The Visual System

ANAT321

Visual illusions reveal a key property of the visual system

http://www.youtube.com/watch?v=dhuUhaNI
WLQ#t=110

Sensation is an abstraction, not a replication,

of the real world.
Vernon Mountcastle

[The problems of vision] the brain is solving are

literally unsolvable An elliptical shape on the retina
could have come from an oval viewed head-on or a
circle viewed at a slant. A patch of gray could have
come from a snowball in the shade or a lump of coal
in the sun. Vision has evolved to convert these illposed problems into solvable ones by adding
premises: assumptions about how the world we
evolved in is, on average, put together.
Steven Pinker

In other words, the visual system is not a camera.

The visual system organizes visual input to

give meaning to visual perceptions.

The visual system actively creates a coherent visual scene.

Part I: The Eye

The Eye
Cornea

Lens

Lens

Vitreous
humor
Vitreous

Choroid

Sclera

Iris

Retina
Optic disc
Optic Disk
Optic nerve

Fovea
Fovea

Retina

Comparison of the Eye and a Camera

Aperture

Lens

Pupil

Lens

The fovea is directly in line with the visual axis. It is the high
acuity region of the retina

Macula

Fovea

The fovea is the high-acuity center of the visual field.

The fovea is the high-acuity center of the visual field.

This is what your eyes are doing when you look at a face.
The rapid eye movements are called saccades.

The optic disk has no photoreceptors, so each eye has a blind spot

Illustration of the Retina by Cajal

The retina has 10 layers

Inner limiting membrane

Nerve fiber layer
Neural retina

Ganglion cell layer

Inner plexiform layer
Inner nuclear layer
Outer plexiform layer
Outer nuclear layer
Outer limiting membrane
Photoreceptor outer and inner segments

Retinal pigment epithelium

Choroid

The Retina

The fovea is the high-acuity center of the visual field.

Morphology of Photoreceptors
Discs

Outer segment

Outer segment
Mitochondria

Inner segment

Inner segment
Nucleus

Soma

Soma
Synaptic terminal

Synaptic terminal

Rod

Cone

Light hyperpolarizes photoreceptor by closing a cGMPgated cation channel.

cGMP Phosphodiesterase

Rod Outer Segment

Rhodopsin

Transducin
(G-Protein)

cGMP

Na+
5GMP

-40 mV
-60 mV

Bipolar cells connect photoreceptors to retinal ganglion cells

Light

Cone

Bipolar Cell

Glutamate (Inhibitory)

Glutamate (Excitatory)

Retinal Ganglion Cell

To Optic Nerve

Cones in the surrounding region are connected to the

central photoreceptor through inhibitory horizontal cells.
Light

Horizontal Cell

Horizontal Cell

These retinal ganglion cells have on-center-off-surround receptive fields,

reflecting convergent input from multiple photoreceptors.

Surround

Center

Surround

Photoreceptors also make excitatory connections with bipolar cells. In this

case, the ganglion cells are off-center-on-surround.

Surround

Center

Surround

Excitatory synapse

In the fovea, each cone forms the center of an on-centered

and an off-centered pathway.

Cone
Glutamate (Inhibitory)

Bipolar Cells

Glutamate (Excitatory)

Off-centered Ganglion cell

OnGanglion cell

Just outside the macula (the area of highest light sensitivity)

thousands of rods may converge on hundreds of bipolar
cells, which, in turn, may converge on a single ganglion cell.

Key Points
Information from multiple photoreceptors converges on
individual retinal ganglion cells.
As a result of this convergence, through the complex lateral
circuitry of the retina, the receptive fields of the retinal ganglion
cells are more complex than the receptive fields of the
photoreceptors.
Retinal ganglion cells are especially good at detecting
edges/boundaries between light and dark regions of visual
space.

Part II: From the Retina to the Cortex

The axons of retinal ganglion cells project as the optic nerves to the lateral
geniculate nuclei. The fibers partially cross at the optic chiasm and continue
as the optic tracts.

Optic Chiasm

Optic nerves

Optic chiasm

Optic tracts

Lateral geniculate

At least two functionally distinct classes of retinal

ganglion cells convey visual information in parallel
pathways.

Cells in the M pathway have large receptive fields and respond

transiently to visual stimuli. The M Pathway is involved in
detecting motion.
Cells in the P pathway have smaller receptive fields and respond
selectively to specific wavelengths of light. The P Pathway is
involved in detecting form and color.

The lateral geniculate nucleus is a six layered structure

Inputs from the two eyes and from the M and P pathways are
segregated in distinct layers of the lateral geniculate nucleus.

Optic nerves
Optic chiasm
Lateral geniculate
nucleus

Optic
tracts

Dorsal

Ventral
M pathway

P pathway

Primary visual
cortex

Lateral geniculate neurons have on-center and off-center

receptive fields, similar to the receptive fields of retinal
ganglion cells.
On-center

Off-center

~ 90% of retinal ganglion axons project to the lateral

geniculate nucleus. The remaining 10% project to the
superior colliculus and to the pretectum.

Projections to the superior colliculs are

involved in saccadic eye movements.

Projections to the pretectum are involved in

pupillary reflexes.

The output from the lateral geniculate nucleus

projects to the primary visual cortex (V1),

Primary visual
cortex (V1)

Optic radiation
Lateral geniculate
nucleus

Projections from the lateral geniculate to V1 are vastly

outnumbered by reciprocal projections coming back from
the cortex.

Part III: Visual Processing in the Cortex

Visual Pathways

Primary Visual Cortex (V1)

Primary visual cortex contains a retanotopic map of the

external visual field.
11

Fovea

5
6
10

13
24

7
8
12

Visual Field

Right

Left

V1

4 8
3 7

Calcarine fissure

12
11

10
9

6 2
5 1

V1

The Visual Field

Left and right eyes

Left eye

Right eye

Layer IV is the main recipient of afferent inputs from

the thalamus.
Golgi stain

Nissl stain

I
II
III

IV
V

VI

Internal granule layer

Lateral geniculate neurons project mainly to layer 4C of

primary visual cortex

http://webvision.med.utah.edu/book/part-ixpsychophysics-of-vision/the-primary-visualcortex/

The M and P pathways remain segregated

through visual cortex.
1

V2 (Thin stripe)
V2 (Interstripe)

2
Blob

Interblob

V2 (Thick stripe)

Ventral stream
Dorsal stream

4A
4B

4B

4C

4C
4C

4C

Superior colliculus, pulvinar, pons

Lateral geniculate, claustrum

M and P connections in V1
Blob

Neuron in layer 4 of V1 have receptive fields similar to those of

retinal ganglion cells and LGN neurons; however, things get more
interesting when we examine the receptive fields of neurons above
and below layer 4.

4A
4B
4C
4C

 V1 neurons in layers above and below layer 4

respond best to light or dark bars of a
particular orientation.
The neurons are organized into vertical
orientation columns. The neurons within a
column respond best to edges with a
particular orientation, located within a
specific region of the visual field.
Cells in visual cortex with these response
properties are referred to as simple cells.

In 1981, Torsten Wiesel (left) and David Hubel (right) won the Nobel Prize for their
discoveries concerning how brain neurons encode visual stimuli. Hubel went to
grade school in Outremont, obtained his undergraduate (in math and physics) and
medical degrees from McGill and studied neurology for three years at the
Montreal Neurological Institute, before moving to the U.S. (Johns Hopkins and
then Harvard).

http://www.youtube.com/watch?v=KE952yue
VLA

The elongated receptive fields of simple cells are built from

convergent input from many layer 4 cells with roughly circular
receptive fields.
_
_ + _
_
_
_ + _
_

_
_ + _
_
_
_ + _
_

Complex cells respond to more complex stimuli,

e.g. a bar of light moving in a particular
direction. The receptive fields of complex cells
are thought to be built up from convergence of
simple cells.

The receptive fields of complex cells are assembled from the

receptive fields of simple cells.

Neurons in adjacent orientation columns in V1 respond to the same region in

visual space, but with systematically different orientation preferences.

The complete cycle orientations columns from a particular region of

visual space forms a spiral on the surface of the cortex.

Superimposed on the orientation columns are columns

corresponding to alternating input from the ipsilateral and
contralateral eyes. These are called ocular dominance columns.
The complete set if orientation columns from a particular region
of space (i.e. from both eyes) is called a hypercolumn.

Primary visual cortex (V1) corresponds to area 17. Areas

18 and 19 comprise extrastriate cortex.

Motion and color activate different regions of

extrastriate cortex.

Visual information flows from V1 through separate dorsal

(where) and ventral (what) streams. The dorsal stream
encodes location and movement. The ventral stream
encodes form and color.

Dorsal (where)

Ventral (what)

Visual information flows from V1 through separate dorsal (where)

and ventral (what) streams. The dorsal stream encodes location and
movement. The ventral stream encodes form and color.

Detection of Motion

Areas MT (medial temporal) and MST (medial superior

temporal) are devoted to processing motion.

Neurons in MT can solve the aperture problem.

Neurons in MT respond to the movement of the entire object in the visual field
rather than to its component parts.

Lesions in monkeys and humans produce deficits

in smooth pursuit movements.

Lesions in monkeys and humans produce deficits

in smooth pursuit movements.

Damage to the dorsal pathways can result in an

inability to perceive motion.

The dorsal stream contributes to pathways from parietal

cortex to premotor cortex that are involved in reaching for
objects.
Sensory-motor transformation
Premotor cortex

Dorsal stream

V4 and other regions of ventral occipital and temporal

cortex are involved in analyzing form and color.

Detection of Form

V4 and other regions of ventral occipital and temporal

cortex are involved in analyzing form and color.

Some neurons in V2 respond to illusory contours

Ventral pathways are involved in

perceiving objects.
e.g. Face neurons in the monkey cortex.

Some ventral stream neurons respond specifically to faces

The Bill Clinton Neuron

The Fusiform Face Area

Prosopagnosia: Loss of the ability to recognize

familiar faces

Part IV: Color Vision

Visible light covers a narrow range of the electromagnetic spectrum.

The wavelengths of radiation within the visible spectrum are
continuous, but we perceive discrete colors.

The human retina contains three types of cones, called S, M and L. Each
type is tuned to respond best to a restricted range of wavelengths within
the visible spectrum.

http://hyperphysics.phy-astr.gsu.edu/hbase/vision/colcon.html

A single type of cones cannot distinguish between different wavelengths

of light, e.g. M cones respond best to green light, but will also respond to
wavelengths covering most of the visible spectrum.

M
http://hyperphysics.phy-astr.gsu.edu/hbase/vision/colcon.html

Naturally occurring substances have complex reflectance functions.

Reflectance

0.6

0.4

0.2
0.0
400

500
600
Wavelength (nm)

700

An orange appears orange over a wide range of ambient lighting.

P-type retinal ganglion cells encode differences in the responses

of S, M and L cones.

L M = red/green

S LM = Yellow/Blue

P-type ganglion cells have red-green or yellow-blue, on- and offcentered receptive fields.

_ + _ _ + _
_
_
On-center

+
_ +
+
+

+
+ _ +
+

Off-center

_ + _ _ + _
_
_
On-center

+
+ _ +
+

+
+ _ +
+

Off-center

Red-Green Color Blindness and Achromatagnosia

The binding problem: How are the different components

of vision (e.g. form, color, location, motion), spread out
over disparate regions of cortex, bound together to form
a unified, coherent percept.

Dorsal (where)

Ventral (what)

Key Ideas
Information destined for the cortex passes through the thalamus (the
lateral geniculate nucleus).
Information is processed at multiple levels (LGN, V1, the dorsal and
ventral streams). Neurons at high levels in the hierarchy have more
complex receptive fields than neurons at low levels in the hierarchy.
Lateral inhibition plays an important role in shaping receptive fields.
Information flows through multiple parallel pathways (the M and P
pathways). Information flowing through the separate pathways is bound
together, through a process that is not fully understood, to create a
unified percept.
Perception of color is a combinatorial process.
Perception is an abstraction not a replication of the external world.

Lesions to different parts of the visual pathway produce

predictable visual deficits