As we navigate through our surroundings, a continuous stream
of light images impinges our eyes, in the back of each eye is a
light-sensitive tissue: the Retina. It converts patterns of light
energy into electrical discharges known as action potentials. These
signals are conveyed along the axons of retinal ganglion cells to
connect for 80% to the LGN (Lateral Geniculate Nucleus) a relay nucleus
in the Thalamus and for 20% to the SC (Superior Colliculus). Most of
the output of the LGN is relayed directly to the Primary Visual Cortex
(V1), and then to surrounding visual association areas."
The Retina is
the innermost layer of the eye, an outgrowth of the Central Nervous
System (CNS), made up of nerve cells and covering most of the inner
wall of the eyeball. When we look directly at an object, its image
falls on the Retina at a fixed spot, along the visual axis, that lies
in a slight depression: the Fovea, this tiny area is responsible for
our central, sharpest vision.
Special cells called Rods and Cones are used to process light, in a human eye, their number is
estimated to be 5-7 million Cones and 110-130 million Rods. Cones are
found mainly in the central area of the Retina (Fovea), while Rods are found in
the Peripheral Retina. The Photoreceptors respond to light, it causes the activation
and consequent breakdown of a photo-sensitive pigment, bound
in large quantities to the membranes of the outer segment. (Image: close-up of hundreds of Rods and 2 Cones:)
Rods: See
in black, white, and shades of gray and tell us the form or shape that
something has. They
are super-sensitive, allowing us to see when it's very dark.
Cones: Sense
color and need more light than Rods to work well. Cones are most
helpful in normal or bright light. There are 3 types of cones
- red, green, and blue - to help you see different ranges of color.
Together, these Cones sense combinations of light waves that enable
our eyes to see millions of colors.
Several layers of
the Retina correspond to different parts of cell groups. Horizontal
Cells connect Photoreceptors together
laterally, their
nuclei lies among the Bipolar cells.
The synaptic layer between Bipolar, Horizontal cells and Photoreceptors
forms the outer plexiform layer of the Retina. Amacrine cells connect
laterally between Ganglion cells, their nuclei also lies amongst those
of the Bipolar cells. These
interactions are important for determining the ways in which patterns
of
light affect the activity in Ganglion cells. The Axons of Ganglion
Cells
sweep across the Retina and meet at the Optic Disc and leave the eye as
the Optic Nerve at the Blind Spot, an area absent of Photoreceptors.
1.
Optic Fibre
Axons of the Ganglion cells. They are unmyelinated until they reach the
optic disc, where they leave the eye as the optic nerve.
2.
Ganglion Cell
Cell bodies of Ganglion cells.
3.
Inner Plexiform
Synapses between interneurones (Bipolar and Amacrine cells) and Ganglion cells.
4.
Inner nuclear
Nuclei and cell bodies
of certain interneurones (e.g. bipolar cells and horizontal cells) as
well as glial cells.
5.
Outer plexiform
Synapses between the photoreceptors and
retinal interneurones. Interneurones include horizontal and bipolar
cells.
6.
Outer Nuclear
The nuclei of the rods and cones.
7.
External Limiting Membrane
Glial cells' tight junction connections with the Photoreceptors.
8.
Receptord
Outer and inner segments of Photoreceptors, arranged in an
organised way so that particular Photoreceptors receive light from a
particular part of the visual field. This orgainisation is in part
maintained by retinal glial cells which are arranged
parallel to the direction of light through the Retina.
9.
Pigment epithelium
Crucial in maintaining the quality of an image, because the pigment absorbs stray light, thus preventing
scatter. It also prevents scatter of light between Rods and Cones by
extending down into the receptor layer. Another function of this
extension into the receptor layer is that contact is maintained between
these two layers.
Photoreceptors are composed of an inner segment and an outer
segment,
as well as a cell body and synaptic terminal. The main difference
between Rods and Cones is in the outer segment, where the visual
pigment is located:
Pigment in Rods: On flattened, internalised discs.
Pigment in Cones: On a region of infoldings of the membrane.
A cycle of events takes place in rhodopsin when light strikes on Rod-cells:
1. In the dark, opsin is bound tightly to retinene. 2.
When light intensity is increased, retinene changes shape.
(This is a
structural change from cis- to trans- form.)
3. Opsin cannot hold onto
retinene, retinene comes off (this process is called bleaching).
4.
Generator potential is produced when the membrane of a Rod is depolarized.
5. Generator potentials summate (add together) to form an action potential. 6. A nerve impulse is fired off to brain via the Optic Nerve. 7.
Rhodopsin is reformed when retinene resumes its original shape using
the energy in mitochondria (the power-house of every cell). This is
light independent (does not need light). 8. Rhodopsin is ready to be bleached again.
Photoreceptors -> Bipolar Cells: The receptors release glutamate on to the Bipolar- and Horizontal
cells, causing either excitation via ionotropic receptors or inhibition
via metabotropic receptors.
IF the Photoreceptor inhibits the Bipolar
cell then the Bipolar cell would be excited
(disinhibited) when the photoreceptor is hyperpolarized: "ON"
response.
OR the Photoreceptor excites the Bipolar cell, then light
would produce an "OFF" response.
Photoreceptors are therefore unusual in
the fact that their stimulus causes a hyperpolarisation, not a
depolarisation.
Photoreceptor
Rod
Cone
Photopigment =opsin + chromophore
rhodopsin
iodopsin
opsin
scotopsin
?
chromophore
retinene
retinene
Light Sensitivity
+++++
+++
Rhodopsin vs. Iodopsin:The light-sensitive
photopigments are made of the protein opsin and the
chromophore retinene. Bleaching
of iodopsin in Cones is similar to rhodopsin in Rods, only Rods contain
a higher concentration of visual
pigment than Cones, so more light is
needed to cause an action potential to be fired in Cones (threshold for
Cones is higher than Rods). In other words, Rods are mainly used for
dim light vision, Cones for bright light vision. A period of time is
needed before our eye can adapt to new light conditions so we can see
properly again, this is called "Adaptation":
Dark Adaptation: When we move from a lit room to a dark room,
we cannot see clearly, because not enough stimulated rhodopsin (peripheral):
rhodopsin is bleached faster than it is reformed in strong light,
insufficient rhodopsin reformed instantaneouslycones are not
stimulated: light intensity too low. It takes about 20 minutes for enough rhodopsin to reform for us to see properly.
Light Adaptation: When
we move from a dark room to a brightly lit room, we feel uncomfortable
from the glare. But after some time, the visual threshold in Cones (foveal)
increases relative to the generator potential. Cones is less
stimulated, and we will see better. This takes about 5 minutes.
3 types of Cones: Blue, Green and Red Sensitive.
This is what allows us to distinguish between
different colours, the iodopsin molecules in these 3 types of Cones are
slightly different from each other.
Overview:
Rods (peripheral)
Cones (foveal)
1. High sensitivity (Night Vision)
2. More photopigment capture more light
3. High amplification, single photon detection
4. Saturate in daylight
5. Low temporal resolution
6. Slow response, long integration time
7. Sensitive to scattered light
1. Lower sensitivity (Day Vision)
2. Less photopigment
3. Less amplification 4. Saturate only in intense light 5. High temporal resolution
6. Fast response, short integration time
7. Sensitive to direct axial rays
Rod System
Cone System
1. Low acuity
2. Highly convergent retinal-pathways
3. Peripheral
4. Contrast and Movement
5. Pigment: rhodopsin
1. High acuity
2. Less convergent retinal-pathways
3. Foveal
4. Chromatic
5. Pigment: 3 iodopsins with different sensitiveties to different parts of the visible spectrum -> 3 types of Cones.
The segregation of motion and feature vision is a pervasive attribute
of brain organization at all levels of the neuraxis, from the Retina to the Frontal Lobe. "Motion-Orientation" and "Colour-Form"
discrimination is carried out by the separate Magnocellular and
Parvocellular pathways, this
segregated
information
then is transmitted from the "LGN" to M- and P-related sublayers and
modules in the "Primary Visual Cortex" (V1). The image (right) shows
where all the sections of the Overlapping Visual Fields are regulated to. (3.1 Magno and Parvo in the LGN )