LECTURE 11:

VISION
Learning Objectives
You should be able to describe the following about the retina
• Major structural features (fovea, macula lutea, optic disk)
• Neuron types
• Retinal circuitry
You should be able to discuss
• Photoreceptors and their role in scotopic, mesopic, and
photopic vision
• The origin of color vision
• Ganglion cell receptive fields
You should be able to
• Discuss the flow of visual information to the primary visual
cortex (V1) and how V1 processes visual information
• Relay the roles of the dorsal and ventral streams in visual
processing after V1
You should know how ganglion cells information is used by
superior colliculus, suprachiasmatic nucleus, and Edinger-
Westphal nucleus
Structure Of The Eye
Overview Of The Retinal Surface
Optic Disk (optic papilla)
• Allows axons of the retina (which comprise
the optic nerve to leave the eye along with
the eye’s blood supply
• DOES NOT have any photoreceptors,
making it light insensitive. This creates a
blind spot in the visual field
Swelling in this region indicates a dangerous
build-up of intracranial pressure. This is used
as a diagnostic during neurological
examination.
Fovea
• Area of the retina that provides the highest
acuity vision
• Contains only cone-type photoreceptors
• Avascular - lacks blood vessels to allow for
tighter packing of photoreceptors
Shown: Inner surface of the retina imaged using an
Macula Lutea ophthalmoscope
• ~3mm diameter area with fovea at the
center
• Contains pigment (xanthophyll) that
protects the retina by filtering out UV light.
The pigment gives the area its yellow color
Retinal Circuitry Has A Laminar Structure
The retina is unique among
sensory systems, because
it is a part of the central
nervous system. It arises
from the diencephalon
during development.

The retina is comprised of 5 neuron types

that are highly organized in 3 lamina
(layers)
• Photoreceptors → Outer nuclear layer
• Bipolar Cells → Inner Nuclear Layer
• Amacrine Cells → Inner Nuclear Layer
• Horizontal Cells → Inner Nuclear Layer
• Ganglion Cells → Ganglion cell layer

Even though photoreceptors are responsible for encoding information about light into neuronal
information, they sit at the back of the retina. Therefore, light has to shine past the ganglion,
bipolar, amacrine, and horizontal cell layers before getting to the photoreceptors.
Tremendous Diversity Exists Within
Retinal Neuronal Types

Masland (2001)

In mammalian retina, there are >50 different neuron subtypes within the 5 neuronal classes,
as classified by the details of their shapes. In many cases, how the neuron functions also
varies, suggesting that these differences are meaningful. Within a neuronal class, for example,
ganglion cells, the different subtypes exist with roughly equivalent frequency.
2 Types of Photoreceptors: Rods & Cones
Rods & Cones Mediate Vision Within Different Ranges Of Light
Luminance (Brightness)

Rods provide low resolution, colorless vision at low levels of light

‣ The absolute threshold for a rod is ~1 photon!
Cones provide high resolution, color vision but require more light

3 Types Of Vision
Scotopic (very low light): Rods Only
Mesopic (low light, like that of a night sky): Rods & Cones contribute
Photopic (Lots of light): Cones mediate vision (rods are saturated)
Rods & Cones Are Inversely Distributed
Throughout The Retina
Cones are concentrated in the fovea to
mediate high-acuity, color vision

Rods are predominant everywhere but

the fovea - mediate low light vision
Phototransduction: Turning Light Into A Neural Signal
Unlike other senses, light does not lead directly to firing of
action potentials.

Instead, greater light hyperpolarizes the photoreceptor leading

to a proportional reduction in neurotransmitter (glutamate)
release by the rod or cone.
Retinal Circuitry For Vision Mediated By Rods
& Cones Is Different
Retinal circuitry is complex, involving many parallel processing neuronal microcircuits and various
levels of feedback (for example, horizontal cell → rod or cone OR amacrine cell → bipolar cell. What
follows is a simplified description of the feedforward paths for rods and cones.

• STEP 1: Both rods and cones synapse onto bipolar cells (neurons)
◦ Bipolar neurons for a rod are of a different, single subtype than those
for cones, which are of 9-11 cone-specific bipolar cell subtypes
◦ Typically, many rods synapse onto one rod bipolar cell, whereas, one
cone will synapse onto one cone bipolar cell
Retinal Circuitry For Vision Mediated By Rods
& Cones Is Different
Retinal circuitry is complex, involving many parallel processing neuronal microcircuits and various
levels of feedback (for example, horizontal cell → rod or cone OR amacrine cell → bipolar cell. What
follows is a simplified description of the feedforward paths for rods and cones.

• STEP 2: Cone Direct Pathway v. Rod Indirect Pathway

◦ Cone Direct Pathway: Cone bipolar cells synapse onto ganglion cells
(the output neurons of the retina to the brain)
◦ Rod Indirect Pathway: Rod bipolar cells synapse onto one specific
subtype of amacrine cell (neuron) which then synapses onto a cone
bipolar cell

Masland (2001)
Cone Diversity Supports Color Vision

Human Color Vision Is Trichromatic (Based On 3 Colors)

• S (blue) cones are only 5-10% of cones in the retina of primates
• M (green) : L (red) ratio can vary from 4:1 to 1:1 across individual humans with
little apparent impact on color perception
◦ Red cones do not exist in many mammals
Each type of cone contains a different photopigment, which provides
different responses to light of a given color
 • Color blindness occurs when the ability to
Color Blindness process 1 of 3 cone photoreceptors paths fails
• Color blindness can have many causes
◦ Genetic: Missing or mutated photopigment
gene
◦ Processing: Damage to “color-processing
areas of the brain
• Red-green color blindness is the most
common form of color blindness
◦ 8% of males v. much lower proportion of
females
‣ Genes for red and green photopigments
are close to one another on the X
chromosome, making these genes
especially vulnerable to deletion or
mutation (due to unequal recombination
of intergenic regions)
‣ Green (1 gene copy) is more vulnerable
than red (1-5 gene copies), but loss of
either abolishes the ability to distinguish
red from green
• Blue-yellow color blindness can result from
loss of the blue photopigment gene on
chromosome 7, but this is much rarer due to the
need to lose or have mutated 2 copies

• Anomalous trichromats also exist - they require

unusual combinations of light intensities at
different colors to make typical color distinctions
Ganglion Cells “See” The World As A Contrast
Of Light & Dark Ganglion cells have
circular receptive fields
and respond to changes
in contrast between the
center of the receptive
field and the annulus
surrounding the center

ON-center ganglion
cells increase spiking
when the center of the
receptive field increases
in intensity relative to the
surround

OFF-center ganglion
cells increase spiking
when the center lowers in
intensity relative to the
surround

The nature of the receptive

field, including size, location in
the visual field, and ON or OFF
centeredness is determined by
the specific connections other
retinal cell types make
upstream of the ganglion cells
Central (Brain) Processing
Of Visual Information
Ganglion Cells Of The Retina Project To
Cortical And Non-Cortical Brain Structures

PrimaryVisual
Cortex Vi
Visual Pathway To Primary Visual Cortex (V1)

Tempord
Temporal

M
Visual Pathway To Primary Visual Cortex (V1)
Notes To Previous Slide
Optic Nerve (A): The axons of the ganglion cells come together into a bundle that
leaves the eye via the optic disk (blind spot). This bundle is called the optic nerve.
It carries all of the visual information from one eye.

Optic Chiasm (B): At the optic chiasm, the optic nerve splits in half, sending
axons from the nasal (medial) half of the retina to the contralateral side of the brain
to meet up with the axons from the temporal (lateral) half of the retina from the
contralateral eye.

Optic Tract (C): The optic tract carries all of the visual information for the
contralateral half of the visual field (for example, the left optic tract has the visual
information from the person’s right visual field) to the dorsal lateral geniculate
nucleus (dLGN) of the thalamus.

Optic Radiation (D): All of the visual information from the contralateral half of the
visual field is relayed from the dLGN of the thalamus to the primary visual cortex
(V1) via the optic radiation. (V1 is in the occipital cortex). The optic radiation splits
into 2 parts. The first carries information from the upper half (superior) of
contralateral visual field in nerve fibers that run through the temporal lobe on their
way to V1. This is called Meyer’s Loop. Information about the inferior (lower half)
visual field goes to V1 by the portion of the optic radiation that runs inner the
parietal lobe.
First Stop: Dorsal Lateral Geniculate
Nucleus (dLGN) - The “Visual Thalamus”
Dorsal

IS

X
Ventral

Different aspects of visual information is carried by 3 different ganglion cell subtypes,

which synapse onto separate groups of relay neurons in the thalamus.
Parvocellular (P) Ganglion Cells
• Smaller receptive fields relative to M ganglion cells
• Respond in sustained fashion to visual stimuli
• Convey color information - each center driven by a different cone type (Blue, green, or red)
than its surround
• Damage to P ganglion cells impairs visual acuity and color perception
Magnocellular (M) Ganglion Cells
• Larger receptive fields than P ganglion cells
• Respond transiently to the presentation of visual stimuli
• Each center and its surround driven by the same cone type - do not provide color
information as a result
• Damage to M ganglion cells impairs ability to perceive rapidly changing stimuli (motion)
First Stop: Dorsal Lateral Geniculate
Nucleus (dLGN) - The “Visual Thalamus”
Dorsal

IS

X
Ventral

Different aspects of visual information is carried by 3 different ganglion cell subtypes,

which synapse onto separate groups of relay neurons in the thalamus.
Koniocellular (K) Ganglion Cells
• Contribution to visual perception not well understood
• Some aspects of color vision, especially related to short-wavelength (blue) cones may be
transmitted by K ganglion cells

Note: Although the dLGN receives inputs from both eyes, the inputs are segregated into
separate layers.
Second Stop: Primary Visual Cortex (V1) -
Where Binocular Vision Is Established
Mixing of visual information from the two eyes first takes place in primary visual cortex.

Visual information for each eye

arrives independently at V1, where
it is mixed. Thus, it is in V1 where
neurons for the first time show
responses to inputs from both eyes.
The relative strength of neuronal
responses to each eye varies in an
anatomically coordinated manner
called ocular dominance columns.
Neurons In Primary Visual Cortex Respond
Strongly To Stimuli With Oriented Edges

Neurons in V1 typically respond vigorously to a bar of light oriented at a particular

angle and less strongly - or not at all - to bars at other orientations.

Neurons in V1 can also have a preferred direction of stimulus motion and preferred
spatial and temporal variation of stimulus contrast

Computational analysis suggests receptive fields with these properties are well
tuned to the structure of natural scenes. Thus, detecting edges provides for an
efficient neural code that maximizes information and minimizes code redundancy.
What Happens After V1? - Processing
Objects & Motion

Anatomical, electrophysiological, and regional damage studies indicate that

visual information from V1 is processed in 2 largely separate paths, one
which feeds information into the medial temporal and parietal lobes (dorsal
stream) and the other which feeds information into the inferior temporal
lobe (ventral stream)
• Dorsal Stream: Responsible for generating the spatial aspects of vision, including motion
and positional relationships between objects in the visual scene (involves a stream of
interconnected areas in medial temporal and parietal lobes)
• Ventral Stream: Generates a high-resolution form of vision for object recognition (relies upon
processing by areas in inferior temporal lobe
Ganglion Cells Of The Retina Project To
Cortical And Non-Cortical Brain Structures
Light-Sensitive Ganglion Cells Help Align
Circadian Rhythm With Day & Night

Retinal
GanglionCell

• Some retinal ganglion cells contain melanopsin,

which makes them light-sensitive.
• These ganglion cells project to and excite the
suprachiasmatic nucleus of the hypothalamus,
which is considered the master circadian oscillator.
• The suprachiasmatic nucleus inhibits the
paraventricular nucleus of the hypothalamus,
which in turn lessens sympathetic drive to the
pineal gland.
• The pineal gland generates melatonin which
promotes sleep. This helps to align the circadian
rhythm to light and dark.
Ganglion Cells Provide Light Information For
Pupil Adjustments

Bilateral projections from ganglion cells in the retina project to a collection of

neurons in the pretectum, which lies between the thalamus and midbrain.

Pretectal neurons then project to the Edinger-Westphal nucleus in midbrain, which

contains the preganglionic parasympathetic neurons that project to the ciliary
ganglion whose neurons control pupil diameter
Specific Ganglion Cells Project To Superior Colliculus
To Drive Coordinated Eye Movements (Saccades)
The superior colliculus
receives visual input from the
retina and sends a command
to the gaze centers to initiate a
saccade (coordinated eye
movement).

The visual layer of the superior

colliculus forms a topographic
map so that each site in this
layer is activated maximally at
a particular point of visual
space.

Activation of visual layer

neurons is enough to generate
saccades, but saccades can
also be generated in the
absence of visual layer activity,
and saccades don’t always
occur as a result of activity in
the visual layer. This suggest
complex control of eye
movements.