive field properties and associated loss of ocular dominance plasticity.

We have investigated the importance of GABAergic inhibition for the receptive field properties and plasticity of cells in the visual cortex of kittens. Osmotic minipumps were used to continuously infuse the GABA-antagonist, bicuculline methiodide (BIC), into striate cortex. Extracellular recordings were made during BIC infusion to assess neuronal response properties during the blockade of inhibition. Recordings were also made from other kittens after concurrent monocular deprivation and BIC infusion to investigate the importance of response selectivity for ocular dominance plasticity. The minipump delivery technique was used to produce a large volume of cortex presumably free of GABA-ergic inhibition. Compared to recordings in saline-infused control hemispheres, about half of the cells in bicuculline-infused hemispheres had abnormally low orientation selectivity. The low selectivity was generally accompanied by marked anomalies in several other receptive field properties. Particularly striking was the large size of the receptive fields. At eccentricities less than 10 deg many receptive fields subtended from 10 to over 30 deg of arc. The less selective neurons also had abnormal responses to flashed stimuli, giving strong transient responses to the onset and offset of large stationary stimuli which filled their receptive fields. These results imply that intracortical inhibition normally suppresses responses to stimuli within a large excitatory zone beyond the classical receptive field. Inhibition is necessary for the normal orientation selectivity of many cells, although the selectivity may be partially established by the cell's excitatory input. Additionally, intracortical inhibition appears to be necessary for the antagonism and segregation of ON and OFF receptive field subregions. In our study of plasticity, we exploited the fact that BIC treatment greatly increases the range of stimuli that activate cortical neurons. Kittens were monocularly deprived for 7 days concurrently with cortical infusion of BIC. After cessation of the drug treatment, physiological recordings were made. Response properties had returned to normal but neurons in BIC-infused hemispheres had a significantly reduced ocular dominance shift compared to neurons in control hemispheres. This is probably related to the reduced selectivity of cells during BIC infusion. The suggestion here is that there is diminished ocular dominance plasticity in BIC-infused hemispheres because of an increased probability of correlated activity between spontaneous discharge from the closed eye and the cortical activity evoked by the open eye afferents.

Animals

Cyclopean visual evoked potentials: a new test of binocular vision.

A new objective method is described to test binocular vision. Identical gratings are presented to each eye, but they are drifted in opposite directions. Uniocular stimulation does not produce a response, but binocular combination of the gratings results in robust visual evoked potentials. Data from cats are presented to illustrate the use of this technique.

Animals

Monocularly deprived cats: binocular tests of cortical cells reveal functional connections from the deprived eye.

Animals that are deprived of vision in one eye during a vulnerable phase of development lose visual function of the eye. Although this phenomenon has been studied extensively, little is known about the mechanism of disconnection of the deprived eye from visual cortex. One fundamental question is whether input remains from that eye. We have examined the hypothesis that there is functional input from a deprived eye to visual cortex that cannot be observed with standard alternate tests of each eye. We have employed a robust visual stimulation procedure in which large sinusoidal gratings are presented to each eye, as well as to both eyes together, at varying relative phases or retinal disparities. Monocular and binocular stimulation was used to test kittens unilaterally deprived for brief, intermediate, or long periods. A fourth group of kittens was studied as normal controls. Standard methods were used to record from single cells in the striate cortex. After initial qualitative exploration of receptive fields, all testing and analysis were quantitative. As expected, monocular tests revealed that, for most cells, the deprived eye was ineffective, i.e., did not activate the unit. This effect was increasingly pronounced as the length of deprivation was increased. However, binocular tests revealed that a large fraction of these cells (30-40%) was clearly influenced by and therefore functionally connected to the deprived eye. This interaction was phase-selective, or suppressive and not selective for phase. There was no indication that the connections that remained were of a specific type, i.e., excitatory or inhibitory. Therefore, excitation and inhibition appear equally resistant to the effects of monocular deprivation. However, with long-term deprivation, we find minimal evidence of functional input from the deprived eye. We conclude that the effects of monocular deprivation occur over a considerably longer time period than was previously thought.

Aging

Binocularly deprived cats: binocular tests of cortical cells show regular patterns of interaction.

If an animal is prevented from receiving visual experience during an early developmental phase, pronounced dysfunctions are observed. Physiological tests reveal gross abnormalities in the striate cortex. Cells in visual cortex are either unresponsive of their response characteristics are erratic. Although fewer than normal numbers of binocular cells are found in cats reared with binocular lid suture, a population remains that can be activated by stimulation through either eye. We have studied cortical cells in binocularly deprived cats in order to specify monocular and binocular response characteristics. The primary hypothesis we have examined is that the abnormal response properties of these cells are a result of an irregular structure or substructure of the receptive fields. Kittens were binocularly lid-sutured soon after birth, and were studied physiologically at ages between 7 and 11 months. Standard techniques were used to record from single cells in striate cortex. Drifting gratings were presented to either eye or to both eyes together. In the latter case, the relative interocular phase was varied between the gratings so that the retinal disparity of the stimuli was changed. We confirmed the expected finding that most cells were either unresponsive or erratic in their response. Of the cells that responded, monocular tuning functions for orientation and spatial frequency of the stimulus were often irregular. However, even in these cases, binocular interaction patterns of cortical responses were nearly always highly regular and displayed phase-specific profiles. A model is presented that explains this finding and suggests how binocular deprivation may result in disorganized receptive-field structure.

Animals