Progressive changes in kitten striate cortex during monocular vision.

Following initial rearing in either total darkness or normal illumination, kittens at different ages were subjected to right-eye closure and various periods of vision through the left eye. After the period of monocular vision, single units in striate cortex were tested for visual responsiveness through each eye. A severe reduction in the proportion of units responsive to the deprived eye occurred over the first few days of monocular vision. Functional abnormalities were variably present after 1 day, marked after 2.5 and 3.5 days, and complete after 10 days. Monocular vision produced very much the same effect on ocular dominance of striate units, provided age and duration of suture were identical, regardless of whether kittens had received prior dark- or light-rearing.     

Cumulative effect of brief daily periods of monocular vision on kitten striate cortex

Summary Eight kittens were subjected to daily 4-h sessions with vision restricted to one eye between the 4th and 9th postnatal weeks. The total duration of monocular exposure ranged from 76 to 152 h. Between monocular exposure periods, four kittens were kept in total darkness, and four experienced binocular vision under normal colony conditions. At the end of the rearing period, all experimental animals and four normally reared controls were studied by means of single-unit recording in primary visual cortex. In kittens with inter-session dark rearing, very few neurons could be driven through the deprived eye. This effect was more extreme than that observed after continuous dark rearing. Visual responses mediated by the experienced eye appeared normal. In contrast, in kittens with inter-session binocular vision, the large majority of visual cortical neurons responded to visual stimulation of either eye. A few neurons appeared to have lost their responsiveness to the deprived eye, but this effect was small. We conclude that monocular vision delivered during brief daily sessions produces a cumulative competitive inactivation of transmission in the pathway from the deprived eye to striate cortex so long as no visual stimulation occurs outside the monocular periods. Binocular visual stimulation received between sessions rapidly and almost completely reverses the effects of monocular vision.     

Vestibulo-ocular reflex and optokinetic nystagmus in adult cats reared in stroboscopic illumination

Summary Cats reared in stroboscopic illumination (strobe reared cats) have been found to have abnormal eye movements. Visual and vestibular evoked compensatory eye movements were inefficient. Vestibuloocular reflex in the dark had a maximum gain of 0.6 (1.0 in normal animals). Optokinetic nystagmus had a mean gain which approached unity only at stimulus velocities around 7 °/s (up to 30 °/s in normal animals). The asymmetry of the Optokinetic nystagmus resulting from monocular stimulation was more pronounced in strobe reared cat than in normal animals. Interaction between vestibulo-ocular reflex and Optokinetic nystagmus to give adequate compensatory eye movements was absent in strobe reared cats: visual suppression of vestibulo-ocular reflex was absent when the animal was rotated in an illuminated environment which remained stationary with respect to the head. Optokinetic nystagmus failed to improve the gain of the vestibulo-ocular reflex when the animal was rotated in a normally lit environment. The deprived animals showed no signs of recovery after 5 months exposure to normal lighting.     

Reversal of the physiological effects of monocular deprivation in the kitten's visual cortex.

1. Twenty-three kittens were monocularly deprived of vision until the age of 4, 5, 6 or 7 weeks. Their deprived eyes were then opened, and their experienced eyes shut for a further 3-63 days. After this time physiological recordings were made in the visual cortex, area 17. Three control kittens, monocularly deprived for various periods, showed that at the time of reverse-suturing, few neurones could be influenced at all from the deprived eye. 2. Following reverse-suturing, the initially deprived eye regained control of cortical neurones. This switch of cortical ocular dominance was most rapid following reverse-suturing at the age of 4 weeks. Delaying the age of reverse-suturing reduced the rate and then the extent of the cortical ocular dominance changes. 3. The cortex of reverse-sutured kittens is divided into regions of cells dominated by one eye or the other. The relative sizes of these ocular dominance columns changed during reversed deprivation. The columns devoted to the initially deprived eye were very small in animals reverse-sutured for brief periods, but in animals that underwent longer periods of reversed deprivation, the columns driven by that eye were larger, while those devoted to the initially open eye were smaller. 4. Clear progressions of orientation columns across the cortex were apparent in many of the kittens, but, in contrast to the situation in normal or strabismic kittens, these sequences were disrupted at the borders of eye dominance columns: the cortical representations of orientation and ocular dominance were not independent. 5. Binocular units in these kittens were rather rare, but those that could be found often had dissimilar receptive field properties in the two eyes. Commonly, a cell would have a normal orientation selective receptive field in one eye, and an immature, unselective receptive field in the other. Cells that had orientation selective receptive fields in both eyes often had greatly differing orientation preferences in the two eyes, occasionally by nearly 90 degrees. 6. During the reversal of deprivation effects, the proportion of receptive fields exhibiting mature properties declined in the initially experienced eye, while the proportion increased in the initially deprived eye. Similarly, the average band width of orientation tuning of receptive fields in the initially deprived eye decreased, while that of receptive fields in the initially experienced eye increased. 7. One kitten was reverse-sutured twice, to demonstrate that cortical ocular dominance may be reversed a second time, even after one reversal of ocular dominance. 8. It is suggested that the sensitive period for cortical binocular development consists of two phases. In the first phase, all cortical neurones may be modified by experience, but the rate at which they may be modified decreases with age. In the second phase, an increasing number of cortical neurones becomes fixed in their properties, while those that remain modifiable are as modifiable as they were at the end of the first phase. 9...     

Blockade of serotonin-2C receptors by mesulergine reduces ocular dominance plasticity in kitten visual cortex

 We have investigated the role of serotonin-2C (5-HT_2C) receptors in modulation of ocular dominance plasticity in kitten visual cortex. A small quantity of the 5-HT_2C receptor blocker, mesulergine, was infused into the visual cortex of one hemisphere of 5- to 7-week-old kittens using osmotic minipumps, while the control hemisphere received vehicle solution. At the same time, one eyelid of the experimental animals was sutured shut. The ocular dominance distributions in the visual cortex (area 17) were assessed using extracellular recording methods after 1 week of combined mesulergine infusion and monocular deprivation. We found that the majority of the neurons remained binocularly responsive in the mesulergine-treated hemisphere, while most of the neurons recorded were either unresponsive or only weakly responsive to the deprived eye in the control hemisphere. Local infusion of mesulergine into the kitten visual cortex thus reduced the shift of ocular dominance that normally occurs in animals of these ages following monocular deprivation. The blocking effect seems to be distance-dependent and therefore dose-dependent: the farther away the recording sites were from the injection site, the fewer binocularly responsive cells were found. These results are relevant to previous findings indicating transient overexpression of 5-HT_2C receptor in visual cortex of kittens at these ages. The data suggest that the 5-HT_2C receptor system may be involved in the formation and modification of ocular dominance columns in the developing visual cortex. Received: 11 December 1995 / Accepted: 9 June 1996     

Stimulus dependence of ocular dominance of complex cells in area 17 of the feline visual cortex

Summary Stimulus dependence of ocular dominance of 31 deep-layer complex cells was assessed from detailed monocular directional tuning curves for motion of bar stimuli or fields of static visual noise, in area 17 of normal adult cats, lightly anaesthetised with N₂O/O₂ supplemented with pentobarbitone. Virtually all cells were binocularly driven, with the anticipated ocular dominance distribution. Interocular differences in directional bias and sharpness of directional tuning for noise were observed in eleven cells, whereas preferred direction and sharpness of tuning for bar stimuli were similar for each eye. In the majority of cells (20/31), any differences between noise and bar tuning in one eye were replicated in the other. Ocular dominance of about half the cells (17/31) for noise and for bar motion was similar, or marginally shifted by up to one ocular dominance group. Substantial shifts in ocular dominance were seen in 14 cells — by up to two ocular dominance groups in 12 cells and by up to three ocular dominance groups in two cells. In three cases these shifts involved a reversal of eye dominance. Notwithstanding these changes, there were no obvious trends in shifts of ocular dominance in favour of the ipsilateral or contralateral eye, nor was there any tendency towards increased binocularity for noise.     

Effects of early monocular deprivation on response properties and afferents of nucleus of the optic tract in the ferret

Summary Effects of early monocular deprivation on visual response properties of neurons in the nucleus of the optic tract (NOT) were studied in six adult ferrets. Retinal input to NOT was investigated by orthodromic electrical stimulation of optic chiasm and optic nerves. Electrical stimulation of the ipsilateral primary visual cortex was applied to reveal the presence of a cortical pathway to NOT. All 75 neurons studied in the NOT displayed the typical strongly direction-specific response to horizontal stimulus motion; they were activated by ipsiversively directed motion (i.e. motion towards the recorded hemisphere) similar to NOT-cells in animals with normal visual experience. When tested binocularly most of the NOT-cells preferred velocities of 10 or 20 deg/s, revealing no significant difference from animals reared with normal binocular experience. The most pronounced effect of monocular deprivation was observed on ocular dominance: In the hemisphere contralateral to the non-deprived eye, NOT-cells were almost exclusively driven through the contralateral eye. In the hemisphere contralateral to the deprived eye, three of the six animals studied showed a marked dominance of the ipsilateral, non-deprived eye. In the other three animals, most neurons were binocularly activated, but over all they were significantly more strongly activated by the ipsilateral eye than found in normal animals. In four animals, dependence of ocular dominance on stimulus velocity was tested in the NOT contralateral to the deprived eye. In one of them, neurons were almost exclusively driven by the ipsilateral, non-deprived eye, irrespective of stimulus velocity. In the other animals, ocular dominance shifted from contralateral to ipsilateral with increasing stimulus velocities. Electrical stimulation of the optic chiasm revealed a mean latency of 5.53 ± 0.48 ms. In both hemispheres, NOT-units could only be activated by stimulation of the contralateral optic nerve. Thus, no significant difference in the retinofugal conduction velocities from the deprived and the normal nerve could be detected. Of 52 cells studied, 28 (= 54%) could be activated by stimulation of primary visual cortex, mean latency being 3.9± 1.7 ms. No significant difference in the percentage of cortically excitable cells between the two hemispheres as well as compared to normal animals was found (contralateral to the deprived eye: 67%, contralateral to the non-deprived eye: 53%). Therefore, cortical projections to NOT seem not to be affected by monocular deprivation. The effects of monocular deprivation in the ferret NOT, especially on ocular dominance and cortical input, are compared to the results previously described for the cat.     

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

Summary 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.     

Substantial reduction of noradrenaline in kitten visual cortex by intraventricular injections of 6-hydroxydopamine does not always prevent ocular dominance shifts after monocular deprivation

Summary Ten kittens had cannulas inserted into their lateral ventricles for daily injections of 6-hydroxydopamine (6-OHDA). At 5–6 weeks of age one eye was sutured shut, and one week later recordings were made from the visual cortex to assay the ocular dominance of a sample of cells. In six kittens the injections of 6-OHDA were continued until the day before recording, while in four kittens the injections were stopped around the time of eye suture, on the assumption that continued injections of 6-OHDA over several days has effects that are not specific to the noradrenaline (NA) system and that the two procedures might show different results. In all animals the concentration of NA in the visual cortex near the site of recording was reduced by approximately 90%. In all animals the ocular dominance histograms recorded from the visual cortex were shifted so that the majority of cells (83 ± 13%) were dominated by the open eye. There were no substantial differences between the two groups of experimental animals or between the experimental animals and two control animals that had cannulas implanted and ascorbate alone injected without 6-OHDA. We conclude that the concentration of NA in the visual cortex can be reduced substantially by injections of 6-OHDA into the lateral ventricle without preventing the shift in ocular dominance that usually occurs after suturing shut the eyelids of one eye.     

Chronic recordings from single sites of kitten striate cortex during experience-dependent modifications of receptive-field properties

1. With the use of chronically implanted floating microelectrodes, we obtained simultaneous single-unit recordings from multiple sites in the kitten striate cortex and followed experience-dependent modifications of receptive-field properties. For induction of experimental modifications, we used the paradigm of monocular deprivation and reverse occlusion. Kittens were implanted when 4-5 wk old. During the following 2 days, receptive-field properties of the recorded units were determined once under light ketamine anesthesia and repeatedly while the kittens were awake and only lightly restrained. Subsequently, one eye was patched, and the resulting changes in neuronal eye preference were followed by repeated measurements of response properties. For investigation of the effects of reverse occlusion, the deprived eye was opened and the previously open eye closed when the neurons had become unresponsive to the initially deprived eye. Alternatively, kittens were monocularly deprived for 1 wk by lid suture before implantation. The closed eye was then opened, the other eye patched, and the effects of reverse occlusion were studied for up to 1 wk by repeated measurements of receptive-field properties. 2. The earliest effect of monocular deprivation was the disappearance of binocular summation, i.e., binocular responses ceased to be superior over monocular responses. Overt changes of ocular dominance were observed as early as 6 h after the beginning of monocular deprivation. These consisted of a gradual decrease of the excitatory response to deprived eye stimulation and, on occasions, of an additional moderate increase of responses to the normal eye. A complete loss of excitatory responses to deprived-eye stimulation was seen as early as 12 h after occlusion. In numerous cells, however, stimulation of the deprived eye continued to evoke inhibitory responses even after excitatory responses had vanished completely. During this shift in ocularity, neurons preserved their orientation and direction selectively. 3. The minimal time required for the manifestation of ocular dominance changes was similar regardless of whether the animals were stimulated continuously or were asleep part of the time, suggesting the existence of an experience-independent consolidation period for ocular dominance changes. 4. The first change after reverse occlusion was a reduction of the response to the newly deprived eye. The time course of this inactivation was similar to that observed after initial deprivation, whereas the recovery of responses to the previously deprived eye had a considerably slower time course.(ABSTRACT TRUNCATED AT 400 WORDS)     

Functional magnetic resonance imaging evidence for binocular interactions in human visual cortex

Using functional magnetic resonance imaging (fMRI), we explored the binocular interactions occurring when subjects viewed dichoptically presented checkerboard stimuli. A flickering radial checkerboard was presented to each eye of the subject, while T2*-weighted images were acquired over the visual cortex with gradient-echo, echoplanar sequences. We compared responses in striate and extrastriate visual cortex under four conditions: both eyes were stimulated at the same time (binocular condition), each eye was stimulated in alternation (monocular condition) or first the one eye then the other eye was stimulated (left eye first - right eye trailing, or vice versa). The results indicate that only the striate area, in and near the calcarine fissure, shows significant differences for these stimulation conditions. These differences are not evident in more remote extrastriate or associational visual areas, although the BOLD response in the stimulation-rest comparison was robust. These results suggest that the effect could be related to inhibitory interactions across ocular dominance columns in striate visual cortex.     

Reversal of the physiological effects of monocular deprivation in kittens: further evidence for a sensitive period

1. It was confirmed that suturing the lids of one eye (monocular deprivation), until only 5 weeks of age, leaves virtually every neurone in the kitten's visual cortex entirely dominated by the other eye. On the other hand, deprivation of both eyes causes no change in the normal ocular dominance of cortical neurones, most cells being clearly binocularly driven.2. Kittens were monocularly deprived until various ages, from 5 to 14 weeks, at which time reverse suturing was performed: the initially deprived right eye was opened and the left eye closed for a further 9 weeks before recording from the visual cortex.3. Reverse suturing at 5 weeks caused a complete switch in ocular dominance: every cell was dominated by the initially deprived right eye. Reverse suturing at 14 weeks, however, had almost no further effect on ocular dominance: most cells were still driven solely by the left eye. Animals reverse sutured at intermediate ages had cortical neurones strongly dominated by one eye or the other, and they were organized into clear columnar groups according to ocular dominance.4. Thus, between 5 weeks and 4 months of age, there is a period of declining sensitivity to both the effects of an initial period of monocular deprivation and the reversal of those effects by reverse suturing.5. The small proportion of binocular cells in reverse sutured kittens (which have never had simultaneous binocular vision) often differed considerably in their receptive field properties in the two eyes. In particular, if the cells were orientation selective in both eyes the two preferred orientations could differ by up to 70 degrees .6. The relative importance of innate and environmental contributions to the properties of cortical cells is discussed.     

Influence of experience on orientation maps in cat visual cortex.

Experience is known to affect the development of ocular dominance maps in visual cortex, but it has remained controversial whether orientation preference maps are similarly affected by limiting visual experience to a single orientation early in life. Here we used optical imaging based on intrinsic signals to show that the visual cortex of kittens reared in a striped environment responded to all orientations, but devoted up to twice as much surface area to the experienced orientation as the orthogonal one. This effect is due to an instructive role of visual experience whereby some neurons shift their orientation preferences toward the experienced orientation. Thus, although cortical orientation maps are remarkably rigid in the sense that orientations that have never been seen by the animal occupy a large portion of the cortical territory, visual experience can nevertheless alter neuronal responses to oriented contours.     

Ocular dominance plasticity restored by NA infusion to aplastic visual cortex of anesthetized and paralyzed kittens

Summary We studied the ocular dominance distribution in visual cortex of kittens which had been monocularly exposed to moving-pattern stimuli under anesthesia and paralysis. 1. We did not obtain any discernible changes in ocular dominance, confirming the previous reports that anesthesia and paralysis prevent ocular dominance plasticity from occurring. 2. The plasticity, however, was restored under the acute experimental condition by a cortical infusion of 1-noradrenaline (1-NA). In the 1-NA-infused visual cortex, the ocular dominance distribution was clearly shifted to the open eye after monocular exposure for about 20–24 h. 3. We also studied how quickly and to what extent the changes were induced when the duration of the combined treatment was varied. The results were: (i) the earliest change was observed in 12 h with disappearance of binocular cells, (ii) the treatment was most effective after 20–24 h in inducing the shift of ocular dominance, and (iii) the treatment longer than 24 h (up to 45 h) did not necessarily enhance the shift, though the state of reduced binocularity was sustained throughout. 4. The effects of the cortical 1-NA infusion combined with monocular exposure became less with increasing the age of experimental animals, suggesting the presence of the susceptible period in the acute experiments. 5. The effects seemed to become smaller toward the end of a given recording session, suggesting that the restored plasticity wanes with time. The present results further support the idea that the direct activation of the NA system enhances cortical plasticity, in principle, independent of general conditions of experimental animals.     

Central core control of developmental plasticity in the kitten visual cortex: II. Electrical activation of mesencephalic and diencephalic projections

Summary Fifteen dark-reared, 4- to 5-week-old kittens were stimulated monocularly with patterned light while they were anesthetized and paralyzed. Six of these kittens were exposed to the light stimuli only, in four kittens the light stimuli were paired with electric stimulation of the mesencephalic reticular formation and in five kittens with electric activation of the medial thalamic nuclei. Throughout the conditioning period, the ocular dominance of neurons in the visual cortex was determined from evoked potentials that were elicited either with electric stimulation of the optic nerves or with phase reversing gratings of variable spatial frequencies. In two kittens, ocular dominance changes were assessed after the end of the conditioning period by analyzing single unit receptive fields. Monocular stimulation with patterned light induced a marked shift of ocular dominance toward the stimulated eye, when the light stimulus was paired with electric activation of either the mesencephalic reticular formation or of the medial thalamus. Moreover, a substantial fraction of cells acquired mature receptive fields. No such changes occurred with light or electric stimulation alone. It is concluded that central core projections which modulate cortical excitability gate experience-dependent modifications of connections in the kitten visual cortex.Part of this work was supported by a grant from the Deutsche Forschungsgemeinschaft SFB50, A14     

Development of the kitten visual cortex depends on the relationship between the plane of eye movements and visual inputs

Summary 1. Previous experiments have demonstrated that eye movements, acting through the extraocular muscle (EOM) proprioceptive afferents, are necessary for the development of orientation selectivity in the cells of the kitten visual cortex. New experiments were carried out to study the effect of the plane of eye movements on the preferred orientation acquired by the visual cortical cells. 2. Darkreared (DR) kittens were operated on at 5–6 weeks of age. In the first series of experiments, 4 out of the 6 EOMs were removed bilaterally in such a way that both eyes could only move in a single plane, either vertical or horizontal. In the second series of experiments, the same operation was performed on one eye which was also sutured shut and, on the other side, the EOM were deafferented by intracranial section of the ophthalmic branch of Vth nerve and the eye left open. 3. 1–4 days after surgery the kittens were given 6 h of visual experience and 12 h later were prepared for visual cell recording in Area 17. 4. In kittens of the first series: orientation selectivity developed in the majority (60–65%) of visual cells, most of which encoded horizontal orientations when the eyes had moved in the vertical plane and vertical orientations when the eyes had moved in the horizontal plane. These results show that the plane of eye movements during early visual experience influences the distribution of preferred orientations with an orthogonal relation. Ocular dominance histograms were strabismic like. 5. In kittens of the second series: orientation selectivity developed in 40–50% of cells, about half of which were tuned for the orientation orthogonal to the direction of movement of the occluded eye, as in experiment I. The seeing, deafferented eye, presumably would have sent normal visual inputs centrally, corresponding to displacements on the retina in every direction since the ocular motility of that eye had not been disturbed. However, proprioceptive information about its movements was suppressed. As only some of the EOMs of the occluded eye were still present and connected, the conclusion is that the observed influence of the plane of eye movements acts through the proprioceptive afferents. The ocular dominance histograms showed: 1) a powerful change in favour of the seeing eye after only 6 h of monocular visual experience; 2) a larger effect of monocular visual experience in the hemisphere contralateral to the seeing eye; 3) a linkage between acquisition of orientation selectivity and shift in ocular dominance. 6. Our results suggest that normal development of orientation selectivity in visual cortical cells results from the close association of visual and EOM afferent inputs. It is suggested that these two signals must occur with a precise temporal relationship.     

Ocular dominance predicts neither strength nor class of disparity selectivity with random-dot stimuli in primate V1

We address two unresolved issues concerning the coding of binocular disparity in primary visual cortex. Experimental studies and theoretical models have suggested a relationship between a cell's ocular dominance, assessed with monocular stimuli, and its tuning to binocular disparity. First, the disparity energy model of disparity selectivity suggests that there should be a correlation between ocular dominance and the strength of disparity tuning. Second, several studies have reported a relationship between ocular dominance and the shape of the disparity tuning curve, with cells dominated by one eye more likely to have disparity tuning of the tuned-inhibitory type. We investigated both of these relationships in single neurons recorded from the primary visual cortex of awake fixating macaques, using dynamic random-dot patterns as a stimulus. To classify disparity tuning curves quantitatively, we develop a new measure of symmetry, which can be applied to any function. We find no evidence for any correlation between ocular dominance and the nature of disparity tuning. This places constraints on the circuitry underlying disparity tuning.     

Role of visual experience in activating critical period in cat visual cortex

Cats were reared in total darkness from birth until 4-5 mo of age (DR cats, n = 7) or with very brief visual experience (1 or 2 days) during an otherwise similar period of dark rearing [DR(1) cats, n = 3; DR(2) cats, n = 7]. Single-cell recordings were made in area 17 of visual cortex at the end of this rearing period and/or after a subsequent prolonged period of monocular deprivation. Control observations were made in normal cats (n = 3), cats reared with monocular deprivation from birth (n = 4), and cats monocularly deprived after being reared normally until 4 mo of age (n = 2). After rearing cats in total darkness, the majority of visual cortical cells were binocularly driven and the overall distribution of ocular dominance was not different from that of normal cats. Orientation-selective cells were very rare in dark-reared cats. Monocular deprivation imposed after dark rearing resulted in selective development of connections from the open eye. Most cells were responsive only to the open eye and the majority of these were orientation selective. These results were similar to, though less severe than, those found in cats reared with monocular deprivation from birth. Monocular deprivation imposed after 4 mo of normal rearing did not produce selective development of connections from the open eye in terms of either ocular dominance or orientation selectivity. In DR(1) cats visual cortical physiology was degraded in comparison to dark-reared cats after the rearing period. Most cells were binocularly driven but there was a higher frequency of unresponsive cells and a reduced frequency of orientation-selective cells. Subsequent monocular deprivation resulted in a further decrease in the number of binocularly driven cells and an increase in unresponsive cells. However, it did not produce a bias in favor of the open eye in terms of either ocular dominance or orientation selectivity. In DR(2) cats there was a high incidence of unresponsive cells and a marked loss of binocularly driven cells after the rearing period. Subsequent monocular deprivation failed to produce any significant changes.(ABSTRACT TRUNCATED AT 400 WORDS)     

Profile of the sensitive period for monocular deprivation in kittens

Summary Thirteen kittens were subjected to 10–12 days of unilateral eye closure at ages spaced regularly through the first 4 months after birth. At the end of each kitten's period of monocular vision, the degree of functional disconnection between the deprived eye and neurons in striate cortex was assessed by means of single-unit recording. When the proportion of cortical cells giving no response to stimulation through the deprived eye was analyzed as a function of the kitten's age at the onset of eye closure, it was found that the effectiveness of monocular deprivation rose prior to postnatal day 28, remained high through day 48 and then subsided gradually, probably persisting at least through the end of the fourth postnatal month. The degree of functional modifiability persisting in the visual cortex of older kittens may be related to the initial ocular dominance of each neuron. Cells responsive exclusively to the deprived eye prior to deprivation probably do not acquire functional input from the nondeprived eye in kittens older than 48 days, for a normal proportion of such cells is encountered after the period of eye closure. Conversely, cells dominated by the nondeprived eye probably are most likely to lose their input from the deprived eye, as indicated by the columnar organization of cells not responsive to the deprived eye.This work was supported by Grant EY01175 and Research Career Development Award EY00092 from the National Eye Institute to R.D.F. A National Institutes of Health Training Grant, GM0748-03, helped to support C.R.O.     

Monocular cells without ocular dominance columns

In many regions of the mammalian cerebral cortex, cells that share a common receptive field property are grouped into columns. Despite intensive study, the function of the cortical column remains unknown. In the squirrel monkey, the expression of ocular dominance columns is variable, with columns present in some animals and not in others. By searching for differences between animals with and without columns, it should be possible to infer how columns contribute to visual processing. Single-cell recordings outside layer 4C were made in nine squirrel monkeys, followed by labeling of ocular dominance columns in layer 4C. In the squirrel monkey, compared with the macaque, cells outside layer 4C were more likely to respond to stimulation of either eye whether ocular dominance columns were present or not. In three animals lacking ocular dominance columns, single cells were recorded from layer 4C. Remarkably, 20% of cells in layer 4C were monocular despite the absence of columns. This observation means that ocular dominance columns are not necessary for monocular cells to occur in striate cortex. In macaques each row of cytochrome oxidase (CO) patches is aligned with an ocular dominance column and receives koniocellular input serving one eye only. In squirrel monkeys this was not true: CO patches and ocular dominance columns had no spatial correlation and the koniocellular input to CO patches was binocular. Thus even when ocular dominance columns occur in the squirrel monkey, they do not transform the functional architecture to resemble that of the macaque.