We take content rights seriously. If you suspect this is your content, claim it here.
Available Formats
Download as PDF or read online on Scribd
You are on page 1/ 8
CHAPTER 23
Color vision is the ability to discriminate a light
stimulus as a function of its wavelength. Various
sensory and cognitive processes combine to result
in the sense of color. The cooperative physical ef-
fects of light and object, the physiologic reaction of
the visual organ to light, and the psychologic con-
text of color perception together produce the pic-
ture of our surroundings, thus influencing our re-
lationship and attitude toward our environment.
This chapter focuses specifically on the reaction of
the retina to light stimulus. The details of physical
events outside of the body known as optical physies,
theories of color, and the higher-order processing,
of the visual signal in the brain to produce visual
experience are beyond the scope of this chapter.
COoLoR AND LIGHT
Electromagnetic energy is wavelengths between
approximately 380 and 760 nm and causes pho-
toreactions on the human retina, which leads to
the experience of vision, Although the perceptions
caused by light waves cannot be directly measured,
optical physics describes the origin of colors as a
breaking down of light into its spectrat con-
stituents.’ A prism, a transparent solid body with a
triangular cross section, causes white, or neutral,
light to be refracted so that it is divided into the
spectrum of rainbow colors. Short-wavelength vis
ible light causes the sensation of violet, and in an
uneven transition, the colors blue, blue-green,
green, yellow-green, yellow, orange, and red are
perceived. For example, red does not noticeably
change in perception from approximately 680 nm
onward. Monochromatic light is colored light of a
single wavelength. The remixing of all colors of
light created by a prism, for example with a convex
COLOR VISION...
THomas P. SAKMAR
578
lens, will create the sensation of white, The sensa~
tion of white can also be created from the mono-
chromatic light rays from the short-, middle-, and
long-wavelength zones of the spectrum. For exam-
ple, mixing blue (435 nm), green (545 nm), and
red (700 nm) light produces white. These three
light rays can also be combined to create any other
color by changing the relative intensities of the in-
dividual components. Thus violet-blue, green, and
red are called the additive primary colors.
The light rays from the mixing of any two thirds
of the spectrum cause the sensations of yellow, ma-
genta-red, and cyan-blue, ‘These three colors are the
so-called subtractive primary colors. Magenta-red,
the mixing of rays of the short- and long-wavelength
ends of the spectrum, does not itself exist in the nat-
ural spectrum. Any two colors are called complemen-
rary colors if their additive mixing forms white.
‘Therefore it follows that blending any one additive
primary color with the corresponding subtractive
primary color forms the whole spectrum, The mix-
ing of color pigments or dyes is an example of sub-
tractive color mixing. The mixing of a cyan-blue
dye and a yellow dye produces green. The cyan-
blue dye absorbs, or subtracts, the long-wavelength.
part of the spectrum; the yellow dye absorbs the
short-wavelength part. Because both dyes reflect the
middle-wavelength light, green color is appreciated.
Color mixing (subtractive, additive, or propor-
tional mixing of pairs of the six primary colors)
produces the nearly limitless range of color hues
that can be perceived. In addition to additive or
subtractive mixing, color can be produced through
the scatter of white light. One example is the deep
blue-colored sky that is apparent at midday in the
summer when the air is clean and dry. The sun's
rays pass vertically through the earth’s atmosphereChapter 25
and the longer wavelengths are scattered. However,
al sunrise and sunset the siua’s rays fall on an acute
angle and their pathlength is longer, giving rise 10
intense red calor depending on prevailing atmo-
spheric conditions. Finally, color can result from in-
terference on thin films. Color perception changes
swith slight changes in the thickness of a film or with
the angle of vision, The mother-of-pearl color of
soap bubbles oF the feathers of iridescent birds are
examples of so-called interference colors.
The human eye differentiates colors according
to the color itself (its wavelength), the brightness,
and the saturation. Therefore a systematic organi-
zation of all colors would be possible only in the
three-dimensional system. There is no perceptual
basis for a hierarchy of colors, but color wheels gen-
erally are used to organize and group chromatic
colors according to their appearance,
The existence of color is at some level a topic
that is equally relevant to philosophy and science.
Throughout recorded history, great philosophers
and scientists have written on theories of color and
color sense and have provided @ rich literature on
the subject. Isaac Newton (1642-1727) carried out
experiments with a prism that transformed the sci-
ence of color from the study of objects to the study
of light, jobann Wollgang von Goethe (1749-1832)
wrote extensively on the experience of color,
‘Thomas Young (1773-1829) proposed a theory of
color vision based on three receptors in the retina
that are sensitive to different spectral regions.
David Brewster (1781-1868) introduced the term
color blindness, which was formerly known as dal-
tonisin, afier John Dalton (3766-1844), who de-
scribed in detail his own inability to distinguish
red. The genetic basis of Dalton’s color blindness
was recently determined, by polymerase chain reac
tion analysis of DNA extracted from his preserved
postmortem eye, to be deuteranopia (see following
discussion)’ Arthur Schopenhauer (1788-1860)
expanded on the work of Goethe and Immanuel
Kant (1724-1804). Joseph Antoine Ferdinand
Plateau (1801-1883) studied afterimages and color
mixing and proposed the Talbot-Plateau law of
color intensity perception. Hermann Ludwig
Ferdinand von Helmholtz (1821-1894) extended
Young's hypothesis, henceforth called the Young-
Helmholiz hypothesis, and devised specteal absorp-
tion curves for three visual photoreceptors.
Herman Rudolf Aubert (1826-1892) was one of the
key contributors to physiologic optics in addition
to measuring absolute visual sensitivity and light
and dark adaptation. He demonstrated that color
ce
LOR VISION 579
perception was largely restricted to the foveal re-
gion and depended on context in other parts of the
retina, James Clerk Maxwell (1831-1879) was
largely responsible for making the study of color vi-
sion 4 quantitative science. He devised methods to
study additive and subtractive color mixing and
color-defective subjects and developed many of the
classifications that are still used today.
BIOCHEMISTRY OF COLOR VISION
All human visual pigments share a common chro-
mophore, which is chemically related to vitamin A,.
‘The 11-cis-structural isomer of the aldehyde of vit-
amin A, reacts with an opsin protein to form a pho-
toreceptor-pigment complex. The rod and cone cell
pigments are all complexes of the same chro-
mophore with different, but related, opsin proteins.
Interestingly, the genes encoding the opsin proteins
ate members of a superfamily of related receptors
called G protein-coupled receptors, These receptors
are involved in different sensory and intercellular
signaling pathways across a wide range of organ-
isms, The receptors are integral membrane proteins
that all share a common structural motif—seven-
transmembrane segments—and they all communi-
cate with cellular biochemical signal-transducer
proteins in the cellular cytoplasm called het-
erotrimeric G proteins, In the case of tod and cone
calls, the G proteins are called transducins. Specific
forms of transducins are found in rods and cones,
although the three cone cell types share a common
form of transducin.
The unique properties of the chromophore in its
opsin-bound state contribute to the key physiologic
properties of vision, including color vision. Spectral
tuning ata molecular level is related to the so-called
opsin-shift. The opsin-shift refers to the change in
the absorption of the chromophore when it be~
comes bound to a particular opsin, The magnitude
of the opsin-shift varies with each particular visual
pigment.
In the biochemical amplification cascade of the
photoreceptor cells of the vertebrate visual system,
the capture of a photon causes irreversible photo-
chemical isomerization of the chromophore in the
visual pigment. A series of protein conformational
changes ensues, and the active pigment becomes a
catalyst that converts transducin into its active
form. Active transducin modulates cyclic guano
sine monophosphate (GMP) phosphodiesterase,