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R D Freeman

Publications and source records attributed to R D Freeman.

At least 73 records · Page 4Linked to original sources
PubMed1988

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↗
PubMed1988

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↗
PubMed1988

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↗
PubMed1988

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↗
PubMed1987

Receptive field properties of cells in area 19 of the cat.

We have recorded extracellularly from single cells in area 19 of the cat for the purpose of providing a quantitative description of response characteristics. A prominent feature of this area is a high incidence of cells that are end-stopped. Drifting sinusoidal gratings were used to determine spatial and temporal characteristics of the discharge region. In addition, we have conducted independent tests to characterize end zones of receptive fields. When a grating patch was used to stimulate the discharge region alone, all of the cells showed a band-pass spatial frequency tuning characteristic. The optimal spatial frequency ranged from 0.1 to 1.13 cycles/deg, and the distribution had a peak at 0.4 cycles/deg. The bandwidth at half peak amplitude ranged widely from 0.7 to 3.3 octaves (mean 2.0 octaves). When gratings were also presented to the end zones, responses to stimulation of the central region were suppressed. The surround was phase-insensitive in that the relative phase between the grating in the two regions generally did not affect the strength of the suppression. To determine spatial characteristics of the end-zone inhibition, the spatial frequency of the end-zone grating was changed while that for the central pattern was fixed. All cells showed a band-pass characteristic for end-zone inhibition, but in each case, the tuning width was broader than that for excitation. The mean spatial frequency bandwidth of end-zone inhibition was 2.7 octaves. The peak of the inhibition generally coincided with the peak of the excitatory spatial frequency tuning of the discharge center. Considered together, these results show that neurons in area 19 share common properties with those in areas 17 and 18, but they exhibit phase-insensitive end-zone inhibition more frequently.

Animals↗
PubMed1987

Dark-reared cats: unresponsive cells become visually responsive with microiontophoresis of an excitatory amino acid.

The visual system of kittens reared in total darkness is grossly abnormal. Although estimates vary, substantial proportions of cells in the visual cortex of these animals are unresponsive to visual stimulation. Additional cells are weakly responsive or erratic. We have considered the possibility that these neurons receive subthreshold input which might be evident if an excitatory neurochemical agent is applied during extracellular recording with a microelectrode. To test this notion, we have recorded from cells in the striate cortex of dark-reared kittens during microiontophoretic application of an excitatory amino acid, DL-homocysteate (DLH). Using this technique, we find that virtually all cells in the visual cortex of dark-reared kittens are responsive to visual stimulation. Prior to application of DLH, 27% of the cells were unresponsive to visual stimuli. Following iontophoresis of DLH, half of these cells responded with excitatory discharge to visual stimuli and the other half exhibited an inhibitory response in that the elevated maintained activity was suppressed during presentation of a visual stimulus. Additional cells from these animals, which were initially visually responsive, were also studied. For some of these units, responses were weak prior to administration of DLH and we were able to obtain a more clear estimate of selectivity for stimulus orientation during microiontophoresis of the drug. In these cases, and for the few cells which were initially responsive and orientation selective, we observed no major differences in selectivity before and after DLH application.

Animals↗
PubMed1987

Dark-reared cats: responsitivity of cortical cells influenced pharmacologically by an inhibitory antagonist.

When studied physiologically, dark-reared kittens exhibit abnormal responses. Specifically, cortical cells respond poorly, if at all, to visual stimulation. In the preceding paper (Ramoa et al. 1986) we showed that iontophoretic application of an excitatory amino acid allows all cells in this preparation to be excited or suppressed by visual stimuli. In the current study, we have pursued this finding by studying additional dark-reared kittens using iontophoretic application of an inhibitory antagonist, bicuculline methiodide (BIC). Responses of most cells studied were affected by application of BIC. Of cells which were originally weak or unresponsive, 76% became clearly responsive with application of BIC. Of cells which responded without BIC and were orientation-selective, only 33% lost selectivity during application of BIC. Considered together, these findings suggest functional inhibitory input in dark-reared animals which appears to differ from that in normal kittens with respect to its role in selectivity for stimulus orientation.

Animals↗
PubMed1987

Dark-reared kittens: GABA sensitivity of cells in the visual cortex.

Most cells in the visual cortex of dark-reared kittens are unselective for stimulus orientation and we examined the notion that this might be due to insufficiently developed gamma-aminobutyric acid (GABA) receptors. We recorded from cortical neurons and examined their sensitivity to iontophoretically applied GABA. As expected, most units were non-selective for orientation, but application of GABA suppressed impulse activity of these cells just as for orientation selective neurons. This result suggests that the development or maintenance of GABA receptors is not critically dependent on visual experience.

Animals↗
PubMed1987

Binocular interaction in the dorsal lateral geniculate nucleus of the cat.

We have investigated binocular interaction in the dorsal lateral geniculate nucleus (LGN) of the cat. Neurons were recorded extracellularly during visual stimulation with sinusoidal gratings which were presented at different interocular phases (disparities). The large majority of cells (91%) exhibited some type of binocular interaction. For 75% and 16% of the total number of cells, the binocular interaction was inhibitory or facilitatory, respectively. For the remaining 9% of cells, no interaction was evident. In marked distinction from visual cortex, the facilitatory and inhibitory interactions in the LGN are independent of the relative interocular phase of the patterns. Neurons in the LGN are therefore insensitive to the stereoscopic depth cue, retinal disparity.

Animals↗
PubMed1987

Visual orientation and spatial frequency discrimination: a comparison of single neurons and behavior.

Neurons in the visual cortex respond selectively to stimulus orientation and spatial frequency. Changes in response amplitudes of these neurons could be the neurophysiological basis of orientation and spatial frequency discrimination. We have estimated the minimum differences in stimulus orientation and spatial frequency that can produce reliable changes in the responses of individual neurons in cat visual cortex. We compare these values with orientation and spatial frequency discrimination thresholds determined behaviorally. Slopes of the tuning functions and response variability determine the minimum orientation and spatial frequency differences that can elicit a reliable response change. These minimum values were obtained from single cells using receiver operating characteristic (ROC) analysis. The average minimum orientation and spatial frequency differences that could be signaled reliably by cells from our sample were 6.4 degrees (n = 22) and 21.3% (n = 18), respectively. These values are approximately 0.20 of the average full tuning width at one-half height of the cells. Although these average values are well above the behaviorally determined thresholds, the most selective cells signaled orientation and frequency differences of 1.84 degrees and 5.25%, respectively. These values are of the same order of magnitude as the behavioral thresholds. We show that, because of slow fluctuations in a cell's responsivity, ROC analysis overestimates response variability. We estimate that these slow response fluctuations elevated our estimates of single cell "thresholds" by, on average, 30%. Our data point to an approximate correspondence between orientation and spatial frequency discrimination "thresholds" determined behaviorally and those estimated from the most selective single cortical cells. Interpretation of this quantitative correspondence is considered in the discussion.

Animals↗
PubMed1987

The effects of contrast on visual orientation and spatial frequency discrimination: a comparison of single cells and behavior.

We have compared the effects of contrast on human psychophysical orientation and spatial frequency discrimination thresholds and on the responses of individual neurons in the cat's striate cortex. Contrast has similar effects on orientation and spatial frequency discrimination: as contrast is increased above detection threshold, orientation and spatial frequency discrimination performance improves but reaches maximum levels at quite low contrasts. Further increases in contrast produce no further improvements in discrimination. We measured the effects of contrast on response amplitude, orientation and spatial frequency selectivity, and response variance of neurons in the cat's striate cortex. Orientation and spatial frequency selectivity vary little with contrast. Also, the ratio of response variance to response mean is unaffected by contrast. Although, in many cells, response amplitude increases approximately linearly with log contrast over most of the visible range, some cells show complete or partial saturation of response amplitude at medium contrasts. Therefore, some cells show a clear increase in slope of the orientation and spatial frequency tuning functions with increasing contrast, whereas in others the slopes reach maximum values at medium contrasts. Using receiver operating characteristic analysis, we estimated the minimum orientation and spatial frequency differences that can be signaled reliably as a response change by an individual cell. This analysis shows that, on average, the discrimination of orientation or spatial frequency improves with contrast at low contrasts more than at higher contrasts. Using the optimal stimulus for each cell, we estimated the contrast threshold of 48 neurons. Most cells had contrast thresholds below 5%. Thresholds were only slightly higher for nonoptimal stimuli. Therefore, increasing the contrast of sinusoidal gratings above approximately 10% will not produce large increases in the number of responding cells. The observed effects of contrast on the response characteristics of nonsaturating cortical cells do not appear consistent with the psychophysical results. Cells that reach their maximum response at low-to-medium contrasts may account for the contrast independence of psychophysical orientation and spatial frequency discrimination thresholds at medium and high contrasts.

Animals↗
PubMed1986

Binocular summation in normal, monocularly deprived, and strabismic cats: visual evoked potentials.

We have studied visual evoked potentials (VEP) in the cat using dichoptically presented sinusoidal gratings. Our goals were to determine if binocular disparity causes differential responses in the VEP, and to examine the effects of monocular deprivation and convergent or divergent strabismus on the degree of binocular summation. Binocular disparity in stimuli causes no regular alterations of visual evoked responses, except at very low spatial frequencies. However, this apparent selectivity is probably due to luminance modulation in the central retina at low frequencies. The insensitivity to binocular disparity establishes that binocular summation in the VEP may be estimated without regard to the relative phase of gratings presented to the two eyes. Binocular summation of the VEP was examined in normal animals. We found that the ratio of the binocularly evoked response to the largest monocular response (averaged across spatial frequency) ranged from 1.27 to 2.12 (4 animals) and had a mean of 1.48. These values fall within the range which has been reported for human subjects. The degree of summation might be expected to be greatly reduced in strabismic and monocularly deprived animals, in which the majority of the cells are functionally monocular. While summation was found to be reduced in 5 esotropic (convergent) animals (range = 1.13-1.24; mean = 1.18) it was approximately normal in three exotropic (divergent) animals (range = 1.29-2.12; mean = 1.61). However, single unit recordings carried out on the same animals show similar reductions of cells that can be driven through either eye for both groups of animals. Recordings from three monocularly deprived animals, on the other hand, show evidence of binocular interaction in the form of suppression. In this case, response amplitudes obtained using binocular stimulation were consistently and substantially smaller than those obtained from the normal eye alone (range = 0.76-0.85; mean = 0.80). We conclude that convergent and divergent strabismus differ substantially in the degree to which binocular summation is exhibited in the VEP, which in the latter condition, is indistinguishable from the normal cat. Monocular deprivation causes an effect which is markedly different from either form of strabismus in that the deprived eye suppresses the response of the normal eye.

Animals↗
PubMed1986

The binocular organization of simple cells in the cat's visual cortex.

We have studied the manner by which inputs from the two eyes are combined in simple cells of the cat's visual cortex. The stimuli for this study are drifting sinusoidal gratings, shown dichoptically at optimal spatial frequency and orientation. The relative spatial phase (disparity) between the gratings for left and right eyes is varied over 360 degrees. Most simple cells show phase-specific binocular interaction such that response amplitudes and phases vary depending on the relative spatial phase. At one phase, response is greater than either of the monocular responses and often greater than the sum of the two. At the phase 180 degrees away from the optimal, the cell's responses are strongly inhibited and often completely suppressed. Phase-specific binocular interaction disappears when the gratings presented to one eye are made orthogonal to the optimal orientation. The degree of binocular interaction does not depend critically on the ocular dominance of the cells. Simple cells that are nearly equally dominated by each eye always exhibit strong phase-specific interaction. The majority of cells that are strongly dominated by one eye, and even those that appear monocular, show phase-dependent changes in responses. We examined the extent of binocular interaction for cells with preferred orientations near vertical compared with those tuned to other optimal orientations. If these cells are conveying information about depth, one might expect a greater degree of binocular phase-specificity for units preferring nearly vertical orientations, which would then be processing horizontal disparities. We find no evidence for this. Predictions of simple-cell responses are derived from a linear model of binocular convergence in which light-evoked neural signals from each eye are summed linearly to determine cell responses. Data from cells generally follow the prediction of the model for both response amplitude and phase. Deviations from predictions of the linear model are found for a minority of cells. This deviation may be accounted for by a threshold mechanism that comes into play after the linear binocular summation. A small proportion of simple cells that appear monocular by alternate tests of each eye show a purely inhibitory influence from the silent eye. This inhibition is not generally dependent on the relative phase of the gratings. We conclude that most binocular interaction in striate simple cells may be accounted for by linear summation of neural signals from each eye.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1986

The binocular organization of complex cells in the cat's visual cortex.

We have studied the manner by which inputs from the two eyes are combined in complex cells of the cat's visual cortex. The stimuli are drifting sinusoidal gratings presented dichoptically at optimal spatial frequency and orientation. The relative phase between the gratings for left and right eyes is varied over 360 degrees. Approximately 40% of complex cells show phase-specific binocular interaction where response amplitudes vary depending on the relative phase of the gratings shown to the two eyes. This interaction is similar to that observed for most simple cells. We devised a test to examine whether the phase-specific interaction in complex cells results from linear convergence of neural signals at subunits of the receptive fields. The data from this test are consistent with a linear combination model. The phase-specific binocular interaction data from complex cells imply that the optimal relative phase of the receptive field subunits is closely matched. Another type of complex cell, approximately 40% of the total, could be driven through either eye, but exhibited non-phase-specific responses to dichoptically presented gratings. This type of interaction is found only in complex cells. Binocularly non-phase-specific complex cells may have subunits whose optimal relative phases are random or monocular. The division of complex cells into these two major groups (binocularly phase specific and non-phase specific) is independent of whether they are standard or special complex-cell types. A small proportion (8%) of complex cells that appear monocular by alternate tests of each eye show a purely inhibitory influence from the silent eye. This inhibition is not generally dependent on the relative phase of the gratings. Unlike simple cells, complex cells are not a homogeneous group. However, nearly half of complex cells show phase-specific binocular interaction that is probably the result of linear convergence. Combined with the results from simple cells, the majority of binocular interaction in the striate cortex may be accounted for by linear summation of neural signals from each eye. This provides a simplified view of the nature of binocular interaction in the visual cortex.

Animals↗
PubMed1986

Orientation discrimination in amblyopia.

Using extended sinusoidal gratings to avoid potential problems of eccentric fixation, the authors have studied orientation discrimination in amblyopia. For all subjects, elevated orientation discrimination thresholds at high spatial frequencies were found. However, raised thresholds decrease with decreasing spatial frequency, and can be normal at low frequencies. Orientation discrimination thresholds for both amblyopic and non-amblyopic eyes are independent of contrast over most of the visible range. Therefore, amblyopic orientation discrimination thresholds cannot be mimicked in non-amblyopic eyes by reducing contrast. Control experiments show that the orientation discrimination deficits are not restricted to vertical stimuli and that they are not a result of exaggerated cyclotorsional eye movements.

Amblyopia↗
PubMed1985

Temporal sensitivity in amblyopia: an explanation of conflicting reports.

Although the spatial characteristics of contrast sensitivity in amblyopia are well established, it is still not clear how temporal vision is affected. A wide range of spatial stimuli have been used to measure flicker sensitivity in amblyopia and we have considered the possibility that reported inconsistancies in results are due to a confounding of spatial and temporal stimulus parameters. To address this question we have measured contrast sensitivity from a sample of eight strabismic/anisometropic amblyopes using (1) stationary sinusoidal gratings, (2) uniform field flicker and (3) counterphased flickering gratings. In each case the contrast sensitivity deficits were predictably determined by the spatial properties of the stimulus (spot size or spatial frequency), and the spatial characteristics of the amblyopia. Results are nearly independent of the temporal properties of the stimulus. Specifically, when the stimulus is sufficiently large, or consists of low spatial frequencies, no temporal deficits are found. We conclude that conflicts between previous reports are attributable primarily to differences in spatial characteristics of the stimuli used.

Amblyopia↗
PubMed1985

Is amblyopia spatial frequency or retinal locus specific?

The visual deficit associated with amblyopia is thought to be both spatial frequency and retinal locus dependent. However, most data that have been obtained can be accounted for by either of these factors or by a combination of both. We have tried to distinguish between these three possibilities by measuring contrast sensitivity at different retinal loci using discrete localized patches of grating. For five of nine amblyopes, we find the contrast sensitivity deficit to be constant across the retina for a given spatial frequency. In only two cases were there substantial changes as a function of retinal eccentricity. Therefore, most of our data suggest that the visual deficit in amblyopia is primarily spatial frequency and not retinal locus specific.

Amblyopia↗