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Eye & Physiology of Vision

LIGHT

Light is the visible part of the electromagnetic spectrum. We "see" because we have
receptors, which are excited by wavelengths between 400-700nm.

THE EYEBALL

OPTICS

The light passes first through the cornea. The cornea and the lens represent the refractive
system of the eye. The cornea is responsible for 70% and the lens for 30% of the refraction.
The refractive system focuses the light upon the retina, where the photosensitive pigment
lies. The image is inverted.

ACCOMODATION

The lens is responsible for accommodation, that is, the adjustment of the focussing system
for near objects. The shape of the lens is determined by the tone of the ciliary muscle. When
it relaxes, as in far vision, the zonular fibres are pulled taut and the lens is under tension and
flat. For near vision, the ciliary muscle contracts, releasing the zonular fibres from tension
and the lens takes up its natural, rounder and more refractive state.

FOCUSSING

Focussing involves:

1. moving the lens towards the back of the eye

2. turning the eyes inward towards the nose (convergence)
3. pupil constriction
4. fattening of the lens
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CORNEA

The cornea has an endothelium (facing the aqueous humour) and an epithelium (facing the
tear film) sandwiching the stroma (fibroblast-like cells and collagen fibrils). The collagen
fibrils exert a high colloid osmotic swelling pressure. The cornea will naturally swell and as
it does so become opaque. This swelling must be counteracted by continual pumping of
water out of the tissue to maintain the correct thickness and transparency.

LENS

The lens is transparent. It will naturally take up water and swell. This must be balanced by
a continuous pumping of (ions and) water out of the lens. Anything that interferes with this
process will cause the lens to swell and opacify - a condition known as cataract. There are
many causes of cataract, ageing being the predominant of these - the so-called "sunlight"
hypothesis for cataract is largely discredited.

ERRORS OF REFRACTION

1. Presbyopia. With age lens thickens and becomes harder (won't accomodate)
2. Myopia. The near sighted eye. Eyeball too long.
3. Hypermetropia. Far-sighted eye. Eyeball too short.
4. Astigmatism. Lens or cornea not smoothly spherical.

AQUEOUS HUMOUR

Secreted by the ciliary epithelium into the posterior chamber of the eye, it diffuses forward
through the pupil and into the anterior chamber. From there it drains through the trabecular
meshwork and into Schlemm's canal. Glaucoma is the raised intraocular pressure that arises
from an imbalance of secretion and drainage, usually due to blocked outflow.

LIGHT REFLEX

In bright light, the parasympathetic nervous system causes the circular sphincter muscle of
the iris to contract and pupil constriction occurs. In dim light, the sympathetic nervous system
causes the radial muscle to contract, dilating the pupil.
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RETINA

These are the main cells in the retina: ·

1. Photoreceptors (rods & cones)

2. Horizontal cells (lateral inhibition at the level of the photoreceptors)
3. Bipolar cells ("on" and "off" cells) connect photoreceptors to retinal ganglion cells
4. Amacrine cells (lateral inhibition at the level of the retinal ganglion cells)

Retinal ganglion cells (the axons of which form the optic nerve)

RECEPTOR CELLS

The retina is the back part of the eye that contains the cells that respond to light. These
specialized cells are called photoreceptors. There are 2 types of photoreceptors in the retina:
rods and cones.
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• The rods are most sensitive to light and dark changes, shape and movement and
only contain one type of light-sensitive pigment. Rods are not good for color vision.
However, in a dim room, we use mainly our rods, but we are "color blind". Rods are
more numerous than cones in the periphery of the retina. There are about 120 million
rods in the human retina.
• The cones are not as sensitive to light as the rods. However, cones are most sensitive
to one of three different colors (green, red or blue). Signals from the cones are sent
to the brain, which then translates these messages into the perception of color.
However, cones only work in bright light. That's why one cannot see color very well
in dark places. So, the cones are used for color vision and are better suited for
detecting fine details. There are about 6 million cones in the human retina. Some
people cannot tell some colors from others - these people are "color blind". Someone
who is color blind does not have a particular type of cone in the retina or one type of
cone may be weak. In the general population, about 8% of all males are color blind
and about 0.5% of all females are color blind.
• Rods are responsible for "scotopic" or low intensity vision. Cones are responsible for
"photopic" or high intensity vision. Both rods and cones contain a photopigment which
absorbs the light. There are 4 photopigments, one in rods and one in each of 3 cones.
The photopigment is called rhodopsin and consists of two parts - a filter called an
opsin and a light sensitive chromophore called retinal. Retinal is common to all four
photopigments. [See graph of distribution of cells across retina]

• In the peripheral retina there is a 1:1000 convergence of photoreceptors (mainly rods)

onto retinal ganglion cells. In the fovea (mainly cones) there is a 1:1 correspondence.

PHOTORECEPTION

In the dark the concentration of cGMP (cyclic guanosine monophosphate) in rods and cones
is high. This opens cyclic nucleotide gated (non-selective cation) channels (CNG channels).
Sodium (Na+) and calcium (Ca2+) enter the cell and this depolarizes, resulting in an increase
in transmitter release.

In the light, a G-protein, transducin, is activated starting a cascade of biochemical events

and resulting in a fall in cGMP. The receptor operated channels close and the cells
hyperpolarise (receptor potential), resulting in a fall in transmitter release.

BIPOLAR CELLS

The release of transmitter either inhibits "depolarising" or (+) bipolar or stimulates

"hyperpolarizing" or (-) bipolar cells which then relay the receptor potential to the retinal
ganglion cells which increase/decrease their firing rate as appropriate. [Therefore, (+)
bipolars lose their transmitter inhibition and depolarize in light, whereas (-) bipolars lose their
stimulus and hyperpolarize in light. Nb. light causes photoreceptors to hyperpolarize and
transmitter release, at synapses with bipolar cells, falls].
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VISUAL CODING

The retina analyses the visual image in terms of (a) colour, (b) form, (c) movement, (d)
luminance and (e) depth.

a. Colour; green, blue and red cones. Cones contain a filter (opsin) and a light-sensitive bit
(chromophore, called retinal). Green cones have green filters, red cones have red filters etc.

• red cones maximally sensitive to 559nm light

• green cones to 531nm
• blue cones to 419nm

Colour opponency. Red/green and blue/yellow "centre-surround" receptive fields of retinal

ganglion cells (see below) mean that there are retinal ganglion cells sensitive to red/green
contrasts and others sensitive to blue/yellow differences. (Yellow is made up of output from
red/green cones combined). This aids in colour contrast definition. Colour-blind people
usually lack either a red or green opsin and have trouble distinguishing red from green (both
appear the same).

b. Form; edge detection, centre-surround receptive fields

Each retinal ganglion cell has a receptive field that corresponds to input from part of the
visual field (a small region of the retina). This receptive field is made up of information from
one to 1000s of photoreceptors arranged in a "centre-surround" fashion (see diagram).

RECEPTIVE FIELDS

"ON" and "OFF" cells

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c. Movement; X (slow, high resolution) and Y (fast, low resolution) cells. These are different
classes of retinal ganglion cells.

• X-cells; these are mainly connected to cones, slow conducting therefore slow
response. Used for high resolution analysis. Found in fovea mainly.
• Y-cells; these are fast responding cells, mainly connected to rods, concerned with
movement and found mainly in the peripheral retina.

d. Luminance; W-cells are another class of retinal ganglion cells that may detect the absolute
levels of luminance and be involved with the light reflex, i.e. pupil constriction/dilation (and
the Edinger-Westphal nucleus).

e. Depth; binocular vision. The visual field of each eye is slightly different. Fusion of the two
images (in the cortex) confers the perception of depth. Cf. the stereoscope - an instrument
presenting one, slightly different, image to each eye, give remarkable appearance of 3-D.

By analysing the visual image in terms of colour, form (edges), luminance, depth and
movement, the retina has achieved a degree of pre-processing of the image before it leaves
for the visual cortex via the optic nerve.
Dark
Biosynthesis Rod Opsin Rhodopsin
Light
Vit A1 NAD+ Retinine Cone Opsin Iodopsin
NADH
Rod Opsin Porphyropsin
Dark
Vit A2 Retinine Cone Opsin Cyanopsin