Lateral feedback from monophasic horizontal cells to cones in carp retina. II. A quantitative model

About half of the monophasic horizontal cells in carp retina receive input from both red- and green-sensitive cones. Since the horizontal cells feed back to cones, the color and feedback pathways result in wavelength- and intensity-dependent changes of the dynamics and of the receptive field amplitude profile of the horizontal cell responses. In this paper we present a quantitative model that describes adequately the color and spatial coding and the dynamics of the responses from monophasic horizontal cells in carp. Lateral feedback plays a distinct role in this model.     

Dynamics of turtle horizontal cell response

The small- and large-field (cone) horizontal cells produce similar dynamic responses to a stimulus whose mean luminance is modulated by a white-noise signal. Nonlinear components increase with an increase in the mean luminance and may produce a mean square error (MSE) of up to 15%. Increases in the mean luminance of the field stimulus bring about three major changes: the incremental sensitivity defined by the amplitude of the kernels decreases in a Weber-Fechner fashion; the waveforms of the kernels are transformed from monophasic (integrating) to biphasic (differentiating); the peak response time of the kernels becomes shorter and the cells respond to much higher-frequency inputs. The dynamics of the horizontal cell response also depend on the area of the retina stimulated. Smaller spots of light produce monophasic kernels of a longer peak response time. The presence of a steady background produces three major changes in the spot kernels: the kernel's amplitude becomes larger (incremental sensitivity increases); the peak response times become shorter; the waveform of the kernels changes in a fashion similar to that observed with an increase in the mean luminance of the field stimulus. A similar enhancement in the incremental sensitivity by a steady background has also been observed in catfish, which shows that this phenomenon is a common feature of the horizontal cells in the lower vertebrate retina.     

Effects of stimulus size and spatial organization on pigeons’ conditional same-different discrimination

In two experiments, we explored the effects of varying the size and the spatial organization of the stimuli in multi-item arrays on pigeons’ same-different discrimination behavior. The birds had previously learned to discriminate a simultaneously presented array of 16 identical (Same) visual items from an array of 16 nonidentical (Different) visual items, when the correct choice was conditional on the presence of another cue: the color of the background (Castro et al., in press). In Experiment 1, we trained pigeons with 7-item arrays and then tested them with arrays containing the same item, but in a variety of sizes. In Experiment 2, we tested the birds with the items grouped in novel locations: the top, the bottom, the left, or the right portions of the display area, which generated different vertical and horizontal alignments. Accuracy scores revealed virtually perfect stimulus generalization across various item sizes and spatial organizations. Reaction times revealed that the birds perceived different sizes of a single icon as the same stimulus (Experiment 1) and that the birds processed vertical arrangements faster than horizontal arrangements (Experiment 2). These results suggest that the pigeons noticed both physical and spatial changes in the stimuli (as shown by their reaction times), but that these changes did not disrupt the birds’ discriminating the sameness or differentness of the multi-item arrays (as shown by their accuracy scores).     

Modeling of the vertebrate visual system. II. Application to the turtle cone retina

The model of the vertebrate cone retina was adapted to the turtle retina with its red cone- and L-channel-dominances. The model consists of an ordering of four spatial organizations of unit hexagons, weighted inputs for all cones in the receptive fields, and linear polarization factors based on data from literature on turtle retina. Data generated by the model for spatial and chromatic patterns of receptive fields, intensity-response curves, dynamic ranges for cones, horizontal and bipolar cells proved remarkably consistent with literature. The model also generates observed phenomena such as near-field enhancement of cones due to stray light effects and electrical coupling of like-cones and far-field decrease in responses due to negative feedback from L-type horizontal cells to cones. Annular stimuli were shown to be more effective than spot stimuli for horizontal cells. The formal approach of the model demonstrates factors which play roles in various observed phenomena and all aspects of model can be displayed and tested both qualitatively and quantitatively.     

Gender-related differences in EEG coherence in Stroop task

Gender-related differences in EEE coherence were studied in young male and female university students during performance of Stroop task, in which color and word could be congruent or incongruent and spatially integrated or separated. The most clear-cut gender-related differences in the EEG coherence were revealed in the alpha2 and beta frequency bands. In the alpha2 band, gender-related differences in the interhemispheric coherence were associated with spatial characteristics of stimuli: left or right presentation and features of mutual localization of relevant and irrelevant stimuli. These differences were observed when information was addressed to the left hemisphere. Gender-related differences associated with spatial organization of stimuli were also observed for intrahemispheric beta1 coherence. Under conditions of the spatial separation of relevant and irrelevant stimuli, only females demonstrated enhancement of coherence in response to the right-side stimulus presentation as compared to the left-side presentation. It was shown that, during the differentiation of the semantic meaning of verbal stimuli, gender-related differences were caused by the features of integration of beta2 oscillators in the posterior cortical regions. The results are indicative of qualitative gender-related differences in organization of both frontoparietal and lateral attention systems in actualization of selective processes.     

The inferior temporal cortex: Architecture, computation, and representation

Neurons in the inferior temporal cortex (IT), an area crucially involved in visual object recognition in monkeys, show the visual response properties and anatomical/chemical nature which are distinct from those in the cortical areas that feed visual inputs to the IT. Earlier physiological studies showed that IT neurons have large receptive fields covering the center and contralateral (often bilateral) visual fields, stimulus selectivity for images of complex objects or shapes, and translation invariance of the stimulus selectivity. Recent studies have revealed new aspects of their properties such as invariant selectivity for shapes despite drastic changes in various physical attributes of stimuli, latent excitatory inputs masked by stimulus-specific GABAergic inhibition, selectivity for binocular disparity and 3-dimensional surface structures, profound effects of learning on the stimulus selectivity, and columnar clustering of neurons with similarstimulus selectivity for shapes and other object features. Another line of research using histological techniques have revealed that pyramidal neurons in the IT are larger in the size of dendritic arbors, in the number of dendritic branches and spines, and in the size and distribution of horizontal axonal arbors than those in the earlier areas, allowing them to integrate a larger population of afferents and process more diverse inputs. The concentration of several neurochemicals including those related to synaptic transmission or plasticity changes systematically towards the IT along the occipitotemporal pathway. Many of the characteristics of IT neurons parallel or explain certain aspects of visual object perception, although the behavioral relevance has yet to be addressed experimentally.     

Spatial summation of heat pain in males and females

Sex differences in pain sensitivity have been found to vary between considerable and negligible. It has appeared that the pain stimulation method is critical in this context. It was assumed this might be due to the different degrees of spatial summation associated with the different pain stimulus modalities. Hence, sex differences were investigated in spatial summation of heat pain in 20 healthy women and 20 healthy men of similar age. Pain thresholds were assessed by a tracking procedure and responses to supra-threshold pain stimulation by numerical ratings. Heat stimuli were administered by a thermode with contact areas of 1, 3, 6 and 10 cm2. Pain thresholds were significantly higher with smaller areas stimulated than with larger ones. No significant effect of area was found for the ratings of the supra-threshold stimuli, the intensities of which were tailored to the individual pain threshold. Consequently, spatial summation of heat pain appeared to result mainly in a shift of the pain threshold on the ordinate and not a change of slope of the stimulus-response function in the pain range. In neither of the two pain parameters were there any sex differences. Therefore, the present study demonstrated that sex differences in spatial summation of heat pain are unlikely.     

A linear chromatic mechanism drives the pupillary response.

Previous studies have shown that a chromatic mechanism can drive pupil responses. The aim of this research was to clarify whether a linear or nonlinear chromatic mechanism drives pupillary responses by using test stimuli of various colours that are defined in cone contrast space. The pupil and accommodation responses evoked by these test stimuli were continuously and simultaneously objectively measured by photorefraction. The results with isochromatic and isoluminant stimuli showed that the accommodative level remained approximately constant (< 0.25 D change in mean level) even when the concurrent pupillary response was large (ca. 0.30 mm). The pupillary response to an isoluminant grating was sustained, delayed (by ca. 60 ms) and larger in amplitude than that for a isochromatic uniform stimulus, which supports previous work suggesting that the chromatic mechanism contributes to the pupillary response. In a second experiment, selected chromatic test gratings were used and isoresponse contours in cone contrast space were obtained. The results showed that the isoresponse contour in cone contrast space is well described (r(2) = 0.99) by a straight line with a positive slope. The results indicate that a /L - M/ linear chromatic mechanism, whereby a signal from the long wavelength cone is subtracted from that of the middle wavelength cone and vice versa, drives pupillary responses.     

Dim light adaptation attenuates acute melatonin suppression in humans

Abstract Studies in rodents with retinal degeneration indicated that neither the rod nor the cone photoreceptors obligatorily participate in circadian responses to light, including melatonin suppression and photoperiodic response. Yet there is a residual phase-shifting response in melanopsin knockout mice, which suggests an alternate or redundant means for light input to the SCN of the hypothalamus. The findings of Aggelopoulos and Meissl suggest a complex, dynamic interrelationship between the classic visual photoreceptors and SCN cell sensitivity to light stimuli, relative to various adaptive lighting conditions. These studies raised the possibility that the phototransductive physiology of the retinohypothalamic tract in humans might be modulated by the visual rod and cone photoreceptors. The aim of the following two-part study was to test the hypothesis that dim light adaptation will dampen the subsequent suppression of melatonin by monochromatic light in healthy human subjects. Each experiment included 5 female and 3 male human subjects between the ages of 18 and 30 years, with normal color vision. Dim white light and darkness adaptation exposures occurred between midnight and 0200 h, and a full-field 460-nm light exposure subsequently occurred between 0200 and 0330-h for each adaptation condition, at 2 different intensities. Plasma samples were drawn following the 2-h adaptation, as well as after the 460-nm monochromatic light exposure, and melatonin was measured by radioimmunoassay. Comparison of melatonin suppression responses to monochromatic light in both studies revealed a loss of significant suppression after dim white light adaptation compared with dark adaptation (p < 0.04 and p < 0.01). These findings indicate that the activity of the novel circadian photoreceptive system in humans is subject to subthreshold modulation of its sensitivity to subsequent monochromatic light exposure, varying with the conditions of light adaptation prior to exposure.     

The Spatial Properties of L- and M-Cone Inputs to Electroretinograms That Reflect Different Types of Post-Receptoral Processing

We studied the spatial arrangement of L- and M-cone driven electroretinograms (ERGs) reflecting the activity of magno- and parvocellular pathways. L- and M-cone isolating sine wave stimuli were created with a four primary LED stimulator using triple silent substitution paradigms. Temporal frequencies were 8 and 12 Hz, to reflect cone opponent activity, and 30, 36 and 48 Hz to reflect luminance activity. The responses were measured for full-field stimuli and for different circular and annular stimuli. The ERG data confirm the presence of two different mechanisms at intermediate and high temporal frequencies. The responses measured at high temporal frequencies strongly depended upon spatial stimulus configuration. In the full-field conditions, the L-cone driven responses were substantially larger than the full-field M-cone driven responses and also than the L-cone driven responses with smaller stimuli. The M-cone driven responses at full-field and with 70° diameter stimuli displayed similar amplitudes. The L- and M-cone driven responses measured at 8 and 12 Hz were of similar amplitude and approximately in counter-phase. The amplitudes were constant for most stimulus configurations. The results indicate that, when the ERG reflects luminance activity, it is positively correlated with stimulus size. Beyond 35° retinal eccentricity, the retina mainly contains L-cones. Small stimuli are sufficient to obtain maximal ERGs at low temporal frequencies where the ERGs are also sensitive to cone-opponent processing.     

Spatial interactions and stimulus-response relations of unit responses evoked by somato-sensory stimuli in the feline caudal medullary reticular formation

Responses of neurons in the bulbar reticular area to separate and simultaneous stimulation of the forelimbs were recorded extracellularly in chloralose-anaesthetized cats. On increasing the stimulus intensity the number of spikes per response increased while the initial latency and interspike intervals decreased in accordance with the functional property of the neuron. Responses evoked by simultaneous stimulation displayed more spikes and a shorter latency than those evoked by separate stimuli of corresponding intensities. The differences in the responses evoked simultaneously and the sums of responses evoked separately showed characteristic distributions as a function of the latter. Three types of distribution were distinguished. The results indicate that stimulus-response relations play a determining role in the mechanism of spatial integration.     

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.     

Attenuation of vibrotactile spatial summation.

Masked and quiet thresholds at several frequencies of vibratory stimuli were measured as a function of contactor area. The test site was the left index finger; the masking site was the left little finger. The quiet threshold data were consistent with previous investigations: Low-frequency stimuli showed no spatial summation, whereas high-frequency stimuli did. In the presence of a masker, spatial summation was reduced or eliminated for high-frequency stimuli, i.e., the masked threshold was, under some conditions, independent of contactor area. Low-frequency stimuli continued to show no spatial summation in the presence of a masker. The attenuation of spatial summation appears to be a direct function of the intensity of the masking stimulus. Additional measurements with the left thenar eminence as the test site showed that spatial summation could be attenuated by a masker placed on a contralateral body site. The implications of the results for quantifying the effectiveness of a masking stimulus, for the duplex mechanoreceptor hypothesis, and for the nature of spatial summation on the skin are discussed.     

Neural mechanisms for color perception in the primary visual cortex

New neurophysiological results show the existence of multiple transformations of color signals in the primary visual cortex (V1) in macaque monkey. These different color mechanisms may contribute separately to the perception of color boundaries and colored regions. Many cells in V1 respond to color and to black-white (luminance) patterns. These neurons are spatially selective and could provide signals about boundaries between differently colored regions. Other V1 neurons that prefer color over luminance respond without much spatial selectivity to colored stimuli, and could be the neural basis for the response to local color modulation within a region. How these different types of color cells combine inputs from cone photoreceptors is what gives them their different spatial selectivities for color.     

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.     

Generation of knock-in mice carrying third cones with spectral sensitivity different from S and L cones

Red-green color vision in primates is unique in the sense that it is mediated by two photoreceptor cells that are indistinguishable in all aspects except for their visual pigments. In order to generate an animal model for investigation of the interaction between red-green inputs at the molecular level, we applied knock-in technology and X-chromosome inactivation machinery to make a mouse model with cone cells possessing visual pigments with different spectral sensitivities. We introduced a S308A point mutation into the Green opsin gene allele on the X-chromosome. This manipulation generated a 24 nm red-shift of absorption maximum in the cone pigment with negligible functional differences in other molecular properties. Amplitudes of responses in ERG and ganglion cell recordings of homozygotes were similar to those of wild-types, although the spectral sensitivities differed. Heterozygotes showed variable spectral sensitivities of ganglion cell responses due to the different integration of the native and the S308A cone inputs on the dendritic fields. In situ hybridization experiments showed that cone cells with respective pigments formed patch-like clusters of specific L cone-types, approximately 30 mum in diameter, which were randomly distributed in the dorsal region of the retinas. Since the patch-like clustering was arranged by X-inactivation, such clustering could be present in the peripheral retinas of New World monkeys with polymorphic L pigments, indicating that our mice would be a suitable model to study evolution of the mammalian color vision system.     

Simulated fovea of the human retina: psychophysical data confirming the model's ability to accurately predict resolution

A series of psychophysical tests were designed to determine whether a computer simulation of the human retina could accurately predict the geometry of various stimuli that were optimally resolved for human foveal vision. Stimuli were used that were of the order of the grain of the cone mosaic, i.e., of the order of 2 × 2. In the first set of experiments, resolution was tested using a two-bar stimulus. In one experimental series the gap between the two bars was varied, and in a second series the gap was kept constant and the width of the bars varied. In a second set of experiments, various block letters and a number of series for each letter were used; in each experimental series a single parameter was systematically varied. The same stimuli were also used as inputs for the computer simulation. When proper controls were used, the psychophysical data and computer simulation gave remarkably comparable results. Care was taken to differentiate between simple detection of a pattern, and resolution, which involved proper identification of the image.     

Mechanisms of cardiac cell excitation with premature monophasic and biphasic field stimuli: a model study.

The mechanisms by which extracellular electric field stimuli induce the (re)excitation of cardiac cells in various stages of refractoriness are still not well understood. We modeled the interactions between an isolated cardiac cell and imposed extracellular electric fields to determine the mechanisms by which relatively low-strength uniform monophasic and biphasic field stimuli induce premature reexcitations. An idealized ventricular cell was simulated with 11 subcellular membrane patches, each of which obeyed Luo-Rudy (phase 1) kinetics. Implementing a standard S1-S2 pulse protocol, strength-interval maps of the cellular excitatory responses were generated for rectangular monophasic and symmetric biphasic field stimuli of 2, 5, 10, and 20 ms total duration. In contrast to previously documented current injection studies, our results demonstrate that a cardiac cell exhibits a significantly nonmonotonic excitatory response to premature monophasic and, to a much lesser degree, biphasic field stimuli. Furthermore, for monophasic stimuli at low field strengths, the cell is exquisitely sensitive to the timing of the shock, demonstrating a classic all-or-none depolarizing response. However, at higher field strengths this all-or-none sensitivity reverts to a more gradual transition of excitatory responses with respect to stimulus prematurity. In contrast, biphasic stimuli produce such graded responses at all suprathreshold stimulus strengths. Similar behaviors are demonstrated at all S2 stimulus durations tested. The generation of depolarizing (sodium) currents is triggered by one or more of the sharp field gradient changes produced at the stimulus edges-i.e., make, break, and transphasic (for biphasic stimuli)-with the magnitude of these edge-induced current contributions dependent on both the prematurity and the strength of the applied field. In all cases, however, depolarizing current arises from the partial removal of sodium inactivation from at least part of the cell, because of either the natural process of repolarization or a localized acceleration of this process by the impressed field.     

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 influence of color stimuli on visually controlled behavior inAglais urticae L. andPararge aegeria L. (Lepidoptera)

Summary In spontaneous-choice experiments on the butterfliesAglais urticae L. (Nymphalidae) andPararge aegeria L. (Satyridae) the spectral effectiveness and spectral sensitivity of various behaviors were investigated and compared.Pilot experiments with colored PVC films showed indications of an intensity dependence of the feeding reaction inP. aegeria. Moreover, they revealed a color discrimination independent of this intensity discrimination:P. aegeria distinguishes red from grey shades as well as from black and white (Fig. 3).According to subsequent spontaneous-choice experiments using monochromatic light stimuli, the various visually controlled functional categories of behavior can be assigned to the following spectral regions: 1. The open-space reaction corresponds to the UV and violet region, ca. 320–420 nm, inP. aegeria (Figs. 4, 7). 2. The feeding reaction corresponds to the blue region, ca. 420–500 nm, inA. urticae (Fig. 1) andP. aegeria (Fig. 4), and the yellow region, ca. 550–590 nm, inA. urticae (Fig. 1) and the orange-red region, ca. 570–670 nm, inP. aegeria (Fig. 4).In these experiments with monochromatic light stimuli the intensity dependence of the reactions is also obvious (Figs. 2, 5, 6).The open-space reaction is elicited inP. aegeria by white light dependent on its UV content (Fig. 8). This is also valid for the feeding reaction inP. aegeria (Fig. 5b). To elicit this reaction it was necessary to offer light stimuli simultaneously with the odour stimulus of honey water. As the latter was of the same quality in combination with all light stimuli the results can be attributed definitely to the different effectiveness of the various light stimuli.Pure wavelength-specific behavior can be ruled out inA. urticae andP. aegeria. Wavelength-specific behavior and color vision are probably present simultaneously.Abbreviation RNQ relative number of quanta Supported by the Deutsche Forschungsgemeinschaft Ko 445/5-3