Neurophysiological correlates of color vision: a model

The tree-receptor theory of human color vision accounts for color matching. A bottom-up, non-linear model combining cone signals in six types of cone-opponent cells in the lateral geniculate nucleus (LGN) of primates describes the phenomenological dimensions hue, color strength, and lightness/brightness. Hue shifts with light intensity (the Bezold-Brücke phenomenon), and saturation (the Abney effect) are also accounted for by the opponent model. At the threshold level, sensitivities of the more sensitive primate cells correspond well with human psychophysical thresholds. Conventional Fourier analysis serves well in dealing with the discrimination data, but here we want to take a look at non-linearity, i.e., the neural correlates to perception of color phenomena for small and large fields that span several decades of relative light intensity. We are particularly interested in the mathematical description of spectral opponency, receptive fields, the balance of excitation and inhibition when stimulus size changes, and retina-to-LGN thresholds...

Spatial sensitivity, responsivity, and surround suppression of LGN cell responses in the macaque.

Responses of cells in the lateral geniculate nucleus (LGN) of the macaque monkey have been measured for different sizes of chromatic and achromatic stimuli, with relative luminance spanning a range of 3-6 log units. Homogeneous illuminated test fields, centered on the receptive field, were used. Responses to these stimuli deviated from what is expected for the grating stimuli used to study the contrast-sensitive mechanisms in the visual pathway. For test fields smaller than the center of the receptive field, both the excitatory and the inhibitory cone-opponent components were present in the response, and the sensitivity to both components increased with the same factor when the test field increased in size (area summation). For test field areas extending into the suppressive surround of the extraclassical receptive field, the excitatory and the inhibitory cone opponents were both suppressed, again by the same factor. This suppression of the cell's responsiveness, as a function of test spot area, was described by a logarithmic function, and the spatial sensitivity of attenuation could therefore be described by a power function of radius. The logarithmic suppression was clear for parvocellular and koniocellular cells but was more prominent for magnocellular cells. The surround field suppression was also found for the prepotential inputs to LGN cells, indicating a retinal origin. The difference of Gaussian (DOG) model has been used successfully to describe the cells' contrast behavior for grating stimuli. However, this model fails to describe the constant excitatory/inhibitory response balance needed to obtain color (hue) stability for light stimuli of different sizes but with the same Commission Internationale de l'Eclairage (CIE) chromaticity and luminance factor. Neither the constant responsiveness found in the center of the receptive field nor the suppressive response in the surround can be described by the DOG model.

Neurophysiological correlates of color vision: a model

The tree-receptor theory of human color vision accounts for color matching. A bottom-up, non-linear model combining cone signals in six types of cone-opponent cells in the lateral geniculate nucleus (LGN) of primates describes the phenomenological dimensions hue, color strength, and lightness/brightness. Hue shifts with light intensity (the Bezold-Brücke phenomenon), and saturation (the Abney effect) are also accounted for by the opponent model. At the threshold level, sensitivities of the more sensitive primate cells correspond well with human psychophysical thresholds. Conventional Fourier analysis serves well in dealing with the discrimination data, but here we want to take a look at non-linearity, i.e., the neural correlates to perception of color phenomena for small and large fields that span several decades of relative light intensity. We are particularly interested in the mathematical description of spectral opponency, receptive fields, the balance of excitation and inhibition when stimulus size changes, and retina-to-LGN thresholds.(AU)

Visual signal processing in the macaque lateral geniculate nucleus.

Comparisons of S- or prepotential activity, thought to derive from a retinal ganglion cell afferent, with the activity of relay cells of the lateral geniculate nucleus (LGN) have sometimes implied a loss, or leak, of visual information. The idea of the "leaky" relay cell is reconsidered in the present analysis of prepotential firing and LGN responses of color-opponent cells of the macaque LGN to stimuli varying in size, relative luminance, and spectral distribution. Above a threshold prepotential spike frequency, called the signal transfer threshold (STT), there is a range of more than 2 log units of test field luminance that has a 1:1 relationship between prepotential- and LGN-cell firing rates. Consequently, above this threshold, the LGN cell response can be viewed as an extension of prepotential firing (a "nonleaky relay cell"). The STT level decreased when the size of the stimulus increased beyond the classical receptive field center, indicating that the LGN cell is influenced by factors other than the prepotential input. For opponent ON cells, both the excitatory and the inhibitory response decreased similarly when the test field size increased beyond the center of the receptive field. These findings have consequences for the modeling of LGN cell responses and transmission of visual information, particularly for small fields. For instance, for LGN ON cells, information in the prepotential intensity-response curve for firing rates below the STT is left to be discriminated by OFF cells. Consequently, for a given light adaptation, the STT improves the separation of the response range of retinal ganglion cells into "complementary" ON and OFF pathways.

Segregation of chromatic and luminance signals using a novel grating stimulus.

Segregation of chromatic and luminance signals in afferent pathways are investigated with a grating stimulus containing luminance and chromatic components of different spatial frequencies. Ganglion cell recordings were obtained from the retinae of macaques (Macaca fascicularis). Cell responses to the 'compound' gratings were compared to responses to standard chromatic and luminance gratings. Parvocellular (PC) pathway cell responses to compound and chromatic gratings were very similar, as were magnocellular (MC) cell responses to compound and luminance gratings. This was the case over a broad range of spatial and temporal frequencies and contrasts. In psychophysical experiments with human observers, discrimination between grating types was possible close to detection threshold. These results are consistent with chromatic and luminance structure in complex patterns being strictly localized in different afferent pathways. This novel stimulus may prove useful in identifying afferent inputs to cortical neurons.

Flicker VEPs reflecting multiple rod and cone pathways.

In an attempt to determine whether the relative contributions of magno-mediated and parvo-mediated inputs to the cortex are significantly altered in the transition from cone to rod vision, VEPs were recorded at different luminance levels (photopic to scotopic) for 2Hz square-wave, isochromatic flicker. The VEP mass response appears capable of reflecting major parvo-mediated contributions even at luminance levels for which responses from individual cells in the parvocellular pathway are reported to be weak. Our findings suggest that parvo-mediated responses are the dominant source of high-contrast isochromatic flicker VEPs at all light levels.

Surround-induced foveal afterimage pulsation: evidence for a long-range neural effect.

We demonstrate that an afterimage resulting from a strong foveal light flash can be made to pulsate by luminance modulation of a surrounding annulus as far as 8 deg away. Afterimage pulsation persists even if all artifacts due to pupil size, stray light and simultaneous contrast are ruled out. This suggests an origin by a long-range neural process acting from the remote surround. The effect is interpreted in terms of an adaptive gain control optimizing the responses of visual cells.

Visual evoked potentials for red-green gratings reversing at different temporal frequencies: asymmetries with respect to isoluminance.

Human visual evoked potentials (VEPs) were recorded for abrupt reversals of 2 cycles/deg (c/deg) square-wave gratings combining high red-green contrast with different levels of luminance contrast. Response characteristics--2nd harmonic amplitudes and peak latencies as a function of luminance contrast--were compared for four different reversal rates ranging from 6.25 Hz to 12.5 Hz. At every reversal frequency, the VEP amplitude and latency plots were nonsymmetrical with respect to isoluminance. The amplitude dropped to a minimum within a region of rapid phase change, always at a red-green luminance contrast for which the green color had the higher luminance, at about 40% or 50% Michelson luminance contrast. The rapid phase shift around this contrast suggested a sudden change in the relative impact of VEP generators with different latencies, possibly dominated by parvocellular or magnocellular input. The most prominent VEP waveform through most of the luminance contrast range, P110, is interpreted in terms of a parvo-mediated response that is attenuated with increasing reversal frequency. Contrast-dependent changes in the P110 amplitude appear to be responsible for the VEP asymmetries reported here.

Binocular contrast inhibition in subjects with age-related macular degeneration.

In subjects with normal vision, binocular contrast sensitivity is generally higher than monocular sensitivity, indicating summation of contrast in the two eyes. We have compared monocular and binocular contrast sensitivity and acuity for a group of 13 subjects with age-related macular degeneration (AMD). Relative to a normal control group, many of the AMD subjects showed reduced binocular contrast summation, and binocular inhibition was found for eight subjects for a narrow or an extended frequency band. A better monocular than binocular function may have practical implications for reading and orientation in AMD.