Comprehensive Physiology Wiley Online Library

Neural Mechanisms of Color Vision

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 Photopigments
1.1 Spectral Absorbance Characteristics
1.2 Univariance of Photopigment Response
2 Receptors
2.1 Prevalence and Retinal Distribution of Different Cone Types
2.2 Cone Responses
2.3 Horizontal Cell Feedback
3 Outer Plexiform Layer
3.1 Horizontal Cells
3.2 Bipolar Cells
4 Ganglion Cells and Lateral Geniculate Cells
4.1 Spectral Properties
4.2 Spatial Properties
5 Cortical Cells
5.1 Area 17
5.2 Cortical Processing of Color Information: Beyond Striate Cortex
6 Physiology and Color Vision
6.1 Relation of Physiology to Color Vision
7 Problems and Unknowns
7.1 Black/White System
7.2 Red/Green vs. Yellow/Blue System
Figure 1. Figure 1.

Diagrammatic representation of luminance profiles of two gratings modulated around some mean level, and of the responses of each of the two long‐wavelength cone populations (P536 and P565) to these gratings. The abscissa is distance along the retina. Because of their broad sensitivities, each cone type responds to each of the gratings, but to differing extents.

Figure 2. Figure 2.

The red and green gratings shown in Figure are combined in each of two phase angles. A: gratings combined out of phase to form a pure‐color grating; luminance is constant throughout, and the only variation is in dominant wavelength. B: gratings combined in phase to form a pure‐luminance grating; wavelength is constant throughout, and the only variation is in luminance. C‐F: results of adding and subtracting receptor responses (shown in Fig. ) to pure‐color and pure‐luminance patterns. If receptor outputs are summed, there is no net response to the color grating, C, but a large response to the luminance grating, D. Differencing the receptor outputs yields no output for the luminance grating, F, but a small output for the color grating, E. Thus it can be seen that luminance information is carried by the receptor sums and color information by the receptor differences. Note also that the color response, E, is considerably smaller than the luminance response, D, even though the wavelengths chosen in Figure were such as to produce about the maximum differentiation between the P536 and the P565 cones. Any smaller wavelength difference would leave the luminance response the same size but would decrease the color response.

Figure 3. Figure 3.

Spectral absorbance curves for primate cone pigments. Each curve is plotted as percentage of maximal absorbance. Curve and data points for P440 generated from a Dartnall nomogram. Curves and data points for the two long‐wavelength photopigments (P536 and P565) from Bowmaker et al. .

Figure 4. Figure 4.

Responses of a P520 turtle cone to stimulation from an annulus of 618 nm (top) and from an annulus of 550 nm (bottom). Stimulus duration is indicated by the marker bar at the top. Response is the change from resting potential of the photoreceptor

Adapted from Fuortes et al.
Figure 5. Figure 5.

Example of S potentials recorded from fish retina. Potentials were recorded from a small micropipette electrode inserted into a horizontal cell. Each vertical trace shows magnitude of change from resting membrane potential when the retina was in darkness. Each trace is the response to a single test wavelength. Note that these graded potentials are in the hyperpolarizing direction (upward) to long wavelengths and depolarizing direction (downward signals) to short wavelengths

From Svaetichin and MacNichol
Figure 6. Figure 6.

Horizontal cell responses in turtle retina. Top: spectral response curves for C‐cells (B/Y, blue/yellow, and R/G, red/green) and L‐cells. Bottom: wiring diagram illustrating the kinds of interactions believed necessary to account for responses of horizontal cells. Vertical ovals represent classes of cones found in this retina. Pluses and minuses (+, −) indicate transmission with and without inversion of polarity. Basic connections responsible for the main properties of the cells are shown by solid lines; dashed lines represent the modifying interactions

Adapted from Fuortes and Simon
Figure 7. Figure 7.

Responses of a + G/‐R spectrally opponent LGN cell to light of various wavelengths. A line is drawn through superimposed records at left to indicate time of light onset; the line at right marks light offset (1‐s duration). Numbers at left: wavelength (in nm) of each light stimulus

From De Valois, Abramov, and Jacobs
Figure 8. Figure 8.

Responses of a + G/‐R lateral geniculate nucleus cell to shifts back and forth between two lights of 620 and 593 nm. Middle record: the two wavelengths are of equal luminance. Top record: 593‐nm light had the same luminance as in the middle record; 620‐nm light was 0.5 log units brighter than in middle record. Bottom record: 593‐nm light had the same luminance as in the middle record; 620‐nm light was 0.5 log units dimmer than in middle record. It can be seen that the cell fires to the 593‐nm light and inhibits to the 620‐nm light despite wide variations in their relative luminances

From De Valois et al.
Figure 9. Figure 9.

Plots of average firing rates of a large sample of cells of each of the six lateral geniculate nucleus cell types. Stimuli were 0.7 log unit incremental flashes of monochromatic light, 1 s in duration, covering the entire receptive field. Horizontal dashed lines, average maintained discharge rates of these cells, as measured in the 1‐s interval before each stimulus. Top and middle panels: results from spectrally opponent cells; bottom panels: results from spectrally nonopponent cells. Note that these are responses to increments of light; in response to decremental flashes of any wavelength the +Bl/‐Wh cells would fire and the +Wh/‐Bl cells would inhibit

Adapted from De Valois, Abramov, and Jacobs
Figure 10. Figure 10.

Chromatic adaptation experiment with a +G/‐R cell. Bottom left, responses of the cell under neutral (white light) adaptation conditions: to 1‐s flashes of light covering the entire receptive field, the cell fires to short wavelengths (maximally at about 500 nm) and inhibits to long wavelengths. Data plotted as open circles in graph at top. With a 510‐nm bleach (bottom center and closed circles at top) inhibition increases and the point of maximum inhibition is shifted to shorter wavelengths. With 660‐nm bleach (bottom right and half‐closed circles at top) amount of excitation is greater and maximum is shifted toward longer wavelengths. It is clear that responses under neutral adaptation conditions are the result of algebraic addition of separate excitatory and inhibitory components, from P536 and P565 cones, respectively

From De Valois
Figure 11. Figure 11.

Threshold (in log number of quanta per stimulus) sensitivity curves for nonopponent ganglion cells in monkey in the presence of intense red (open symbols) and intense blue (closed symbols) adaptation

From Gouras
Figure 12. Figure 12.

Map of cone input and maps of receptive field (RF) for a spectrally opponent cell that also shows spatial opponency. Left: cone inputs from the two long‐wavelength cones feed into this cell in opposite directions, and into the center and surround, respectively. Such a cell responds to both luminance and color changes but with entirely different receptive field maps for the two types of stimuli. Top right: when mapped with a luminance change, the cell shows a spatial antagonism between center and surround. Bottom right: mapped with a color change, the cell shows center‐surround synergism, firing everywhere in the field to a shift toward green, and inhibiting everywhere to a shift toward red

From De Valois and De Valois
Figure 13. Figure 13.

Averaged responses of 23 spectrally opponent lateral geniculate nucleus cells, including cells from each of the opponent cell classes to luminance and color lines of various widths, centered on the receptive field. Luminance responses are those given to a log unit change of 0.7 in luminance of a white light, either an increment or a decrement depending on type of cell. Color responses are those given to an equal‐luminance change in color, from a wavelength that inhibited the cell to one that excited it

From De Valois et al.
Figure 14. Figure 14.

Contrast sensitivity for a simple cortical cell, responding to red/green pure‐color gratings. Note that the cell shows sharp low and high spatial frequency attenuation. It is much more narrowly tuned than the overall monkey behavioral contrast sensitivity function, also shown.

Figure 15. Figure 15.

Responses of a macaque striate cortex cell to white, small filled circles, black, large filled hexagons, and red‐on‐green, small open circles, pure‐color bars moved across the cells' receptive field at various orientations. In this polar plot, number of spikes to bars of each orientation are shown by distance out and along that radius.

R. L. De Valois, E. W. Yund, and N. K. Hepler, unpublished data
Figure 16. Figure 16.

Polar plots of combined orientation and spatial‐frequency selectivity of typical lateral geniculate nucleus (LGN) cells (A) and cortical cells (B). In these figures spatial frequency increases from center outward, and various orientations are the radii around the circle. It can be seen that, in response to gratings of various spatial frequencies and orientations, LGN cells respond to a disk‐shaped region in this orientation frequency space, since they respond to any orientation and to all spatial frequencies up to some cutoff point (chosen as that frequency at which response drops to half‐maximum). The cortical cells, on the other hand, have both orientation and frequency selectivity and respond to stimuli only within a small region in this orientation frequency space. Two neighboring cortical cells are shown, each with the same orientation selectivity but with different spatial frequency selectivities. The limit of vision is the highest spatial frequency that the animal can discriminate under these conditions

From De Valois et al.
Figure 17. Figure 17.

Contrast sensitivities of a human subject for luminance‐modulated (Δ) and red/green color‐modulated (•) gratings of the kind diagramed in Figure . Note that humans are sensitive over different spatial frequency bands with these two types of stimuli

Adapted from Granger and Heurtley
Figure 18. Figure 18.

Representation of ability of humans to discriminate chromatic stimuli of various spatial and temporal frequency ranges. At low spatial and temporal frequencies trichromatic color vision is present; at higher spatial and temporal frequencies, the yellow/blue system drops out and only dichromatic vision is present; at the highest spatial and temporal frequencies the red/green system drops out as well, and only monochromatic color vision is present.

Figure 19. Figure 19.

Response of a spectrally opponent lateral geniculate nucleus cell to pure‐luminance (◯) and pure‐color (▴) gratings of various spatial frequencies, which were drifted across the cells' receptive field. The cell responds to the two types of stimulus patterns over different spatial frequency ranges.

Based on unpublished data from H. von Blanckensee


Figure 1.

Diagrammatic representation of luminance profiles of two gratings modulated around some mean level, and of the responses of each of the two long‐wavelength cone populations (P536 and P565) to these gratings. The abscissa is distance along the retina. Because of their broad sensitivities, each cone type responds to each of the gratings, but to differing extents.



Figure 2.

The red and green gratings shown in Figure are combined in each of two phase angles. A: gratings combined out of phase to form a pure‐color grating; luminance is constant throughout, and the only variation is in dominant wavelength. B: gratings combined in phase to form a pure‐luminance grating; wavelength is constant throughout, and the only variation is in luminance. C‐F: results of adding and subtracting receptor responses (shown in Fig. ) to pure‐color and pure‐luminance patterns. If receptor outputs are summed, there is no net response to the color grating, C, but a large response to the luminance grating, D. Differencing the receptor outputs yields no output for the luminance grating, F, but a small output for the color grating, E. Thus it can be seen that luminance information is carried by the receptor sums and color information by the receptor differences. Note also that the color response, E, is considerably smaller than the luminance response, D, even though the wavelengths chosen in Figure were such as to produce about the maximum differentiation between the P536 and the P565 cones. Any smaller wavelength difference would leave the luminance response the same size but would decrease the color response.



Figure 3.

Spectral absorbance curves for primate cone pigments. Each curve is plotted as percentage of maximal absorbance. Curve and data points for P440 generated from a Dartnall nomogram. Curves and data points for the two long‐wavelength photopigments (P536 and P565) from Bowmaker et al. .



Figure 4.

Responses of a P520 turtle cone to stimulation from an annulus of 618 nm (top) and from an annulus of 550 nm (bottom). Stimulus duration is indicated by the marker bar at the top. Response is the change from resting potential of the photoreceptor

Adapted from Fuortes et al.


Figure 5.

Example of S potentials recorded from fish retina. Potentials were recorded from a small micropipette electrode inserted into a horizontal cell. Each vertical trace shows magnitude of change from resting membrane potential when the retina was in darkness. Each trace is the response to a single test wavelength. Note that these graded potentials are in the hyperpolarizing direction (upward) to long wavelengths and depolarizing direction (downward signals) to short wavelengths

From Svaetichin and MacNichol


Figure 6.

Horizontal cell responses in turtle retina. Top: spectral response curves for C‐cells (B/Y, blue/yellow, and R/G, red/green) and L‐cells. Bottom: wiring diagram illustrating the kinds of interactions believed necessary to account for responses of horizontal cells. Vertical ovals represent classes of cones found in this retina. Pluses and minuses (+, −) indicate transmission with and without inversion of polarity. Basic connections responsible for the main properties of the cells are shown by solid lines; dashed lines represent the modifying interactions

Adapted from Fuortes and Simon


Figure 7.

Responses of a + G/‐R spectrally opponent LGN cell to light of various wavelengths. A line is drawn through superimposed records at left to indicate time of light onset; the line at right marks light offset (1‐s duration). Numbers at left: wavelength (in nm) of each light stimulus

From De Valois, Abramov, and Jacobs


Figure 8.

Responses of a + G/‐R lateral geniculate nucleus cell to shifts back and forth between two lights of 620 and 593 nm. Middle record: the two wavelengths are of equal luminance. Top record: 593‐nm light had the same luminance as in the middle record; 620‐nm light was 0.5 log units brighter than in middle record. Bottom record: 593‐nm light had the same luminance as in the middle record; 620‐nm light was 0.5 log units dimmer than in middle record. It can be seen that the cell fires to the 593‐nm light and inhibits to the 620‐nm light despite wide variations in their relative luminances

From De Valois et al.


Figure 9.

Plots of average firing rates of a large sample of cells of each of the six lateral geniculate nucleus cell types. Stimuli were 0.7 log unit incremental flashes of monochromatic light, 1 s in duration, covering the entire receptive field. Horizontal dashed lines, average maintained discharge rates of these cells, as measured in the 1‐s interval before each stimulus. Top and middle panels: results from spectrally opponent cells; bottom panels: results from spectrally nonopponent cells. Note that these are responses to increments of light; in response to decremental flashes of any wavelength the +Bl/‐Wh cells would fire and the +Wh/‐Bl cells would inhibit

Adapted from De Valois, Abramov, and Jacobs


Figure 10.

Chromatic adaptation experiment with a +G/‐R cell. Bottom left, responses of the cell under neutral (white light) adaptation conditions: to 1‐s flashes of light covering the entire receptive field, the cell fires to short wavelengths (maximally at about 500 nm) and inhibits to long wavelengths. Data plotted as open circles in graph at top. With a 510‐nm bleach (bottom center and closed circles at top) inhibition increases and the point of maximum inhibition is shifted to shorter wavelengths. With 660‐nm bleach (bottom right and half‐closed circles at top) amount of excitation is greater and maximum is shifted toward longer wavelengths. It is clear that responses under neutral adaptation conditions are the result of algebraic addition of separate excitatory and inhibitory components, from P536 and P565 cones, respectively

From De Valois


Figure 11.

Threshold (in log number of quanta per stimulus) sensitivity curves for nonopponent ganglion cells in monkey in the presence of intense red (open symbols) and intense blue (closed symbols) adaptation

From Gouras


Figure 12.

Map of cone input and maps of receptive field (RF) for a spectrally opponent cell that also shows spatial opponency. Left: cone inputs from the two long‐wavelength cones feed into this cell in opposite directions, and into the center and surround, respectively. Such a cell responds to both luminance and color changes but with entirely different receptive field maps for the two types of stimuli. Top right: when mapped with a luminance change, the cell shows a spatial antagonism between center and surround. Bottom right: mapped with a color change, the cell shows center‐surround synergism, firing everywhere in the field to a shift toward green, and inhibiting everywhere to a shift toward red

From De Valois and De Valois


Figure 13.

Averaged responses of 23 spectrally opponent lateral geniculate nucleus cells, including cells from each of the opponent cell classes to luminance and color lines of various widths, centered on the receptive field. Luminance responses are those given to a log unit change of 0.7 in luminance of a white light, either an increment or a decrement depending on type of cell. Color responses are those given to an equal‐luminance change in color, from a wavelength that inhibited the cell to one that excited it

From De Valois et al.


Figure 14.

Contrast sensitivity for a simple cortical cell, responding to red/green pure‐color gratings. Note that the cell shows sharp low and high spatial frequency attenuation. It is much more narrowly tuned than the overall monkey behavioral contrast sensitivity function, also shown.



Figure 15.

Responses of a macaque striate cortex cell to white, small filled circles, black, large filled hexagons, and red‐on‐green, small open circles, pure‐color bars moved across the cells' receptive field at various orientations. In this polar plot, number of spikes to bars of each orientation are shown by distance out and along that radius.

R. L. De Valois, E. W. Yund, and N. K. Hepler, unpublished data


Figure 16.

Polar plots of combined orientation and spatial‐frequency selectivity of typical lateral geniculate nucleus (LGN) cells (A) and cortical cells (B). In these figures spatial frequency increases from center outward, and various orientations are the radii around the circle. It can be seen that, in response to gratings of various spatial frequencies and orientations, LGN cells respond to a disk‐shaped region in this orientation frequency space, since they respond to any orientation and to all spatial frequencies up to some cutoff point (chosen as that frequency at which response drops to half‐maximum). The cortical cells, on the other hand, have both orientation and frequency selectivity and respond to stimuli only within a small region in this orientation frequency space. Two neighboring cortical cells are shown, each with the same orientation selectivity but with different spatial frequency selectivities. The limit of vision is the highest spatial frequency that the animal can discriminate under these conditions

From De Valois et al.


Figure 17.

Contrast sensitivities of a human subject for luminance‐modulated (Δ) and red/green color‐modulated (•) gratings of the kind diagramed in Figure . Note that humans are sensitive over different spatial frequency bands with these two types of stimuli

Adapted from Granger and Heurtley


Figure 18.

Representation of ability of humans to discriminate chromatic stimuli of various spatial and temporal frequency ranges. At low spatial and temporal frequencies trichromatic color vision is present; at higher spatial and temporal frequencies, the yellow/blue system drops out and only dichromatic vision is present; at the highest spatial and temporal frequencies the red/green system drops out as well, and only monochromatic color vision is present.



Figure 19.

Response of a spectrally opponent lateral geniculate nucleus cell to pure‐luminance (◯) and pure‐color (▴) gratings of various spatial frequencies, which were drifted across the cells' receptive field. The cell responds to the two types of stimulus patterns over different spatial frequency ranges.

Based on unpublished data from H. von Blanckensee
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Russell L. De Valois, Gerald H. Jacobs. Neural Mechanisms of Color Vision. Compr Physiol 2011, Supplement 3: Handbook of Physiology, The Nervous System, Sensory Processes: 425-456. First published in print 1984. doi: 10.1002/cphy.cp010310