Chapter 5. Vision

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Vision
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Module 5.1 Visual Coding

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.” That
principle was probably the first scientific discovery in psychology
(Steffens, 2007)
What was Ibn al-Haytham’s evidence that we see only because light
enters the eyes, not by sending out sight rays? → First, you can see
distant objects such as stars far faster than we could imagine any sight rays
reaching them. Second, when light strikes an object, we see only the light
rays that reflect off the object and into the eyes.
General Principles of Perception

How does your brain make sense of that information? The 17th-century
philosopher 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 1838, Johannes Müller described this insight as the law of specific
nerve energies. Müller held that whatever excites a particular nerve
establishes a special kind of energy unique to that nerve.
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1/25/24, 2:38 PM Chapter 5. Vision
If someone electrically stimulated the auditory receptors in your ear,

what would you perceive? → Because of the law of specific nerve
energies, you would perceive it as sound, not as shock. (Of course, a
strong enough shock might spread far enough to excite pain receptors also.
If it were possible to flip your entire brain upside down, without breaking
any of the connections to sense organs or muscles, what would happen
to your perceptions of what you see, hear, and so forth? → Your
perceptions would not change. The way visual or auditory information is
coded in the brain does not depend on the physical location within the
brain. Seeing something as “on top” or “to the left” depends on which
neurons are active but does not depend on the physical location of those
neurons
The Eye and Its Connections to the Brain

Light enters the eye through an opening in the center of the iris called the
pupil
Retina, the rear surface of the eye, which is lined with visual receptors
Remember, the visual system does not duplicate the image. It codes it by
various kinds of neuronal activity.
Route within the Retina

In the vertebrate retina, however, messages go from the receptors at the
back of the eye to bipolar cells, located closer to the center of the eye
(see Figure 5.2). 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.
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
What makes the blind spot of the retina blind? → The blind spot has no
receptors because it is occupied by exiting axons and blood vessels.
Fovea and Periphery of the Retina

the fovea (meaning “pit”), a tiny area specialized for acute, detailed vision
More importantly 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.
The ganglion cells in the fovea of humans and other primates are called
midget ganglion cells because each is small and responds to just a single
cone.
Header 2
Visual Receptors: Rods and Cones
The vertebrate retina contains two types of receptors: rods and cone
The 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.
Although rods outnumber cones by about 20 to 1 in the human retina,

cones provide about 90 percent of the brain’s input
Remember the midget ganglion cells: In the fovea, each cone has its own
line to the brain. In the periphery (mostly rods), each receptor shares a line
with tens or hundreds of others. Overall, 120 million rods and 6 million
cones converge onto 1 million axons in the optic nerve, on average
Both rods and cones contain photopigments, chemicals that release
energy when struck by light.
You sometimes find that you can see a faint star on a dark night better if
you look slightly to the side of the star instead of straight at it. Why? → If
you look slightly to the side, the light falls on an area of the retina with more
rods and more convergence of input.
If you found a species with a high ratio of cones to rods in its retina, what
would you predict about its way of life? → We should expect this species
to be highly active during the day and seldom active at night
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
fish, and insects have visual receptors sensitive to what we call ultraviolet
radiation
In some species of birds, the male and female look alike to us, but different
to birds, because the male reflects more ultraviolet light.
The Trichromatic (Young-Helmholtz) Theory

Thomas Young (1773–1829). 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
According to the trichromatic theory, we discriminate among wavelengths
by the ratio of activity across the three types of cones. For example, light
at 550 nm excites the medium wavelength and long-wavelength receptors
about equally and the short-wavelength receptor almost not at all.
The smaller the dot, the farther you have to move it into your visual field—
that is, the part of the world that you see—before you can identify the
color
The Opponent-Process Theory

Ewald Hering, a 19th-century physiologist, proposed the opponent-
process theory: 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. After you stare at one color in one location long enough, you fatigue
that response and swing to the opposite
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
This demonstration suggests that afterimages depend on the whole
context, not just the light on individual receptors. The cerebral cortex must
be responsible, not the bipolar or ganglion cells.
Examine Figure 5.9. According to the trichromatic theory, what causes

you to perceive red? → Activity of the long-wavelength cone is not
sufficient. In fact, notice that the long-wavelength cone responds to what
we call yellow more than to what we call red. A perception of red occurs
only if the long-wavelength cone has a high ratio of response relative to
the other two types of cone
According to the opponent-process theory, under what circumstance

would you perceive a white object as blue? → If you stared at a bright
yellow object for a minute or so and then looked at a white object it would
appear blue.
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.
To account for color and brightness constancy, Edwin Land proposed the
retinex theory The cortex compares information from various parts of the
retina to determine the brightness and color for each area.
When a television set is off, its screen appears gray. When you watch a
program, parts of the screen appear black, even though more light is
actually showing on the screen than when the set was off. What
accounts for the black perception? → The black experience arises by
contrast with the brighter areas around it.
Figure 5.9 shows light at about 510 nm as green. Why should we
nevertheless not call it “green light”? → Color perception depends not just
on the wavelength of light from a given spot but also the light from
surrounding areas. As in Figure 5.13, the context can change the color
perception.
Color Vision Deficiency

One of the first discoveries in psychology was colorblindness, better
described as color vision deficiency
Color deficiency results because people with certain genes fail to develop
one type of cone, or develop an abnormal type of cone
the most common form of color deficiency, people have trouble
distinguishing red from green because their long- and mediumwavelength
cones have the same photopigment instead of different ones
Why is color vision deficiency a better term than color blindness?

→Very few people see the world entirely in black and white. The more
common condition is difficulty discriminating red from green.
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.
Header 1
Module 5.2 How the Brain Processes Visual
Information
Vision is complicated. We shall go through it in some detail, for two
reasons:
First, without vision and other senses, you would have no more mental
experience than a tree does. Everything in psychology starts with
sensations.
Second, neuroscientists have investigated vision in more detail than
anything else that the brain does. Examining the mechanisms of vision
illustrates what it means to explain something in biological terms.
An Overview of the Mammalian Visual System

The rods and cones of the retina make synapses with horizontal cells and
bipolar cells
Most ganglion cell axons go to the lateral geniculate nucleus, part of the
thalamus. (The term geniculate comes from the Latin root genu, meaning
“knee.” To genuflect is to bend the knee.
The lateral geniculate, in turn, sends axons to other parts of the thalamus
and the visual cortex. Axons returning from the cortex to the thalamus
modify thalamic act
Where does the optic nerve start and where does it end? → It starts with
the ganglion cells in the retina. Most of its axons go to the lateral geniculate
nucleus of the thalamus, but some go to the hypothalamus and superior
colliculus
Processing in the Retina

To understand how the wiring diagram of your retina highlights those
patterns, let’s start by exploring one example in detail: lateral inhibition
Lateral inhibition is the retina’s way of sharpening contrasts to emphasize
the borders of objects.
light striking the rods and cones decreases their spontaneous output, and
the receptors make inhibitory synapses onto the bipolar cells. Therefore,
light on the rods or cones decreases their inhibitory output.
lateral inhibition, the reduction of activity in one neuron by activity in
neighboring neurons (Hartline, 1949). Lateral inhibition heightens contrast.
Lateral inhibition is important for many functions in the nervous system.
When light strikes a receptor, does the receptor excite or inhibit the
bipolar cells? What effect does it have on horizontal cells? What effect
does the horizontal cell have on bipolar cells? → The receptor excites
both the bipolar cells and the horizontal cell. The horizontal cell inhibits the
same bipolar cell that was excited plus additional bipolar cells in the
surround.
If light strikes only one receptor, what is the net effect (excitatory or
inhibitory) on the nearest bipolar cell that is directly connected to that
receptor? What is the effect on bipolar cells to the sides? What causes
that effect? → It produces more excitation than inhibition surround. for
the nearest bipolar cell. For surrounding bipolar cells, it produces only
inhibition. The reason is that the receptor excites a horizontal cell, which
inhibits all bipolar cells in the area.
Examine Figure 5.17. You should see grayish diamonds at the crossroads
among the black squares. Explain why → In the parts of your retina that
look at the in the area. long white arms, each neuron is inhibited by white
input on two of its sides (either above and below or left and right). In the
crossroads, each neuron is in hibited by input on all four sides. Therefore,
the response in the crossroads is decrease compared to that in the arms.
Further Processing
Each cell in the visual system of the brain has a receptive field, an area in
visual space that excites or inhibits it.
The receptive field of a rod or cone is simply the point in space from which
light strikes the cell.
Receptive fields The receptive field of any neuron in the visual system is
the area of the visual field that excites or inhibits it. Receptors have tiny
receptive fields and later cells have progressively larger receptive fields
Prime ganglion cells fall into three categories:
The parvocellular neurons, with small cell bodies and small receptive fields,
are mostly in or near the fovea. (Parvocellular means “small celled,” from
the Latin root parv, meaning “small.”)
The magnocellular neurons, with larger cell bodies and receptive fields, are
distributed evenly throughout the retina. (Magnocellular means “large
celled,” from the Latin root magn, meaning “large.” The same root appears
in magnify.)
The koniocellular neurons have small cell bodies, similar to the parvocellular
neurons, but they occur throughout the retina. (Koniocellular means “dust
celled,” from the Greek root meaning “dust.” They got this name because of
their granular appearance.)
The high sensitivity to detail and color relates to the fact that parvocellular
cells are located mostly in and near the fovea, which has many cones
Magnocellular neurons are found throughout the retina, including the
periphery.
Axons from the ganglion cells form the optic nerve, which proceeds to the
optic chiasm, where half of the axons (in humans) cross to the opposite
hemisphere. Most of the axons go to the lateral geniculate nucleus of the
thalamus.
As we progress from bipolar cells to ganglion cells to later cells in the

visual system, are receptive fields ordinarily larger, smaller, or the same
size? Why? → They become larger because each cell’s receptive 16. field
is made by inputs converging at an earlier level
What are the differences between the parvocellular and magnocellular

systems? → Neurons of the parvocellular system have small cell 17. bodies
with small receptive fields, are located mostly in and near the fovea, and
are specialized for detailed and color vision. Neurons of the magnocellular
system have large cell bodies with large receptive fields, are located in all
parts of the retina, and are specialized for perception of large patterns and
movement
The Primary Visual Cortex

Information from the lateral geniculate nucleus of the thalamus goes to the
primary visual cortex in the occipital cortex, also known as area V1 or the
striate cortex because of its striped appearance.
If you see an optical illusion, the activity in area V1 corresponds to what you
think you see, not what the object really is
People with damage to area V1 report no conscious vision, no visual
imagery, and no visual images in their dreams In contrast, adults who lose
vision because of eye damage continue to have visual imagery and visual
dreams.
blindsight, the ability to respond in limited ways to visual information
without perceiving it consciously
The research supports two explanations for blindsight:

First, in some cases, small islands of healthy tissue remain within an
otherwise damaged visual cortex, not large enough to provide conscious
perception but enough to support limited visual function.
Second, the thalamus sends visual input to several other brain areas,
including parts of the temporal cortex.
Even if your brain is intact, you can experience something like blindsight
under certain circumstances .
In this procedure, known as continuous flash suppression, a viewer is
conscious of the rapidly changing stimuli and not the steady picture
Simple and Complex receptive fields

In the 1950s, David Hubel and Torsten Wiesel (1959) inserted thin
electrodes to record activity from cells in cats’ and monkeys’ occipital
cortex while they shined light patterns on the retina
They wondered why cells were unresponsive, they knew the occipital
cortex was essential for vision. Then they noticed a big response while they
were moving a slide into place. They quickly realized that the cell was
responding to the edge of the slide.
Their research, for which they received a Nobel Prize, has often been called
“the research that launched a thousand microelectrodes” because it
inspired so much further research.
Figure 5.19 illustrates the receptive field of a simple cell. A simple cell has
a receptive field with fixed excitatory and inhibitory zones. The more light
shines in the excitatory zone, the more the cell responds. The more light
shines in the inhibitory zone, the less the cell responds.
complex cells, located in areas V1 and V2, do not respond to the exact
location of a stimulus. A complex cell responds to a pattern of light in a
particular orientation (e.g., a vertical bar) anywhere within its large
receptive field (see Figure 5.20). Most complex cells respond most strongly
to a stimulus moving in a particular direction—for example, a vertical bar
moving horizontally. The best way to classify a cell as simple or complex is
to present the stimulus in several locations. A cell that responds to a
stimulus in only one location is a simple cell. One that responds equally
throughout a large area is a complex cell
End-stopped, or hypercomplex, cells resemble complex cells with one
exception: An end-stopped cell has a strong inhibitory area at one end of
its bar-shaped receptive field. The cell responds to a bar-shaped pattern of
light anywhere in its broad receptive field, provided the bar does not
extend beyond a certain point
How could a researcher determine whether a given neuron in the visual

cortex is simple or complex? → First identify a stimulus, such as a
horizontal line, that stimulates the cell. Then present the stimulus in several
locations. If the cell responds strongly in only one location, it is a simple
cell. If it responds in several locations, it is a complex cell.
The Columnar Organization of the Visual Cortex
Cells with similar properties group together in the visual cortex in columns
perpendicular to the surface.
For example, cells within a given column might respond to only the left eye,
only the right eye, or both eyes about equally. Also, cells within a given
column respond best to lines of a single orientation.
Electrode path B, which is not perpendicular to the surface of the cortex,
crosses through columns and encounters cells with different properties. In
short, the cells within a given column process similar information.
Are Visual Cortex Cells Feature Detectors?
feature detectors—neurons whose responses indicate the presence of a

particular feature.
Supporting the idea of feature detectors is the fact that prolonged
exposure to a given visual feature decreases sensitivity to that feature, as if
it fatigued the relevant detectors.
For example, if you stare at a waterfall for a minute or more and then look
to the side, the rocks and trees next to the waterfall appear to flow upward
This waterfall illusion suggests that you have fatigued the neurons that
detect downward motion, leaving unopposed the detectors for the
opposite motion
Development of the Visual Cortex

How do cells in the visual cortex develop their properties?
Waves of spontaneous activity sweep over the developing retina,
synchronizing the activity of neighboring receptors and enabling
appropriate combinations of receptors to establish connections with cells in
the brain
What about connections beyond the primary visual cortex? A study of
people who were born without eyes found that the connections from the
primary visual cortex to its main targets were more or less normal (Bock et
al., 2015). Evidently, certain axon paths develop automatically, without any
need for guidance by experience. Nevertheless, visual experience after
birth modifies and fine-tunes many of the connections
Deprived Experience in One Eye

What would happen if a young animal could see with one eye but not the
other? When a kitten opens its eyes at about age 9 days, each neuron
responds to areas in the two retinas that focus on approximately the same
point in space—a process necessary for binocular vision.
If an experimenter sutures one eyelid shut for a kitten’s first 4 to 6 weeks of
life, synapses in the visual cortex gradually become unresponsive to input
from the deprived eye. After the deprived eye is opened, the kitten does
not respond to it.
After an eye deprived of vision in adults is reopened, cells gradually return

to their previous levels of responsiveness
Deprived Experience in Both Eyes

If both eyes stayed shut for the first few weeks, what would you expect?
You might guess that the kitten would become insensitive to both eyes, but
it does not. When just one eye is open, the synapses from the open eye
inhibit the synapses from the closed eye
If the eyes remain shut still longer, the cortical responses start to become
sluggish and lose their well-defined receptive fields
For each aspect of visual experience, researchers identify a sensitive
period, when experiences have a particularly strong and enduring influence
The sensitive period depends on inhibitory neurons
a prolonged experience— such as a full week without visual stimulation to
one eye— produces a smaller but measurable effect on the visual cortex
What is the effect of closing one eye early in life? What is the effect of
closing both eyes → If one eye is closed during early development, the
cortex becomes unresponsive to it. If both eyes are closed, cortical cells
remain somewhat responsive for several weeks and then gradually become
sluggish and unselective in their responses
Uncorrelated Stimulation in the Two Eyes

Stereoscopic depth perception requires the brain to detect retinal
disparity, the discrepancy between what the left and right eyes see.
Experience fine-tunes binocular vision, and abnormal experience disrupts
it.
Imagine a kitten with weak or damaged eye muscles so that its eyes do not
point in the same direction. Both eyes are active, but no cortical neuron
consistently receives messages from one eye that match messages from
the other eye. Each neuron in the visual cortex becomes responsive to one
eye or the other, and few neurons respond to both
A similar phenomenon occurs in humans. Certain children are born with

strabismus (or strabismic amblyopia), also known as “lazy eye,” a condition
in which the eyes do not point in the same direction
A promising therapy for lazy eye is to ask a child to play three-dimensional
action video games that require attention to both eye
What early experience would cause a kitten or human child to lose

stereoscopic depth perception? → If the eye muscles cannot keep both
eyes focused in the same direction, the developing brain loses the ability
for any neuron in the visual cortex to respond to input from both eyes.
Instead, each neuron responds to one eye or the other. Stereoscopic depth
perception requires cells that compare the input from the two eyes.
Early Exposure to a Limited Array of Patterns

If a kitten spends its entire early sensitive period wearing goggles with
horizontal lines painted on them nearly all its visual cortex cells become
responsive only to horizontal lines. Even after months of later normal
experience, the cat does not respond to vertical lines.
What happens if human infants are exposed mainly to vertical or horizontal
lines instead of both equally? They become more sensitive to the kind of
line they have seen. You might wonder how such a bizarre thing could
happen
About 70 percent of all infants have astigmatism, a blurring of vision for
lines in one direction (e.g., horizontal, vertical, or one of the diagonals),
caused by an asymmetric curvature of the eyes. Normal growth reduces
the prevalence of astigmatism to about 10 percent in 4-year-old child
Impaired Infant Vision and Long-Term Consequences

Newborn infants have that experience, and we assume they have no idea
what they are seeing.
At first, these children have only a limited idea of what they are seeing. In
one study, children looked at a picture of a toy building block, and another
picture with two blocks. The task was to point to the block in the second
picture that matched the first. Children did well on this task, indicating that
they could see. However, when the task was to feel a building block and
point to which of two choices was the picture of that block, performance
was only a little better than chance. They could see the pictures, but they
didn’t understand them. A week later, without any special training, they did
much better on this task
Within weeks they could start recognizing faces. With much practice, they
began to develop hand–eye coordination. Seeing well enough to ride a
bicycle took a year and a half
One man had normal vision in early childhood until age 3½, when a
chemical explosion destroyed one eye and damaged the cornea of his
other eye so badly that he could see nothing more than light versus dark.
By adulthood, he had no visual memories and no visual imagery.
At age 43, a corneal transplant enabled him to recover vision. Immediately
he could see colors and he could soon identify simple shapes. Eventually
he learned to recognize common household objects, but unlike most people
who identify objects immediately, he had to think about it more carefully
Even 10 years later, he could not identify whether a face was male or
female, happy or sad
Various other aspects of vision remained impaired. For example, when
viewing something like Figure 5.26, he reported seeing three objects,
instead of a partly transparent object overlapping a second one
What causes astigmatism? → Astigmatism results when the eyeball is not

quite spherical. As a result, the person sees one direction of lines more
clearly than the other.
If an infant is born with dense cataracts on both eyes and they are
surgically removed years later, how well does the child see at first → The
child sees well lines more clearly than the other. enough to identify
whether two objects are the same or different, but the child doesn’t
understand what the visual information means. In particular, the child
cannot answer which visual display matches something the child touches.
However, understanding of vision improves with practice
SUMMARY
Understanding Vision by Understanding the Wiring Diagram
Your eyes are bombarded with a complex pattern of light emanating from
every source in front of you. Out of all this, your brain needs to extract the
most useful information. The nervous system from the start identifies the
borders between one object and another through lateral inhibition. It
identifies lines and their locations by simple and complex cells in the
primary visual cortex. Researchers have gone a long way toward mapping
out the excitatory and inhibitory connections that make these cells
possible. The visual experiences you have at any moment are the result of
an awe-inspiring complexity of connections and interactions among a huge
number of neurons. Understanding what you see is also the product of
years of experience.
The optic nerves of the two eyes join at the optic chiasm, where half of the
axons from each eye cross to the opposite side of the brain. Most of the
axons then travel to the lateral geniculate nucleus of the thalamus, which
communicates with the visual cortex
Lateral inhibition is a mechanism by which stimulation in any area of the
retina suppresses the responses in neighboring areas, thereby enhancing
the contrast at light–dark borders
Lateral inhibition in the vertebrate retina occurs because receptors
stimulate bipolar cells and also stimulate the much wider horizontal cells,
which inhibit both the stimulated bipolar cells and those to the sides.
Each neuron in the visual system has a receptive field, an area of the visual
field to which it is connected. Light in the receptive field excites or inhibits
a neuron depending on the light’s location, wavelength, and movement
The mammalian vertebrate visual system has a partial division of labor. In

general, the parvocellular system is specialized for perception of color and
fine details; the magnocellular system is specialized for perception of
depth, movement, and overall patterns
After damage to area V1, people report no vision, even in dreams. However,
some kinds of response to light (blindsight) can occur after damage to V1
despite the lack of conscious perception.
Within the primary visual cortex, neuroscientists distinguish simple cells
with fixed excitatory and inhibitory fields, and complex cells that respond
to a light pattern of a particular shape regardless of its exact location
Neurons within a column of the primary visual cortex have similar
properties, such as responding to lines in the same orientation.
Understanding what you see requires much more than just adding up
points and lines. Vision is an active process based partly on expectations
During infancy, the cells of the visual cortex have nearly normal properties.
However, experience is necessary to maintain and fine-tune vision.
Abnormal visual experience can change the properties of visual cells,
especially if the experience occurs early in life
Cortical neurons become unresponsive to axons from an inactive eye
because of competition with the active eye. If both eyes are closed, each
cortical cell remains somewhat responsive to axons from one eye or the
other, although that response becomes sluggish and unselective as the
weeks of deprivation continue
To develop good stereoscopic depth perception, a kitten or human child
must have experience seeing the same object with corresponding portions
of the two eyes early in life. Otherwise, each neuron in the visual cortex
becomes responsive to input from just one eye
If a kitten sees only horizontal or vertical lines during its sensitive period,
most of the neurons in its visual cortex become responsive to such lines
only. For the same reason, a child with astigmatism may have decreased
responsiveness to one kind of line or another
Some people have cataracts removed after years of cloudy vision. Vision,
useless at first, improves with practice but remains imperfect in several
ways
Module 5.3 Parallel Processing in the

Visual Cortex
It is natural to assume that anyone who sees an object sees everything
about it—the shape, color, location, and movement. However, one part of
your brain sees its shape, another sees color, another detects location, and
another perceives movement
The Ventral and Dorsal Paths

The primary visual cortex (V1) sends information to the secondary visual
cortex (area V2), which processes the information further and transmits it
to additional areas.
The connections in the visual cortex are reciprocal.
For example, V1 sends information to V2, and V2 returns information to V1.
From V2, the information branches out in several directions for specialized
processing.
Researchers distinguish between the ventral stream and the dorsal stream.
They call the ventral stream through the temporal cortex the perception
pathway or the “what” pathway, because of its importance for identifying
and recognizing objects.
The dorsal stream through the parietal cortex is the action pathway or the
“how” pathway, because of its importance for visually guided movement
ventral stream problem. A woman known as patient DF was exposed to

carbon monoxide, causing damage mainly to the ventral stream—that is,
the temporal cortex and its connections with the primary visual cortex
(Bridge et al., 2013). She cannot name the objects she sees, cannot
recognize faces, and cannot even distinguish a square from a rectangle.
When she was shown a slot in the wall, she could not say whether it was
horizontal or vertical. Nevertheless, when she was asked to put an
envelope through the slot, she aimed it correctly at once. When asked to
guess the size of an object she sees, she performs at chance levels.
However, when asked to pick up the object, she reaches out correctly,
adjusting her thumb and finger before touching the object.
Several other patients with temporal lobe damage have similar problems.
One man could not say where objects were in his room, but he could take a
walk, accurately avoiding obstacles in his way. He could reach out to grab
objects, and he could reach out to shake hands.
Another patient had such trouble recognizing objects by sight that she
attached distinctive colored tapes to the important objects she needed to
find in her house.
In short, people with temporal lobe damage can use vision to guide their
actions, but they cannot identify what the objects are.
People with damage to the dorsal stream (parietal cortex) have somewhat
the opposite problem: They see objects but they don’t integrate their vision
well with their arm and leg movements.
They can read, recognize faces, and describe objects in detail but they
cannot accurately reach out to grasp an object. While walking, they can
describe what they see, but they bump into objects, oblivious to their
location. Although they can describe from memory what their furniture
looks like, they cannot remember where it is located in their house
One patient had dorsal stream damage only in his left hemisphere. He
showed low accuracy at aiming his right arm or leg toward an object on the
right side of his body. However, his accuracy was normal when aiming his
left arm or leg toward either side, or when aiming his right arm toward the
left side
So his problem is not with attention, and not exactly vision either. It is
specifically a problem of using vision to control certain arm and leg
movements
Although the distinction between ventral and dorsal pathways is useful, we
should not overstate it. Ordinarily you use both systems in coordination
with each other
Suppose someone can describe an object in detail but stumbles and

fumbles when trying to walk toward it and pick it up. Which is probably
damaged, the dorsal path or the ventral path? → The inability to guide
movement based on vision implies damage to the dorsal path.
Detailed Analysis of Shape

In Module 5.2, we encountered simple and complex cells of the primary
visual cortex (V1). As visual information goes from the simple cells to the
complex cells and then to other brain areas, the receptive fields become
larger and more specialized. In the secondary visual cortex (V2), just
anterior to V1 in the occipital cortex, most cells are similar to V1 cells in
responding to lines, edges, or sine wave gratings, except that V2 receptive
fields are more elongated some V2 cells respond best to corners, textures,
or complex shapes
The Inferior Temporal Cortex

Cells in the inferior temporal cortex (see Figure 5.27) learn to recognize
meaningful objects
Another study considered the phenomenon of object permanence. Children
as young as age 3½ months show evidence of understanding that an
object continues to exist after it goes behind an object that prevents a child
from seeing it
. Studies of the inferotemporal cortex show a possible basis. A monkey saw
an object, and then saw some other object move in front and occlude the
first object. When the occlude moved away, either the original object
reappeared, or a new object appeared in its place. Some cells in the
inferotemporal cortex responded strongly whenever an original object
reappeared, and some responded strongly whenever a new, “surprising”
object appeared
As we might expect, damage to the ventral pathway of the cortex leads to

specialized deficits. Visual agnosia (meaning “visual lack of knowledge”) is
an inability to recognize objects despite otherwise satisfactory vision. It is
a common result from damage in the temporal cortex.
For example, one patient, when shown a key, said, “I don’t know what that
is. Perhaps a file or a tool of some sort.” When shown a stethoscope, he
said that it was “a long cord with a round thing at the end.” When he could
not identify a smoker’s pipe, the examiner told him what it was. He then
replied, “Yes, I can see it now,” and pointed out the stem and bowl of the
pipe. Then the examiner asked, “Suppose I told you that the last object was
not really a pipe?” The patient replied, “I would take your word for it.
Perhaps it’s not really a pipe”
According to fMRI studies as people viewed pictures, most objects do not

activate one brain area more than another. That is, the brain does not have
a specialized area for seeing flowers, fish, birds, clothes, food, or rocks.
However, three types of objects do produce specific responses. One part of
the Para hippocampal cortex (next to the hippocampus) responds
strongly to pictures of places, and not so strongly to anything else
Part of the fusiform gyrus of the inferior temporal cortex, especially in the
right hemisphere
The brain is amazingly adept at detecting biological motion—the kinds of

motion produced by people and animals.
Recognizing Faces
Face recognition is an important skill for humans. For civilization to

succeed, we have to know whom to trust and whom to distrust, and that
distinction requires us to recognize people that we haven’t seen in months
or years
Human newborns come into the world predisposed to pay more attention
to faces than other stationary displays
Newborns showed a strong preference for a right-side-up face over an
upside-down face, regardless of whether the face was realistic (left pair) or
distorted (central pair). When confronted with two right-side-up faces
(right pair), they showed no significant preference between a realistic one
and a distorted one. Evidently, a newborn’s concept of face requires the
eyes to be on top, but the face does not have to be realistic.
Through childhood and the early teenage years, connections strengthen
between the fusiform gyrus, especially in the right hemisphere, and part of
the inferior occipital cortex known as the occipital face area. The occipital
face area responds strongly to parts of a face, such as the eyes and
mouth. The fusiform gyrus responds strongly to a face viewed from any
angle, as well as line drawings and anything else that looks like a face
People vary considerably in their ability to recognize faces, and the reason
is not just that some people don’t care or don’t pay attention. People with
severe problems are said to have prosopagnosia (PROSS-oh-pag-NOH-
see-ah), meaning impaired ability to recognize faces. That problem can
result from damage to the fusiform gyrus, or from a failure of that gyrus to
develop fully.
In contrast, if you can recognize faces more easily than average, it may be
that you have richer than average connections between fusiform gyrus and
occippital cortex
People with prosopagnosia can read, so visual acuity is not the problem.
They recognize people’s voices, so their problem is not memory .
Furthermore, if they feel clay models of faces, they are worse than other
people at determining whether two clay models are the same or different .
Their problem is not vision, but something that relates specifically to faces
When people with prosopagnosia look at a face, they can describe each
element of a face, such as brown eyes, big ears, a small nose, and so forth,
but they do not recognize the face as a whole. You would have a similar
difficulty if you viewed faces quickly, upside down.
One study found that children who devoted at least an hour a day to some
special interest, such as watching soccer or looking at pictures of space
travel, showed fusiform gyrus responses to images related to that interest
Motion Perception
Moving objects often merit immediate attention. A moving object might be
a possible mate, something you could hunt and eat, or something that
wants to eat you. If you are going to respond, you need to identify what the
object is, where it is going, and how fast. The brain is set up to make those
calculations quickly and efficiently
The brain has no specialized areas for perceiving flowers, clothes, or

food. For what items does it have specialized areas → The temporal
cortex has specialized areas for perceiving places, faces, and bodies,
including bodies in motion
The ability to recognize faces correlates with the strength of

connections between which brain areas? → Ability to recognize faces
correlates with the strength of connections between the occipital face area
and the fusiform gyrus
The Middle Temporal Cortex

Two areas especially important for motion perception are area MT (for
middle temporal cortex), also known as area V5 and an adjacent region,
area MST (medial superior temporal cortex)
These areas receive input mostly from the magnocellular path which
detects overall patterns, including movement over large areas of the visual
field. Given that the magnocellular path is color insensitive, MT is also color
insensitive
Most cells in area MT respond selectively when something moves at a
particular speed in a particular direction. MT cells detect acceleration or
deceleration as well as the absolute speed, and they respond to motion in
all three dimensions. Area MT also responds to photographs that imply
movement, such as a photo of people running
In short, MT activity is apparently central to the experience of seeing

motion. Cells in the dorsal part of area MST respond best to more complex
stimuli, such as the expansion, contraction, or rotation of a large visual
scene
That kind of experience occurs when you move forward or backward or tilt
your head.
In short, MT and MST neurons enable you to distinguish between the result
of eye movements and the result of object movement
Motion Blindness
Given that areas MT and MST respond strongly to moving objects, and
only to moving objects, what would happen after damage to these areas?
The result is motion blindness, being able to see objects but unable to see
whether they are moving or, if so, which direction and how fast
People with motion blindness are better at reaching for a moving object
than at describing its motion but in all aspects of dealing with visual
motion, they are far behind other people
Motion blindness in the absence of other dysfunction is a rare condition.
The best described case, “LM,” reported that she felt uncomfortable when
people walked around because they “were suddenly here or there but I
have not seen them moving.
People would seem to appear or disappear suddenly, even when she was
trying to keep track of them. Someone who was walking would appear to
her as “restless,” but she would not know which direction the person was
going.
She could not cross a street without help, because she could not tell which
cars were moving, or how fast.
Pouring coffee became difficult. The flowing liquid appeared to be frozen
and unmoving, so she did not stop pouring until the cup overfilled
Several patients were reported who apparently became motion blind as a
result of brain damage, but most scientists ignored or disbelieved those
reports.
You do not see your own eyes move because area MT and parts of the
parietal cortex decrease their activity during voluntary eye movements,
known as saccade an activity does not decrease while your eyes are
following a moving object
The brain areas that monitor saccades tell area MT and the parietal cortex,
“We’re about to move the eye muscles, so take a rest for the next split
second.” Neural activity and blood flow in MT and part of the parietal
cortex begin to decrease 75 milliseconds (ms) before the eye movement
and remain suppressed during the movement
The general point is that different areas of your brain process different
kinds of visual information, and it is possible to develop many kinds of
disability
When you move your eyes, why does it not seem as if the world is
moving? → Neurons in areas MT and MST respond strongly when an object
moves relative to the background, and not when the object and
background move in the same direction and speed
Under what circumstance does someone with an intact brain become

motion blind, and what accounts for the motion blindness? → People
become motion blind shortly. before and during a saccade (voluntary eye
movement), because of suppressed activity in area MT.
SUMMARY
Researchers distinguish between the ventral visual stream, responsible for
perceiving objects, and the dorsal stream, responsible for visual guidance
of movements.
The inferior temporal cortex detects objects and recognizes them despite
changes in position, size, and so forth
A circuit including the fusiform gyrus of the temporal cortex is specialized
for recognizing faces. People with impairments in this circuit experience
prosopagnosia, a difficulty in recognizing faces despite nearly normal
vision in other regards
Although the fusiform gyrus is important for recognizing faces, it also

contributes to other types of visual expertise.
The middle temporal cortex (including areas MT and MST) is specialized
for detecting the direction and speed of a moving object. People with
damage in this area experience motion blindness, an impairment in their
ability to perceive movement
Even people with an intact brain experience a brief period of motion
blindness beginning about 75 ms before a voluntary eye movement and
continuing during the eye movement

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Module 5.1 Visual Coding

General Principles of Perception

If someone electrically stimulated the auditory receptors in your ear,

The Eye and Its Connections to the Brain

Route within the Retina

Fovea and Periphery of the Retina

Although rods outnumber cones by about 20 to 1 in the human retina,

Visible light consists of electromagnetic radiation within the range from

The Trichromatic (Young-Helmholtz) Theory

This theory, later modified by Hermann von Helmholtz, is now known as

The Opponent-Process Theory

Examine Figure 5.9. According to the trichromatic theory, what causes

According to the opponent-process theory, under what circumstance

The Retinex Theory

Color Vision Deficiency

Why is color vision deficiency a better term than color blindness?

An Overview of the Mammalian Visual System

Processing in the Retina

As we progress from bipolar cells to ganglion cells to later cells in the

What are the differences between the parvocellular and magnocellular

The Primary Visual Cortex

The research supports two explanations for blindsight:

Simple and Complex receptive fields

How could a researcher determine whether a given neuron in the visual

The Columnar Organization of the Visual Cortex

Are Visual Cortex Cells Feature Detectors?

feature detectors—neurons whose responses indicate the presence of a

Development of the Visual Cortex

Deprived Experience in One Eye

After an eye deprived of vision in adults is reopened, cells gradually return

Deprived Experience in Both Eyes

Uncorrelated Stimulation in the Two Eyes

A similar phenomenon occurs in humans. Certain children are born with

What early experience would cause a kitten or human child to lose

Early Exposure to a Limited Array of Patterns

Impaired Infant Vision and Long-Term Consequences

What causes astigmatism? → Astigmatism results when the eyeball is not

The mammalian vertebrate visual system has a partial division of labor. In

Module 5.3 Parallel Processing in the

The Ventral and Dorsal Paths

ventral stream problem. A woman known as patient DF was exposed to

Suppose someone can describe an object in detail but stumbles and

Detailed Analysis of Shape

The Inferior Temporal Cortex

As we might expect, damage to the ventral pathway of the cortex leads to

According to fMRI studies as people viewed pictures, most objects do not

The brain is amazingly adept at detecting biological motion—the kinds of

Face recognition is an important skill for humans. For civilization to

The brain has no specialized areas for perceiving flowers, clothes, or

The ability to recognize faces correlates with the strength of

The Middle Temporal Cortex

In short, MT activity is apparently central to the experience of seeing

Under what circumstance does someone with an intact brain become

Although the fusiform gyrus is important for recognizing faces, it also

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