NLM Citation: Gouras P. Color Vision.

2005 May 1 [Updated 2009 Jul

1]. In: Kolb H, Fernandez E, Nelson R, editors. Webvision: The
Organization of the Retina and Visual System [Internet]. Salt Lake City
(UT): University of Utah Health Sciences Center; 1995-.
Bookshelf URL: https://www.ncbi.nlm.nih.gov/books/

Color Vision
Peter Gouras
Created: May 1, 2005; Updated: July 1, 2009.

Introduction
Color vision is an illusion created by the interactions of billions of neurons in our brain. There is no color in the
external world; it is created by neural programs and projected onto the outer world we see. It is intimately linked
to the perception of form where color facilitates detecting borders of objects (Fig. 1).
Color is created by utilizing two properties of light, energy and frequency of vibration or wavelength. How our
brain separates these two properties of light, energy and wavelength, and then recombines them into color
perception is a mystery that has intrigued scientists through the ages. We know much about the nature of light
and the subjective impressions of color, definable by physical standards (1) but ultimately color should be
explained at the level of single cells in our brain. Examination of the responses of single neurons or arrays of
such neurons provides the best insights into the physiology of color vision. Ultimately our understanding of this
process will allow us to model the neural circuits that underlie the perception of color and form. Although still
beyond reach, progress is being made in deciphering these clever circuits that create our perception of the
external world.
We start by describing the nature of the photoreceptors that convert light energy into neural signals. Then we
consider the parallel channels leading from the retina to the thalamus carrying information into visual cortex,
where color is ultimately determined. Lastly we use our current understanding to speculate on how visual cortex
uses neural circuits to create the perception of color and form.

The Photoreceptors
Photoreceptors are neurons specialized to detect light. The detection occurs in an organelle called the outer
segment, a membranous structure where light absorbing proteins, opsins, are embedded. There are two major
types of photoreceptors in most vertebrate eyes, rods and cones. Rods are very sensitive but slow and their
response saturates at light levels where cones function optimally. Rods are not used much in modern society
where artificial illumination adequate for cone vision is ubiquitous. Cones are less sensitive but are fast and can
adapt to the brightest lights, being almost impossible to saturate. Cones evolved before rods undoubtedly in
areas of strong sunlight where vision was a great advantage. In broad sunlight, shadows are strong and more
important to detect than increments of light in the struggle for survival. Shadows depolarize cones leading to a
release of a transmitter that influences second order retinal neurons. The appearance of light hyperpolarizes
cones leading to a curtailing of this transmitter.

Copyright: © 2023 Webvision

2 Webvision

Figure 1. A floral scene with and without color.

A cone responds only to the energy it absorbs (2). All wavelength of light can produce identical responses from a
cone if the energy absorbed by the cone is the same for these wavelengths (Fig. 2). Cones are therefore color
blind producing a univariant response reflecting only the amount of energy they absorb. Detecting objects by the
energy reflected from their surfaces, however, can fail when objects reflect a similar amount of energy as their
background. Here is where color vision becomes important. Wavelength contrast can detect objects when energy
contrast is absent or minimal. An object can reflect the same energy but seldom reflects the same wavelength
composition as its background. Color vision combines both energy and wavelength contrasts to detect objects
and this advantage must have evolved early in the evolution of vision.
In order to detect objects by differences in spectral reflectance, two or more different types of cones are needed.
This is an important concept for understanding color vision. For divariant color vision, two cone types must
exist and be sensitive to different parts of the visible spectrum, preferably as different as possible. The range of
the visible spectrum depends on the ability of light to penetrate the eye and be absorbed by the photoreceptors.
Ultra-violet light is absorbed by the anterior segment of our eyes and seldom reaches the photoreceptors. Infra-
red light penetrates our eye readily but its quantal energy may be too small to activate opsins. Therefore early in
the evolution of color vision, opsins sensitive to the middle of our visible spectrum evolved, near spectral yellow,
and a short wavelength opsin evolved in a second type of cone, near spectral blue (Fig. 3). These have been called
L (long wavelength sensitive) and S (short wavelength sensitive) cones, respectively, and this was a first step in
the evolution of color vision.
In animals with large eyes, an interesting strategy evolved. L-cones were used to detect both energy and
wavelength contrast but S-cones were used only for wavelength contrast. This was due to chromatic aberration.
Color Vision 3

Short wavelength images are out of focus when longer wavelength images are in focus on the photoreceptor
mosaic. Chromatic aberration increases greatly at short wavelengths, which led to the L cone system dominating
energy contrast. As a result there are many more L than S cones in many mammals in order to gain spatial
resolution by achromatic contrast detectable by L cones.
In animals with small eyes, like mice and rats, ultra-violet light can reach the photoreceptor mosaic and in this
case ultra-violet sensitive cone opsins have evolved to widen the spectral range of vision and if combined with L
cones could allow color vision (3-5). Chromatic aberration is reduced in these small, highly spherical eyes which
have outer segments as long as animals with large eyes. This increases their depth of focus minimizing chromatic
aberration, an advantage of being small. Their retinal images are less magnified, however, than those in large
eyes.

Chromatic and Achromatic Contrast

In order to establish chromatic contrast it is necessary to compare the responses of a group of cones of one type
with the responses of a group of cones of another type within the same area of visual space. One cannot compare
only two neighboring cones such as the S and the L cone in Fig. 4A.
If an image of gray and yellow were to cover these two cones in an inappropriate way as done in Fig. 4A the
brain would reach an erroneous conclusion about this border because the S cone would not be affected by the
yellow side and the L cone would be strongly affected by the gray side. The brain might consider this to be a
black/yellow border rather than a yellow/gray border. Even if a slit of light imaged on only the more numerous L
cones (Fig. 4B), there would be ambiguity about the colors of adjacent images. If only L cones are affected by
both yellow and gray light, the brain would not see any border here. Only when a larger image covers a number
of S and L cones (Fig. 4C) can an unequivocal decision be made about the color of these bordering images. In
this case the left side, which is yellow, would strongly affect L but not S cones and therefore be judged to be
yellow. The right side of this border, which is gray, would strongly affect both L and S cones and therefore would
appear white or gray. For achromatic contrast, smaller images can be analyzed for border contrasts and in most
cases only two neighboring cones would be sufficient to distinguish a light/dark border. This is why a unit area of
achromatic space is smaller than a unit area of chromatic space.

Horizontal Cells
Cones receive an antagonistic input from horizontal cells, whose cell bodies reside in the outer nuclear layer and
whose processes contact the spherules of rods and the pedicles of cones (6). There are at least two varieties of
cone horizontal cells (Fig. 5).
One variety (H1) only contacts L cones; the other variety contacts both S and L cones (H2). Cone horizontal
cells receive an excitatory input from cones and send back an inhibitory input to cones. This is a type of negative
feedback. When a cone is hyperpolarized by an increment or depolarized by a decrement of light, it receives an
opposing input from horizontal cells after a brief synaptic delay. This dampens the response and can also reduce
the effects of scattered light by minimizing cone responses outside of the focal image on the retina. In color
vision the horizontal feedback also acts to narrow the action spectrum of cone bipolar cells. In divariant color
vision this can narrow the action spectrum of S cone bipolar cells. This occurs because the processes of the H2
horizontal cell that reach L cones are only post-synaptic to S cones. Therefore L cones can send an antagonistic
signal to S cones which can reduce the effectiveness of wavelengths, absorbed by both L and S cones and this
narrows the action spectrum of the S cone channels (7).
4 Webvision

Figure 2. Different wavelengths (colors) producing identical responses from cones.

Bipolar and Ganglion Cells

The retina is composed of three layers of neurons, the outermost being the photoreceptors, rods and cones,
which for divariant color vision are L and S cones. A second layer of bipolar cells transmits the signals of the
photoreceptors to a third layer of neurons, the ganglion cells whose axons form the optic nerve. The ganglion
cells serving the two photoreceptor systems are quite different. The L cones synapse with unique bipolar cells,
called "midget" bipolar. These single cone detectors were discovered by Stephan Polyak (8) in monkey retina
using the Golgi method of silver impregnation. Because of their small size they were called "midget" bipolar cells.
This provides the brain with the ultimate in spatial resolution, a single cone, and in addition isolates the signals
of L cones which can be used for color vision. A feature of this L cone bipolar system is the presence of two types
of cells (Fig. 6). One type of L cone bipolars depolarizes whenever the cone or cones they synapse with
hyperpolarize; this set is called on-bipolars because they are excited (turned on) by light. The other set is
depolarized whenever the cones they synapse with detect decrements of light; this set is called off-bipolars
because they are excited by darkness and inhibited (turned off) by light. These two sets of cone bipolar cells
synapse with separate sets of on and off ganglion cells at two levels in the inner plexiform layer of the retina, a
more external off-lamina and a more internal on-lamina (9).
These parallel channels, transmitting lightness and darkness from local retinal areas, are maintained throughout
the visual pathway to visual cortex. This "push-pull" system of neurons is thought to increase the dynamic range
for detecting decrements and increments of light in local areas of the retinal image. Only L cones seem to be
connected to both on and off bipolar cells while S cones are connected to only on-bipolar cells. The reason for
this may be that the latter are only involved in chromatic vision while the former are involved in both chromatic
and achromatic vision. Achromatic vision involves the detection of lightness and darkness while chromatic
Color Vision 5

Figure 3. The normalized absorption spectra of the S and L cone opsins mediating divariant color vision. Strong absorption by the S
cone opsin alone induces "blue"; strong absorption by L cone opsin induces "yellow" and absorption by both induces "white or gray"
depending on achromatic contrast of light and dark. The 4th root makes the curves identical in shape.

vision involves the detection of color. Not only do S cones lack an off-bipolar system but they have a much
different route to the ganglion cell output layer. The S cone on bipolar excite the internal arbor of a bistratified
ganglion cell while a wide-field L cone off bipolar excites the external arbor of this same ganglion cell (10). This S
cone system is absent in the very center of the fovea where achromatic contrast is mediated by the L cone midget
system. Away from the fovea the L cone midget system begins to contact more than one cone and therefore loses
spatial resolution.
There is a second ganglion cell system that plays no role in color vision but is also connected to only L cones.
These are parasol ganglion cells. They are larger cells with faster conduction velocities and they target the
magno-cellular layers of the lateral geniculate nucleus (LGN). They appear to play a role in the detection of
movement and possibly slow tracking movements. There is some evidence that they might receive an input from
S cones but I have never been able to detect such an input. Barry Lee's group considers these cells play a role in
the detection of luminance (11).

Divariant Blue/Yellow Color Vision

The ganglion cells in Fig. 6 are thought by most to be the essential channels for divariant color vision. The
bistratifid S cone on-cells and the L cone midget cells monitor the light absorption of S and L cones respectively.
The signals from these two channels must be compared in the same areas of visual space for color vision. The
6 Webvision

Figure 4. The difficulty determining the color of small objects.

cone bipolar cells driving the bistratified S cone on-ganglion cells have co-extensive fields (Fig. 7), which is ideal
for comparing differences between two cone mechanisms in the same area of visual space.
These two different bipolar inputs to the bistratified ganglion cell do not appear to oppose each other because
these cells are excited by white light. Another more indirect input to these cells comes from the H2 horizontal
cell which transmits antagonistic signals from L to S cones. This input provides spectral antagonism to the
bistratified ganglion cell because light activation of L cones will produce a depolarizing signal in S cones that
counteracts the hyperpolarization produced by short wavelength light. The strong response to white light,
however, implies that this H2 mediated antagonism is relatively weak.
The channel transmitting L cone signals for spectral contrast in visual cortex has been considered to be the
midget system, at least in primates. These midget ganglion cells are considered to receive no input from S cones,
either synergistic or antagonistic, although there are connections to S cones through H2 horizontal cells. In
trivariant monkey retina there is no evidence of S cone input to either the midget or the parasol ganglion cell
systems which implies that H2 horizontal cells are only post- and not pre- synaptic to L cones. The H1 horizontal
cell only contacts L cones and therefore provides only spatial antagonism to neighboring L cones and does not
produce spectral antagonism.
Color Vision 7

Figure 5. Two varieties of horizontal cells and their relation to L and S cones in a divariant retina.

This previous picture is not accepted by everyone. There is evidence that a L cone on/S cone off ganglion cell
exists to provide an opponent L cone on input to visual cortex (12-15). It has been difficult to find such a
ganglion cell in primate retina but they they have been reported in the koniocellular layers of the primate lateral
geniculate nucleus. These geniculate cells have been traced back to their retinal ganglion cell inputs, which have
their dendritic arbors in the on-lamina of the inner plexiform layer, quite different than the bistratified arbors of
S on/ L off ganglion cells. This produces a curious difference in the blue/yellow channels of color vision in
primates. In small animals, ground squirrels, guinea pigs and mice, there appears to be a more symmetrical
system of S cone on/Lcone off and M cone on/S cone off opponent ganglion cells, both of which send their
dendrites to the on-lamina of the inner plexiform layer. This implies that the horizontal cells and/or intervening
amacrine cells are involved in their unique opponent organization, perhaps more similar to the rod system. The
existence of this retinal L cone on/S cone off channel would preclude the midget cells depicted in Fig. 7 from
contributing to color vision in animals with divariant color vision.
A more iconoclastic hypothesis recently proposed by the Neitz group (13) considers the bistratified S cone on/L
cone off retinal ganglion cell to play no role in color vision proposing that its signals go to the brain stem where
they mediate unconscious visual functions together with light sensitive retinal melanopsin ganglion cells. They
propose that surround antagonism from the H2 horizontal cell turns L cone off-center midget bipolar cells into S
cone-on/L cone-off cells and L cone-on/S cone-off cells. This hypothesis brings S cone signals into the midget
ganglion cell system designed for high spatial resolution. Evidence that the bistratified S cone on-cell does affect
conscious vision comes from an observation of W.S. Stiles (16) that his S cone mechanism, isolated
psychophysically, affects conscious experience because it exhibits a curious behavior; its sensitivity declines
8 Webvision

Figure 6. The currently understood retinal ganglion cells responsible for rod (achromatic) and cone achromatic and chromatic signals
being sent to the lateral geniculate nucleus. We assume that the L cone off bipolar signals darkness but it could also be blue.

when a long wavelength adapting light, which has no effect on S cones, is turned off. The bistratified S-cone
ganglion cell exhibits this same behavior implying that its signals do reach visual cortex and consciousness. It is
also possible, however, that the unusual midget cells proposed by the Neitz team (13) also exhibit this
phenomenon, which has been labeled "transient tritanopia".
Regardless of which of these three hypotheses is correct, there must be a way for the visual cortex to compare L
with S cone activity in the same area of visual space in order to produce color vision. In Fig. 8 we use two of the
ideas that are demonstrated in Fig. 7 to illustrate how striate cortex uses these retino-geniculate inputs to
construct cells that are more responsive to color.
Fig. 9 shows a logical way to extract the signals from the cone mosaic into parallel channels mediating
achromatic contrast with high spatial resolution and chromatic contrast with lower spatial resolution from the
same mosaic.
The perception of color mixes both achromatic with chromatic signals to create the combined experience of
color. In this arrangement the S cone system only plays a role in chromatic vision while the L cone system
contributes to both chromatic and achromatic vision. The repertoire of colors that are produced and their
relationship to the activity of the two cone mechanisms involved are shown in Fig. 10; these colors are shown at
borders of minimum achromatic contrast where chromatic contrast becomes most important.
Color Vision 9

Figure 7. Two options being considered for the ganglion cell system providing chromatic and achromatic information to the lateral
geniculate nucleus for divariant color vision.

Color Constancy and Double Opponency

The schemes of the previous figures neglect a problem called "ch22color constancy". We see colors as unchanged
even when there are large changes in the spectral properties of an illuminant. The colors in a scene illuminated
by fluorescent lights, which generate much short wavelength energy or by tungsten filament lights, which
generate much long wavelength energy are not significantly altered by such changes. In other words we should
see things "bluer" in fluorescent light and redder in tungsten light but we don't. This constancy of colors despite
changes in illumination intrigued Edwin Land, the founder of the Polaroid Corporation, who spent years
investigating this phenomenon and demonstrating significant global aspects of color vision (17). Local objects
can reflect identical spectral components from their surfaces but will appear of different color because of the
influence of the entire scene. He proposed a model in which the signals from each cone mechanism are
normalized over the entire visual scene before being compared with each other locally to generate the perception
of the color of an object scene. Fig. 11 illustrates why this idea is reasonable. Here two lights beams, one white,
the other yellow illuminate a screen.
An arrow blocks a portion of the yellow beam and this produces a shadow that appears "blue". If one determines
the light energy being absorbed by the L and S cones from the screen, it is apparent that there is more light
affecting the L than the S cones coming from the shadow, which would suggest that the shadow should look
yellow and not blue. But if one normalizes the light coming from the entire screen to 100%, then the relative
effect on the S cones becomes greater than that on the L cones and the shadow appears "blue". This supports the
idea that responses from each cone mechanism across the entire visual scene are first normalized before being
10 Webvision

Figure 8. How retino-geniculate inputs are used to build cells with spectral antagonism, i.e., selectivity for certain wave-bands and
ultimately color. This figure illustrates interaction between the achromatic system that determines the lightness or darkness of an object
and the chromatic system that detects "hue". Together they create color.

compared with each other locally to produce the perception of color from specific objects in the scene. Such
normalization could occur if S cone and L cone transmitting neurons were to inhibit each other following the
rules outlined in Fig. 12. This could reduce the responses to global illumination that affected one cone
mechanism more strongly than the other and support color constancy. If, in addition, the excitation of these
cone specific channels were organized so that each local area of color space would be enhanced by neighboring
spectral contrast between cone mechanisms it would lead to simultaneous color contrast.
By the rules of Fig. 12, a unit area of color space A would appear "blue" if there were short wavelengths and no
long wavelengths in area A; the converse would lead to "yellow". This blueness or yellowness would be enhanced
if the surrounding areas of color space were activated by the opposite color, i.e. simultaneous color contrast. Such
cells have been called "double opponent" cells because they have one form of antagonism between cone
mechanisms mediating a local area of color space and an opposite arrangement in neighboring areas of color
space. Charles Michaels (18) found such cells in striate cortex of non-human primates following up on
observations of Nigel Daw (19) in goldfish retina. Double opponency tends to separate spectral from energy
contrast.

Trivariant Color Vision

In primates, high resolution vision and trivariant color vision evolved to enhance survival. A fovea formed to
facilitate high resolution achromatic vision and a third opsin evolved from the original mammalian L cone opsin
Color Vision 11

Figure 9. Parallel pathways extract chromatic and achromatic contrast from the same cone mosaic. The unit area of achromatic space is
smaller than that of chromatic space.

to create a new dimension of color in higher primates (20). The gene for the L cone opsin duplicated itself and
one of the paired genes developed polymorphisms to absorb further into the long wavelength region of the
spectrum. The original long wave sensitive L cone now became an M cone being the partner of an even longer
wavelength sensitive L cone. Fig. 13 shows that a trivariant system can detect a larger variety of spectral
reflectances (21). In Fig. 13 above right, one sees that a divariant color vision system can detect spectral contrasts
that reflect more at one end than at the other end of the spectrum; these reflectances tend to tilt the solar
spectrum.
If, however, a surface reflects less from both sides of the visible spectrum, i.e. bending the solar spectrum, it
might be invisible to a divariant system because both cone types could be absorbing the same amount of light
from the object and its background (Fig. 13 lower left). A trivariant system detects this object because it is
impossible for all three cones to be absorbing the same amount of light from the object and its background (Fig.
13 lower right). More complex reflecting surfaces might confuse even a trivariant system but they are probably
very rare in the natural world. This change split the yellow region of the original spectrum and created two new
chromatic percepts, red and green (Fig.14). It is noteworthy that a rise in the beta-band absorption of this new L
cone pigment also provided a long wave influence at the short wavelength region of the spectrum.
12 Webvision

Figure 10. The variety of colors seen by a divariant color vision system using only L and S cones. Each of these colors is related to a
certain combination of on and off cell systems.

Fig. 15 shows how a trivariant system facilitates the distinguishing of red and green objects which would remain
indistinguishable for a divariant observer.
In the fovea, the same midget cells which had been contacting only single L cones for divariant color vision now
contacted either an L or an M cone (Fig. 16), providing the brain with separate channels for these two different
cones and another opportunity for chromatic contrast.
In Fig. 16 it is assumed that the bistratified S cone on cell conveys S cone signals from the retina and the midget
cell system conveys the signals of L or M cones for color vision (6). The midget cell system is currently assumed
to play a role as a "double duty" detector contributing to both high spatial resolution achromatic vision and
lower spatial resolution chromatic vision.
Fig. 17 (upper) shows the responses of a midget ganglion cell receiving an excitatory L cone input. This cell is
continuously excited by the red adapting field; under these conditions the cell does not respond to the small red
spot. A blue adapting field stops this continuous discharge and under these circumstances the cell responds to
the small red spot. Fig. 17 (lower) shows the responses of an L cone on midget ganglion cell (large amplitude)
and a M cone on midget-like ganglion cell (small amplitude) responding to a small red spot in the presence of
the same adapting fields. The red adapting field continuously excites the L cone cell and inhibits the M cone cell;
under these conditions the small red spot inhibits the L cone cell, an inhibition mediated by M cones.
Color Vision 13

Figure 11. An experiment of Edwin Land. Two projectors cast a mixture of white and yellow light on a screen and an object (arrow) is
placed in the yellow beam which leads to the appearance of a "blue" shadow of the arrow on the screen. This somewhat paradoxical
appearance of the shadow can be explained if one assumes that the effects of the separate cone mechanisms viewing this screen have
their responses normalized across the screen before being compared with each other to produce color.

Fig. 18 shows an S cone on-cell responding to spectrally narrow stimuli.

The cell is excited by the blue spot on the yellow background and is profoundly excited by the turning off of the
yellow adapting light. The cell is not responsive to the red spot in the presence of the yellow adapting light but is
inhibited by the red spot in the absence of the adapting light.
Fig. 19 illustrates how these midget cells could participate in both chromatic and achromatic vision. The same
concentrically organized retinal and geniculate cells feed two different circuits in striate cortex. One achromatic
with a large number of orientation selective neurons capable of high spatial resolution and the other chromatic
built up from the same center/surround organized midget-like cells. How center/surround cells can build up a
circuit which is more co-extensively organized more suitable for chromatic processing remains enigmatic.
David Hubel (22) and the late Bob Rodieck (23) have suggested that the midget system may not be involved in
mediating chromatic vision but another much less common ganglion cell type with a receptive field in which L
and M cones driving these cells are co-extensively rather than concentrically organized in the receptive field of
these cells. It has been difficult to obtain any evidence for such a cell.
The evolution of trivariant color vision in higher primates increased the repertoire of colors we perceive and the
power of spectral contrast to detect objects. The original blue/yellow form of color vision was now accompanied
by a parallel system of red/green colors occurring in the yellow region of the spectrum where brightness is
maximal (24). Where the dynamic range of achromatic contrasts are maximal is where chromatic contrast can
contribute most to detecting borders when achromatic contrast is minimal. Fig. 20 shows how the activity of this
trivariant cone system contributes to the variety of major colors we see. The borders of effective energy contrast
for each of these colors are minimal, which is where color contrast is especially important.
14 Webvision

Figure 12. How double opponent cells can be constructed. The central two on cells, one S on and the other L on, inhibit each other. The
surround L on cells excite the central S on cell and inhibit the central L on cell. the surround S on cells excite the central L on cell and
inhibit the central S cone on cell (not shown).

The Cone Mosaic of Trivariant Color Vision

The division of a single L cone type into two spectrally different L and M cones produced a mosaic of L, M and S
cones from which each ganglion cell picks out the input of one or the other of these cone mechanisms to
transmit to the brain. Adaptive optics has revealed significant differences in the number of L versus M cones in
normal subjects (25) (Fig. 21).
Some subjects have 15 times as many L than M cones and others have twice as many M than L cones but all have
normal color vision. This reinforces the idea that a unit area of color space involves comparisons of large groups
of cones. What remains problematic is how midget-like ganglion cells in the parafovea select only L or only M
cones to form synaptic contact within this variegated cone field.

A Parallel System of Achromatic Ganglion Cells

In addition to midget and midget-like ganglion cells described there is another parallel system of larger ganglion
cells with less representation in the fovea than the midget cell system (26). These ganglion cells also have: on-
and off-varieties mediated by a separate set of cone bipolar cells and are known to be the "parasol" ganglion cells
found by the Golgi method of silver impregnation. They seem to receive their inputs from only L and M cones
through a different set of bipolar cells than those serving the midget cell system. Fig. 22 shows two retinal
ganglion cells in rhesus monkey retina.
The cell with smaller amplitude is a phasic on-cell that responds at on to all three wavelengths with a shorter
latency than the tonic S cone on cell which is excited only by the deep blue light (419 nm) and inhibited by the
longer wavelengths (610 and 509 nm). It is thought that a separate relatively wide field bipolar cell feeds these
Color Vision 15

Figure 13. An Object 1 that just "tilts" the solar spectrum but causes as much light absorption as the background will not be detectable
by a retina with only 1 photopigment (opsin) in one class of cones (1). In order to detect this Object 1 by its spectral contrast it is
essential to have a second type of cone (2) which will always recognize this Object 1 by its spectral reflectance relative to its
background. If an object arises such as Object 2 that "bends" the solar reflectance by having a reflectance maxima in the middle of the
spectrum it could become invisible relative to its Background for both cones 1 and 2. For such reflectances three types of cones are
needed to detect this object under all conditions.

phasic cells. These cells receive similar inputs from both L and M cones and therefore cannot play a role in "red/
green" color vision and probably no role in blue/yellow color vision. Fig. 23 is evidence that they receive no input
from S cones. The threshold spectral function obtained on a strong yellow adapting field shows no evidence of
an S cone input. The S cone on-cell has a threshold spectral function that shows it lowest threshold to blue light.
The phasic cells have their lowest threshold resembling that of the photopic luminosity function.
The retinal latencies at of these two types of cells show a significant difference between the tonic and phasic cells
indicating a faster transmission of the latter through the inner nuclear layer by the phasic system. In addition the
larger size of the axons of the phasic system allows this system to transmit its signals to visual cortex much
quicker than the tonic system. The role of the phasic "parasol" ganglion cell system in vision is not entirely clear.
It seems to play a role in the detection of motion and to target different areas of visual cortex. Whether it plays
any role in color vision is moot.

The Lateral Geniculate Nucleus (LGN)

The LGN reorganizes the parallel ganglion cell systems serving local retinal areas into separate layers from where
their axons project to specific layers in striate cortex (12, 15, 27). Signals from nasal retina of the contralateral
16 Webvision

Figure 14. The normalized absorption spectra of the three cone mechanisms that mediate primate trivariant color vision. The original
L cone system of divariant color vision split into two slightly different long wavelength sensitive cones straddling the yellow region of
the visible spectrum.

eye are separated into different layers from those coming from the temporal area of the ipsilateral eye. This is
needed for stereopsis where changes in eye position require comparing signals from different corresponding
retinal regions and accordingly different ganglion cells. Binocular interactions begin for the first time in visual
cortex. The parallel channels coming from the same retina are also separated into different layers. The phasic
achromatic ganglion cell system goes to the magno-cellular layers, one layer for each eye. The midget and
midget-like retinal ganglion cells transmitting signals for high resolution achromatic vision and "red-green"
color vision are placed in four parvo-cellular layers, two for each eye. The S-cone mediating retinal ganglion cells
which are involved in "blue-yellow" color vision go to the konio-cellular layers. These parallel systems of
ganglion cells project to specific layers of striate cortex where form, color, movement, direction and stereopsis
are processed.
An interesting but unknown role must be played by the numerous centrifugal axons that arise from striate
cortical layers and impinge upon the cells in the LGN. Whether this shapes or influences chromatic and
achromatic contrasts that are involved in form vision is unknown.

Striate Cortex
The LGN transmits its signals to striate cortex, the first visual area (V1) to process visual signals in cerebral
cortex. There is a retinotopic order to the projection of these producing a map of the visual field. The map is
distorted because the fovea occupies a relatively large area compared to the more peripheral retina. There are
columns of cells extending from the upper to the lower layers, one column receiving signals from one eye and a
Color Vision 17

Figure 15. Trivariant color vision distinguishes red from green, expanding the repertoire of colors.

neighboring column receiving signals from the other eye (Fig. 24). Within each of these "ocularity" columns
there are micro-columns of cells favoring a particular orientation of an extended light stimulus. Together these
sets of columns have been called a "hyper-column". Within each hyper-column there is a local area that receives
inputs from chromatic selective cells, i.e. the S cone on and off cells and the L and M cone midget and midget-
like cells; these areas have been called "blobs", and are thought to be where color is processed.
18 Webvision

Figure 16. Four varieties of ganglion cells considered to mediate trivariant color vision in primates. The L cone on/ S cone off cells
reported to be in the koniocellular layers of the lateral geniculate nucleus are not included here.

Figure 17. Midget-like ganglion cell responses. Figure 17(upper) shows an L cone on midget-like ganglion cell being stimulated by a
red and blue adapting light while the center of its receptive field is being stimulated with a small red spot. The blue light suppresses the
discharge but the cell responds to the red spot. The red adapting light excites the cell tonically but the cell no longer responds to the red
spot. Figure 17 (lower) shows the responses of two midget-like ganglion cells. The one of small amplitude responds to the red spot on
the red background and is excited tonically by the blue background. The one of large amplitude is excited tonically by the red
background light and is inhibited by the red spot. It is suppressed by the blue adapting light but is then excited by the red spot. The
latter is a L cone on cell with an antagonistic M cone surround; the former cell is the converse.

There are six distinct layers extending from an upper layer 1 down to a lowermost layer, 6. The projections from
the functionally different layers of the LGN target different layers to synapse on cortical cells (Fig. 24).
Color Vision 19

Figure 18. An S cone on/L+M off cell responding to a blue spot (above) and a red spot (below) in the presence and absence of a yellow
adapting light.

The L and M midget and midget-like cells, projecting from the parvo-cellular layers, synapse in layer 4Cbeta
from which their post-synaptic target cells appear to send axons to layer 3 within "blobs". The S cone system,
projecting from the konio-cellular layers, appears to send axons directly to layer 3 within blobs. The fact that the
parvo-cellular input, incorporating the L and M cone midget and midget-like cells, makes an intermediary
synapse may be due to the double duty role these cells play in handling both achromatic and chromatic vision.
The area surrounding blobs process achromatic contrast, undoubtedly receiving their input from the parvo-
cellular layers. The magno-cellular layers of the LGN target cells in layer 4Calpha from which postsynaptic cells
send axons to layer 4B. Therefore there is segregation of cell systems, some playing a role in chromatic contrast
and others playing a role in achromatic contrast and movement. The existence of "double opponent" cells in
blobs supports the idea that this zone within each hyper-column is devoted to detecting chromatic contrast for
color vision. There are many unknowns in this structure that can only be solved by more single cell physiology
combined with anatomical insights into the circuitry. What seems clear is that each unit area of visual space is
processed in parallel by achromatic and chromatic mechanisms (28).

Pure Spectral Contrast

Double opponent cells tend to eliminate the influence of energy contrast establishing pure spectral contrast as an
indicator of the borders of objects. With trivariant color vision this can be formed by comparing L with M cone
responses and/or by S cone versus L&M cone responses. Fig. 25 illustrates the organization of a "red/green"
double opponent cell. The system uses spectral contrast.
20 Webvision

Figure 19. L and M cone midget-like cells can be used to extract achromatic high spatial resolving vision and chromatic "red/green"
chromatic vision of lower spatial resolution.

Fig. 25 shows how concentrically organized double opponent cells can be constructed by having identical (red)
neighboring cells inhibit a central cell (red) and have opposite neighboring cells (green) excite the central cell
(red) making it most sensitive to a red central object in a green background and unresponsive to the converse.
Such a cell responds to spectral rather than energy contrast as an indicator of a border. This behavior is
illustrated by a cell detected in visual cortex that responds exclusively to spectral and not to energy contrast (Fig.
26). The stimuli with maximum energy contrasts such as a red or green bar on a black background or a black bar
on a green or red background produce no response from the cell. But a green bar on a red background where
energy contrast is minimal generates a large response. The brain has now separated the two major forms of
contrast, spectral versus energy contrast to use them as independent variables to create color.

Simple, Complex and Hypercomplex Double Opponent Cells

One of the remarkable discoveries in visual cortex has been that of a unique body of cells that exhibit definable
patterns of responses all based on orientation selectivity. One group called "simple" cells show orientation
selectivity which can be predicted by the organization of responses produced within their receptive field.
Another group called "complex" cells also show orientation selectivity but are less obviously related to the
distribution of responses within their receptive field. A third group of "hypercomplex" cells are orientation-
selective but are "end-stopped" being inhibited if the oriented stimulus is too long. These three types of cells are
thought to contribute to form vision based on energy contrast. Charles Michael (18) has shown that a similar
Color Vision 21

Figure 20. The major colors produced by trivariant color vision in primates. The figure indicates the synaptic pattern in which the
three cone inputs of on and off cells make to produce particular colors.

group of double opponent cells exist in striate cortex of monkeys. Fig. 27 shows such a simple cell responding
best to strong spectral contrast.
Fig. 27 shows that this cell is excited when an oriented red bar moving upward crosses the red selective area of its
receptive field and inhibited when it crosses the green selective area; it produces an off-response when it leaves
this green selective area. A green bar produces excitation when it crosses the green selective area only. A green-
red and a red-green bar produce strong responses but the responses occur at different points in the receptive
field and are therefore totally out of phase with each other. This cell responds poorly to white bars being
spectrally selective. Fig. 28 shows a complex cell that responds to spectral and not energy contrasts. It is both
orientation and directionally selective and refuses to respond to energy but responds strongly to spectral "red/
green" contrast.
Hypercomplex double opponent cells (Fig. 29) are also detectable in striate cortex completing the variety based
on energy contrasts. Therefore the same repertoire of orientation selective cells responsive to energy contrasts in
striate cortex can also be found for spectral contrast. It is thought that both groups of cells play a role in form
vision. Parallel systems of cells are detecting the forms of objects, one based on energy and the others based on
spectral contrast. Because energy contrasts allow a higher spatial resolution, more of visual cortex is dedicated to
achromatic than chromatic contrast detection.
22 Webvision

Figure 21. An adaptive optics view of the mosaic of L (red), M (green) and S (blue) cones in four human subjects with normal color
vision. The ratio of S to L and M cones is constant but thaf of L to M cones varies from 1:2.7 (M:L) to 16.5:1 (L:M). (Adapted from
Williams).

Fig. 30 shows how these two parallel contrast detecting systems could work in detecting a red object. Both
simple cells of different orientation selectivity can detect the border of this cross, one set based on energy
contrast and with a higher spatial resolution and the other set based on spectral contrast and with a lower spatial
resolution. Sharp corners in the object would be detected by end-stopped hypercomplex cells. This creates two
different views of the object which may be fused into one colored object just as two similar stereoscopic views
are fused into an object in depth. Together they create shapes in full color (Fig. 30).
Color Vision 23

Figure 22. A magno-cellular on cell (small amplitude) and an S cone on cell (large amplitude) responding to three different parts of the
spectrum (419, 509 and 610 nm). The calibration, lower right , indicates 200 microvolts vertically and 20 msec horizontally. The lower
trace shows the duration of the light stimulus. Positivity is upwards.

A similar system for using spectral contrast created by double opponent S cone versus L cones or L+M cones
must create another spectral view of objects by border contrasts (Fig. 31 and Fig. 32). In Fig. 31 groups of double
opponent simple cells detect a circular blue form on a yellow background. In Fig. 32, similar simple cells detect a
border of contrast between yellow and white, allowing the detection of an object of minimum energy contrast.
This highlights an important role of the S cone mechanism in vision to detect border contrasts of white versus
yellow which are difficult for the achromatic system which is mediated only by L and M cones. This scheme
implies that for color vision three different views of the same object are formed and then fused to provide colors
that represent mixtures of the red/green and the blue/yellow spectral contrasts systems. If there is no activity in
the spectral contrast detectors then only the energy (achromatic) system is active and the object is pure white,
gray or black. There are some different views on these ideas. Single opponent cells are thought to detect surface
color while double opponent cells detect borders (29). Others have suggested that neurons involved in the
analysis of color should retain their chromatic properties in the face of changes in other components of
stimulation, orientation, size and contrast and this may not be the case for color selective neurons found in
striate cortex (30).
24 Webvision

Figure 23. The threshold (left) in log relative quanta of 3 on-center phasic cells (filled symbols) and 7 S cone on/L cone off cells (open
symbols) and their latencies at threshold (right) on a reciprocal scale of milliseconds. The dashed line indicates the Stiles' ¹ 1 and 4
mechanisms. The continuous line is the CIE (Commission Internationale de l' Eclairage) luminosity function.

The Stabilized Retinal Image

If an image is made stationary on the retina it will disappear in seconds. Fig. 33 illustrates what is perceived by a
subject presented with a stabilize image of a red/green boundary based on the research of the Russian
physiologist Yarbus (31) and others (32). Initially the subject sees the red/green scene but in several seconds it
fades into a featureless field. If a pale blue light is added to both sides only a blue field is seen without the red/
green scene. This also fades within seconds. If the pale blue field is removed the subject sees a faint red/green
scene but this fades again. This indicates that we only see in transients determined by change in the retinal
image.
If the retinal image does not change in space and/or time, it will disappear. Constant small micro-movements of
the eye continue to produce transient changes at the borders of objects that are able to maintain an
undiminished view of the external world. This supports the idea that objects are seen because of their borders, as
depicted by the orientation selective border contrast cells illustrated previously, which must be continuously
moved across the retina to maintain their visibility. Fig. 34 illustrates how the responses of on and off cells
contribute to maintaining strong responses to movements of borders of contrast. When a long wavelength (red)
surface enters the receptive fields of on and off cells, the on-cell (L off) responds more strongly than the off-cell if
Color Vision 25

Figure 24. S cone on and off cells of the koniocellular layers and the four varieties of L and M midget-like cells of the parvo-cellular
layers of the LGN target different layers of striate cortex. The midget system must project to both the achromatic areas as well as the
"blobs" involved in chromatic vision. The parasol cells from the magno-cellular layers of the LGN target a different layer than the
previous cells.

there is an increment in energy contrast. But the off-cell (M off) will respond more strongly if there is a
decrement in energy contrast. These two systems are showing their ability to respond to increments and
decrements of energy contrast but also manage to favor long over other shorter wavelengths, which is a key for
their role in color vision. M on and L off -cells would respond weakly to such a moving border.

Color Vision Beyond Striate Cortex

An intriguing but poorly understood aspect of visual cortex is the remapping of visual space into multiple
adjacent visual areas where a retinotopic order is maintained. Fig. 33 shows how V1 projects to V2, a second
visual area juxtaposed to V1 where there continues to be a segregation of functionally different cells. The "blob"
areas of V1 appear to project to "slabs" of cells in V2, which project to "globs" of cells in V4 (Fig. 35).
26 Webvision

Figure 25. Concentrically organized double opponent cells can be constructed by having identical (red) neighboring cells inhibit a
central cell (red) and have opposite neighboring cells (green) excite the central (red) making it more sensitive to a red central object in
a green background and unresponsive to the converse.

Within these three interconnected areas spectral contrast is processed, presumably to determine its contribution
to color, i.e., hue. Adjacent areas processing energy (achromatic) contrasts contribute to determining the
saturation and brightness in colors. The achromatic areas are larger because of the higher spatial resolution of
energy contrasts. The receptive field sizes of cells in these higher visual areas tend to be larger than they are in V1
suggesting that, several blobs in V1 project to a single slab in V2 and several slabs in V2 project to a single glob
in V4 (33-35). The functional roles of these higher areas could be to recognize objects better as they come closer
to the observer when they cast a larger retinal image. It is known that the perception of color increases with the
size of an object over sizes as large as 20 degrees of visual angle. Under ideal conditions i.e. a well lit surface, we
can perceive at least a million different colors. Fig. 36 shows how more basic hues determined by single cells in
blobs could be used to build higher order colors that combine the inputs of the more basic hue determining cells.
As one develops more experience with colors even higher order cells could be formed allowing the
distinguishing of many different shades of color.
There is a question as to whether a single cell is committed to detecting a particular color or whether a group of
cells are involved. This might depend on the practice one has in working with colors such as artists who may
form single cells that are sensitive to a particular color and capable of contributing to even higher order colors
while those who are less involved have fewer such cells and depend on less fastidious interactions.
There has been a controversy about whether color processing is exclusively segregated to V4 while achromatic
processing occurs in another complete area of visual space. Evidence implying this type of organization involves
rare individuals who have lost all color vision but can see objects with high spatial resolution in black and white.
Color Vision 27

Figure 26. A cell selective for orientation based on pure spectral contrast. It fails to respond to strong energy contrast such as occurs
with the movement of the colored bar on a dark background (above) and a dark bar on a color background (below).

This hypothesis was championed by Semir Zeki (36) but the existence of "globs" related to spectral contrast in V4
surrounded by larger areas of achromatic contrast detecting cells tends to weaken this idea. Because "globs" are
relatively large it was perhaps possible for an investigator making a limited number of penetrations to conclude
that all of V4 is devoted to chromatic vision. Nevertheless there must be an explanation of why in these very rare
patients color can be lost without a loss in spatial resolution which depends entirely on achromatic contrast. The
enigma may be explicable by damage to higher areas of the brain where interpretation and language are involved
linking vision to external communication.

Redness at short wavelengths

It is well known that short wavelengths can produce a reddish sensation that makes blue appear violet. This
affect appears to arise from the beta-band of L cone opsin which makes this opsin more sensitive to light than M
cone opsin at this region of the spectrum (see Fig. 14). This effect is best observed in midget-like cone opponent
retinal ganglion cells which are excited by long wavelengths and inhibited at wavelengths at the middle of the
spectrum. In many of these cells excitation appears with short wavelength stimulation implying that the L cone
response is overpowering the M cone antagonism. This is supported by the fact the L cone response is
28 Webvision

Figure 27. A simple cell responding best to strong spectral contrast.

strengthened by a blue adapting light, which selectively depresses the responses of M cones. Had this short
wavelength excitatory input been due to S cones, it would have been weakened by the blue adapting light.

Hering's Theory of Color Vision

An insightful theory was proposed by Ewald Hering (37) in Prague in 1874 which was resisted by many leading
scientists in the field, such as Herman von Helmholtz (38). Hering noted that the sensations of blue and yellow
Color Vision 29

Figure 28. A complex cell responding to spectral contrast but not to energy contrasts.

opposed each other so that when seen simultaneously, the original colors were lost and a new sensation of a
totally different color "white" was seen. Similarly the colors red and green exhibited a similar opposing
relationship. When mixed they lose all traces of the original colors to create a new sensation "yellow". Hering
suggested that there were two pairs of opponent processes underlying human color vision. This contrasted with
the fact that there are only three variables underlying human color vision. This three dimensional system was
suggested by Thomas Young in 1802 (39) and proven by James Clerk Maxwell in 1872 (2) and strongly
promulgated by Helmholtz in 1852 (38). The Hering theory suggested that there are two pairs of basic colors that
have a unique opponent relationship for each pair. He was correct in this analysis although his idea that the
opponency between colors was occurring in the photoreceptors was incorrect.
At Hering's time the understanding of neuro-physiology was poor and the concepts of excitation and inhibition
of neurons in unique and complex ways was unknown. The first suggestion that antagonism between colors
could be due to excitation and inhibition between nerve cells, that I have found, was in a paper by the Swedish
physiologist Gustaf F. Gothlin in 1944 (40). He had followed Sherington's work on the spinal cord where the idea
of excitation and inhibition between nerve cells first evolved. Fig. 37 shows how prophetic Gothlin's ideas were.
Here he creates an antagonistic balance between blue and yellow processes as a first stage and then hangs a scale
30 Webvision

Figure 29. A double opponent orientation and directionally selective hypercomplex cell, characteristically failing to respond if the red
bar is elongated.

on the yellow side of the blue/yellow balance to show an antagonistic relationship between red and green. He
predicts that these antagonistic interactions were due to excitation and inhibition between neurons. It is
interesting that the pioneer of exploring nerve cell responses to spectral stimuli in the retina, the Nobel Prize
winner, Ragnar Granit, also a Swedish physiologist, made no reference of Gothlin's concept in any of his writings
on retinal responses to color.
Color Vision 31

Figure 30. Two systems, one based on achromatic contrast with finer spatial resolution and a second based on spectral contrast, detect
the borders of an object using simple cells and end-stopped cells with orientation selectivity (30).

Granit (41) proposed in 1947 a dominator/modular theory of color vision in which there were some retinal
neurons which responded to only a narrow part of the visible spectrum, "modulators", and another group that
responded to a broad band of the spectrum, "dominators". He thought the modulators were more involved in
color vision but never mentioned the concept of excitation and inhibition between colors or cone mechanisms as
Gothlin did. The first American studies of single visual neuron responses to monochromatic stimuli were
obtained in the monkey's lateral geniculate nucleus by Russel DeValois and associates in 1958 (42). They
emphasized the presence of narrow-band responding cells resembling the modulators of Granit but they also
noted some cells that gave on responses to red and off responses to green light but again Hering's concept of
opponent colors was not mentioned. The experiments that brought Hering's opponent color theory to the
forefront were done in fish retina by another Swede, Gunter Svaetichin (1956) using glass instead of metal
electrodes, enabling him to record intra-cellularly from horizontal cells. This allowed him to see both
depolarizing and hyperpolarizing responses, the latter not obvious with extra-cellular recordings, where one sees
only a silence or perhaps an off response from inhibition (hyperpolarizing) which could easily be overlooked.
Svaetichin's results gave rise to a resurgence in Hering's opponent color theory (43, 44). This quickly led to the
idea that there were three retinal channels representing Hering's color opponent channels, red opposing green,
yellow opposing blue and white opposing black. This idea, however, was short lived. First of all it became
apparent that the red/green opponent cells had concentrically organized receptive fields with the color opponent
inputs coming from adjacent retinal regions. One would expect the color opponent signal should involve the
same area of visual space and not neighboring areas. In addition the red-green opponent ganglion cells seemed
to be the midget cell system which was the logical mediator of high spatial resolution. This created a dilemma in
having a red/green color opponent channel carrying achromatic information for high spatial resolution. In
32 Webvision

Figure 31. Double opponent simple cells with orientation selectivity detect a blue object in a yellow background. A population of such
cells with slightly different orientation preferences detects a circular object.

addition the S cone channel was opposed by yellow light but not strongly enough to stop responses to white. The
S cone on ganglion cells appeared to be transmitting a signal that said that the S cones were absorbing light but
not that this light was "blue"; it could be white, grey as well. The so called white/black channel was receiving no
input from S cones which is not what is expected for a channel signaling white. The contribution of activation of
the S cone retinal ganglion cells is illustrate by the after images produced in Fig. 38.
Therefore the link between Hering's opponent color theory and retinal ganglion cell signals was scrapped.
Because the neurons in the lateral geniculate nucleus responded similarly to what was found in the retina,
attention went to striate cortex to explain opponent color theory by physiology. The initial ideas of channels
signaling color at the retinal level were wrong; the opponent signals detectable in the retina were cone opponent
and not color opponent. Color vision turned out to be more sophisticated, more arcane but more realistic.

The Future
Certainly the future will involve anatomical definition of how the projections from the lateral geniculate synapse
in striate cortex and begin to define the neurocircuitry of this hexalaminar cortical structure. The dendritic input
and axonal output of each functionally unique cell must be defined as has been done to a large degree in the
retina. This strategy must be repeated in higher areas of visual cortex, perhaps the most intriguing phase of this
crusade. New optical techniques that combine both anatomical identification and physiological responses of the
same cells could expedite this enormous task. But this is the only way to understand how the most complex
Color Vision 33

Figure 32. Spectral contrast based simple cells with orientation selectivity detect a white object in a yellow background. A population
of cells with slightly different orientation preferences detects a circular object. Note the synergism of L & S cones at one edge of the
simple cell.

machine in our universe, the cerebral cortex works. For this color vision is an ideal starting point because it has
carried us further into this organ than any other neural process.
34 Webvision

Figure 33. A stabilized retinal image of a red/green scene changes with time and its modification by additional stimulation (from
Iarbus, 1967).
Color Vision 35

Figure 34. A moving border affects neighboring midget-like ganglion cells when there is a change in wavelength as well as a large
change in energy contrast. When there is an increment in energy contrast the L on cell responds more strongly; when there is a
decrement of energy contrast the M off cell responds more strongly. Both contribute to the sensation or red.
36 Webvision

Figure 35. "Blobs" of V1 target slabs of cells in V2 which target "globs" of cells in V4 producing a hierarchy of chromatic processing
embedded in areas involved in achromatic vision (33-35).
Color Vision 37

Figure 36. More basic hue responsive cells could be used to create higher order hue responsive cells.
38 Webvision

Figure 37. Gothlin's concept of a two stage representation of the three component theory of man's color vision on the assumption that
the fundamental colors are red, green and blue but are organized into two antagonistic pairs. The color depicted here is an unsaturated
purple.
Color Vision 39

Figure 38. The after image of a white object is the opposing color to blue which is yellow. By fixating on the black spot on the left for a
minute and then glancing over to the right, one sees yellow, not black, candles on the tree.
40 Webvision

About the Author

Peter Gouras was born in Brooklyn, New York. He received an MD degree from Johns Hopkins University
School of Medicine and proceeded directly into a career in eye research at the National Institutes of Health, as
part of the Public Health Service, at first in the Neurological Institute and the Division of Blindness and later in
the National Eye Institute. Dr. Gouras is renowned in three areas of vision research, all of which he has been a
pioneer and leader. For most of his career, he has been active in the fields of basic and clinical research on the
ERG. In the late 1960s, he pioneered recordings of primate ganglion cells in the fovea and did pivotal work on
the analysis of color pathways from the retina through LGN to the visual cortex. On his moving to the faculty of
Harkness Eye Institute at Columbia Medical School, New York City in the mid-1970s, Dr. Gouras made daring
strides into the area of pigment epithelium transplantation for the treatment of retinal and macular disease. He is
the author of several hundred papers and many chapters and books, particularly concerning color vision and
retinal information processing.

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