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.     

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 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

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.     

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.     

Spectral Characteristics of Phase Sensitivity and Discharge Rate of Neurons in the Ascending Tectofugal Visual System

Drifting gratings can modulate the activity of visual neurons at the temporal frequency of the stimulus. In order to characterize the temporal frequency modulation in the cat’s ascending tectofugal visual system, we recorded the activity of single neurons in the superior colliculus, the suprageniculate nucleus, and the anterior ectosylvian cortex during visual stimulation with drifting sine-wave gratings. In response to such stimuli, neurons in each structure showed an increase in firing rate and/or oscillatory modulated firing at the temporal frequency of the stimulus (phase sensitivity). To obtain a more complete characterization of the neural responses in spatiotemporal frequency domain, we analyzed the mean firing rate and the strength of the oscillatory modulations measured by the standardized Fourier component of the response at the temporal frequency of the stimulus. We show that the spatiotemporal stimulus parameters that elicit maximal oscillations often differ from those that elicit a maximal discharge rate. Furthermore, the temporal modulation and discharge-rate spectral receptive fields often do not overlap, suggesting that the detection range for visual stimuli provided jointly by modulated and unmodulated response components is larger than the range provided by a one response component.     

Different Stimuli,Different Spatial Codes: A Visual Map and an Auditory Rate Code for Oculomotor Space in the Primate Superior Colliculus

Maps are a mainstay of visual, somatosensory, and motor coding in many species. However, auditory maps of space have not been reported in the primate brain. Instead, recent studies have suggested that sound location may be encoded via broadly responsive neurons whose firing rates vary roughly proportionately with sound azimuth. Within frontal space, maps and such rate codes involve different response patterns at the level of individual neurons. Maps consist of neurons exhibiting circumscribed receptive fields, whereas rate codes involve open-ended response patterns that peak in the periphery. This coding format discrepancy therefore poses a potential problem for brain regions responsible for representing both visual and auditory information. Here, we investigated the coding of auditory space in the primate superior colliculus(SC), a structure known to contain visual and oculomotor maps for guiding saccades. We report that, for visual stimuli, neurons showed circumscribed receptive fields consistent with a map, but for auditory stimuli, they had open-ended response patterns consistent with a rate or level-of-activity code for location. The discrepant response patterns were not segregated into different neural populations but occurred in the same neurons. We show that a read-out algorithm in which the site and level of SC activity both contribute to the computation of stimulus location is successful at evaluating the discrepant visual and auditory codes, and can account for subtle but systematic differences in the accuracy of auditory compared to visual saccades. This suggests that a given population of neurons can use different codes to support appropriate multimodal behavior.     

Orientational and directional selectivities of visual neurons in the superior colliculus of the cat

Based on quantitative analyses of the response characteristics of visual neurons in the superior colliculus to moving optical bar stimuli, it is demonstrated for the first time that the visual neurons in superior colliculus of the cat have, to some extent, orientational selectivity. The significance of this selectivity is discussed in reference to its morphological substrate and physiological functions. In addition, both the directional and orientational selectivities in the superior colliculus are relatively weak when compared with those in the primary visual cortex, and the majority of the neurons prefer upward or downward motion in the visual field.     

Hemispheric asymetry of the evoked electrical activity of the cerebral cortex to letter and non-verbal stimuli]

Average evoked potentials (AEP) were recorded in practically healthy subjects to "meaningless" figures and letters, presented to different halves of the visual field. Analysis of the amplitudes of AEP late components to verbal and non-verbal stimuli reveals hemispheric asymmetry. A higher amplitude of the late positive evoked response (P300) to a "direct" stimulation both by verbal and non-verbal stimuli (in the contralateral field of vision) is recorded in the left hemisphere than in the right one. Similar stimulation of the right hemisphere does not reveal sucha difference. In the left hemisphere the P300 wave is of a clearly greater amplitude to a "direct" stimulation (contralateral visual field) than to an "indirect" one (ipsilateral visual field), regardless of the nature of the stimulus. No such difference is observed in the right hemisphere. The magnitude of the late negative wave (component N200) to non-verbal stimuli is greater in the right hemisphere both in response to "direct" and "indirect" stimulations. No intrahemispheric difference has been found in the amplitude of late evoked responses of the cerebral cortex to verbal and non-verbal stimuli.     

Figure-ground activity in V1 and guidance of saccadic eye movements.

Every day we shift our gaze about 150.000 times mostly without noticing it. The direction of these gaze shifts are not random but directed by sensory information and internal factors. After each movement the eyes hold still for a brief moment so that visual information at the center of our gaze can be processed in detail. This means that visual information at the saccade target location is sufficient to accurately guide the gaze shift but yet is not sufficiently processed to be fully perceived. In this paper I will discuss the possible role of activity in the primary visual cortex (V1), in particular figure-ground activity, in oculo-motor behavior. Figure-ground activity occurs during the late response period of V1 neurons and correlates with perception. The strength of figure-ground responses predicts the direction and moment of saccadic eye movements. The superior colliculus, a gaze control center that integrates visual and motor signals, receives direct anatomical connections from V1. These projections may convey the perceptual information that is required for appropriate gaze shifts. In conclusion, figure-ground activity in V1 may act as an intermediate component linking visual and motor signals.     

Neural representation of cutaneous aftersensations by spinothalamic tract neurons.

Temporal summation of second pain and long-lasting tactile-evoked aftersensations are examples of sensory phenomenons that cannot be explained on the basis of responses of primary afferents. Two distinct classes of monkey spinothalamic tract neurons have responses to controlled natural stimuli that parallel and thus could account for the above phenomenons. One class, termed wide-dynamic-range, receives excitatory effects from sensitive mechanoreceptive afferents and from various nociceptive afferents including Adelta and C mechanothermal nociceptive afferents. Another class, termed nociceptive-specific, receives excitatory effects exclusively from primary nociceptive afferents. Both classes respond with an early and late response to a single noxious heat pulse (peak temperature = 51 C). The late response, unlike C nociceptive afferents but like second pain, summates in magnitude with each successive heat pulse. Gentle moving tactile stimuli evoke long-lasting (20-56 sec) after-discharges only in wide dynamic range neurons, and are similar in duration to the tactile after-sensation evoked by similar stimuli. Both the after-discharges and after-sensations can be abruptly terminated by rubbing the affected region. Temporal summation of second pain and cutaneous after-sensations are at least partly subserved by spinal cord mechanisms within the dorsal horn and are manifested in the output of spinothalamic tract neurons.     

United we sense, divided we fail: context-driven perception of ambiguous visual stimuli

Ambiguous visual stimuli provide the brain with sensory information that contains conflicting evidence for multiple mutually exclusive interpretations. Two distinct aspects of the phenomenological experience associated with viewing ambiguous visual stimuli are the apparent stability of perception whenever one perceptual interpretation is dominant, and the instability of perception that causes perceptual dominance to alternate between perceptual interpretations upon extended viewing. This review summarizes several ways in which contextual information can help the brain resolve visual ambiguities and construct temporarily stable perceptual experiences. Temporal context through prior stimulation or internal brain states brought about by feedback from higher cortical processing levels may alter the response characteristics of specific neurons involved in rivalry resolution. Furthermore, spatial or crossmodal context may strengthen the neuronal representation of one of the possible perceptual interpretations and consequently bias the rivalry process towards it. We suggest that contextual influences on perceptual choices with ambiguous visual stimuli can be highly informative about the neuronal mechanisms of context-driven inference in the general processes of perceptual decision-making.     

Single units and conscious vision.

Figures that can be seen in more than one way are invaluable tools for the study of the neural basis of visual awareness, because such stimuli permit the dissociation of the neural responses that underlie what we perceive at any given time from those forming the sensory representation of a visual pattern. To study the former type of responses, monkeys were subjected to binocular rivalry, and the response of neurons in a number of different visual areas was studied while the animals reported their alternating percepts by pulling levers. Perception-related modulations of neural activity were found to occur to different extents in different cortical visual areas. The cells that were affected by suppression were almost exclusively binocular, and their proportion was found to increase in the higher processing stages of the visual system. The strongest correlations between neural activity and perception were observed in the visual areas of the temporal lobe. A strikingly large number of neurons in the early visual areas remained active during the perceptual suppression of the stimulus, a finding suggesting that conscious visual perception might be mediated by only a subset of the cells exhibiting stimulus selective responses. These physiological findings, together with a number of recent psychophysical studies, offer a new explanation of the phenomenon of binocular rivalry. Indeed, rivalry has long been considered to be closely linked with binocular fusion and stereopsis, and the sequences of dominance and suppression have been viewed as the result of competition between the two monocular channels. The physiological data presented here are incompatible with this interpretation. Rather than reflecting interocular competition, the rivalry is most probably between the two different central neural representations generated by the dichoptically presented stimuli. The mechanisms of rivalry are probably the same as, or very similar to, those underlying multistable perception in general, and further physiological studies might reveal much about the neural mechanisms of our perceptual organization.     

Two types of binocular convergence of visual information in the cat superior colliculus

During binocular stimulation of different sectors of the retina the amplitude of the two first postsynaptic components of the evoked potential in the superior colliculus to the second stimulus varies with the time delay between the testing and conditioning stimuli. Correlation is shown between the form of the evoked potential arising in response to the conditioning stimulus and the character of convergence of visual impluses in the superior colliculus. Qualitative differences are found in binocular interaction between sensory impulses depending on the way in which the conditioning impulses reach the region of the superior colliculus tested. An attempt is made to assess interaction between sensory volleys in the superior colliculus quantitatively.Institute of the Brain, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 5, No. 2, pp. 133–137, March–April, 1973.     

The effect of gaze angle and fixation distance on the responses of neurons in V1, V2, and V4.

What we see depends on where we look. This paper characterizes the modulatory effects of point of regard in three-dimensional space on responsiveness of visual cortical neurons in areas V1, V2, and V4. Such modulatory effects are both common, affecting 85% of cells, and strong, frequently producing changes of mean firing rate by a factor of 10. The prevalence of neurons in area V4 showing a preference for near distances may be indicative of the involvement of this area in close scrutiny during object recognition. We propose that eye-position signals can be exploited by visual cortex as classical conditioning stimuli, enabling the perceptual learning of systematic relationships between point of regard and the structure of the visual environment.     

Auditory properties of the superior colliculus in the horseshoe bat,Rhinolophus rouxi

Summary Auditory response properties were studied in the superior colliculus (SC) of the echolocating horseshoe bat Rhinolophus rouxi, a long CF-FM bat, by the use of stationary, dichotic stimuli.The most striking finding in the horseshoe bat was an enormous overrepresentation of neurons with best frequencies in the range of the constant frequency component of the species specific echolocation call (72% of the auditory neurons). These neurons had response thresholds as low as 0 dB SPL and were narrowly tuned with Q_{10 dB} — values up to 400, just as in the nuclei of the primary auditory pathway in this species. This overrepresentation may suggest the importance of the superior colliculus in the context of echolocation behavior.While noise stimuli were not particularly effective, other auditory response properties were similar to those described in other mammals. 65% of the SC neurons in the horseshoe bat responded only to monaural stimulation of one ear, primarily the contralateral one. 32% of the neurons received monaural input from both ears. The proportion of neurons responsive to ipsilateral stimulation (41%) was rather high. Mean response latency was 8.9 ms for contralateral stimulation.A tonotopic organization is lacking, but high-frequency neurons are less frequent in rostral SC.Abbreviations CF constant frequency component of echolocation call; - >CF frequencies above range of CF-component - FM frequency modulated component of echolocation call - <FM frequencies below range of FM-component - RF resting frequency of an individual bat - Rh.r. Rhinolophus rouxi - SC superior colliculus     

Working Memory for Low-Level Visual Features: Sensory Mechanisms for Detecting the Mismatch between the Current Orientations and Those Stored in Memory

The amplitudes of the P100 and N150 early components of evoked potentials in the visual cortex have been analyzed on 33 volunteers with normal vision during matching between the current orientation and that stored in memory. An increase in the P100 response in the occipital and parietal cortical areas was identified as an informative indicator of mismatch between the current and stored-in-memory orientations. This effect was not found for more complex stimuli, namely, spatial patterns. The N150 component demonstrated a similar effect, but in contrast to P100 it was not stimulus specific. Thus, in the first 100 ms, a signal of mismatch between the current and stored-in-memory orientations arises in the early visual areas that represents a mechanism for early implicit response to changes in the basic characteristics of the visual space.     

Inhibitory components in the neuronal response of the lateral suprasylvian cortical area in the cat

Inhibitory components in the response evoked by presentation of mobile visual stimuli in neurons belonging to the lateral suprasylvian area of the cerebral cortex were investigated in cats. It was demonstrated by comparing poststimulus histograms of neuronal response to movement in two opposite directions that the location of discharge centers within the receptive fields changed in relation to movement direction. No spatial area giving rise to the inhibitory component of response could be found in any of the neurons with monotone stationary structure of their receptive fields. Findings from experiments involving techniques of stimulating a test area of the receptive field separately indicated that inhibitory components of response in neurons of the lateral suprasylvian area with monotone organization of the receptive field could represent inhibitory after-response following the neuronal excitation produced by the visual stimulus traveling across this field.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 19, No. 3, pp. 299–308, May–June, 1987.     

Ensemble cortical responses to rival visual stimuli: Effect of monocular transient

Binocular rivalry is a fascinating perceptual phenomenon that has been characterized extensively at the psychophysical level. However, the underlying neural mechanism remains poorly understood. In particular, the role of the early visual pathway remains controversial. In this study, we used voltage-sensitive dye imaging to measure the spatiotemporal activity patterns in cat area 18 evoked by dichoptic orthogonal grating stimuli. We found that after several seconds of monocular stimulation with an oriented grating, an orthogonal stimulus to the other eye evoked a reversal of the cortical response pattern, which may contribute to flash suppression in perception. Furthermore, after several seconds of rival binocular stimulation with unequal contrasts, transient increase in the contrast of the weak stimulus evoked a long-lasting cortical response. This transient-triggered response could contribute to the perceptual switch during binocular rivalry. Together, these results point to a significant contribution of early visual cortex to transient-triggered switch in perceptual dominance.     

Perisaccadic Remapping and Rescaling of Visual Responses in Macaque Superior Colliculus

Visual neurons have spatial receptive fields that encode the positions of objects relative to the fovea. Because foveate animals execute frequent saccadic eye movements, this position information is constantly changing, even though the visual world is generally stationary. Interestingly, visual receptive fields in many brain regions have been found to exhibit changes in strength, size, or position around the time of each saccade, and these changes have often been suggested to be involved in the maintenance of perceptual stability. Crucial to the circuitry underlying perisaccadic changes in visual receptive fields is the superior colliculus (SC), a brainstem structure responsible for integrating visual and oculomotor signals. In this work we have studied the time-course of receptive field changes in the SC. We find that the distribution of the latencies of SC responses to stimuli placed outside the fixation receptive field is bimodal: The first mode is comprised of early responses that are temporally locked to the onset of the visual probe stimulus and stronger for probes placed closer to the classical receptive field. We suggest that such responses are therefore consistent with a perisaccadic rescaling, or enhancement, of weak visual responses within a fixed spatial receptive field. The second mode is more similar to the remapping that has been reported in the cortex, as responses are time-locked to saccade onset and stronger for stimuli placed in the postsaccadic receptive field location. We suggest that these two temporal phases of spatial updating may represent different sources of input to the SC.