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.

Effects of hyperthermic seizures on the developing primary visual cortex of the rat.

Febrile seizures occurring during childhood have been shown to interfere with the development of cognitive functions. However, an alteration of the developing sensory systems might also result from febrile seizures. In order to test this hypothesis, seizures were induced by hyperthermia in Long Evans rats on postnatal day 10. Extracellular single neuron recordings were carried out from postnatal days 15 to 30 and at adulthood. The response of neurons in the primary visual cortex to drifting sinusoidal gratings was recorded in anaesthetized rats. As soon as postnatal day 15, the neurons of rats having experienced a hyperthermic seizure showed significantly lower optimal spatial frequencies (SF), broader directional and temporal bandwidths, as well as higher contrast thresholds than did neurons recorded in normal rats. At adulthood, significantly broader spatial bandwidths and lower optimal temporal frequencies (TF) were obtained from neurons of rats subjected to hyperthermia. These results suggest that febrile seizures during infancy could affect the development of spatio-temporal receptive field properties of neurons in primary visual cortex. Such alterations of a sensory system might contribute to the cognitive deficits associated with early-onset febrile seizures.

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.

Responses of inferior collicular cells to species-specific vocalizations in normal and enucleated rats.

The inferior colliculus (IC) is an obligatory relay for the ascending and descending auditory pathways. Cells in this brainstem structure not only analyze auditory stimuli but they also play a major role in multi-modal integration of auditory and visual information. The aim of the present study was to determine whether cells in the central nucleus of the inferior colliculus (CNIC) of normal rats respond selectively to complex auditory signals, such as species-specific vocalizations, and compare their responses to those obtained in neonatal bilateral enucleated (P2-P3) adult rats. Extra-cellular recordings were carried out in anesthetized normal and enucleated rats using auditory stimuli (pure tones, broadband noise and vocalizations) presented in free field in a semi-anechoic chamber. The results indicate that most cells in the CNIC of both groups respond selectively to species-specific vocalizations better than to the same but inverted sounds. No significant differences were found between the normal and enucleated rat groups in their responses to broadband noise and pure tones.

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.

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.

Blind subjects process auditory spectral cues more efficiently than sighted individuals.

The goal of the present study was to investigate how monaural sound localization on the horizontal plane in blind humans is affected by manipulating spectral cues. As reported in a previous study (Lessard et al. 1998), blind subjects are able to calibrate their auditory space despite their congenital lack of vision. Moreover, the performance level of half of the blind subjects was superior to that of sighted subjects under monaural listening conditions. Here, we first tested ten blind subjects and five controls in free-field (1) binaural and (2) monaural sound localization tasks. Results showed that, contrary to controls and half the blind subjects, five of the blind listeners were able to localize the sounds with one ear blocked. The blind subjects who showed good monaural localization performances were then re-tested in three additional monaural tasks, but we manipulated their ability to use spectral cues to carry out their discrimination. These subjects thus localized these same sounds: (3) with acoustical paste on the pinna, (4) with high-pass sounds and unobstructed pinna and (5) with low-pass sounds and unobstructed pinna. A significant increase in localization errors was observed when their ability to use spectral cues was altered. We conclude that one of the reasons why some blind subjects show supra-normal performances might be that they more effectively utilize auditory spectral cues.

Putting order into the development of sensitivity to global motion.

We studied differences in the development of sensitivity to first-versus second-order global motion by comparing the motion coherence thresholds of 5-year-olds and adults tested at three speeds (1.5, 6, and 9 degrees s(-1)). We used Random Gabor Kinematograms (RGKs) formed with luminance-modulated (first-order) or contrast-modulated (second-order) concentric Gabor patterns with a sinusoidal spatial frequency of 3c deg(-1). To achieve equal visibility, modulation depth was set at 30% for first-order Gabors and at 100%, for second-order Gabors. Subjects were 24 adults and 24 5-year-olds. For both first- and second-order global motion, the motion coherence threshold of 5-year-olds was less mature for the slowest speed (1.5 degrees s(-1)) than for the two faster speeds (6 and 9 degrees s(-1)). In addition, at the slowest speed, the immaturity was greater for second-order than for first-order global motion. The findings suggest that the extrastriate mechanisms underlying the perception of global motion are different, at least in part, for first- versus second-order signals and for slower versus faster speeds. They also suggest that those separate mechanisms mature at different rates during middle childhood.

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.

Longer VEP latencies and slower reaction times to the onset of second-order motion than to the onset of first-order motion.

We compared visual evoked potentials and psychophysical reaction times to the onset of first- and second-order motion. The stimuli consisted of luminance-modulated (first-order) and contrast-modulated (second-order) 1 cpd vertical sine-wave gratings drifting rightward for 140 ms at a velocity of 6 degrees /s. For each condition, we analysed the latencies and peak-to-baseline amplitudes of the P1 and N2 peaks recorded at Oz. For first-order motion, both P1 and N2 peaks were present at low (3%) contrast (i.e., depth modulations) whereas for second-order motion they appeared only at higher (25%) contrasts. When the two types of motion were equated for visibility, responses were slower for second-order motion than for first-order motion: about 44 ms slower for P1 latencies, 53 ms slower for N2 latencies, and 76 ms slower for reaction times. The longer VEP latencies for second-order motion support models that postulate additional processing steps for the extraction of second-order motion. The slower reaction time to the onset of second-order motion suggests that the longer neurophysiological analysis translates into slower detection.

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 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.

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.

Contrast dependency of VEPs as a function of spatial frequency: the parvocellular and magnocellular contributions to human VEPs.

The present study investigated the contrast dependency of visual evoked potentials (VEPs) elicited by phase reversing sine wave gratings of varying spatial frequency. Sixty-five trials were recorded for each of 54 conditions: 6 spatial frequencies (0.8, 1.7, 2.8, 4.0, 8.0 and 16.0 c deg(-1)) each presented at 9 contrast levels (2, 4, 8, 11, 16, 23, 32, 64 and 90%). At the lowest spatial frequency, the waveform contained mainly one peak (P1). For spatial frequencies up to 8 c deg(-1), P1 had a characteristic magnocellular contrast response: it appeared at low contrasts, increased rapidly in amplitude with increasing contrast, and saturated at medium contrasts. With increasing spatial frequency, an additional peak (N1) gradually became the more dominant component of the waveform. N1 had a characteristic parvocellular contrast response: it appeared at medium to high contrasts, increased linearly in amplitude with increasing contrast, and did not appear to saturate. The data suggest the contribution of both magnocellular and parvocellular responses at intermediate spatial frequencies. Only at the lowest and highest spatial frequencies tested did magnocellular and parvocellular responses, respectively, appear to dominate.

Neurons in the posteromedial lateral suprasylvian area of the cat are sensitive to binocular positional depth cues.

Single units in the posteromedial lateral suprasylvian area of the cat are known to be very sensitive to movement. A proportion of these cells can encode movement in depth, but it is unclear whether posteromedial lateral suprasylvian cells only rely upon motion cues to evaluate stimulus depth or whether they can also code for spatial cues. The present study aims at assessing the sensitivity to spatial disparity of binocular cells, in the postero-medial lateral suprasylvian area, in order to determine whether these units are tuned to positional depth cues. A total of 126 single cells located in the posteromedial lateral suprasylvian area of anesthetized, paralyzed cats were examined. As recordings were performed in the central visual field representation, receptive fields were small. A third of the receptive fields were surrounded by an inhibitory region and almost three-quarters of the cells were direction-selective. Most cells (110/114) were binocular, and a large proportion of single neurons responded to stimuli appearing on the fixation plane by increasing (tuned excitatory cells, 43%) or decreasing (tuned inhibitory cells, 14%) their response rate. A smaller proportion of cells increased their firing rate in response to crossed (near cells, 10%) or uncrossed (far cells, 6%) spatial disparities, hence demonstrating respective preference for stimuli presumably appearing in front of or behind the fixation plane. As compared to primary visual cortex, the proportion of disparity-sensitive cells in posteromedial lateral suprasylvian area is similar, but selectivity is significantly coarser. As the posteromedial lateral suprasylvian area can code for both spatial and temporal aspects of stimuli, this area might be involved in the spatiotemporal integration of depth cues, a process that may also participate in the control of accommodation and vergence.

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.

Cellular response to texture and form defined by motion in area 19 of the cat.

The present study examined the neuronal sensitivity in area 19 of the cat to a motion-defined bar and to texture. Sensitivity was tested in normal, lesioned (areas 17-18) and split-chiasm cats using a kinematogram, as well as a textured bar drifting on a uniform light background and a light bar drifting on a stationary textured background. Texture density was varied. The results indicate that almost all cells of area 19 recorded in the three groups of cats responded to a motion-defined bar or to its edges. Texture density influenced the responses in that the discharge rate increased as density decreased. However, the majority of cells were sensitive to the highest texture density kinematogram. Moreover, the neural responses of all cats were either independent of the density of the textured bar or background, or were modulated by it. These results show that cells in area 19 can signal the presence of a kinetic bar and that the density of either the textured bar, the background or both can influence figure-ground detection. The results are interpreted with respect to how various inputs influence the function of area 19.

Responses of cells to stationary and moving sound stimuli in the anterior ectosylvian cortex of cats.

The azimuthal, directional and angular speed sound selectivities of single units were examined in the posterior part of the anterior ectosylvian cortex. Broadband noise bursts and simulated moving sounds were delivered from 16 loudspeakers fixed on the horizontal plane in a quasi-anechoic sound-isolation chamber. The activity of 78 neurons was recorded and quantitatively analyzed. Most cells responded to at least the static sound. The relative strengths of their responses suggested that the cells could be classed as omnidirectional (37.2%), contralateral hemifield (29.5%), ipsilateral hemifield (2.5%) and azimuth (7.7%) selective. The remaining 23.1% could not be classified. All cells responded to a simulated moving sound displaced at five different speeds. A majority (88%) of them showed some directional preference in that they discharged at least twice as strongly for one direction as for the other for at least one speed. 14.7% displayed angular speed selectivity. Different patterns of neuronal discharges were evoked. For static sounds, most of the cells gave ON-type responses. A large proportion (60%) of the cells responded in a sustained manner to maintained stimulation. Among these, 68% also gave sustained discharges to moving sounds. The spatial tuning and the directional and angular speed selectivity of neurons in the posterior part of the AEC suggest that this area is involved in the processing of static and moving sounds.

Binocular interactions and spatial disparity sensitivity in the superior colliculus of the Siamese cat.

In Siamese cats, a genetically determined massive misrouting of retinal ganglion cells toward the contralateral hemisphere, as well as an accompanying strabismus, is believed to underlie the extreme paucity of binocular cells in the primary visual cortex. However, binocular cells have been shown to be present in more important numbers at the collicular level. The present study aims at investigating binocular interactions and sensitivity to spatial disparity in the superior colliculus of the Siamese cat. The activity of single units was recorded in the superficial layers of paralyzed and anesthetized Siamese cats. Although most collicular cells were monocularly driven, a significant proportion could be driven through both eyes (34/216 or 16%). Upon isolation of a binocular cell, the receptive fields were separated, then simultaneously stimulated with two light bars. A temporal delay was introduced between the arrival of the bars in the receptive fields to generate spatial disparities (-3 degrees to +3 degrees, in 0.5 degrees or 1 degree steps). Results showed that some binocular cells presented disparity tuning profiles similar to the tuned excitatory (12/34), tuned inhibitory (2/34), near (2/34) and far (3/34) cells found at various cortical levels in the normal cat. These interactions might allow for coarse binocular fusion as well as play a role in the initiation of vergence and the fixation of the eyes upon the appropriate plane of vision.

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.