CHAPTER 12Vision 197Old World monkeys, apes, and humans are trichromats, with
separate middle- and long-wave pigments—in all probability
because there was duplication of the ancestral long-wave gene
followed by divergence.
There are variations in the red, long-wave pigment in humans.
It has been known for some time that responses to the Rayleigh
match, the amounts of red and green light that a subject mixes to
match a monochromatic orange, are bimodal. This correlates
with new evidence that 62% of otherwise color-normal individu-
als have serine at site 180 of their long-wave cone opsin, whereas
38% have alanine. The absorption curve of the subjects with
serine at position 180 peaks at 556.7 nm, and they are more sen-
sitive to red light, whereas the absorption curve of the subjects
with alanine at position 180 peaks at 552.4 nm.
NEURAL MECHANISMS
Color is mediated by ganglion cells that subtract or add input
from one type of cone to input from another type. Processing in
the ganglion cells and the lateral geniculate nucleus produces im-
pulses that pass along three types of neural pathways that project
to V1: a red–green pathway that signals differences between L-
and M-cone responses, a blue–yellow pathway that signals dif-
ferences between S-cone and the sum of L- and M-cone respons-
es, and a luminance pathway that signals the sum of L- and M-
cone responses. These pathways project to the blobs and the
deep portion of layer 4C of V1. From the blobs and layer 4, color
information is projected to V8. However, it is not known how
V8 converts color input into the sensation of color.
OTHER ASPECTS OF
VISUAL FUNCTION
DARK ADAPTATION
If a person spends a considerable length of time in brightly
lighted surroundings and then moves to a dimly lighted envi-
ronment, the retinas slowly become more sensitive to light as
the individual becomes “accustomed to the dark.” This decline
in visual threshold is known as dark adaptation. It is nearly
maximal in about 20 minutes, although some further decline
occurs over longer periods. On the other hand, when one pass-
es suddenly from a dim to a brightly lighted environment, the
light seems intensely and even uncomfortably bright until the
eyes adapt to the increased illumination and the visual thresh-
old rises. This adaptation occurs over a period of about 5 min-
utes and is called light adaptation, although, strictly speaking,
it is merely the disappearance of dark adaptation.
The dark adaptation response actually has two components.
The first drop in visual threshold, rapid but small in magni-
tude, is known to be due to dark adaptation of the cones
because when only the foveal, rod-free portion of the retina is
tested, the decline proceeds no further. In the peripheral por-
tions of the retina, a further drop occurs as a result of adapta-
tion of the rods. The total change in threshold between the
light-adapted and the fully dark-adapted eye is very great.
Radiologists, aircraft pilots, and others who need maximal
visual sensitivity in dim light can avoid having to wait 20 min-
utes in the dark to become dark-adapted if they wear red gog-
gles when in bright light. Light wavelengths in the red end of
the spectrum stimulate the rods to only a slight degree while
permitting the cones to function reasonably well. Therefore, a
person wearing red glasses can see in bright light during the
time it takes for the rods to become dark-adapted.
The time required for dark adaptation is determined in part
by the time required to build up the rhodopsin stores. In
bright light, much of the pigment is continuously being bro-
ken down, and some time is required in dim light for accumu-
lation of the amounts necessary for optimal rod function.
However, dark adaptation also occurs in the cones, and addi-
tional factors are undoubtedly involved.CRITICAL FUSION FREQUENCY
The time-resolving ability of the eye is determined by measur-
ing the critical fusion frequency (CFF), the rate at which
stimuli can be presented and still be perceived as separate
stimuli. Stimuli presented at a higher rate than the CFF are
perceived as continuous stimuli. Motion pictures move be-
cause the frames are presented at a rate above the CFF, and
movies begin to flicker when the projector slows down.VISUAL FIELDS & BINOCULAR VISION
The visual field of each eye is the portion of the external world
visible out of that eye. Theoretically, it should be circular, but
actually it is cut off medially by the nose and superiorly by the
roof of the orbit (Figure 12–21). Mapping the visual fields is
important in neurologic diagnosis. The peripheral portions of
the visual fields are mapped with an instrument called a pe-
rimeter, and the process is referred to as perimetry. One eye
is covered while the other is fixed on a central point. A smallFIGURE 12–20 Absorption spectra of the three cone
pigments in the human retina. The S pigment that peaks at 440 nm
senses blue, and the M pigment that peaks at 535 nm senses green.
The remaining L pigment peaks in the yellow portion of the spectrum,
at 565 nm, but its spectrum extends far enough into the long wave-
lengths to sense red. (Reproduced with permission from Michael CR: Color
vision. N Engl J Med 1973;288:724.)
500 600 700 nm
Wavelength400Absorption