[How Sound Modulates Response to Light when Using Various Temporal Intervals between Stimuli of Different Modalities].

Extracellular activity of 34 primary visual cortex neurons of the rabbit was recorded in response to audiovisual stimulation with stimulus onset asynchronies (SOA) in range from -750 to + 150 ms (15 ranges in total). Auditory stimulus: 2000 Hz, 70 dB, 40 ms. Visual stimuli: two weak intensities 0.3 and 1 cd/m2. Sound by itself did not elicit a neuronal response. Factor analysis ANOVA revealed that SOA significantly influence on neuronal responses. Neuronal reaction included responses to increase (on-response) and decrease (off-response) of light intensity. Earliest phase of response (40-100 ms from visual stimuli substitution) is most affected by sound. Neuronal reactions of the every interval of SOA comprised both increase and decrease of discharge in response to addition of a sound. We used a Wilcoxon signed-rank test to show the differences between reactions in response to visual and audio-visual stimuli. Audio-visual on-responses statistically exceeded the responses to visual stimuli at -150, -40 and 0 ms SOA for the all neurons. Two groups of neurons were revealed. The first group (n = 16) showed dependence of on-re- sponses on sound in a wide range of SOA: -150, -40, -20, 0, +20, +50 and +100 ms. Also the first group showed maximum increase of spike number (18-28%) in response to audio-visual stimulation. For the second group of neurons there were no significant SOA for on-responses. We haven't found a significant decrease of audio-visual response compared to a visual response. However, we found the tendency to reduction of audio-visual discharge at intervals SOA -750 and -80 ms (p < 0.07) for the first group and at SOA -500 and +20 ms (p < 0.1) for the second group of neurons. Also we revealed that on-responses are more influenced by sound than off-responses. We have researched the audio-visual interaction in the second phase of neuronal discharges (120-160 ms and later, n = 23). Sound influence on a second phase is weaker than on a first phase. Significant SOA for on-responses: 0 ms; for off-responses: +100 and +150 ms. This study has revealed similarities of audio-visual interaction range for animal and psychophysical researches. Our results allows. to research cross-modal integration in more detail.

[Sound affects the discrimination of weak intensities of light in the visual cortex of the rabbit depending on time intervals between sound and light].

In this paper, we researched an influence of sound (2000 Hz, 70 dB, 40 ms) to the discrimination of low-light intensities (0.3 and 1 cd/m2) in the visual cortex of the rabbit. We used a recording of evoked potentials from the visual cortex of awaked rabbits in chronic experiments. The sound was switched on with different time slots before and after the replacement of the light intensities at each other (range from -750 to +150 ms). Sound itself caused no response. In 42 experiments on 3 rabbits we revealed that' he sound has a significant modulating effect on the discrimination of low-light intensities in the range of time shifts from -300 to +50 ms. Maximum sound effect was manifested in the transition of light from a high-intensity (1 cd/m2) to lower (0.3 cd/m2). Analyses of the phases of visual evoked potentials revealed that significant influence of sound to the light occurs in the intervals -300, -100, -60, -40, -20, 0, -20 and + 50 ms. We found that phase P2 (120-150 ms from the moment of replacement of the light stimuli) is most affected by sound in response to the replacement of low-light intensities both in the number of significant (p < 0.05) time slots (7) and the impact of sound on the light response. In phase P2 the impact of sound was almost exclusively facilitating (by 19-36%) compared with the responses to the light, whereas in phases N1 80-110 ms) and N2 (180-250 ms) were only 2-3 intervals with the significant influence of the sound. And the degree of response facilitation to light was ranged by 8-12%. We assumed that the effect of sound on the light response in visual cortex is delayed that caused by the passage of auditory signal through the auditory,parietal cortex, superior colliculus.

[Specific modulation of neuronal responses to light of different intensities by sound in the rabbit's primary visual cortex].

Changes in activity of 92 neurons in the primary visual cortex of four rabbits (Orictolagus cuniculus) were analyzed. In the first series of experiments, we recorded discharges of 63 neurons in response to replacement of visual stimuli in pairs (pairs of 0.28 - 1, 1 - 3, 3 - 6, 6 - 8.5, 8.5 - 14, 14 - 17, 17 - 20 cd/m2). Then the same stimuli were presented simultaneously with sound (70 dB, 2000 Hz, 40 ms). Neurons did not respond directly to the sound. Two groups of neurons were found. In the first group of neurons (31%), responses to the complex "light and sound" (40-100 ms from the moment of substitution of stimuli) increased on average by 41% (p < 0.0001) under conditions of the lowest stimuli intensities. With increasing light intensities, discharges to the complex were reduced to the background level of responses to light and even lower. The second group of neurons (19%) showed the opposite properties: at low intensities, responses to the complex were comparable to responses to light (or even lower). At high intensities (14-20 cd/m2), discharges to the complex were significantly (p < 0.05) different from the responses to light (20% and higher, up to 39%). In the second series of experiments, we reconstructed vector sensory spaces on the basis of responses of 29 neurons to light of different intensities and eight complexes of "light and sound." It was found that the sound had also a dual effect on the sensory space of complexes. Some neurons showed an enhancement of the angular distance between the two lowest light intensities (0.28 and 1 cd/m2). Other neurons showed an increase in the angular distance between the highest intensities. Such changes in the space structure are consistent with the groups of neurons revealed in the first two series of the experiments. Comparison of the dynamics of neuronal responses and the amplitudes of evoked potentials under the same conditions of stimulation revealed their considerable similarity. Thus, modulation of neuronal activity in the visual cortex by sound is a complex nonlinear process.

[Sound improves distinction of low intensities of light in the visual cortex of a rabbit].

Electrodes were implanted into cranium above the primary visual cortex of four rabbits (Orictolagus cuniculus). At the first stage, visual evoked potentials (VEPs) were recorded in response to substitution of threshold visual stimuli (0.28 and 0.31 cd/m2). Then the sound (2000 Hz, 84 dB, duration 40 ms) was added simultaneously to every visual stimulus. Single sounds (without visual stimuli) did not produce a VEP-response. It was found that the amplitude ofVEP component N1 (85-110 ms) in response to complex stimuli (visual and sound) increased 1.6 times as compared to "simple" visual stimulation. At the second stage, paired substitutions of 8 different visual stimuli (range 0.38-20.2 cd/m2) by each other were performed. Sensory spaces of intensity were reconstructed on the basis of factor analysis. Sensory spaces of complexes were reconstructed in a similar way for simultaneous visual and sound stimulation. Comparison of vectors representing the stimuli in the spaces showed that the addition of a sound led to a 1.4-fold expansion of the space occupied by smaller intensities (0.28; 1.02; 3.05; 6.35 cd/m2). Also, the addition of the sound led to an arrangement of intensities in an ascending order. At the same time, the sound 1.33-times narrowed the space of larger intensities (8.48; 13.7; 16.8; 20.2 cd/m2). It is suggested that the addition of a sound improves a distinction of smaller intensities and impairs a dis- tinction of larger intensities. Sensory spaces revealed by complex stimuli were two-dimensional. This fact can be a consequence of integration of sound and light in a unified complex at simultaneous stimulation.

[Responses of rabbit's visual cortex neurons to changes in intensity and orientation of visual stimuli].

Changes in the number of spikes in the early phasic discharge (50-90 ms from stimuli replacement) of neurons in the rabbit's primary visual cortex were studied under conditions of an instant change in a flashing-line pattern. We used three type of stimulation: lines with different orientations (0-90 degrees) but constant intensity; lines with constant orientation but different intensities; complex stimuli with different intensities and different orientations of lines. Factor analysis made it possible to reconstruct two-dimensional sensory spaces of orientations in 13 of 43 analyzed neurons (30%). In 5 of 30 analyzed neurons (16.6%), both two-dimensional spaces of orientations and two-dimensional spaces of intensities were revealed. Achromatic spaces were reconstructed during changes in the lines of varying intensities but constant orientation. In experiments with complex stimuli, the intensity of lines with orientations varying from 0 to 38.58 degree was 5 cd/m2. The intensity of lines with orientations varying from 51.44 to 90 degrees was 15 cd/m2. In the sensorial space, stimuli with different intensities were located on the plane formed by the first and second significant factors in opposite quadrants, whereas within each quadrant, the stimuli were arranged closely to their orientation from minimum to maximum. We suggest that this type of sensory space reflects the interaction between intensity and orientation attributes of visual stimuli with the factor of intensity prevailing over the factor of orientation. Only 7 (12%) neurons with such complex spaces were found.

[Visual evoked potentials in rabbit's visual cortex reflect variations in orientation and intensity of lines].

Amplitude variations of visual evoked potentials (VEP) in rabbit were studied using changes of three types of stimuli: lines with different orientations (0-90 degree) but constant in intensity, lines with different intensities but constant in orientation, and complex stimuli with different intensities and different orientations. Factor analysis of component N85 of VEP to stimuli with varying orientation and constant intensity revealed two-dimensional sensorial space of orientations. Also, the two-dimensional achromatic sensorial space was revealed for stimuli different only in intensities. In experiments with complex stimuli, two versions of stimulation were used. In the first version, four stimuli with varying orientation from 0 to 38.58 degrees were of 5 cd/m2 in intensity, the remaining four stimuli (from 51.44 to 90 degrees) were of 15 cd/m2 in intensity. In the sensorial space, stimuli with different intensities were disposed on the plane formed by the first two significant factors in the opposite quadrants, whereas within each quadrant, stimuli were arranged according to their orientation from minimum to maximum degrees. It is suggested that, in this version of stimulation, the interaction between two characteristics (intensity and orientation) took place with clear-cut dominance of the factor of intensity. In the second version of the experiments, also eight complex stimuli were used. Each stimulus was characterized by specific combination of orientation (one of the eight directions in the range from 0 to 90 degrees) and intensity (one of the eight intensities in the range from 5 to 21 cd/m2). Three or four significant factors were revealed in each test. Only the plane of intensity of sensorial space (XIX2) formed by two significant factors was revealed in the overwhelming majority of cases. On this plane, stimuli were arranged according to their intensities. This fact can be explained by specific (twilight) type of rabbit's vision with predominant discrimination of intensities. However, some findings suggest that visual evoked potentials reflect simultaneous processing of two characteristics of visual stimuli (intensity and orientation) being indicative of a possibility of the complex stimuli analysis in the primary visual cortex.

[Estimation of color and brightness differences by neurons in the rabbit superior colliculus].

Changes in activity of 83 neurons in the rabbit colliculus superior evoked by the replacement of eight color and eight achromatic stimuli in pairs were analyzed. It was found out that neurons displayed the early and late phasic responses (within 50-90 and 120-300 ms respectively, after the replacement) and long-term tonic response component, which depended on stimuli intensity. Analysis of phasic component revealed three neuronal groups. The first group (n=25, 30%) selected on the basis of the earliest component, was specialized to differentiate stimuli only by intensities. The perceptual spaces of these neurons reconstructed on the basis of spike discharge in the earliest response were two-dimensional. The second group of neurons (n=16, 19%) selected on the basis of the late phasic component demonstrated four-dimensional structure of perceptual space. Neurons of the third group (n=4, 5%) possessed a two-dimensional structure of perceptual space reconstructed by the analysis of the early component, whereas analysis of the late response revealed a four-dimensional structure. We suggest that information about differences between stimuli in color and intensity coming from cortical neurons is necessary for the reconstruction of four-dimensional space. The structure of perceptual spaces reconstructed on the basis of phasic responses of neurons in the colliculus superior was similar to the spaces of neurons in the primary visual cortex and lateral geniculate nucleus. The structure of perceptual space reconstructed on the basis of neuronal spikes was also similar to the space calculated from the N85 component of the visual evoked potential recorded under similar conditions. This finding confirms the general principle of vector coding in the visual system.

[Computation of responses to color and brightness differences of neurons in the rabbit lateral geniculate nucleus].

Changes in activity of 51 neurons in the rabbit lateral geniculate nucleus evoked by the replacement of eight color and eight achromatic stimuli in pairs were analyzed. It was found that neurons displayed the earliest phasic (within 50-90 ms after the replacement) and tonic response components. The earliest component strongly correlated with differences between stimuli, whereas the tonic component depended on stimuli intensity. Analysis of phasic component revealed two neuronal populations: the first group of cells was specialized for stimuli differentiation only by their intensities, and, and the second group could measure differences in colors and intensities. Neuronal perceptual spaces were reconstructed using the average of the earliest response component as a measure of differences between stimuli. Spaces of 44 neurons (86%) were two-dimensional with brightness and darkness axes. Such neurons had the same structures of space for color and achromatic stimuli. Spaces of 7 neurons (14%) were four-dimensional with two chromatic and two achromatic axes. The structures of perceptual space reconstructed from neurons in the lateral geniculate nucleus were identical to the spaces calculated from the neurons in the primary visual cortex. The structure of the perceptual space reconstructed from neuronal spikes was also similar to space calculated from the N85 visual evoked potential component recorded under similar conditions and to another space reconstructed on the basis of rabbit's instrumental learning. This fact confirmed the general principle of vector coding in the visual system. The tonic component of the most of neurons in the lateral geniculate nucleus showed a linear correlation with changes in intensities, thereby these neurons could be characterized as pre-detectors for cortical selective detectors.

[Limited plasticity of the rabbit visual cortex and hippocampal neurons in the oddball test].

The activity of 41 visual cortex and 20 hippocampal neurons from field CA1 was registered in experiments using oddball-stimulation with different color stimuli varied in intensity. 34% cortical and 37% hippocampal neurons demonstrated plasticity reactions. The significant increase of latest phases of neuronal activity (200-500 and 200-1000 ms after stimulation for cortical neurons and 300-550 ms for hippocampal neurons) was shown in responses to rare deviant stimuli, which had a less intensity than frequently standards. The quantity of the earliest neuronal phase of activity (40-120 ms after stimulation) was stabilized in responses to deviants and standards during the experiment. We propose that such increase of the latest phases of neuronal activity (the limited plasticity) may reflect the mechanisms of orienting reaction.

[Computation of color and brightness differences by neurons in the rabbit visual cortex].

Changes in activity of 54 neurons in the rabbit visual cortex evoked by the replacement of eight color and eight achromatic stimuli in pairs were analyzed. The diffused stimuli generated by color SVGA monitor were used in the experiments. The earliest response of phasic neurons (50-90 ms after the replacement) was strongly correlated with differences between stimuli in color or intensity. This response ("the signal of differences") was used as a basis of a matrix (8 x 8) constructed for each neuron. Such matrices included mean numbers of spikes per second in responses to changes of different stimuli pairs. All matrices were subjected to factor analysis, and the basic axes (the main factors) of sensory spaces were revealed. It was found that 16 neurons (30%) detected only achromatic differences between stimuli. Perceptual spaces of these neurons were two-dimensional with brightness and darkness orthogonal axes. The spaces of 12 neurons (22%) were four-dimensional with two chromatic and two achromatic axes. The structure of the perceptual space reconstructed from neuronal spikes was similar to the space calculated from the early VEP components recorded under similar conditions and to another space reconstructed on the basis of rabbit's instrumental learning. The fundamental coincidence of color spaces revealed by different methods may reflect the general principle of vector coding in the visual system and suggests the coexistence of two independent cortical mechanisms of the detection of chromatic and achromatic differences.

[The phases of visual cortex and hippocampal evoked potentials in rabbit reflect the orienting reaction in case of visual stimuli intensity changes]. / Otrazhenie orientirovochnogo refleksa v fazakh vyzvannykh potentsialov v zritel'noi kore i gippokampe krolika pri zamene intensivnosti stimulov.

The oddball paradigm was applied in experiments with waking rabbits using rare (deviant) and frequent (standard) stimuli, which were similar in color but different in intensity, deviant ones being of lower intensity. In addition, the VEPs to the single deviants were averaged. Such single deviants (without standard stimuli) were given at the beginning and at the end of the stimulation. The positivity of cortical and hippocampal visual evoked potentials to deviant stimuli increased in comparison to responses to standards and single deviants. The VEP-peaks P1 and P2 in the visual cortex and the VEP P1, N1 and P2 peaks in hippocampus increased. The most prominent significant changes were demonstrated for cortical VEP P2 (P130) peak. It is suggested that the increase of positivity to oddball deviants is due to the orienting reflex arising in response to rare stimuli. The increase of P2-peak can be connected with the transfer of information signaling changes of light intensity. It was demonstrated that the most clear and contrast differences in the VEPs to deviants and standards took place for the case of minimal distinction in their intensities. This fact may reflect the orienting reactionto threshold stimuli.

[Orienting reflex: "targeting reaction" and "searchlight of attention"]. / Orientirovochnyi refleks: "reaktsiia pritselivaniia" i "prozhektor vnimaniia".

The concept of orienting reflex based on the principle of vector coding of cognitive and executive processes is proposed. The orienting reflex to non-signal and signal stimuli is a set of orienting reactions: motor, autonomic, neuronal, and subjective emphasizing new and significant stimuli. Two basic mechanisms can be identified within the orienting reflex: a "targeting reaction" and a "searchlight of attention". In the visual system the first one consists in a foveation of a target stimulus. The foveation is performed with participation of premotor neurons excited by saccadic command neurons of the superior colliculi. The "searchlight of attention" is based on the resonance of gamma-oscillations in the reticular thalamus selectively enhancing responses of cortical neurons (involuntary attention). The novelty signal is generated in novelty neurons of the hippocampus, which are selectively tuned to a repeatedly presented standard stimulus. The selective tuning is caused by the depression of plastic synapses representing a "neuronal model" of the standard stimulus. A mismatch of the novel stimulus with the established neuronal model gives rise to a "novelty signal" enhancing the novel input. The novelty signal inhibits current conditioned reflexes (external inhibition) contributing to redirecting the behavior. By triggering the expression of early genes the novelty signal initiates the formation of the long-term memory connected with neoneurogenesis.

[The reconstruction of the perceptual spaces of brightness and color on the basis of visual evoked potentials and their comparison with the data from behavioral trials]. / Rekonstruktsiia pertseptivnykh prostranstv iarkosti i tsveta na osnove zritel'nykh vyzvannykh potentsialov i ikh sravnenie s dannymi povedencheskikh opytov.

The visual evoked potentials to a change in color stimuli were studied. The amplitude of the N85 component was correlated with color discrimination. In cases when the brightness of one color was fixed and that of the other one changed, the amplitude dynamics of N85 was V-shaped with the minimum corresponding to the point of equal brightness of both colors. The N85 amplitude in this point serves as a measure of discrimination between stimuli chromaticity. The perceptual color space in rabbit (possessing two cone pigments) was constructed in accordance with the amplitudes of N85 to color change. This space represented a hypersphere in the four-dimensional Euclidean space. The perceptual spaces for brightness and color reconstructed on the basis of conditioning probabilities and N85 amplitudes evoked by replacement of colored and achromatic stimuli were shown to coincide. This suggests the common mechanism of the vector color coding.

[A model of color vision in trichromats and protans]. / Model' tsvetnogo zreniia trikhromatov i protanov.

A model which explains the human vision protanopic deficiency and its biologic prototype with the absence of red-absorbing pigment (rabbit) was constructed from neuron-like elements. In behavioral experiments and by means of evoked potential technique it was shown that the rabbit's color space is characterized by a spherical four-dimensional with a reduction of red-coding area. Similar spherical four-dimensional structure of color space is characteristic for a group of protanopic human subjects. The perceptive space of another group of protanopic subjects (protanomals) is characterized by a reduction of both parts of the red-green opponent axis. These disorders are reproduced in the model either by a loss of some color-coding elements (the absence of the red-absorbing pigment as in protanops) or a shift of the spectral characteristics of the red pigment towards those of the green one (protanomals).

[The dynamic localization of a dipole source of the alpha rhythm in the human brain]. / Dinamicheskaia lokalizatsiia dipol'nogo istochnika al'fa-ritma v mozge cheloveka.

In 11 subjects the dynamic localization (with 2-ms step) of the equivalent current dipole of alpha rhythm during travelling alpha waves was studied using one-dipole 3-layered spherical head model and MRI. The dipole was localized in the occipital cortex and during the development of a single alpha wave it shifted in one or another direction while dipole's moment revealed fan-like rotation mainly in sagittal and horizontal planes. The results obtained indicate changing localization of the alpha-rhythm source in the region corresponding to striate cortex. They seem to prove scanning hypothesis that is still under discussion.

[The brightness components of the visual evoked potential to color stimuli in the rabbit]. / Iarkostnye komponenty zritel'nogo vyzvannogo potentsiala na tsvetovye stimuli u krolika.

Two earliest components of visual evoked potentials (N85 and P130) which were related with substitution of stimuli for those identical in spectra but different in brightness were detected in rabbits. This finding suggests an analogy between the N85 and P139 in rabbits and N87 and P120 in humans.

[Evoked potentials in assessing the sensory, activating and cognitive functions of the normal human brain and in cortical and subcortical types of vascular dementia]. / Vyzvannye potentsialy v otsenke sensornykh, aktiviruiushchikh i kognitivnykh funktsii mozga cheloveka v norme i pri sosudistoi dementsii korkovogo i podkorkovogo tipa.

Characteristics of the auditory EP under conditions of stimulus recognition (the P300 technique) and parameters of the visual "nonspecific" EP to light flash were studied in healthy subjects of the wide age range (18-78 years old) and patients with vascular dementia of different severity. It was shown that P300 parameters depended on age in healthy patients and on expression of cognitive dysfunction in demented patients. The changes were more pronounced in dementia of the cortical type. The P300 to nonrelevant stimulus was not age-dependent and slightly changed at cognitive dysfunction. Parameters of the nonspecific visual responses (the latency and duration--the index of homeostatic regulation) and maximal amplitude (reflecting the extent of activation processes) were not age-dependent and changed both in subcortical and cortical dementia types (increase in response latency and duration). In cognitive dysfunctions of subcortical origin homeostatic changes in the form of increase in time of homeostatic regulation and deterioration of stabilization of brain electrical processes were more pronounced.

[The perceptive color space of the rabbit]. / Pertseptivnoe tsvetovoe prostranstvo krolika.

Discrimination of colors was studied using instrumental learning paradigm in three rabbits (Oryctolagus cuniculus). The rabbits were able to discriminate all but red stimuli by their color. The red stimulus could not be discriminated from the black one. The confusion matrix composed of probabilities of instrumental reactions was subjected to factor analysis in order to reveal the basic axes of the color perceptual space. The four-dimensional spherical structure of perceptual color space was obtained, which was different from that of trichromatic species in a reduction of color axes in red and yellow parts. The evidence characterizes rabbit as a protanopic animal.

[The characteristics of the color perceptive space in protanomals]. / Osobennosti tsvetovogo pertseptivnogo prostranstva protanomalov.

Color vision of three protanomal subjects was studied by means of direct paired comparison technique using 25 colors with different brightness. It was shown that the characteristics of their color vision could be completely and adequately described in the frames of the four-dimensional spherical model of color perception. The spatial axes could be identified as the two color-opponent mechanisms (red-green and blue-yellow) and the two achromatic mechanisms (brightness and darkness). Deformation of the color axes in protanomals (as compared with the normal trichromatics) was demonstrated in the "red" and "yellow" spectral region. The visual disturbance in protanomal subjects involves not only color but also achromatic mechanisms. This is manifested in the a deformation of perceptual brightness scale. In comparison with normal trichromatic subjects, the protanomals perceive the red and adjacent colors as achromatic while green, yellow-green, and orange as more bright but low-saturated colors.

[The discrimination by the rabbit of light intensities and the plotting of its achromatic space]. / Razlichenie krolikom intensivnostei sveta i postroenie ego akhromaticheskogo prostranstva.

Elaboration of differential instrumental conditioned reflexes in rabbits showed that probabilities of responses to differential stimuli were inversely related to differences in intensity between conditioned and differential stimuli. Factor analysis of a response probability matrix revealed two orthogonal axes. They could be interpreted as the axes of brightness and darkness in achromatic space of a rabbit. Lights of different intensities were located on a semicircle in accordance with their intensities. The experiments imply that light intensities are coded by excitation vectors composed of responses of brightness and darkness neurons.