Neuroscience 106: Lecture 10 - The Retina

Assignments: In Bear et al. study Chapter 9.

TODAY'S LECTURE:
I. Introduction to SYSTEMS neuroscience.
A. Thus far the focus of the course has been on the cellular/molecular level.
B. The rest of the course will be focused on systems level. We will apply what we have already
learned to the systems level.
C. Systems are circuits of neurons that interact to perform some function (for example the visual
system, the auditory system, the pain sensing system, memory systems, motor systems, etc.).
D. Problems in understanding systems.
The unknowns are greater. The complexity is vastly greater. The operation of many systems is
highly counterintuitive.
E. Neural Codes: A fundamental concept. The essential idea is that the meaning of activity (e.g.,
action potentials) by a neuron depends on the system it is part of. Action potential by a neuron in a
pain system would yield the perception of pain; action potential by a neurons in the visual system
would yield the perception of sight; action potentials by a neuron in a motor systems would cause
movement; action potential by a neuron in a system underlying emotion might cause the feeling of
fear or another emotion.
1. Also, systems have subsystems and neurons in different subsystems code for different
functions. For example, in the visual system, different areas and groups of neurons perform
different functions. Some neurons are responsible for detecting the form of a visual object; others
are responsible for detecting color (they code for color), other for movement (they code for
movement), others for location (they code to location of an object in the visual field), etc. The
action potential produced by these different neurons mean different things, i.e. they "code" for
different perceptions/meanings etc.
2. What is a neuron coding? What attribute of the visual world is a neuron in the visual system
encoding? All that you measure when you record membrane voltage is action potentials, but
action potentials in different neurons mean different things. How do neuroscientists determine the
function of a neuron or of a system of neurons?

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II. The Visual System: The retina
A. The retina consists of several thin layers of cells distributed across the inside of the eye.
B. The fovea is the portion of the retina where light falls from an object that you are looking directly
at. It is the portion of the retina with the highest acuity, the ability to resolve fine detail and patterns
of light. Note: Acuity and sensitivity are different.
C. The optic disk is the retinal location where axons from a type of retinal cell collect and exit the
eye and form the optic nerve. This is the blind spot because there are no photoreceptors in the optic
disk.
D. Below is a diagram depicting the eye and how a visual image is "mapped" onto the retina.

Retina
A

Fovea

E. A neuron’s receptive field is the location in the environment (or the surface of the body) from
which an appropriate stimulus will change that cell's activity. For example light at 'A' (the tip of the
flame in the diagram above) will affect the activity of retinal cell in location 'a' in the retina. Cells in
different locations in the retina have receptive fields in different locations in the visual field.
F. There are five cell types in the retina.
1. Photoreceptors – The first stage in the visual system.
a. Photoreceptors - the only cell type in the visual system that is directly sensitive to light.
b. There are two types of photoreceptors.

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 1) Rods
a) There are about 120 million rods in the human retina.
b) Rods are highly sensitive to light and are responsible for vision in very dim light.
c) Rods are bleached in bright light and thus unresponsive in bright light.
d) Rods are not responsible for high acuity vision (not good for fine detail).
e) Rods are achromatic (insensitive to colors).
f) Rods only exist outside of the fovea.
2) Cones
a) There are about 6 million cones in the human retina.
b) Cones are less sensitive to light intensity and are inoperative in dim light.
c) Cones are sensitive to color. There are three subtypes, selectively sensitive to red,
blue, and green wavelengths of light.
d) Cones are most concentrated in the fovea.
c. Photoreceptors project to the bipolar cells.
2. Bipolar cells (BPs)
3. Retinal ganglion cells (RGCs)
a. RGCs are the only output cell type in the retina.
b. RGCs are the only means by which information from the eye gets to the rest of the visual
system and their axons form the optic tract.
4. Horizontal cells (HCs)
5. Amacrine cells (ACs)

See figures 9.11, 9.12 and 9.13

G. Why is it that only the RGCs have axons?

1. Axons are needed for long-distance transfer of information. In the retina, the cells are very
close together and so don’t need action potentials or axons. Also, communication by PSPs may
be able to convey information that is more subtle than can be conveyed by the AP frequency code.
2. Only the RGCs and ACs generate action potentials. The rest of the cell types use graded
depolarization to release neurotransmitter to the next cell. A depolarization increases
neurotransmitter release. Small depolarizations cause small release of neurotransmitter; large
depolarizations cause large release of neurotransmitter.
H. The relationship between different cell types in the retina.

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 1. The retina is "inside-out" with the photoreceptors furthest away from the light (at the very back
of the eye) and the RGCs the closest to the light. Thus light must pass through the other cell types
to reach the photoreceptors.
2. This works because all the cells in the eye, except the photoreceptors, are translucent.
3. Also, at the foveal pit all cell types, except the photoreceptors, are pushed out of the way (see
9.15).

III. How light is absorbed by photoreceptors.

A. Phototransduction is how light energy leads to a change in membrane potential.
B. Below is a diagram depicting how light changes photoreceptor membrane potential.

Also, see Figure 9.17

DARK: RMP ~ -30 mV

Photoreceptor
membrane
voltage

Dim Moderate Bright

-30mV

Highest release

Glutamate release < as light increases

Lowest glutamate release

-65mV

Time (msec)

1. The resting membrane potential of photoreceptors is -30 mV and this is in the dark.
2. The maximum hyperpolarization is down to -65 mV and this is produced by bright light.
3. Glutamate is the neurotransmitter used by photoreceptors. The greater the intensity of light, the
less neurotransmitter released.

III. How does light produce the graded hyperpolarization?

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See Figure 9.19

1. The ligand-gated Na+ channels in the outer segment membrane are open in the dark, causing
depolarization (to the “resting membrane potential” of -30 mV).
2. These ligand-gated channels are like receptors, but they are "inside-out", meaning that they
bind their ligand cGMP to a binding site on the intracellular face of the Na+ channel and this
opens the channel.
A. How does light decrease the concentration of cGMP?
1. The photopigment, which is highly concentrated in the membrane of the disks in the outer
segment of the photoreceptors, is purple in the dark. When it absorbs light the photopigment is
bleached to a pale yellow.
2. The photopigment is called rhodopsin, and it consists of two parts...
a. Opsin is a protein.
b. Retinal, which is the only light sensitive molecule anywhere in the visual system. The
precursor of retinal is vitamin A. Retinal exists in 2 conformations.
1) In the dark it is 11-cis-retinal.
2) A photon of light will switch it to the all trans-retinal conformation.

See Figure 9.18.

a. Opsin passes through the membrane seven times.

b. The release of retinal from opsin allows opsin to change shape and this activates a G-
protein (transducin).

See Figure 9.19.

c. The G-protein (G) dissociates and travels along the membrane and activates an enzyme
(cGMP phosphodiesterase).
d. cGMP phosphodiesterase converts cGMP to GMP, and thus lowers the concentration of
cGMP.

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 e. In the dark, cGMP is bound to the Na+ channel. Light decreases the concentration of
cGMP, causing cGMP to disassociate from the channel. Consequently, the channels close
and the photoreceptor cells hyperpolarize.

B. Why do we have this type of system (what is the advantage)?

a. The increased surface area and increased photopigment produced by having the
photopigment molecules on the stacked disks, instead of on the Na+ channels, increases the
chance of the light being detected by a rod. This system is so sensitive that a single photon
can produce a detectable change in membrane potential of a rod type photoreceptor.
b. The use of G-proteins allows for amplification; each molecule of opsin can activate many
G-proteins, each of which, in turn, can activate many enzymes of cGMP phosphodiesterase,
each of which can, in turn, convert many molecules of cGMP into GMP.

Neuroscience 106: Lecture 11 - The Retina, continued

Assignment: In Bear et al. study Chapters 9 and 10.

TODAY'S LECTURE: The Visual System

I. Rods versus Cones
A. Rods contain rhodopsin and are more light sensitive than cones (a single photon of light may be
detectable by rods but not by cones).
1. One reason for this is that the process of signal amplification is greater in rods.
2. Cones have coneopsin instead of rhodopsin.
a. There are three different types of coneopsin: red, green, and blue. Meaning that they are
most sensitive to light in the red, or green or blue wavelengths.
b. The difference between the three forms of coneopsin is small. The difference is a small
change in the amino acid sequence allowing for maximal sensitivity to different wavelengths of
light. As an interesting tidbit, there are 2 versions of the red coneopsin gene in men and these
produce red opsins that vary by a single amino acid. This difference results in the two red
coneopsins being maximally sensitive to slightly different wavelengths of red light. Thus, men
with different versions of this opsin do not perceive the same thing when they see a red object.
This answers the philosophical question about whether different people see objects, colors etc.
in the same way. They don't.
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 3. Another reason that rods are more sensitive to light is that the outer segment of rods is larger;
therefore they have a larger surface area to absorb light.
4. Rods also have more photopigment densely packed into the membrane of the optic disks, so
they absorb more light.
B. In bright light the photopigment in rods (but not cones) is saturated (bleached). Therefore rods
aren't functional in bright light, while cones are.
1. This means that we have two parallel visual systems, one for bright light and one for very dim
light.
2. At night colors appear to be muted. But the same spectral frequencies exist in bright and dim
light. The reason we don’t perceive colors in dim light is that cones don't work in dim light, so
we don’t perceive color because we’re using a part of the visual system that is color blind.

II. Receptive fields of cells.

A. How do RGCs code information that results in perception of the visual stimulus?
B. Below is a diagram exemplifying receptive fields of RGCs.

Photoreceptors

RGC

Receptive
Fields of
RGC in
different
locations

1. A neuron’s receptive field is the location in the environment (or the surface of the body) from
which an appropriate stimulus will change that cell's activity. The term Receptive Field applies to

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 cells in the visual system other that RGCs and also applies to neurons in other sensory modalities
(e.g., touch, auditory).
2. Different RGCs have receptive fields in different locations in the visual field/retina.
3. RGCs in the fovea have exquisitely tiny receptive fields and the size of the receptive fields
increases in RGCs with distance from the fovea.
4. Light anywhere within the cell’s receptive field will change the cell's activity.
C. Some features of receptive fields of RGCs:
1. The receptive fields of RGCs are circular.
2. The receptive fields of RGCs vary in size.
3. Receptive fields of adjoining RGCs may overlap.
D. What are the mechanisms that account for the differences in size of receptive fields?
1. RGCs on the edges of the retina (so their receptive fields are in the periphery of the visual field)
collect information from a greater number of photoreceptors than do RGCs closer to the fovea.
Therefore, their receptive fields are larger.
2. Convergence is a situation where many neurons converge onto a few neurons. (There are about
120-130 million photoreceptors and only 1 million RGCs; therefore there is a lot of convergence
of photoreceptors onto RGCs).

This is shown in the diagram below.

Photoreceptors

RGC

Convergence of synaptic input to No convergence =

RGC = large receptive field. small receptive field
Mostly in periphery of retina. Mostly in fovea.

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 3. Divergence is few (or one unit) projecting onto many units.

Divergence: One or a few units projecting to many units.

4. In the fovea there is less convergence of photoreceptors onto RGC (via bipolar cells) than in the
periphery of the retina and this is a mechanism that explains the differences in size of the
receptive fields of RGCs.

E. Convergence also helps to explain light sensitivity. There is less convergence in the cone system,
therefore RGCs receiving input from cones are not as sensitive to light.

F. Receptive fields overlap and the consequence of this is that light from one point in the visual field
will affect many different RGCs.

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 These photoreceptors
project to both RGCs

ON Center RGC because it

generates AP with light onto
receptive field center.

OFF Center RGC because it stops generating AP when light is

shined onto its receptive field center.

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F. Receptive fields of retinal cells are modeled as concentric circles.
1. There are two types of concentric fields, on center and off center (for both BPs and RGCs).
2. These two RGCs are defined by their response to light in the center of their receptive fields.
On center RGCs are turned on (generate AP) by light and Off Center RGCs are turned off by
light in there receptive field centers.
3. Off center RGCs generate more action potential generation in the dark than do on center RGCs.
4. These concentric fields have antagonistic centers and surrounds because light falling in the
center of their receptive fields has the opposite effect of light in their receptive field surrounds.
5. Below are some examples of the effects of light falling on different portions of an ON and an
OFF center receptive field of a RGC.

[See power point figure].

a. Antagonistic… effect of light on the center is opposite to the effect of light on the surround,
and if light covers both the center and the surround, the effects cancel out.
G. Center/surround properties.

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 1. That our visual system is set up with centers and surrounds that are antagonistic to each other
suggests that the visual system is more sensitive to contrasts in the intensity of illumination than
to total brightness.
2. See the PowerPoint figure showing the responses of RGCs with different receptive fields just
outside versus just inside the border of the light reflected off of a sheet of white paper on a black
background..

For an On Center RGC, when the light completely fills the center and only part of the
surround, the net effect is an increase in action potentials.
At a corner the response is greater because the center is still completely filled and less of the
surround is filled.
For an On Center RGC, light falling only on the receptive field surround actually suppresses
the ongoing rate of AP production.
When light either fills both the center and the surround or it is dark there is little to no net
effect on action potential production. The action potential rate is nearly the same as the
baseline condition.
3. On center and off center RGCs overlap completely in the retina and their response to light is
opposite to each other.
H. Why is the visual system organized to detect contrasts?
1. The logical answer is that contrasts in light intensity are more informative than the overall
illumination. Where are the contrasts in the figure below?
2. Another way to say this is that the amount of light reflected by an object can vary dramatically
depending upon ambient illumination, but we do not want our perceptions to vary dramatically in
different lighting conditions.

I. Mechanisms for on center versus off center receptive field properties.

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 1. It is because of a difference in receptor type expressed by BP cells. The receptor expressed by
the On Center BP cell produces an IPSP in response to glutamate and the Off Center BP cell
produces an EPSP in response to glutamate.

Left Side: Responses of On Center Right Side: Responses of Off Center

Bipolar and RGC to light shining Bipolar and RGC to light shining onto
onto receptive field center. receptive field center.

Glutamate is excitatory here: In light,

< glutamate from PR cell produces
Glutamate inhibitory here: In light, hyperpolarization in BP cell.
< glutamate from PR cell produces #3
depolarization in BP cell.

Glutamate excitatory here: In

light, > glutamate from BP cell Glutamate is excitatory here:
depolarizes RGC. In light, < glutamate from
#1 #2
BP cell hyperpolarizes RGC.

2. When is a neurotransmitter excitatory? Answer: When its presence produces depolarization.

#1 Glutamate, which is a transmitter released by the bipolar cells, is excitatory here because an
increase in neurotransmitter release results in a depolarization.
#2 Glutamate is excitatory here because a decrease in neurotransmitter release results in a
hyperpolarization.
#3 Glutamate in inhibitory here because a decrease in neurotransmitter release results in a
depolarization.
(A little hint: If the direction of polarization (“synaptic sign”) is the same in two neurons where
one is being driven by the other, then the neurotransmitter between them is excitatory. In
contrast, if the direction of polarization is opposite, the neurotransmitter is inhibitory. Also,

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 remember that neurotransmitter release by a presynaptic terminal is increased by depolarization
and decreased by hyperpolarization).
3. So, the same neurotransmitter (glutamate) is released onto both ON and OFF center BP cells
but there are different receptors for that neurotransmitter the BP cells (glutamate inhibits the
receptor expressed by the ON center BP cell and glutamate excites the receptor on the OFF center
BP cell) and this accounts for the difference between on and off center receptive fields.
4. Why do we have both ON and OFF center systems? The on center system is most sensitive to
increases in illumination and off center system is most sensitive to decreases in illumination.

Neuroscience 106: Lecture 12 - Retinal Processing and Outputs.

ANNOUNCEMENTS: From this point on, much of lecture material will not be found in the book, so it
is particularly important that you come to class!!!

TODAY'S LECTURE: The Visual System

I. Mechanisms for the formation of an antagonistic surround.

A. The key mechanism in the formation of the antagonistic surround is the horizontal cell.
B. The diagram below shows the role of a horizontal cell (HC) when light falls on a
photoreceptor (PR) in a bipolar cell's (BP) and RGC’s surround.

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Responses when light shines
to center of ON center
receptive field.

PR PR

HC

BP
Responses to
light onto
surround of
On center RF.

RGC

C. How does light falling on the center and surround of a BC or RGC receptive field produce
antagonistic effects? Light in the surround of a bipolar cell's receptive field produces the opposite effect
on the membrane potential of a photoreceptor than light falling in the center. This is because of the
horizontal cell’s synaptic connection between photoreceptors in the surround and photoreceptors in the
center of a bipolar cell's and retinal ganglion cell’s receptive field.
D. Figure 9.22 in Bear et al. is an excellent depiction of a single bipolar cell, with synaptic inputs from
photoreceptors from its center, surround, and outside of its receptive field.

II. What happens to the output from the retina? (Where do axons of retinal ganglion cells
(RGCs) go?)
A. Below is a diagram illustrating general features of serial and parallel processing.

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 Serial Processing

Parallel processing + serial processing

The next diagram is an overview conception of the organization of the visual systems. Note that there is
massive parallel and serial processing of the retinal output.

V3

Lateral
Eye Geniculate V1 V2 V4
Nucleus
(LGN)

V5
Superior
Colliculus
(SC)
These are all regions in
visual (V) cortex
Suprachiasmatic n.
(SCn)

Different portions of the visual system (or visual systems) have different functions. For example:
1. The superior colliculus is a tracking system for orienting the eyes towards visual (and also auditory
and somatosensory) stimuli in the environment.
2. The suprachiasmatic nucleus of the hypothalamus (SCn) is like the brain's clock. Neurons in the
SCn have intrinsic firing frequencies of about 24 hours. Things such as sleep/wake cycles are controlled
by the SCn. Light input to the SCn from the retina resets the SCn clock to the rising and the setting of
the sun.
B. In most of the components of the visual system, visual information is mapped in an

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orderly point to point fashion: These are retinotopic and visuotopic maps.
1. Visuotopic map: every point in the visual world is mapped in a point to point fashion onto various
brain areas in the visual system (including the retina).
2. Retinotopic map: every point in the retina in mapped in a point to point fashion onto various brain
areas in the visual system.
Visuotopic and retinotopic maps are essentially the same. What is different is whether the visual field or
the retina is used as the initial point of reference. The visual world is repeatedly represented by
visuotopic maps in different brain regions (to date there are 25-30 known visuotopic maps in the
brain!!).
C. How do these orderly representations (maps) occur? The short answer is that the orderly location of
the RGC in the retina is maintained by orderly point-to-point neuronal projections from visual brain
area to visual brain area.

The diagram below shows how the retinas are divided into halves at the fovea and how the RGCs from
the temporal hemiretinas project to the ipsilateral (same side) of the brain while the nasal hemiretinas
project to the contralateral (opposite) side of the brain.

Nasal
Temporal Temporal
hemiretinas
hemiretina hemiretina

1. The retinas are vertically divided in half through the fovea.

2. The two halves of the retina have different patterns of projections.
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a. Nasal hemiretinas project to the opposite side of the brain (contralateral
projections).
b. Temporal hemiretinas project to the same side of the brain (ipsilateral projections).

D. Below is a diagram depicting how a visuotopic map is produced. However, the general answer is
that the orderly representations of the visual field in the brain are produced by orderly patterns of
anatomical connections. (Also, see book Figure 10.8)

Your Visual Field

A B C D E
F

E B
Eyes/Retinas A F
D B C
C B E D

D C

E B

F A

Superior Colliculus

1. This diagram represents a bird’s eye view.

2. Light from objects in the extreme periphery of the visual field (A and F) to the contralateral eyes is
blocked by the nose, so they are detected only in the ipsilateral eye.
3. The superior colliculus on each side of the brain contains a complete visual representation of the

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contralateral visual field.
4. So, the mechanism for the formation of the visuotopic map is that the nasal
hemiretina’s RGCs project in an orderly manner to the contralateral side of the brain and the temporal
hemiretina’s RGCs project in an orderly manner to the ipsilateral side of the brain. Also, the axons
from RGCs in either eye that are activated by light from the same object converge on the same neurons
in the superior colliculus.
5. Therefore, if you were to record from neurons at point B in the superior colliculus, the cells there
could be driven by inputs from the right eye, the left eye, or both eyes.
6. At F or A locations in the superior colliculus the cells respond to light input only from the
contralateral hemiretina.

III. Similarly, our nervous systems sometimes integrate information using the same general types of
mechanisms, even when the information comes from different sensory modalities (i.e. from visual and
auditory). Visual and auditory information enters the eyes and ears, which are obviously separate. How
do we integrate the information from both organs?
A. It happens because information from the different sources is brought together in the brain in the
superior colliculus.
B. Below is a diagram depicting how information from different sensory modalities is mapped in the
superior colliculus.

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 Cross section through superior colliculus showing
C location specific multimodal responses of neurons

B B
A
C
A

Respond to sight

Respond to sound

Respond to touch

Cause movement

1. There are several functionally distinct layers in the superior colliculus, and cells in each layer respond
to different sensory modalities.
a. Cells in the visual layer of the superior colliculus respond to visual information
from a single location. Neurons the auditory layer of the superior colliculus respond to auditory from a
single location, and cells in the somatosensory layer of the superior
colliculus respond to touch from a single location. These modality specific maps are aligned with each
other. For example, a visual stimulus at location B (in the visual field) will cause cells in location B in
the superior colliculus to fire. If that visual stimulus makes a noise (like a train whistle) then that will
drive neurons at location B (in the auditory field).
b. Stimulating neurons in location B of the motor layer of the superior colliculus will cause the eyes and
head to move to look towards location B in the visual field.
2. So one way we integrate information from different modalities is that we bring the information
together in the brain.

Neuroscience 106: Lecture 13 - The Superior Colliculus and Primary Visual Cortex
Assignments: Read Zeki, The Visual Image in Mind and Brain (in Reader).
In Bear et al. understand figure 10.5.

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TODAY'S LECTURE: The Visual System
I. The role of the superior colliculus in the visual system.
A. The superior colliculus acts as a tracking mechanism; it points the eyes and head toward a visual
object.
B. Below is a diagram depicting the superior colliculus and its inputs from the retina.

Your Visual Field

A B C D E F

Eyes/Retinas E B
A F
D B C
C B E D

Optic nerve

Optic chiasm

Optic tract
D C

E B

F A

Superior Colliculus

1. Neurons in the anterior portion of the superior colliculus respond to a visual image on the
center of gaze from both eyes (but only from the contralateral visual field).
2. Neurons in the posterior portion of the superior colliculus respond to a visual image in the
periphery of the visual field on the contralateral side.

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 3. Neurons in the center portion (at B) in the superior colliculus receive synaptic input from
retinal ganglion cells (RGCs) from both eyes that respond to light at position B in the contralateral
visual field.
4. This anatomical pattern of connections is the mechanism for generation of retinotopic maps in
the superior colliculus. (See Figure 10.7)
C. Below is a cross section diagram of the superior colliculus showing how visual, auditory,
somatosensory and motor maps are aligned with each other.

C Cross section through superior colliculus showing

location specific multimodal responses of neurons

B
B
A
C
A

Respond to sight

Respond to sound

Respond to touch

Cause movement

Eye muscle Oculomotor n. Neck muscle

1. Neurons in different layers in the superior colliculus respond to input from different sensory
modalities: visual, auditory, and somatosensory.
2. Each layer contains a map of a particular sensory modality, and these maps are all aligned (or in
register) with each other.

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 3. The motor layer contains motor neurons, not sensory neurons. Stimulating neurons at position B in
the motor layer of the superior colliculus causes the head and eyes to move to look towards location
B in the visual field.
a. Axons from the motor layer of the superior colliculus terminate in the oculomotor nucleus and
neurons from the oculomotor nucleus project to muscles that move the eye.
b. Axons from the motor layer of the superior colliculus also terminate onto motor neurons which
project to muscles of the neck.
D. This sensory tracking, eye pointing, subsystem is important for survival. If you hear something
unfamiliar, this system will cause your head and eyes to orient towards the sound to visually identify
what it was you heard. This occurs without conscious awareness.
E. The existence of this visual subsystem explains (and is the reason for) a phenomena called
blindsight.
1. Blindsight is “sight” that remains when a person is cortically blind. (The eyes and optic nerve
work properly, but the visual cortex is damaged so that the person cannot perceive visual images).
2. A person that is cortically blind cannot “see” or identify visual objects, but can localize and
track objects in visual space and they do this without awareness.
3. The superior colliculus is homologous to the optic tectum, which is the main visual brain area
in non-mammalian vertebrates. Apparently, our superior colliculus became specialized for
tracking and lost the ability to contribute to object identification, as that function expanded in
visual cortex.
F. You now have an example of a complete neural circuit, starting from photoreceptors in the eyes to
the muscles that produce movement of the eyes and head.

II. The cortical visual subsystem.

A. Thus far we have discussed two visual subsystems.
1. The superior colliculus which is involved in object tracking.
2. The suprachiasmatic nucleus, which is the brain's clock. The suprachiasmatic nucleus is
located dorsal to (above) the optic chiasm. Spontaneous daily patterns of activity of neurons in
this region are the reason that people experience jet lag and the reason people get over jet lag
(visual input into the suprachiasmatic nucleus resets the brain's clock).
B. Another visual subsystem is the cortical portion of the visual system, which is the part used in the
perception of a visual stimulus.

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C. Below is a diagram of a side view of a human brain depicting some of the major pathways in the
cortical subsystem of the visual system in a simplified form.

Lateral Geniculate
Nucleus of the “Dorsal stream”
Thalamus (LGN)
(LGn)
V5 V3

Eyes
V2 V1
V4
Inferotemporal
Cortex (IT) “Ventral stream”

1. Neurons in different regions of the visual cortex (V) have different functions.
a. Neurons in V5 are highly sensitive to movement but not color of objects in the visual field.
b. Neurons in V4 are highly sensitive to color but not movement of objects in the visual field.
2. Perception of visual objects is dependent upon activity of neurons in these cortical areas (V3,
V4, V5...). For example, if a person’s V4 is destroyed then that person’s visual world will be
black and white and memories are also in black or white.
3. Each of these cortical areas are retinotopically mapped (V1, V2,....). The major exception is
the inferotemporal cortex (IT). Neurons in IT cortex are not retinotopically mapped and don't
respond to color or movement, but respond to complex forms, such as faces, hands, chairs, etc.
D. The diagram below shows the pattern of nasal and temporal hemiretina projections to the 6
layers of the lateral geniculate nucleus of the thalamus (LGN).

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 Your Visual Field

A B C D E F

Left nasal
hemiretina Right temporal
Eyes/Retinas E B
A F hemiretina
D B C
C B E D

LGN (6 layers)

A B C L6
D E F L6 B C L5
D E L5 A B C L4
D E F L4 B C L3
D E L3 B C L2
D E L2 A B C L1
D E F L1

Primary visual cortex

1. The basic form of projections from the retinal ganglion cells to the LGN is similar to that of the
retinal ganglion cells to the superior colliculus, so that neurons in the LGN respond only to visual
images in the contralateral visual field.
2. The LGN has six layers and each layer is retinotopically mapped, with neurons in each layer
receiving input from the RGCs from only one hemiretina. Consequently, the neurons in the LGN

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are monocularly (respond to input from one eye) driven. Each of these maps are aligned with each
other.
3. Why this repetition of visuotopic maps in the LGN? The cells in the different layers are
processing different aspects of the visual image.
4. There are physical differences in the cells of the different layers.
a. The cells in layers 1 and 2 are larger and are called magnocellular. Cells here respond to
movement and are achromatic (do not respond to color).
b. The cells in layers 3 through 6 are smaller and are called parvocellular. Cells here are color-
sensitive and are more sensitive to detail.
This is an example of parallel processing. The idea is that different aspects of a visual object
(aspects like color, movement, shape) are coded by different parallel pathways.
5. Projections to the primary visual cortex (V1) from the LGN are solely ipsilateral. Therefore,
neurons in V1 respond exclusively to visual input from the contralateral visual field. This means
that if the right V1 is destroyed, you would be blind to input in the left visual field (but still have
“blindsight” due to an intact superior colliculus).

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 E. The diagram below illustrates the visuotopic/retinotopic map in primary visual cortex.

The visual field divided into quadrants that are coded by letters, numbers and colors

1 3
A C
B D
2 4

Calcarine fissure

D 4 2 B

C A
3 1

Left and right primary visual cortex showing the visuotopic map.
That is, neurons at location “A” respond only to objects at
location “A” in the visual field.

1. Although there is a one-to-one relationship between locations in the visual field and locations in the
cortical areas that encode them, almost 1/2 of the neurons in V1 code for the fovea region even though
this is only a very tiny portion of the visual field. Apparently, this is because it takes a lot of neural
machinery to decode the complex and detailed information that we can perceive from this area of
highest visual acuity

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