Abolition of optokinetic nystagmus in the cat.

Combining a behavioral and a surgical manipulation, namely complete visual deprivation with surgical section of the optic chiasm, results in the abolition of optokinetic nystagmus in the cat. This basic optomotor reflex remains relatively unaffected by either of these manipulations performed singly.

Receptive field properties and sensitivity to edges defined by motion in the postero-lateral lateral suprasylvian (PLLS) area of the cat.

The present study investigated the spatial properties of cells in the postero-lateral lateral suprasylvian (PLLS) area of the cat and assessed their sensitivity to edges defined by motion. A total of one hundred and seventeen (117) single units were isolated. First, drifting sinusoidal gratings were used to assess the spatial properties of the cells' receptive fields and to determine their spatial frequency tuning functions. Second, random-dot kinematograms were used to create illusory edges by drifting textured stimuli (i.e. a horizontal bar) against a similarly textured but static background. Almost all the cells recorded in PLLS (96.0%) were binocular, and a substantial majority of receptive fields (79.2%) were end-stopped. Most units (81.0%) had band-pass spatial frequency tuning functions and responded optimally to low spatial frequencies (mean spatial frequency: 0.08 c./degree). The remaining units (19.0%) were low-pass. All the recorded cells responded vigorously to edges defined by motion. The vast majority (96.0%) of cells responded optimally to large texture elements; approximately half the cells (57.3%) also responded to finer texture elements. Moreover, 38.5% of the cells were selective to the width of the bar (i.e., the distance between the leading and the trailing edges). Finally, some (9.0%) cells responded in a transient fashion to leading and to trailing edges. In conclusion, cells in the PLLS area are low spatial frequency analyzers that are sensitive to texture and to the distance between edges defined by motion.

Auditory responses in the visual cortex of neonatally enucleated rats.

A number of studies on humans and animals have demonstrated better auditory abilities in blind with respect to sighted subjects and have tried to define the mechanisms through which this compensation occurs. The aim of the present study, therefore, was to examine the participation of primary visual cortex (V1) to auditory processing in early enucleated rats. Here we show, using gaussian noise bursts, that about a third of the cells in V1 responded to auditory stimulation in blind rats and most of these (78%) had ON-type responses and low spontaneous activity. Moreover, they were distributed throughout visual cortex without any apparent tonotopic organization. Optimal frequencies determined using pure tones were rather high but comparable to those found in auditory cortex of blind and sighted rats. On the other hand, sensory thresholds determined at these frequencies were higher and bandwidths were wider in V1 of the blind animals. Blind and sighted rats were also stimulated for 60 min with gaussian noise, their brains removed and processed for c-Fos immunohistochemistry. Results revealed that c-Fos positive cells were not only present in auditory cortex of both groups of rats but there was a 10-fold increase in labeled cells in V1 and a fivefold increase in secondary visual cortex (V2) of early enucleated rats in comparisons to sighted ones. Also, the pattern of distribution of these labeled cells across layers suggests that the recruitment of V1 could originate at least in part through inputs arising from the thalamus. The ensemble of results appears to indicate that cross-modal compensation leading to improved performance in the blind depends on cell recruitment in V1 but probably also plastic changes in lower- and higher-order visual structures and possibly in the auditory system.

Age-related audiovisual interactions in the superior colliculus of the rat.

It is well established that multisensory integration is a functional characteristic of the superior colliculus that disambiguates external stimuli and therefore reduces the reaction times toward simple audiovisual targets in space. However, in a condition where a complex audiovisual stimulus is used, such as the optical flow in the presence of modulated audio signals, little is known about the processing of the multisensory integration in the superior colliculus. Furthermore, since visual and auditory deficits constitute hallmark signs during aging, we sought to gain some insight on whether audiovisual processes in the superior colliculus are altered with age. Extracellular single-unit recordings were conducted in the superior colliculus of anesthetized Sprague-Dawley adult (10-12 months) and aged (21-22 months) rats. Looming circular concentric sinusoidal (CCS) gratings were presented alone and in the presence of sinusoidally amplitude modulated white noise. In both groups of rats, two different audiovisual response interactions were encountered in the spatial domain: superadditive, and suppressive. In contrast, additive audiovisual interactions were found only in adult rats. Hence, superior colliculus audiovisual interactions were more numerous in adult rats (38%) than in aged rats (8%). These results suggest that intersensory interactions in the superior colliculus play an essential role in space processing toward audiovisual moving objects during self-motion. Moreover, aging has a deleterious effect on complex audiovisual interactions.

Greater losses in sensitivity to second-order local motion than to first-order local motion after early visual deprivation in humans.

We compared sensitivity to first-order versus second-order local motion in patients treated for dense central congenital cataracts in one or both eyes. Amplitude modulation thresholds were measured for discriminating the direction of motion of luminance-modulated (first-order) and contrast modulated (second-order) horizontal sine-wave gratings. Early visual deprivation, whether monocular or binocular, caused losses in sensitivity to both first- and second-order motion, with greater losses for second-order motion than for first-order motion. These findings are consistent with the hypothesis that the two types of motion are processed by different mechanisms and suggest that those mechanisms are differentially sensitive to early visual input.

Binocular Interactions in the Lateral Suprasylvian Visual Area of Strabismic Cats Following Section of the Corpus Callosum.

Visually responsive neurons have been recorded in the lateral suprasylvian area (LSA) of cats raised with either a convergent or a divergent strabismus. In contrast to areas 17 and 18, where many studies have documented a profound loss of binocularly activated neurons following early strabismus, in the LSA the majority of cells could still be binocularly driven. Acute or chronic section of the splenium of the corpus callosum reduced but did not abolish binocularity in the LSA. We propose that the widespread callosal connections, the large size of the receptive fields and the peculiar internal circuitry of the LSA all concur in permitting the maintenance of binocular coding in spite of early misalignment of the eyes.

Somatosensory receptive field properties of corpus callosum fibres in the raccoon.

Anatomical studies in a number of species have shown that most areas of the somatosensory cortex are callosally interconnected. This is also true for the raccoon, at least for those parts representing proximal and axial body regions. Electrophysiologically, studies carried out in cats and monkeys have demonstrated that all sensory sub-modalities cross in the callosum. Moreover, cells representing the paws and fingers, though occupying a large portion of areas SI and SII, seem to send proportionately fewer axons through the callosum than axial structures. No comparable study has been carried out in the raccoon. The purpose of the present experiment was therefore to investigate the functional organization of the callosal system in this animal by examining the receptive field properties of the somatosensory fibres crossing in the callosum. Axonal activity was recorded directly through tungsten microelectrodes in the corpus callosum of eight raccoons. Results indicated that somatosensory information is transmitted in its rostral portion. Most receptive fields concerned axial and proximal body regions and the head and face. Some receptive fields represented para-axial regions of the body and a few concerned the hands and fingers. Slowly and rapidly adapting fibres were found, as were all the sensory sub-modalities tested. A substantial proportion of the axons had bilateral receptive fields. These results are discussed in relation to those obtained in other species, with particular reference to: (1) the midline fusion hypothesis of callosal function; (2) the representation within this structure of the distal extremities, and (3) the origin of the bilateral receptive fields.

Stereopsis in the cat: behavioral demonstration and underlying mechanisms.

The neural substrates subserving stereopsis were investigated behaviorally and electrophysiologically in the cat. In one set of studies, we examined behaviorally the ability of normal cats to perceive depth on the sole basis of spatial disparity using random-dot stereograms. Results showed that the animals were able to carry out this discrimination. We then evaluated the contribution of the optic chiasm, the corpus callosum and the primary visual cortex to this function. Results indicated that: (1) chiasma transection drastically reduced the ability of the animals to solve the random-dot problem; (2) a callosal split had little or no effect on their ability to relearn the same discrimination; (3) a section of both the corpus callosum and optic chiasm abolished this ability; and (4) bilateral lesions of areas 17-18 also abolished it. In another set of studies, we examined electrophysiologically the properties of neurons in the various visual cortical areas where disparity-based depth discrimination processes are presumed to take place. We recorded from areas 17, 18 and 19 of normal and split-chiasm cats. Results showed that: (1) the primary visual cortex of the normal cat contained cells sensitive to stimulus disparity; (2) these disparity sensitive neurons were also present in area 19 although in a much lower proportion and were more widely tuned than those in areas 17-18; and (3) following the section of the optic chiasm, there was a significant decrease in the number of disparity sensitive cells in areas 17-18, whereas in area 19 they were nearly completely absent. The results obtained from the lesion studies and from the single unit recording experiments indicate that stereoscopic depth perception is highly dependent in the cat upon the integrity of the through-the-chiasm geniculo-striate pathway and its target primary visual cortex.

Unilateral and bilateral temperature comparisons in acallosal and split-brain subjects.

Therapeutic section of the corpus callosum in adult epileptic patients typically results in their incapacity to carry out interhemispheric comparisons of lateralized information. The fact that acallosal and early split-brain subjects display few of these symptoms when tested in the tactile modality has led to the suggestion that these patients may use ipsilateral projections of the somatosensory system more effectively. Compensation, however, is limited by the fact that the lemniscal pathway is strongly lateralized, especially for the distal parts of the body, where few ipsilaterally projecting fibres have been demonstrated. The pathway carrying temperature information has a larger ipsilateral component. Bilateral comparisons within the same hemisphere in subjects who are lacking the corpus callosum should be more common and the development of compensatory mechanisms in early-sectioned or acallosal subjects should be more likely. The objective of the present experiment was to evaluate differential thresholds for thermal stimuli applied on a number of regions either on the same side or on corresponding sites on opposite sides of the body. One subject callosotomized as an adult and one split-brain subject who underwent callosotomy in childhood, as well as three acallosal subjects, were compared to IQ-matched and normal-IQ control subjects. The fingers, forearm and trunk were tested. The comparison temperature was 30 degrees C and the other was varied in an ascending or descending fashion using a modified method of limits. Differential thresholds were similar for within- and between-side comparisons, and comparable to those of the IQ-matched subjects. The results indicate that comparisons involving temperature discrimination for stimuli applied to the two sides of the body do not require the integrity of the corpus callosum.

Binocular fusion/suppression to spatial frequency differences at the border of areas 17/18 of the cat.

As shown by various human psychophysical studies, interocular spatial frequency disparities can yield a variety of percepts. In order to examine how binocular fusion is affected by spatial frequency differences, we have recorded cells in the border region of areas 17/18 of anesthetized cats. The optic axes of the eyes were deviated onto cathode-ray screens, and the optimal spatial frequency of each eye was assessed by monocular stimulations using drifting sinusoidal gratings. The optimal relative phase using identical spatial frequencies in both eyes was first determined. Spatial frequency differences were then introduced by keeping the optimal spatial frequency constant in one eye and varying the spatial frequency in the other. Results indicate that cells (39%) responded with an increased firing rate (facilitation) to similar spatial frequencies in each eye and with a gradual attenuation (occlusion or suppression) when spatial frequency differences were increased. However, binocular facilitation did not always occur to the presentation of identical stimuli. For 16% of the cells, maximal responses were observed when lower spatial frequencies than the optimal one were presented in one eye while higher spatial frequencies produced suppression. The opposite pattern was observed only for two cells. These findings are discussed in terms of binocular fusion and suppression.

Spatial properties and direction selectivity of single neurons in area 21b of the cat.

The receptive field properties of single units were assessed in area 21b of the cat visual cortex. Visual cells in this area were binocular and showed relatively large receptive fields. Most cells were strongly sensitive to the direction of drifting gratings. The mean value of the half-widths of the direction tuning curves (32 degrees ) suggests broader direction tunings than are typically found in other visual areas. The spatial frequency tuning functions were either band-pass or low-pass. Cells responded optimally to low spatial frequencies (mean =0.08c/deg) and also showed low spatial resolution (mean =0.29c/deg.). The estimated values of spatial bandwidths (mean=2.2 octaves) suggest that area 21b cells act as relatively good spatial filters. Although some cells exhibited a low contrast threshold, most cells began to respond at intermediate or high contrast values (mean threshold =15.5%). Temporal frequency tuning functions were mostly band-pass and usually broad (mean temporal bandwidth=3.3 octaves). Cells were found that responded optimally to various temporal frequencies (mean optimal temporal frequency=3.2Hz), although the majority preferred a temporal frequency below 4Hz.These results suggest that visual properties (receptive fields sizes, spatial resolution and orientation/direction selectivity) of cells in area 21b differ from those of cells previously observed in the adjoining area 21a. These differences provide evidence in support of functional distinction between these two visual areas.

Phase- and position-disparity coding in the posteromedial lateral suprasylvian area of the cat.

The posteromedial lateral suprasylvian area of the cat is known to be involved in the analysis of motion and motion in depth. However, it remains unclear whether binocular cells in the posteromedial lateral suprasylvian area rely upon phase or positional offsets between their receptive fields in order to code binocular disparity. The present study aims at clarifying more precisely the neural mechanisms underlying stereoperception with two objectives in mind. First, to determine whether cells in the posteromedial lateral suprasylvian area code phase disparities. Secondly, to examine whether the cells sensitive to phase disparity are the same as those which code for position disparities or whether each group represent a different sub-population of disparity-sensitive neurons. We investigated this by testing both types of disparities on single neurons in this area. The results show that the vast majority of cells (74%), in the posteromedial lateral suprasylvian area, are sensitive to relative interocular phase disparities. These cells showed mostly facilitation (95%) and a few (5%) summation interactions. Moreover, most cells (81%) were sensitive to both position and phase disparities. The results of this study show that most binocular cells in the posteromedial lateral suprasylvian area are sensitive to both positional and phase offsets which demonstrate the importance of this area in stereopsis.

Spatial and temporal matching of receptive field properties of binocular cells in area 19 of the cat.

The spatial and temporal properties of single neurons were investigated in area 19 of the cat. We evaluated the matching of binocular receptive field properties with regard to the respective strength of the ipsilateral and contralateral inputs. Results indicate that most cells in area 19 are well tuned to spatial and temporal frequencies and exhibit relatively low contrast threshold (mean=6.8%) when assessed using optimal parameters and tested through the dominant eye. Spatial resolution (mean=0.75 c/degree), optimal spatial frequencies (mean=0.16 c/degree) were relatively low and spatial bandwidths (mean=2.1 octaves) were broader as compared to those of cells in area 17 but comparable to those of cells in other extrastriate areas. On the other hand temporal resolution (mean=10.7 Hz), optimal temporal frequency (mean=4.5 Hz) and temporal bandwidths (mean=2.9 octaves) were higher and broader than in primary visual cortex. A significant relationship exists between most of the cell's properties assessed through either eye. For some parameters, such as spatial and temporal resolution, ocular dominance was shown to be significantly related to the extent of matching between the two eyes. For these parameters, binocular cells that exhibited a balanced ocular dominance were generally well matched with regard to the receptive field properties of each eye whereas the largest mismatches were found in cells that were more strongly dominated by one eye. These results suggest that visual input contributes to the activation of cells in area 19 in a redundant manner, possibly attesting to the multiplicity of parallel pathways to this area in the cat.

Interhemispheric transfer of visual pattern discriminations: evidence for a bilateral storage of the engram.

The present experiment was carried out to determine whether the memory trace of a pattern discrimination learned with one hemisphere is also transmitted to the second hemisphere via the corpus callosum or whether the trace is limited to the trained hemisphere and becomes accessible to the second during recall via this route. Split-chiasma cats learned two pattern discriminations with one eye (and hemisphere), then were subjected to a mid-sagittal transection of their corpus callosum, followed by learning with the other untrained eye (and hemisphere). Ten cats were separated into two groups: one group learned the discrimination to criterion (the non-overtrained group) while the other received 1600 overtraining trials over and beyond those needed to reach criterion (the overtrained group). Results indicated that there was little bilateral storage in the non-overtrained group (as determined by the number of trials needed to attain criterion with the second eye). Most subjects from the overtrained group showed chance performance during the first transfer session but learned the pattern discrimination much more rapidly with the second eye than with the first. These results are taken as indicating that memory transcription is possible through the callosum but that this route is slower and its readout is possibly contaminated by secondary non-specific factors which affect the initial utilization of the trace.

Receptive field properties of somatosensory callosal fibres in the monkey.

The corpus callosum is the principal neocortical commissure which transmits lateralized information between the hemispheres. The aim of the present experiment was to study the receptive field properties of somatosensory callosal fibres in rhesus macaque monkeys. The callosum was approached under direct visual control and axonic responses were recorded using tungsten microelectrodes. All sensory submodalities which could be examined with the available instruments were found (light touch, medium and deep pressure, joint movement and light pinches). Most fibres had receptive fields concerned with the trunk, followed by the head, with only a few responding to stimulation of the extremities. The medial borders of the unilateral receptive fields situated on the trunk and the head extended to the midline. The results are interpreted in terms of the roles of the corpus callosum in midline fusion and interhemispheric transfer.

Somatosensory receptive fields of fibres in the rostral corpus callosum of the cat.

The corpus callosum is the principal neocortical commissure which transmits lateralized information between the hemispheres. The aim of the present experiment was to study the receptive field (RF) properties of somatosensory callosal fibres in the cat. The callosum was approached under direct visual control and axonic responses were recorded under N2O anaesthesia using tungsten microelectrodes or, mostly, glass micropipettes. RFs representing all the sensory submodalities tested (light touch, medium and deep pressure, joint movement and light pinches) were found to be present in the axons which travelled through the callosum. Rapidly adapting units were more common than slowly adapting ones. The axial and para-axial portions of the body accounted for about three-fifths of all RFs, followed by the head (about one-fifth), with the rest responding to stimulation of the extremities. The medial borders of most of the unilateral RFs situated on the trunk and, to a lesser degree, the head, extended to the mid-line. The results are interpreted in terms of the roles of the corpus callosum in mid-line fusion and interhemispheric transfer.

Spatial and temporal frequency tuning and contrast sensitivity of single neurons in area 21a of the cat.

The spatial and temporal selectivities of single neurons in area 21a of the adult cat were investigated using sinusoidal gratings. Optimal spatial frequencies and visual acuity (high cut-off frequency) were fairly low and spatial bandwidth was mainly narrow. Contrast threshold was generally low but a substantial number of cells were only excited by high contrast stimuli. The temporal selectivity suggests that cells responded to a wide range of temporal frequencies.

Spatial disparity sensitivity in area PMLS of the Siamese cat.

Previous studies of the visual system of Siamese cats have shown that binocular cells are scarce in areas 17, 18 and 19, yet significantly more abundant in suprasylvian areas such as the postero-medial lateral suprasylvian area (PMLS). The present study aims at evaluating the sensitivity to spatial disparity of PMLS binocular cells in paralyzed and anesthetized Siamese cats. Centrally located receptive fields were mapped, separated using prisms and then stimulated simultaneously using two luminous bars optimally adjusted to the size of the excitatory receptive fields. Delays were introduced in the arrival of the luminous bars in the receptive fields so as to create the desired spatial disparities. Results indicate that approximately a third of PMLS units are binocular and that these binocular cells can detect spatial disparity cues. Indeed, although the sample was relatively small, cells of the tuned excitatory (14/34), tuned inhibitory (2/34), near (6/34) and far (1/34) types were identified. The spatial selectivity, as measured by the width at half height of the tuning curves of the excitatory and inhibitory cells and the slopes of the near and far cells, was similar to that obtained in PMLS of normal cats but not as precise as that found for primary visual areas in these animals. This suggests that these cells might serve as a substrate for coarse stereopsis.

Bilateral interaction in the second somatosensory area (SII) of the cat and contribution of the corpus callosum.

There are indications in the literature that convergent ipsilateral and contralateral input to the second somatosensory area (SII) may interact. Single unit activity of SII bilateral cells was studied to evaluate the impact of simultaneous bilateral stimulation of the receptive fields (RF) on neural discharge. The cellular responses to unilateral ipsilateral and contralateral, as well as to bilateral stimulation were compared. 22% of bilateral cells showed interaction, usually facilitation. Bilaterally evoked responses were found to be as great as 250% of the strongest unilateral response. Only bilateral responses stronger or weaker than the dominant unilateral response by at least 50% were considered as interactive. The great majority of interactive cells had their RF on the forelimb and were responsive to deep stimulation. The corpus callosum appears to be responsible for part of the observed interaction since in callosotomized cats only 5% of bilateral cells were interactive. A non-callosal ipsilateral pathway must be postulated because both bilaterality and bilateral interaction persist to some degree after callosotomy. A putative role for bilateral interaction in sensory-motor integration is discussed.

Comparative computations of spike synchronization in visual cortex of cats.

In recent years it has been proposed that synchronous activity between neurons is a putative mechanism to bind together various trigger features of an image. Thus the measure of synchronization becomes an important issue since it may be an electrophysiological sign of visual perception. This paper describes and compares six techniques of computing synchronization strength, that is, the central peak of a cross-correlogram. Data were obtained in anesthetized cats prepared for electrophysiological recordings in a conventional fashion. Results indicate that: (1) eye fits are misleading. Visual inspection of cross-correlograms, may be interesting if one needs to estimate approximately synchronization strength and the presence of oscillations in the cross-correlograms, however it may be misleading if one wants to compare different cross-correlograms; (2) regression analysis to compare one method against the others yields a relatively poor correlation suggesting that methods are not directly comparable; (3) the sensitivity of each computational method is unequal. The results may indicate that some functional connections are either under- or over-evaluated depending upon the strategy employed to measure synchronization.