CHAPTER 5.
1 - VISUAL CODING surface of the eye, which is lined with visual
 How far an ant sees, or how far you see, depends
on how far the light travels before it strikes the eyes.
You do not send out “sight rays.”
 Ibn al-Haytham
- an arab philosopher
- observed that when you open your eyes at night,
you immediately see the distant stars. He reasoned
that if you saw by sending out sight rays, they
couldn’t get to the stars that fast. Then he
demonstrated that light rays bounce off an object in
all directions, but you see only those rays that reflect
off the object and strike your retina.
 When you see a tree, for example, your perception
is not in the tree. It is in your brain. You see
something only when light from the object alters your
brain activity. Even if you did send out rays from your
eyes— and you don’t—when they struck some
object, you wouldn’t know about it, unless they
bounced back and returned to your eyes. Similarly,
you feel something only when touch information
reaches your brain. When you feel something with
your fingers, you don’t feel it in your fingers. You feel
it in your brain.
GENERAL PRINCIPLES OF PERCEPTION
 You see an object when it emits or reflects light that
stimulates receptors that transmit information to your
brain.
 René Descartes
- believed that the nerves from the eye would send
the brain a pattern of impulses arranged like a
picture of the perceived object, right side up. In fact,
your brain encodes the information in a way that
doesn’t resemble what you see.
 The brain codes information largely in terms of which
neurons are active, and how active they are at any
moment.
 Law Of Specific Nerve Energies
- by Johannes Müller
- Müller held that whatever excites a particular nerve
establishes a special kind of energy unique to that
nerve. In modern terms, the brain somehow
interprets the action potentials from the auditory
nerve as sounds, those from the olfactory nerve as
odors, and so forth.
THE EYE AND ITS CONNECTIONS TO THE BRAIN
 Light enters the eye through an opening in the
center of the iris called the pupil.
 It is focused by the lens (adjustable) and cornea (not
adjustable) and projected onto the retina, the rear
receptors.
 Light from the left side of the world strikes the right
half of the retina, and vice versa. Light from above
strikes the bottom half of the retina, and light from
below strikes the top half.
ROUTE WITHIN THE RETINA
 In the vertebrate retina, messages go from the
receptors at the back of the eye to bipolar cells,
located closer to the center of the eye.
 The bipolar cells send their messages to ganglion
cells, located still closer to the center of the eye.
 The ganglion cells’ axons join together and travel
back to the brain.
 Additional cells called amacrine cells get
information from bipolar cells and send it to other
bipolar, amacrine, and ganglion cells. Amacrine cells
refine the input to ganglion cells, enabling certain
ones to respond mainly to particular shapes,
directions of movement, changes in lighting, color,
and other visual features.
 One consequence of this anatomy is that light
passes through the ganglion, amacrine, and bipolar
cells en route to the receptors.
 The ganglion cell axons join to form the optic nerve
that exits through the back of the eye. The point at
which it leaves (also where the blood vessels enter
and leave) is a blind spot because it has no
receptor.
FOVEA AND PERIPHERY OF THE RETINA
 Fovea - a tiny area specialized for acute, detailed
vision.
 For perceiving detail, each receptor in the fovea
connects to a single bipolar cell, which in turn
connects to a single ganglion cell that has an axon
to the brain.
 Midget ganglion cells - the ganglion cells in the
fovea of humans and other primates.
 Many bird species have two foveas per eye, one
pointing ahead and one pointing to the side. Hawks
and other predatory birds have a greater density of
visual receptors on the top half of their retinas
(looking down) than on the bottom half (looking up).
Conversely, many prey species such as rats have
most of their receptors on the bottom half of the
retina, enabling them to see up better than they see
down.
 Toward the periphery of the retina, more and more
receptors converge onto bipolar and ganglion cells.
As a result, the brain cannot detect the exact
location or shape of a peripheral light source.
 In short, foveal vision has better acuity (sensitivity to
detail), and peripheral vision has better sensitivity to
dim light.
 In the periphery, your ability to detect detail is limited
by interference from other nearby objects.
VISUAL RECEPTORS: RODS AND CONES
The vertebrate retina contains two types of receptors:
rods and cones.
 Rods - abundant in the periphery of the human
retina, respond to faint light but are not useful in
daylight because bright light bleaches them.
 Cones - abundant in and near the fovea, are less
active in dim light, more useful in bright light, and
essential for color vision. Because of the distribution
of rods and cones, you have good color vision in the
fovea but not in the periphery.
 Although rods outnumber cones by about 20 to 1 in
the human retina, cones provide about 90 percent of
the brain’s input.
 People vary substantially in the number of axons in
their optic nerve and the size of the visual cortex,
largely for genetic reasons.
 Both rods and cones contain photopigments,
chemicals that release energy when struck by light.
 Photopigments consist of 11-cis-retinal (a derivative
of vitamin A) bound to proteins called opsins, which
modify the photopigments’ sensitivity to different
wavelengths of light.
 Light converts 11-cis-retinal to all-trans-retinal, thus
releasing energy that activates second messengers
within the cell.
COLOR VISION
 Visible light consists of electromagnetic radiation
within the range from less than 400 nm (nanometer,
or 10–9 m) to more than 700 nm.
 We perceive the shortest visible wavelengths as
violet. Progressively longer wavelengths are
perceived as blue, green, yellow, orange, and red.
 We call these wavelengths “light” only because the
receptors in our eyes are tuned to detecting them.
 Many species of birds, fish, and insects have visual
receptors sensitive to what we call ultraviolet
radiation.
THE TRICHROMATIC (YOUNG-HELMHOLTZ)
THEORY
 Thomas Young
- Young was the first to start deciphering the Rosetta
stone. He also founded the modern wave theory of
light, defined energy in its modern form, founded the
calculation of annuities, introduced the coefficient of
elasticity, discovered much about the anatomy of the
eye, and made major contributions to other fields.
- Young recognized that color required a biological
explanation. He proposed that we perceive color by
comparing the responses across a few types of
receptors, each of which was sensitive to a different
range of wavelengths.
 This theory, later modified by Hermann von
Helmholtz, is now known as the trichromatic
theory of color vision, or the Young-Helmholtz
theory.
- According to this theory, we perceive color through
the relative rates of response by three kinds of
cones, each one maximally sensitive to a different
set of wavelengths. (Trichromatic means “three
colors.”)
- How did Helmholtz decide on the number three?
He found that people could match any color by
mixing appropriate amounts of just three
wavelengths. Therefore, he concluded that three
kinds of receptors—we now call them cones—are
sufficient to account for human color vision.
 According to the trichromatic theory, we discriminate
among wavelengths by the ratio of activity across
the three types of cones.
 Your perception of color depends on the frequency
of response in each cell relative to the frequency of
other cells.
 The nervous system determines the color of the light
by comparing the responses of different types of
cones.
 Long- and medium-wavelength cones are far more
abundant than short-wavelength (blue) cones.
 Consequently, it is easier to see tiny red, yellow, or
green dots than blue dots.
 In the retina’s periphery, cones are so scarce that
you have no useful color vision.
 Visual field - the part of the world that you see
before you can identify the color.
THE OPPONENT-PROCESS THEORY
 Stare at an object (in figure 5.11) it under a bright
light, without moving your eyes, for a minute. (The
brighter the light and the longer you stare, the
stronger the effect.) Then look at a plain white
surface, such as a wall or a blank sheet of paper.
Keep your eyes steady. You will see a negative
color afterimage, a replacement of the red you had
been staring at with green, green with red, yellow
and blue with each other, and black and white with
each other.
 Opponent-Process Theory
- proposed by Ewald Hering
- We perceive color in terms of opposites. That is,
the brain has a mechanism that perceives color on a
continuum from red to green, another from yellow to
blue, and another from white to black.
- Part of the explanation for this process pertains to
the connections within the retina. For example,
imagine a bipolar cell that receives excitation from a
short-wavelength cone and inhibition from long- and
medium-wavelength cones. It increases its activity in
response to short-wavelength (blue) light and
decreases it in response to yellowish light. After
prolonged exposure to blue light, the fatigued cell
decreases its response. Because a low level of
response by that cell usually means yellow, you
perceive yellow.
THE RETINEX THEORY
 The trichromatic theory and the opponent-process
theory cannot easily explain color constancy, the
ability to recognize colors despite changes in
lighting.
 If you wear green-tinted glasses or replace your
white light bulb with a green one, you still identify
bananas as yellow, paper as white, and so forth.
Your brain compares the color of one object with the
color of another, in effect subtracting a certain
amount of green from each.
 To illustrate, examine Figure 5.13 (Purves & Lotto,
2003). Although different colors of light illuminate the
two objects at the top, you easily identify the squares
as red, yellow, blue, and so forth. Note the result of
removing context. The bottom part shows the
squares that looked blue in the top left part and
yellow in the top right part. Without the context that
indicated yellow light or blue light, all these squares
look gray. For this reason, we should avoid talking
about the color of a wavelength of light. A certain
wavelength of light can appear as different colors
depending on the background.
 Similarly, we perceive the brightness of an object by
comparing it to other objects. The object in the
center appears to have a dark gray top and a white
bottom. Now cover the border between the top and
the bottom with a finger. You see that the top of the
object has exactly the same brightness as the
bottom!

 To account for color and brightness constancy,
Edwin Land proposed the retinex theory (a
combination of the words retina and cortex): The
cortex compares information from various parts of
the retina to determine the brightness and color for
each area.
 Dale Purves and colleagues have expressed a
similar idea in more general terms: Whenever we
see anything, we make an inference. For example,
when you look at the objects in Figures 5.13 and
5.14, you ask yourself, “On occasions when I have
seen something that looked like this, what was it
really?” You go through the same process for
perceiving shapes, motion, or anything else. That is,
visual perception requires reasoning and inference,
not just retinal stimulation.
COLOR VISION DEFICIENCY
 Color vision deficiency - colorblindness
 Color is in the brain, not in the light or the object
itself.
 Color deficiency results because people with certain
genes fail to develop one type of cone, or develop
an abnormal type of cone.
 In red-green color deficiency, the most common
form of color deficiency, people have trouble
distinguishing red from green because their long-
and medium-wavelength cones have the same
photopigment instead of different ones. The gene
causing this deficiency is on the X chromosome.
 Women with one normal gene and one color-
deficient gene—and that includes all women with a
red-green color-deficient father—are slightly less
sensitive to red and green than the average for other
people.
 What would happen if people had a fourth type of
cone? Actually, some women do, in a way. The long-
wavelength cone shows genetic variation. At one
point in the protein, most genes code for the amino
acid serine but 16 to 38 percent of the genes
(depending on people’s ethnic background) produce
instead the amino acid alanine. Because the gene is
on the X chromosome, a man has only one or the
other. However, because women have two X
chromosomes, some women have one long-
wavelength receptor with serine and one with
alanine. Those two versions of the long-wavelength
receptor differ slightly in their responsiveness to
light. Women with different versions of that receptor
make somewhat finer distinctions between one color
and another, compared to other people. Because
some women have two types of long-wavelength
receptors and others have just one, women’s
performance on color vision tests is more variable
than men’s is.
SUMMARY:
 You see because light strikes your retina, causing it
to send a message to your brain. You send no sight
rays out to the object.
 According to the law of specific nerve energies, the
brain interprets any activity of a given sensory
neuron as representing a particular type of sensory
information.
 Sensory information is coded so that the brain can
process it. The coded information bears no physical
similarity to the stimuli it describes.
 Light passes through the pupil of a vertebrate eye
and stimulates the receptors lining the retina at the
back of the eye.
 The axons from the retina loop around to form the
optic nerve, which exits from the eye at a point
called the blind spot.
 Visual acuity is greatest in the fovea, the central
area of the retina. Because so many receptors in the
periphery converge their messages to their bipolar
cells, our peripheral vision is highly sensitive to faint
light but poorly sensitive to detail.
 The retina has two kinds of receptors: rods and
cones. Rods, more numerous in the periphery of the
retina, are more sensitive to faint light. Cones, more
numerous in the fovea, are more useful in bright
light.
 People vary in their number of axons from the retina
to the brain. Those with more axons show a greater
ability to detect brief, faint, or rapidly changing
stimuli.
 According to the trichromatic (or Young-Helmholtz)
theory of color vision, color perception begins with a
given wavelength of light stimulating a distinctive
ratio of responses by the three types of cones.
 According to the opponent-process theory of color
vision, visual system neurons beyond the receptors
respond with an increase in activity to indicate one
color of light and a decrease to indicate the opposite
 color. The three pairs of opposites are red-green,
yellow-blue, and white-black.
 According to the retinex theory, the cortex compares
the responses across the retina to determine
brightness and color of each object.
 For genetic reasons, certain people are unable to
distinguish one color from another. Red-green color
deficiency is the most common type.