The Visual System

General plan for visual system material:

How the visual input is received and transduced at
the retina by photoreceptors (rods and cones)
How that input is modified by processing in the
retina, concept of receptive fields
How that input is projected to the central nervous
system (to the lateral geniculate nucleus (LGN)
and cortex), the anatomy and processing within
that pathway
 The Retina
Images focused by
the lens go through
the liquid within the
eyeball and fall on
the retina. In most
of the retina, layers
of neurons lie over
the photoreceptors,
and light must go
through them.
Fig 15-1
 The Retina
At the fovea, the overlying cells are displaced, so the visual image
is clearest there. There are more photoreceptors at the fovea.
The Retina
classes of neurons
Light goes through other
layers of neurons before being
transduced at the rod and cone
photoreceptors
Photoreceptors send the light
information to the bipolar
cells, which send it to the
ganglion cells, and the
information exits the retina to
the brain
Horizontal cells and amacrine
cells modulate the information
Fig 15-2
Photoreceptors of the Retina
Rods and cones

Fig 15-3
 Photoreceptors of the Retina
Rods and cones are specialized
Rods are highly sensitive to light, and thus are good for
dim light or night vision. They are able to capture more
light, but they do not respond well to moving stimuli
because their response time is slow.
Cones are not as sensitive to dim light, and are able to
respond quickly, and therefore transduce moving stimuli.
They are used more in daytime vision, and also are able to
detect color because three classes of cones express three
different photopigments. Cones are more concentrated in
the fovea of the retina.
 Photoreceptors of the Retina
The outer
segments
contain
stacks of
membranes,
where the
light-
absorbing
molecules
are found

Fig 15-7
Light response
Light causes a
hyperpolarization
In darkness, the rod
membrane potential is
relatively depolarized.
Light causes a
hyperpolarization by
reducing a Na
permeability; the current
is through a cGMP-
dependent cation
channel.
Light response
Light causes a
hyperpolarization
Light causes closure
of a cation-permeable
channel, via a process
using cGMP as a second
messenger. If more
cGMP-dependent cation
channels are closed, the
membrane potential is
more hyperpolarized.
Light transduction
Light is detected by
retinal
Rhodopsin (from rods)
is a combination of
retinal, the light-
absorbing molecule, and
a large protein called
opsin. The opsins are
closely related in
structure to the G-
protein-coupled
receptors.

Fig 15-9
Light transduction
Light changes
retinal conformation
When retinal absorbs
light, it undergoes a
conformational change.
This change moves the
position of the opsin
molecule, which
activates a G protein
(transducin) and begins
a second messenger
Fig 15-8
pathway.
 Phototransduction - cGMP changes
Transducin activates a phosphodiesterase
In the dark, phosphodiesterase activity is quite low, thus levels of
cGMP are high; the cGMP-dependent cation channels are gated
open. When light activates rhodopsin, the levels of cGMP are
reduced by the phosphodiesterase, and the channels are closed.
 Phototransduction - dark current
In the dark, Vm is approximately -40 mV
In the rod, the cGMP channels
are located in the outer segment,
and a K channel is present in the
inner segment. In the dark, when
the cGMP concentrations are
high, current flows in through the
cGMP-gated channels and out
through the K channels.
 Phototransduction - dark current
Light hyperpolarizes the photoreceptor
With light stimulation, the cGMP
channels close.
Pathway: light rhodopsin
rhodopsin transducin
transducin phosphodiesterase
phosphodiesterase cGMP
cGMP cGMP-gated channel
 Phototransduction - dark current
Light hyperpolarizes the photoreceptor
With light stimulation, the cGMP
channels are closed.
Pathway: light rhodopsin
rhodopsin transducin
transducin phosphodiesterase
phosphodiesterase cGMP
cGMP cGMP-gated channel

Fig 15-11
 Phototransduction - dark current
AMPLIFICATION!
One rhodopsin molecule absorbs one photon
500 transducin molecules are activated
500 phosphodiesterase molecules are activated
105 cyclic GMP molecules are hydrolyzed
250 cation channels close
106-107 Na+ ions per second are prevented from entering the
cell for a period of ~1 second
rod cell membrane is hyperpolarized by 1 mV
Retinal Processing
Horizontal cells mediate
lateral interactions
Horizontal cells are
post-synaptic to many
photoreceptors, and in
the dark, receive a large
glutamate stimulus; thus
they are depolarized in
the dark. Light causes
hyperpolarization. Via
electrical synapses (gap
junctions) they also then
hyperpolarize their
neighboring cells.
Fig 15-13
 Processing in the retina
Cones send information in parallel pathways
Bipolar cells are a direct
pathway from the
photoreceptors to ganglion
cells. In the dark, cones are
Excitatory Inhibitory
depolarized and release
glutamate, which has opposite
effects on two types of bipolar
cells that mediate the parallel
direct pathways to ganglion
cells. When a cone is
stimulated by light, it reduces
transmitter release.
Retinal Processing
Off-type bipolar cells
hyperpolarize with light
In the dark, cones release
glutamate onto off-center
bipolar cells and open a Off-
ionotropic cation channel. type
This depolarizes the bipolar
cells until a light stimulus.
The bipolar cell then
hyperpolarizes, and reduces
the firing of the off-center
ganglion cell that is next in
the sequence.
Retinal Processing
On-type bipolar cells
depolarize with light
The on-center bipolar cells
are inhibited by glutamate
(by two different On-
mechanisms). With light type
stimulation, the glutamate
concentration from the cone
is decreased and the bipolar
cells are depolarized. This
increases the firing of the
on-center ganglion cell next
in line.
Retinal Processing
Horizontal cells influence a
wide distribution of cones.
At the photoreceptor-horizontal
cell synapse, in the dark, glutamate
will depolarize horizontal cells, and
horizontal neuron GABA will
hyperpolarize cones. With light, the
horizontal cell is hyperpolarized and
less GABA is released.
Light stimulus at one point of the
horizontal cell receptive field causes
a depolarization in other -
Fig 15-17 +
photoreceptors in the receptive field.
Retinal Processing
Synaptic interactions increase
the receptive field of bipolar cells
Off-center bipolar cells respond to
a spot of light by hyperpolarizing,
because of reduced cone glutamate.
Horizontal cells hyperpolarize with
light, reducing their inhibitory
transmitter output onto cones; with
less hyperpolarization, the cones
secrete more transmitter and bipolar
cell is depolarized. With the
inverted influence of the horizontal
cell, the bipolar cell response is a
combination of the two.
Fig 15-16