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PHYSIOLOGY
OF
VISION
The retina contains the photosensitive cells of the eye, along with
a number of other neural processing cells, and is oriented to
other structures of the eye in the following way:
Sclera
Iris
EYE
Vitreous
Humor
Pupil
Lens
Optic Nerve
Optic Chiasm
Optic Tract
Lateral Geniculate Nucleus
Cornea
Aqueous
Humor
Retina
Optic
Nerve
Suspensory
Ligaments
Circular and
Meridional
Fibers
Right Hemisphere
The cornea and lens bend the incident light, with the cornea
doing most of the bending and the lens providing fine tuning and
focus:
Original light
Retina
Reflected
Light
Cornea
Object
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We can accommodate for near or far objects using the circular and
meridional fibers attached to the suspensory ligaments.
Contracting these fibers causes tension on the ligaments to be
reduced, allowing the lens to take on a more spherical shape. This
bends light more, enabling us to focus on nearer objects (reading).
Distant focus (gazing) is accomplished by relaxing the fibers,
which tightens the ligaments and elongated the lens - resulting in
less bending of light:
More bending of light
Contracted
Relaxed
Inserting a concave lens in front of the eye will diverge the incident
light before it is bent by the cornea and lens. This forces the cornea
and lens to bend the light more than without the concave lens, and
allows the near-sighted eye to focus on distant objects:
When the lens doesnt bend light correctly, we get blurred vision:
Myopia - near-sighted
The lens bends too strongly, focussing the image in front of the
retina because the lens cant be elongated enough. The image,
where it strikes the retina, is now blurry. Because the lens still
can bend light maximally, you can still focus on near objects.
Myopia - near-sighted
Hypermetropia - Far-sighted
Distant objects are still in focus, but the assisting lens is needed for
near objects (e.g. reading)
Also with age, we may experience protein damage in the lens, and
resulting calcium precipitation. This is called a cataract, which
will block some light entry and deflect other light from its
straight line path in the lens, This results in diminished and
blurred vision.
It is corrected by removing the lens entirely and replacing it with
an artificial lens. This results in a fixed focal length for the eye.
Convex portion
for gazing
Concave portion
for reading
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We have seen that the cornea and lens are necessary for bending
light and focusing on the retina. Now, why have an iris inserted in
the light path?
This means that we must allow more light into the eye in dark
situations, and when trying to see more distant objects.
The iris acts like a shutter, allowing more or less light to reach the
retina depending on the intensity of the light source.
The pupil is the central opening in the iris, and it dilates when the
iris constricts:
Too little light and the photoreceptors will not be activated, while too
much light will activate all photoreceptors fairly evenly (such as in
snow blindness).
NOTE: We see an object because of the light reflected off of it
(reflected), or because of light emitted by the object (Luminescence).
Dark objects reflect less light toward the retina and are more difficult
to see. Similarly, light from distant objects has more time to diverge,
so fewer photons will strike our retina (even from a brightly lit
object) from a distance.
RETINAL STRUCTURE
Once the incident light is focussed on the retina, how is the
energy of the photons converted (transduced) into a
frequency of action potentials?
The retina consists of combinations of 5 primary cell types
which work together to encode incoming light.
Pigment epithelium
Sclera
Horizontal Cell
Rod or
Cone
Inner
Limiting
Membrane
Bipolar Cell
Light path
from lens
The cell types are Rods and Cones for transduction; Horizontal,
Bipolar, and Amacrine cells for processing; and Ganglion cells
for transmission of the action potentials to the brain. The axons
of the ganglion cells become the optic nerves.
Ganglion Cell
Amacrine Cell
Rod cells are used for black and white vision, while cone cells
provide us with color vision. We will talk about how they can do
this in a few minutes.
There is a skewed distribution of Rods and Cones across the
retina. Cones, which are smaller than rods, are most concentrated
directly behind the pupil, in an area called the macula. The fovea,
located in the center of the macula, is the region of greatest visual
acuity and is predominantly comprised of cone cell transducers:
Retina
Light path
Optic
nerve
Fovea
Macula
So, cone cells are smaller, and therefore better able to discriminate
between two separate light sources (acuity). They are
concentrated in direct line with wherever we direct our vision,
thereby giving us our greatest acuity where we direct our vision.
However, cone cells need a greater light intensity to be activated
than do rods. This means that in low light conditions they may not
be activated. You have experienced this at night when you see
something out of the corner of your eye, but when you turn your
head or eyes to stare at it you see nothing.
The blind spot that we have in our vision results from the location
where the ganglion cell axons congregate and leave the eye to form
the optic nerve. This mass of axons prevents any rod or cone cells to
be located at the optic disk, and light striking that area cannot be
transduced.
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Now lets look at the different functions of the retinal structures and
cells which result in our visual impression of the world around us:
Incident light
400
Rod cell
400
500
600
(wavelength in nm)
700
Pigment
Epithelium
Outer Segment
Inner Segment
Nucleus
Electrotonic connections to
horizontal and bipolar cells
500
600
(wavelength in nm)
700
Synaptic Body
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F
A
S
T
+
S
L
O
W
1) So, the energy in the photon of light brings about the chemical
change of rhodopsin to metarhodopsin II. What happens next?
NOTE 1: Photopsins work like rhodopsin does, but they are about
300-times less likely to be activated by a photon (explaining why
they are less effective for seeing in the dark).
NOTE 2: The incident of 1 photon striking a rod cell triggers a very
rapid response that lasts for a prolonged period. A subsequent
excitation can trigger a response before the preceding one has
finished. In that case we dont see a blinking or flashing light, but a
constant image. The Critical Frequency For Fusion is how close
together in time two flashes of light need to be for us not to percieve
a flicker. This frequency is used in movies and television, such that
we see constant rather than discontinuous images.
NOTE 3: In prolonged dark, most retinene is present in the form of
rhodopsin, so the rod cell membrane is loaded with potential photon
targets and a single photon has an increased chance of being
perceived.
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Notice how a single rod (or cone) affects multiple ganglion cells
because of the presence of the horizontal cells. On the preceding
diagram we could see this effect to the left and right. Keep in mind
that the effect would exist as a circle around the activated rod. If
looking down from the top of the previous diagram, we would see a
field of ganglion cells which surround the affected rod cell:
- +- - -
+
+ -- +
+ +
Why have lateral inhibition at all? It accentuates edges, and our eyes
are very good at detecting edges. In fact, our primary visual
information is encoding of the edges of whatever we are looking at.
In the absence of lateral inhibition we would have:
AP frequency
These fields are overlapping each other all across the retina,
allowing us to accentuate edges of different objects:
AP frequency
+
+
+
+
+
+
+
+
It turns out that a single ganglion cell receives input from multiple
rods or cones. All of the rods or cones influencing a ganglion cell
are encoding similar information (e.g. edges, luminosity, red, green
or blue), so a single ganglion cell is transmitting information about
only one attribute.
Edge information
Red information
Luminosity
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BECOMES:
Cortical columns