Compare the Properties of Rods and Cones, and their Retinal Pathways.

When we investigate vision, we see that it can be divided into two distinct
phases – the conversion of light, which has been taken in through the cornea, into
electrical signals and the processing of these impulses by higher areas of the brain.
The retina, through specialised neurons called photoreceptors, is responsible for
the first of these events.
Lining the back of the eyeball is a thin layer of neuronal cells, where action
potentials resulting from incoming light instigate a nervous response that is
relayed to the brain for image resolution. This is the retina. Comprising five
separate layers, the retina have photoreceptive cells, termed rods and cones, on
their outer nuclear later, which are responsible for our interpretation of our visual
surroundings. Inside this layer is the outer plexiform layer, which contains the
dendrites of horizontal cells as well as the synapses between these, bipolar cells,
rods and cones. Then comes the inner nuclear layer, which comprises cell axons of
bipolar cells and the cell bodies of amacrine, bipolar and horizontal cells.
Dendrites of amacrine, bipolar and ganglion cells are found in the next layer, the
inner plexiform layer. The innermost layer, the ganglion cell layer, consists of the
cell bodies of ganglion cells, which are concerned with the output of the neural
impulses in the retina to the brain.
Histologically, rods and cones can be seen to have an inner and an outer
segment. At the inner segment, one will find the organelles associated with most
cells e.g. Mitochondria, nucleus and ribosomes while the outer segment consists
of a comb-shaped group of folds of the cell membrane. In rods, these often isolate
forming free-floating disks inside the rod whereas in cones the disks are attached
to the outer membrane. The large surface area, granted by these folds is ideal for
collecting photons with the photosensitive transmembrane proteins and thus one
can understand why in rods, which are more sensitive in dim light (i.e. Times
when every last photon needs to be gathered), the outer segments are much larger.
Together, rods and cones account for our concept of vision. Rods are
responsible for our night sight and therefore function in periods of dim light
whereas cones function in day light. Thus, in the absence of rods one would have
only night-blindness but loss of cones results in legal blindness. Although rods are
found in much more abundance than cones (20 times as much), there is only one
type of this photoreceptor while cones come in three variations, each one having
greatest sensitivity at a different wavelength of light. Both photoreceptive cells
share the same foundation pigment, that is a photosensitive pigment made of
retinal, a light-absorbing molecule, attached to a membrance protein called opsin.
The variation between the different photoreceptive cells, lies in the type of opsin
molecule bound as these differ in their peak light absorption energy. The three
types of cones exist as those which are excited by light with short, middle and
long wavelengths. The first concerns blue light (437 nm), the second green (533
nm) and the third red (564 nm) while rods have only one type with a peak
absorpency at 498
nm, provided by the pigment rhodopsin.
In terms of their actual detection, the photosensitive pigment found in the
rod cells is much more sensitive, capturing more light and also amplifying
incoming impulses to a greater extent than cones. Baylor and his colleagues
showed that a single photon had the ability to evoke a response from a rod but
hundreds of photons were required to excite a cone cell. However, in the rods
fewer photons are needed to elicit the maximum possible response relative to the
cone cells. The ability of spatial and temporal resolution derives primarily from
the cones for two major reasons. One, a high concentration of the cones can be
found in the fovea, a
site where vision is most acute due to displacement to the side of all cells except
the photoreceptors themselves hence reducing visual distortion. Secondly, a group
of rods generally synapse onto the same bipolar interneuron illustrating the
concept of convergence. Although the different impulses are capable of
reinforcing each other and thus generating an increased overall signal, the
precision of a given impulse is also decreased. This arises because the
convergence of several different responses means that the signal from these rods is
averaged when meeting at the relay neuron and so transmission of spatial
variations is relatively poor. Temporal resolution of a visual image is also much
more acute in cone cells. Impulses provided by the photoreceptor cells does not
rely on action potentials but rather the membrane potential of these cells varies
depending on the light present. The response by rods is relatively slow and
photons, which are absorbed, in a given time frame (approx. 100ms) will
generally summate and this aids in the detection of limited amounts of light during
periods of dim light. However in cones, the response is much quicker and thus a
greater degree of temporal resolution is attained.
Before the incoming light stimulus can be transmitted through the optic
nerve, it must be transduced into an electrical signal which is able to be
propagated. This begins with the absorption of a photon by one of the pigment
molecules. Whereas neurons often rely on a stimulus providing a depolarisation,
the resting membrane of these photoreceptive cells is depolarised near 40 mV and
activation generally leads to a hyperpolarisation of around 70 mV. In rods,
rhodopsin absorbs light and causes excitation leading to a conformational change
in the molecule retinal. Normally this is found in a 11-cis isomer however the
structural alteration produces a reaction where it is converted entirely into the all-
trans shape. Rhodopsin can not bind the all-trans type of retinal and so the
molecule becomes detached. During this, metarhodopsin II, a quite stable
intermediate, is formed and this activates the G protein transducin, which in turn
stimulates a phosphodiesterase thus converting cyclic GMP to 5' GMP. The newly
reduced levels of cGMP intracellularly results in a decreased influx of Na+ via a
cGMP-gated channel. Therefore we can see that a constant influx of Na+ occurs
while in the dark due to increased intracellular cGMP concentration. This flux is
known as the dark current. The retinal, which has been detached, is then recylced
into the pigment epithelium, a lining adjacent to the outer nuclear layer of the
retina, and converted to all-trans retinol or vitamin A. This molecule is the
precursor to 11-cis retinal, which can not be synthesised in vivo, and therefore one
can see that a lack of vitamin A may lead
to night-blindness.
Cones have a much greater range of photosensitivity which is achieved
through adaptation of the light. Transduction of the light into an electrical signal
also reduces the ability a cGMP-gated Ca 2+ channel. Thus cation inhibits the
enzyme guanylate cyclase and so a reasonably steady concentration of cGMP is
maintained, using this negative feedback system, despite possible changes in
surrounding brightness.
Having responded to incoming light, impulses generated by the
photoreceptor cells must be passed onto the higher centres of the brain. This is
done though synapses between a sequence of cells. Cones firstly synapse onto
bipolar horizontal cells, the former synapsing directly onto ganglion cells, the
output cells of the retina. The horizontal cells along with the amacrine cells, are
relay neurons, the first one affecting the impulses passed between the
photoreceptors and the bipolar cells and the second one affecting the information
between the bipolar and the ganglion cells. The horizontal cells synapse back onto
the cones supplying them and have an inhibitory impact but also are joined to
other horizontal cells via electrical synapses, creating a sheet which receives input
from cones. Cones therefore are constantly experiencing inhibition by
neighbouring cells, giving rise to the receptive fields which are distinct in the
early stages of the visual system. The cones also release the neurotransmitter
glutamate which has varying effects depending on the type of bipolar cell
addressed. In an on-centre bipolar cell, the molecule opens Na+ channels resulting
in a depolarisation while in off-centre bipolar cells the glutamate closes these
same channels and conversely open K+ channels generating a hyperpolarisation.
Action potentials are not fired until the stage of the ganglion cells, which the
bipolar cells synapse onto. Instead the bipolar cells use the notion of a fluctuating
membrane potential, which occurs as a result of the electrotonic conduction.
In conditions of sufficient light i.e. Photopic vision, the rods synapse onto
nearby cone cells through gap junctions however the adequate amount of light
bleaches their pigment so that their impact on vision can be disregarded.
Conversely, in scotopic vision, these same gap junctions close off and the rods
instead synapse onto neighbouring bipolar cells, which then connect with AII
amacrine cells. A2 amacrine cells will synapse with off-centre ganglion cells and
on-centre bipolar cells, which synapse onto on-centre ganglion. Thus the visual
input ultimately reaches the higher centres of the brain.
The importance of the rod and cone systems can be observed in pathologies
that affect these networks. The common condition colour blindness is a result of
one or more cone pigments being absent or defective leading to incomplete
interpretation of visual input. While this may seem like a rather insignificant
disease, the existence of much more substantial effects such as diabetic
retinopathy, where the elevated blood glucose levels can lead to severe damage in
the retina and thus distort vision massively, demonstrates the sheer importance of
the individual retinal components in contributing to our idea of vision.