BioPsy C5

Chapter 5
Vision
© Cengage Learning 2016 © Cengage Learning 2016

5.1 Visual Coding
• How far one sees is dependent on how far

light travels before it strikes one’s eyes
• Perception of vision is not in the eyes; it’s
in the brain
© Cengage Learning 2016

General Principles of Perception
• Each of our senses has specialized

receptors that are sensitive to a particular
kind of energy
• Law of specific nerve energies states that
activity by a particular nerve always
conveys the same type of information to
the brain
– Example: impulses in one neuron indicate light;
impulses in another neuron indicate sound

The Eye and Its Connections to the Brain
• Light:
– Enters the eye through an opening in the
center of the iris called the pupil
– Is focused by the lens and the cornea onto the
rear surface of the eye known as the retina,
which is lined with visual receptors
– From the left side of the world strikes the right
side of the retina and vice versa
– From above strikes the bottom half of the
retina and vice versa
Route Within the Retina – Bipolar Cells
• Cells, located closer to the center of the

eye, that receive messages from visual
receptors at the back of the eye
• These cells send messages to ganglion
cells that are even closer to the center of
the eye
– The axons of ganglion cells join one another to
form the optic nerve that travels to the brain

Route Within the Retina – Amacrine Cells
• Additional cells that receive information

from bipolar cells and send it to other
bipolar, ganglion, or amacrine cells
• Control the ability of the ganglion cells to
respond to shapes, movements, or other
specific aspects of visual stimuli

Cross Section of a Vertebrate Eye

Visual Path Within the Eye

The Optic Nerve
• Consists of the axons of ganglion cells that

band together and exit through the back of
the eye and travel to the brain
• Leaves the back of the eye; the point at
which it leaves is called the blind spot
because it contains no receptors

Illustration of the Blind Spot

The Fovea, Part 1
• Is the central portion of the retina and

allows for acute and detailed vision
– Packed tight with receptors
– Nearly free of ganglion axons and blood
vessels

The Fovea, Part 2
• Each receptor in the fovea attaches to a

single bipolar cell and a single ganglion cell
known as a midget ganglion cell
• Each cone in the fovea has a direct line to
the brain which allows the registering of the
exact location of input
• Our vision is dominated by what we see in
the fovea

The Placement of Receptors on the Retina

The Periphery of the Retina
• In the periphery of the retina, a greater

number of receptors converge into
ganglion and bipolar cells
– Detailed vision is less in peripheral vision
– Allows for the greater perception of much
fainter light in peripheral vision

Convergence of Input onto Bipolar Cells

The Arrangement of Visual Receptors
• Highly adaptive
– Example: predatory birds have a greater
density of receptors on the top of the eye; rats
have a greater density on the bottom of the
eye

The Difference Between Foveal and
Peripheral Vision
Characteristic Foveal vision Peripheral vision
Receptors Cones Proportion of rods increases
toward periphery
Convergence of Each ganglion cell excited by a Each ganglion cell excited by

Input single cone many receptors
Brightness Distinguishes among bright lights; Responds well to dim light; poor
sensitivity responds poorly to dim light for distinguishing among
bright lights
Sensitivity Good detail vision because each Poor detail vision because many
to detail cones own ganglion cell sends a receptors converge their input onto
message to a given ganglion cell
the brain
Color Vision Good (many cones) Poor (few cones)

Visual Receptors: Rods and Cones, Part 1
• The vertebrate retina consists of two kinds

of receptors
– Rods: most abundant in the periphery of the
eye and respond to faint light (120 million per
retina)
– Cones: most abundant in and around the fovea
(6 million per retina)
• Essential for color vision and more useful in bright
light

Visual Receptors: Rods and Cones, Part 2
• Though cones are outnumbered, they

provide about 90% of the brain’s input
• On average, 120 million rods and 6 million
cones converge onto 1 million axons in the
optic nerve
• The ratio of rods to cones is higher in
species that are more active at dim light

A Comparison of Rods and Cones

Photopigments
• Chemicals contained by both rods and

cones that release energy when struck by
light
– Consist of 11-cis-retinal bound to proteins
called opsins
– Light energy converts 11-cis-retinal quickly into
all-trans-retinal
– Light is thus absorbed and energy is released
that activates second messengers within the
cell
Color Vision
• Visible light is a portion of the

electromagnetic spectrum
• The perception of color is dependent upon
the wavelength of the light
• “Visible” wavelengths are dependent upon
the species’ receptors
• Humans perceive wavelengths between
400 and 700 nanometers (nm)

Visible Light on the Electromagnetic
Spectrum

Specificity of Color Vision
• Depends on specific receptors within the

eye
• Two major interpretations of color vision
– Trichromatic theory/Young-Helmholtz theory
– Opponent-process theory

Trichromatic Theory, Part 1
• Color perception occurs through the

relative rates of response by three kinds of
cones
– Short wavelength
– Medium-wavelength
– Long-wavelength

Wavelength-Sensitivity Functions

Trichromatic Theory, Part 2
• Each cone responds to a broad range of

wavelengths, but some more than others
• The ratio of activity across the three types
of cones determines the color
• More intense light increases the brightness
of the color but does not change the ratio
• Three kinds of cones are unevenly
distributed

Distribution of Cones in Two Human Retinas

The Opponent-Process Theory
• Suggests that we perceive color in terms of

paired opposites
• The brain has a mechanism that perceives
color on a continuum from red to green and
another from yellow to blue
– A possible mechanism for the theory is that
bipolar cells are excited by one set of
wavelengths and inhibited by another

An Afterimage as an Effect of Context

Limitations of Color Vision Theories
• Both the opponent-process and

trichromatic theory have limitations
– Color constancy, the ability to recognize color
despite changes in lighting, is not easily
explained by these theories
• Retinex theory suggests the cortex
compares information from various parts of
the retina to determine the brightness and
color for each area

The Context of Color Perception

Brightness Constancy

Color Vision Deficiency
• An impairment in perceiving color

differences
– Gene responsible is contained on the X
chromosome
– Caused by either the lack of a type of cone or
a cone that has abnormal properties
– Most common form is difficulty distinguishing
between red and green
• Results from the long- and medium-wavelength
cones having the same photopigment
5.2 How the Brain Processes Visual
Information
• Senses, such as vision, provide
psychological experiences
• Neuroscientists have developed a relatively
detailed understanding of vision
• Understanding the mechanisms of vision
provides a model of what it means to
explain something in biological terms

An Overview of the Mammalian Visual
System, Part 1
• Rods and cones of the retina make
synaptic contact with horizontal cells and
bipolar cells
• Horizontal cells are cells in the eye that
make inhibitory contact onto bipolar cells
• Bipolar cells make synapses onto amacrine
cells and ganglion cells
• Different cells are specialized for different
visual functions
An Overview of the Mammalian Visual
System, Part 2
• Ganglion cell axons form the optic nerve
• The optic chiasm is the place where the
two optic nerves leaving the eye meet
• In humans, half of the axons from each eye
cross to the other side of the brain
• Most ganglion cell axons go to the lateral
geniculate nucleus, a smaller amount to
the superior colliculus, and fewer to other
areas
The Vertebrate Retina

The Path of Visual Input

Processing in the Retina
• The lateral geniculate nucleus

– Part of the thalamus
– Specialized for visual perception
– Destination for most ganglion cell axons
– Sends axons to other parts of the thalamus
and to the visual areas of the occipital cortex
• The cortex and thalamus constantly feed
information back and forth to each other

Lateral Inhibition in the Retina
• Sharpens contrasts to emphasize the

borders of objects
• The reduction of activity in one neuron by
activity in neighboring neurons
• The response of cells in the visual system
depends upon the net result of excitatory
and inhibitory messages it receives

An Illustration of Lateral Inhibition

Further Processing
• The receptive field refers to the part of the

visual field that either excites or inhibits a
cell in the visual system of the brain
• For a receptor, the receptive field is the
point in space from which light strikes it
• For other visual cells, receptive fields are
derived from the visual field of cells that
either excite or inhibit
– Example: ganglion cells converge to form the
receptive field of the next level of cells
An Example of a Receptive Field

Primate Receptive Fields
• Ganglion cells of primates generally fall

into three categories
– Parvocellular neurons
– Magnocellular neurons
– Koniocellular neurons

Parvocellular Neurons
• Mostly located in or near the fovea

• Have smaller cell bodies and small
receptive fields
• Highly sensitive to detect color and visual
detail

Magnocellular Neurons
• Distributed evenly throughout the retina

• Have larger cell bodies and visual fields
• Highly sensitive to large overall pattern and
moving stimuli

Koniocellular Neurons
• Have small cell bodies

• Found throughout the retina
• Have several functions, and their axons
terminate in many different places

Characteristics of Receptive Fields
• Cells of the lateral geniculate have a

receptive field similar to those of ganglion
cells:
– An excitatory or inhibitory central portion and a
surrounding ring of the opposite effect

The Primary Visual Cortex, Part 1
• The primary visual cortex (area V1)

receives information from the lateral
geniculate nucleus and is the area
responsible for the first stage of visual
processing
• Some people with damage to V1 show
blindsight: an ability to respond to visual
stimuli that they report not seeing

The Primary Visual Cortex, Part 2
• Hubel and Weisel (1959, 1998)

distinguished various types of cells in the
visual cortex
– Simple cells
– Complex cells
– End-stopped/hypercomplex cells

Simple Cells
• Fixed excitatory and inhibitory zones

• The more light that shines in the excitatory
zone, the more the cell responds
• The more in the inhibitory zone, the less
the cell responds
• Bar-shaped or edge-shaped receptive
fields with vertical and horizontal
orientations outnumbering diagonal ones

Complex Cells
• Located in either V1or V2

• Have large receptive field that can not be
mapped into fixed excitatory or inhibitory
zones
• Responds to a pattern of light in a
particular orientation and most strongly to a
moving stimulus

Simple and Complex Receptive Fields

The Receptive Field of a Complex Cell

End-Stopped or Hypercomplex Cells
• Similar to complex cells but with a strong

inhibitory area at one end of its bar shaped
receptive field
• Respond to a bar-shaped pattern of light
anywhere in its large receptive field,
provided the bar does not extend beyond a
certain point

The Receptive Field of an End-Stopped Cell

Properties of Simple, Complex, and End-
Stopped Cells
Characteristic Simple Cells Complex Cells End-Stopped Cells

Location V1 V1 andV2 V1 and V2
Binocular input Yes Yes Yes
Size of receptive Smallest Medium Largest
field
Receptive field Bar- or edge-shaped, Bat- or edge-shaped, Same as complex cell but
with fixed excitatory and without fixed excitatory or with strong inhibitory zone
inhibitory zones inhibitory zones at one end

Columnar Organization of the Visual Cortex,
Part 1
• In the visual cortex, cells are grouped
together in columns perpendicular to the
surface
• Cells within a given column process similar
information
– Respond either mostly to the right or left eye,
or respond to both eyes equally
– Do not consistently fire at the same time

Columnar Organization of the Visual Cortex,
Part 2

Visual Cortex Cells as Feature Detectors,
Part 1
• Cells in the visual cortex may be feature
detectors, neurons whose response
indicate the presence of a particular
feature/stimuli
• Prolonged exposure to a given visual
feature decreases sensitivity to that feature

Visual Cortex Cells as Feature Detectors,
Part 2

Development of the Visual Cortex
• Animal studies have greatly contributed to

the understanding of the development of
vision
• Early lack of stimulation of one eye: leads
to synapses in the visual cortex becoming
gradually unresponsive to input from that
eye
• Early lack of stimulation of both eyes:
cortical responses become sluggish but do
not cause blindness
Critical Periods in Development
• Sensitive/critical periods are periods of

time during the lifespan when experiences
have a particularly strong/enduring effect
– Ends with the onset of chemicals that inhibit
axonal sprouting
– Changes that occur during critical period
require both excitation and inhibition of some
neurons
• Cortical plasticity is greatest in early life,
but never ends
Stereoscopic Depth Perception
• A method of perceiving distance in which

the brain compares slightly different inputs
from the two eyes
– Relies on retinal disparity or the discrepancy
between what the left and the right eye sees
– The ability of cortical neurons to adjust their
connections to detect retinal disparity is
shaped through experience

Strabismus
• A condition in which the eyes do not point

in the same direction
– Usually develops in childhood
– Also known as “lazy eye”
• If two eyes carry unrelated messages,
cortical cell strengthens connections with
only one eye
• Development of stereoscopic depth
perception is impaired
Two Examples of Lazy Eye
Early Exposure to a Limited Array of
Patterns
• Leads to nearly all of the visual cortex cells
becoming responsive to only that pattern
• Astigmatism refers to a blurring of vision for
lines in one direction caused by an
asymmetric curvature of the eyes
– 70% of infants have astigmatism

Restricting Early Visual Experience

An Informal Test for Astigmatism

Long-Term Consequences of Impaired Infant
Vision
• Study of people born with cataracts but had
them removed at age 7 or 12 indicate that
vision can be restored gradually, but
problems persist
– Difficulty in recognizing objects
– Unable to tell that components are part of a
whole
– Best prognosis is for children whose vision
problems are corrected early in life

5.3 Parallel Processing in the Visual Cortex
• Neuroscientists have identified at least 80

brain areas that contribute to vision in
different ways
• One part of your brain sees its shape,
another sees color, another detects
location, and another perceives movement

The Ventral and Dorsal Paths
• The secondary visual cortex (area V2)

receives information from area V1,
processes information further, and sends it
to other areas
• Information is transferred between area V1
and V2 in a reciprocal nature

Visual Pathways in the Monkey Cerebral Cortex
The Ventral and Dorsal Streams, Part 1
• The ventral stream refers to the path that

goes through temporal cortex
– The “what” path
– Specialized for identifying and recognizing
objects
• The dorsal stream refers to the visual path
in the parietal cortex
– The “how” path
– Important for visually guided movements.
The Ventral and Dorsal Streams, Part 2
• Normal behavior makes use of both

pathways in collaboration
• Damaging either stream will produce
different deficits
– Ventral stream damage: can see where objects
are but cannot identify them
– Dorsal stream damage: can identify objects but
not know where they are

Detailed Analysis of Shape
• Receptive fields become larger and more

specialized as visual information goes from
simple cells to the complex cells and then
to other brain areas
• The inferior temporal cortex contains cells
that respond selectively to complex shapes
but are insensitive to distinctions that are
critical to other cells
• Cells in this cortex respond to identifiable
objects
Transformations of a Drawing

Visual Agnosia
• The inability to recognize objects despite

satisfactory vision
– Caused by damage to the pattern pathway
usually in the temporal cortex

Face Recognition – The Fusiform Gyrus

Recognizing Faces
• Face recognition occurs relatively soon

after birth
– People with cataracts removed at 2-6 months
develop nearly normal vision but have slight
difficulties in distinguishing faces
– Newborns show strong preference for a right-
side-up face and support idea of a built-in face
recognition system
• Facial recognition continues to develop
gradually into adolescence
Amount of Time Infants Spend Looking at
Patterns

How Infants Divided Their Attention Between
Faces

Prosopagnosia
• The impaired ability to recognize faces

– Occurs after damage to the fusiform gyrus of
the inferior temporal cortex
– The fusiform gyrus responds much more
strongly to faces than anything else

Color Perception
• Dependent on both the light reflected on an

object and how it compares with objects
around it
– Area V4 may be responsible for color
constancy and visual attention
– Color constancy: the ability to recognize
something as being the same color despite
changes in lighting

Motion Perception
• Involves a variety of brain areas in all four

lobes of the cerebral cortex
– The middle-temporal cortex (MT/V5) responds
to a stimulus moving in a particular direction
– Cells in the dorsal part of the medial superior
temporal cortex (MST) respond to expansion,
contraction, or rotation of a visual stimulus
– Both receive input from the magnocellular
path; color-insensitive

Stimuli That Excite the Dorsal Part of Area
MST

Motion Blindness
• The inability to determine the direction,

speed and whether objects are moving
– Likely caused by damage in area MT
• Some people are blind except for the ability
to detect which direction something is
moving
– Area MT probably gets some visual input
despite significant damage to area V1

Saccades
• Several mechanisms prevent confusion or

blurring of images during eye movements
– Saccades are a decrease in the activity of the
visual cortex during quick eye movements
– Neural activity and blood flow decrease 75
milliseconds before and during eye
movements

BioPsy C5

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Chapter 5

© Cengage Learning 2016 © Cengage Learning 2016

• How far one sees is dependent on how far

© Cengage Learning 2016

• Each of our senses has specialized

© Cengage Learning 2016

• Cells, located closer to the center of the

© Cengage Learning 2016

• Additional cells that receive information

© Cengage Learning 2016

© Cengage Learning 2016

© Cengage Learning 2016

• Consists of the axons of ganglion cells that

© Cengage Learning 2016

© Cengage Learning 2016

• Is the central portion of the retina and

© Cengage Learning 2016

• Each receptor in the fovea attaches to a

© Cengage Learning 2016

© Cengage Learning 2016

• In the periphery of the retina, a greater

© Cengage Learning 2016

© Cengage Learning 2016

© Cengage Learning 2016

Convergence of Each ganglion cell excited by a Each ganglion cell excited by

© Cengage Learning 2016

• The vertebrate retina consists of two kinds

© Cengage Learning 2016

• Though cones are outnumbered, they

© Cengage Learning 2016

© Cengage Learning 2016

• Chemicals contained by both rods and

• Visible light is a portion of the

© Cengage Learning 2016

© Cengage Learning 2016

• Depends on specific receptors within the

© Cengage Learning 2016

• Color perception occurs through the

© Cengage Learning 2016

© Cengage Learning 2016

• Each cone responds to a broad range of

© Cengage Learning 2016

© Cengage Learning 2016

• Suggests that we perceive color in terms of

© Cengage Learning 2016

© Cengage Learning 2016

• Both the opponent-process and

© Cengage Learning 2016

© Cengage Learning 2016

© Cengage Learning 2016

• An impairment in perceiving color

© Cengage Learning 2016

© Cengage Learning 2016

© Cengage Learning 2016

• The lateral geniculate nucleus

© Cengage Learning 2016

• Sharpens contrasts to emphasize the

© Cengage Learning 2016

© Cengage Learning 2016

• The receptive field refers to the part of the

© Cengage Learning 2016

• Ganglion cells of primates generally fall

© Cengage Learning 2016

• Mostly located in or near the fovea