VM8054 Veterinary Histology

Lab 11

Special Senses I: The Eye

Author: Dr. Thomas Caceci
Objectives For This Exercise
Laboratory Exercise on the Ear
Special Senses Introductory Page

The Eye
Let's first consider the overall
structure of eyes. The vertebrate
eye is constructed pretty much
the same way in all species of
domestic mammals, in birds, and
in primates, though there do
appear to be some specific
structural and functional
adaptations in each of these
groups. To a large extent,
however, similarity in structure is

to be expected. After all, the same

laws of optics apply to everybody,
and it's therefore likely that all of
the final designs would be pretty
much variations on the same
theme. In a mechanistic sense
designing a functional eye is
really an engineering problem,
and there are only a few ways in
which that problem can be
solved.
For a more comprehensive
treatment of the
physiology of the eye and the
nature of vision, click here.
Terminology and Orientation
A bit of terminology is in order at
this point. Since the eye is
globular, it's customary to refer to
its structures with respect to 1)

its optical axis, and 2) its

imaginary central point. To speak
of a structure as being "inner" or
"outer" means it's nearer to (or
farther away from) the physical
central point of the spherical
eyeball. The structural asymmetry
resulting from the placement of
the lens and the retina defines the
"front" and "back." A section of
the eyeball taken parallel to the
optical axis is a "horizontal"
section; one at right angles to the
optical axis is a "vertical" section.
General Structure
Most of this exercise will be done
on slide 268. This is a horizontal
section, and if you will examine it
off the microscope, you'll be able
to make out the general shape
and several major structures.

The Three Tunics of the Eye

Now turn to the microscope with
slide 267 or 268. The wall of the
eyeball is made up of three coats,
or tunics, one inside the other.
From outermost to innermost,
these are the corneoscleral tunic,
the uvea, and the retina.
Corneoscleral Tunic: Sclera
The first and largest part of
the corneoscleral tunic is the
sclera, the tough, collagenous,
outer part of the eyeball. This is
what we call the "whites of the
eyes," and this tightly woven bag
of collagen fibers forms a flexible,
watertight, and protective
covering. It's also the site of
attachment of muscles that rotate
the eye.

Corneoscleral Tunic: Cornea

The second part of this tunic, the
cornea, is the principal light
refracting structure. It forms the
anterior part of the corneoscleral
tunic, joining the sclera at the
limbus or corneoscleral junction.
To see the structure of the cornea,
click here.
The cornea is exposed to the
environment on its outer surface,
and is epithelial in nature. The
outermost part of the cornea is a
stratified squamous layer, the
corneal epithelium. This layer
isn't normally keratinized, of
course. The curvature of the
cornea is what refracts light
coming into the eye and provides
for most of the focusing;

alterations in the shape of the

cornea can have profound effects
on vision, and surgical procedures
to correct vision problems take
advantage of this fact. It is
possible to remove the cornea,
freeze it, and actually machine it
in a small lathe-like device to
alter its curvature, then stitch it
back into place, with different
focusing characteristics!
The cells of the corneal
epithelium sit on a thick basal
lamina, Bowman's membrane,
(named for Sir William Bowman,
1816-1892, the English
ophthalmologist, anatomist and
physiologist).
The corneal epithelium varies in
thickness among different
species, and if you have trouble

finding in on this slide, try slide

268, where it should be very
evident. The bulk of the tissue in
the cornea is the substantia
propria, a thick collagenous CT in
the form of regular lamellae. The
inner surface of the cornea has a
very thin, single layer of
epithelium, and between these
cells and the substantia propria
you will find Descemet's
membrane, their basal lamina
(after Jean Descemet, 1732-1810,
a French physician).
Uveal Tunic
The uvea is the middle tunic, and
it's composed of three elements.
These are the choroid, which
comprises most of it, the ciliary
body, and the iris.

Uveal Tunic: Choroid

The choroid is the highly
pigmented and vascular layer of
the uveal tunic that underlies the
sclera. It provides nutritive
support, and so there are
numerous blood vessels and large
lymphatic channels visible in it. In
most cases the melanin
pigmentation in very obvious; the
heavy pigmentation serves as a
"light trap" in much the same way
the black matte finish of the
inside of a camera does. It
minimizes internal reflections and
increases contrast by absorbing
stray light passing through the
retina, thus increasing the acuity
of vision.
To see the choroid, click here.

Uveal Tunic: Ciliary Body

The ciliary body is part of the
uveal tunic, too. It's located in the
angle between the end of the
cornea and the lens, and runs
circularly around the anterior part
of the eye.
Most of the ciliary body is
composed of smooth muscle
whose contractions work the lens
accommodation mechanism via
the tension imposed on the zonule
fibers.
To see the relationship between
ciliary body, zonule fibers, and the
lens, click here.
In addition to its role in the
mechanism of accommodation
(see below), the ciliary body is a
secretory structure. The inner

surface of the ciliary body is lined

on its inner surface with a double
layer of cells. These are
continuous with the retinal
pigmented epithelium and are
derived from the optic cup. This
ciliary epithelium is the site of
production of the fluid which fills
the anterior and posterior
chambers of the eye, the aqueous
humor.
To see the ciliary epithelium, click
here.
The epithelium on the finger-like
projections of the ciliary body into
the posterior chamber (the ciliary
processes) produce the aqueous
humor, secreting it into the
posterior chamber. It then passes
through the opening in the iris
(the pupil) and is drained away via

a series of channels in the angle

between the iris and the cornea.
The principal site of drainage is
the canal of Schlemm (Friedrich
S. Schlemm, 1795-1858, a German
anatomist). The canal is easily
seen in the iridio-corneal angle as
an irregularly shaped opening
lined with simple squamous
epithelium. It looks much like a
small lymphatic duct or vein, but
of course you would not expect to
see cells in it!
To take a ride on the Canal of
Schlemm, click here.
The continued production of the
aqueous humor and its removal is
important in function; the
intraocular pressure thus
produced helps to maintain the
normal shape and size of the

eyeball, which is vital to keeping

an image in focus. Blurry vision
and "seeing stars" are
premonitory symptoms of
increased intraocular pressure,
which can result from
overproduction of aqueous humor,
or damage to the drainage system,
and this is the condition of
glaucoma. Uncorrected, pressure
can build up and damage the eye,
causing eventual blindness.
Uveal Tunic: Iris
Projecting from the ciliary body is
the iris. This forms a pigmented,
diaphragm-like structure around
the eye. It has a hole of variable
aperture in it, the pupil, through
which light passes. The iris also
serves to divide the eye into

anterior and posterior chambers

in front of the lens.
The iris contains a fair amount of
muscle, and can contract in bright
light or expand in dim light to
provide the light sensitive retina
with the proper level of
illumination. The iris forms the
colored part of the eye.
Sometimes the formation of the
iris is incomplete, a
developmental defect called
coloboma iridis. It's frequently
associated with an incomplete
choroid coat (coloboma
choroidea) and it may be the
result of trauma or surgical
intervention.
The Retina

The retina is the innermost tunic

of the eye, in which the light
sensitive elements are located and
on which the image is formed. The
vertebrate retina has nine true
layers, and a tenth pigmented
layer is closely associated with it.
The vertebrate retina is
"reversed." That is, the image is
actually formed at the back (the
outer part) and the nervous
impulse it generates is brought
inward by the converging axons
that form the optic nerve. The
optic nerve physically passes back
through the entire thickness of
the retina, and thence on to the
brain. This situation results from
the way the eye forms in the
embryo, by invagination of the
optic vesicle into a double walled
optic cup.

Layers of the Retina: Pigment

Layer
We'll look at the retina's layers
from the outermost to the
innermost, in order of their
occurrence.
To see these layers of the retina,
click here.
The cells of the pigment layer are
impregnated with melanin, and
they also contain lipofuscin. The
light sensitive parts of the retina
are continually "turned over" to
maintain optimum function. The
pigment layer cells phagocytose
the ends of the rods and cones as
they are renewed: hence the
accumulation of lipofuscin. The
pigmentation this activity
produces helps to increase the
contrast of the visual image by

absorbing light that would

otherwise be reflected back
inwards towards the rods and
cones. These cells comprise a
layer of cuboidal cells that have
round nuclei and an aggregation
of melanin and lipofuscin at their
inner ends.
The pigment layer is not
considered by all authorities to be
part of the retina proper, because
its embryological origin is
different from the retina's. It
comes from the outer wall of the
optic cup, not the inner one. But
the association is so close and so
vital to normal retinal function
that a strong case can be made for
including it as a part of the retina.
Layers of the Retina: Rods and
Cones

The outermost layer of the retina

proper is the bacillary layer or the
layer of rods and cones. These are
the actual light sensitive
elements.
Rods and cones are cellular
structures derived from cilia, and
they represent another example of
the modification of cilia for
sensory purposes. Each is a fairly
large structure, enclosing a stack
of light sensitive membranes.
The names "rod" and "cone"
reflect the general shape each
type of light receptor takes, and
in good preparations for the LM,
the shapes are generally visible.
Remember that each of these is
actually only part of a cell; there
is a narrow "waist" between the
rod or cone and the cell body of

which it's a part. Rods and cones

are transducers and their function
is to take the physical signal of
electromagnetic radiation and
transform it into a neural one.
Rods are associated with low light
vision, and are most numerous in
nocturnal animals. Cones are
required for color vision (there
are three types, one sensitive to
each of the primary colors) and
are most numerous in diurnal
species. You would expect this,
since the spectral range of
daylight is much broader than the
light of the moon or the stars is.
Although Benjamin Franklin had
something else in mind when he
made the remark, "In the dark, all
cats are gray," nevertheless the
statement is true in and of itself.
It's extremely difficult, even for

animals with good color vision, to

make out colors in low light
environments. Humans have
excellent color vision, and you can
prove this to yourself by going out
on a moonlit night and trying to
distinguish between different
colors.
Color vision is best in the birds
and primates. You will frequently
hear the statement made that
domestic animals are "color
blind," but this has yet to be
definitively proven. As a matter of
fact, there is no way we can tell
exactly what some other animal
sees, and such data can only be
derived from indirect
experimentation. There seems to
be no doubt that birds can see
colors quite well (though whether
they see them exactly as we do is

open to debate) and from

behavioral studies there's some
evidence that other mammals
besides primates have at least
rudimentary color vision. There is
fairly good evidence that deer can
see blaze orange, and seeing eye
dogs can certainly tell red traffic
signals from green ones.
To finish off this train of thought,
I'll note that cones are present in
animals that have long been held
to be "color blind" like dogs and
cats, though they are less
numerous than in primates. If
cones are part of the color vision
process, as they seem certainly to
be, it's a bit premature to assume
that any animal that has them
"can't see color."

Layers of the Retina: Outer

Limiting Membrane
Moving inward, you'll find the
next layer, the outer limiting
membrane. This isn't really a
membrane. It's a place where the
rod and cone cells are intersected
by the "feet" of the quasi-glial
element, the Mller cells. This has
the appearance of a membrane in
the light microscope because
everything is closely packed
together. The Mller cells form
junctions with the photoreceptors
and with other Mller cells. This
"membrane" may be a way to
isolate the subretinal space,
creating a space for the outer
segments of the rods and cones
and limiting diffusion of
nutrients, growth factors, and
waste products between the

choriocapllaris and the outer

segments.
Layers of the Retina: Outer
Nuclear Layer
Immediately inwards of the outer
limiting membrane, you'll find a
group of nuclei. This layer is the
location for the nuclei and cell
bodies of the rod and cone cells
(whose sensitive elements we saw
projecting outwards as the
bacillary layer). Collectively the
nuclei of these cells constitute the
outer nuclear layer.
Layers of the Retina: Outer
Plexiform, Inner Nuclear, and
Inner Plexiform Layers
Inwards of the outer nuclear layer
you'll find a relatively clear zone.

This is the site of numerous

synapses between the rod and
cone cells and the processes of
various integrator neurons. This is
the outer plexiform layer. While
the initial neural impulse is
generated in the rods and cones,
they must pass the signal along to
other neural elements for
processing and further handling.
The dendrites of these cells and
the axons of the rod and cone
cells constitute the outer
plexiform layer.
The integrating elements are the
horizontal, bipolar, and amacrine
cells. They pass along the signal
to the next neural element in the
chain. The cell bodies and nuclei
of these integrator neurons are
located in the inner nuclear layer,
next inwards from the outer

plexiform layer. The bipolar cells

are particularly important, as they
have one pole thrust outwards
into the outer plexiform layer, and
the othertheir axonthrust
inwards to the next layer, the
inner plexiform layer.
The inner plexiform layer, like the
outer one, is a region of synapses.
Here the bipolar cell processes
synapse with the processes of
ganglion cells. Ganglion cells are
also neurons, and they are
specifically those neurons whose
axons will collect to form the optic
nerve. Thus they form the final
intra-retinal element in the
neuron chain, and their axons run
back into the visual centers of the
brain proper.

Layers of the Retina: Ganglion

Cell Layer
While the ganglion cells have
synapses with bipolar cells in the
inner plexiform layer, their cell
bodies aren't located there. The
cell bodies and nuclei of the
ganglion cells are located in the
next retinal layer, the ganglion
cell layer, which has far fewer
nuclei than the inner or outer
nuclear layers. Like any other
neurons, these ganglion cells have
axons, and the axons form the
next layer inwards.
Layers of the Retina: Nerve Fiber
Layer
The axons of ganglion cells are
bundled together to form the
nerve fiber layer. These axons are

afferent fibers from the ganglion

cells, and after running radially
around the inner surface of the
retina, they all come together at
the site where the optic nerve
penetrates the retina (see below).
This is the "blind spot" of the
vertebrate eye.
Layers of the Retina: Inner
Limiting Membrane
The innermost layer is the inner
limiting membrane, which again
isn't really a membrane, but is
instead a place where the Mller
cell feet are in close
approximation to each other.
Remember that most of the cells
of the retina are in fact neural
elements; they are either
transducers (the rods and cones),

integrators (bipolar and

horizontal cells) or the terminal
sensory neuron of the chain (the
ganglion cells). As is the case with
other neurons, all of these cells
are pretty much helpless to
defend themselves against the
insults of the outside world. They
have to be cosseted and coddled
by neuroglia, or at least their
equivalent. The cells involved in
the formation of the inner
limiting membrane are Mller
cells, tall cells which (like
astrocytes in the CNS) serve to
support and protect the neurons.
The Mller cells run from the
outer limiting membrane to the
inner limiting membrane, and are
sealed at both ends. At their outer
ends they are fused to the "waists"

of the rod and cone cells, and at

the inner end to each other.
The Optic Nerve and the Ora
Serrata
You may be able to see a section
of the optic nerve on your slide
267, if there is one in your box.
The optic nerve begins at the
optic disc, the point where all the
axons of the ganglion cells come
together, and projects back into
the main portion of the brain.
This is the "blind spot," of course,
because no light sensitive
elements can located at the point
of perforation.
To see the optic nerve, and the
"blind spot" click here.
There is another non light
sensitive portion of the retina,

that part between the ciliary body

and the posterior part of the iris.
The margin between the sensitive
and insensitive parts of the retina
is the ora serrata, normally visible
only in gross anatomy
preparations.
The Fovea
Vertebrate eyes have a region of
most acute vision, the fovea. The
retina is thinner here, and
receptors more crowded together.
The fovea may not be present on
your slide, but some slides in the
set do show one. It's an area
where the thickness of the retina
is reduced but the elements in it
are noticeably more orderly and
tightly packed.

At first thought you might guess

that the fovea would be centrally
located, and that gazing straight
at an object would put its image
in the area of most acute vision,
but that isn't the case. The fovea
is slightly off the optical center of
the eye, and rays that come in at
an angle to the central axis are
focused directly on it.
Why should this be so? The reason
is simple. This arrangement gives
an animal much better peripheral
vision, which is of real survival
value. Think in terms of an animal
who lives in fear of attack from
predators and relies on vision for
warning of an approach. That
animal is using its eyes for other
things as well, and any predator
who knows his job will normally
sidle up alongside his prey, hoping

to avoid detection. By arranging

the visual organ so that the most
acute area is off the main axis,
any object moving through the
edge of an animal's visual field
will more readily be detected.
An old trick, long taught in the
military, is to tell sentries to keep
their eyes moving and not to stare
out at one spot. If they perceive
movement and are interested in
seeing some small or distant
object, they are taught not to look
straight at it, but rather to look
past it.
The Lens
The lens is the largest and most
obvious part of the eye. It's a
biconvex, transparent structure
composed of numerous layers of

cells. It's located between the

large vitreous chamber of the eye
and the smaller posterior
chamber. The anterior chamber is
demarcated by the cornea on the
outside surface, and by the iris
between it and the lens.
To see the relationship of the lens,
iris, and anterior chamber, click
here.
The lens, because of its dense
construction, is usually poorly
infiltrated by paraffin, and
histological sections usually show
some damage and considerable
shrinkage; however, you should be
able to make out the major
features of its histology.
The lens is covered by a thin lens
capsule, an amorphous layer that
is really the basement membrane

of the underlying lens epithelium.

The capsule is thickest on that
side of the lens facing the iris,
and thinnest on the posterior
surface. The lens epithelium that
produced it is a single layer of
cuboidal cells, which are the
source not only of the capsule, but
of the lens cells that constitute
the bulk of the lens' material. At
the equator of the lens these
epithelial cells proliferate,
elongate, and transform
themselves into lens cells.
The lens grows in size throughout
life, and a constant production of
new lens cells is necessary.
Mitoses of the lens epithelium
and differentiation provide for
this. At the equator of the lens the
proliferating cells begin to
elongate and send out long

processes that "dig" under the

adjacent lens epithelium. The
nuclei of these cells remain in the
region of the equator, and you will
see them scattered there. The
extrusion of processes causes new
layers to be added to the lens
continually. Eventually the nuclei
of the transformed lens cells will
fragment and disappear, but their
cytoplasmic boundaries can
usually be made out fairly easily.
The lens is responsible for
accommodation, i.e. the
adjustment of the eye for
observing near or far objects. It
does this by changing its shape.
This changes the focal length of
the lens, and permits refraction of
the rays coming from near or far
objects.

The lens is suspended from the

muscular ciliary body by a number
of delicate zonule fibers, which
you should be able to make out as
wispy strands between the edge of
the lens and the pigmented ciliary
body. Contraction of the muscles
of the ciliary body tugs at the
fibers and changes the shape of
the lens. In the relaxed condition
(that is, with no tension on the
fibers beyond that needed to
support the lens) the shape is
such as to refract rays from
distant objects, and the eye is
focused at infinity.
Accommodation for close vision
requires tension to be exerted to
deform the eye.
This system makes a lot of sense
for an organ with the role of a
long range warning system: most

predators (and prey) spend their

time scanning the far horizon,
looking for something to eat (or
for something that might want to
eat them). By having long
distance vision as the "default"
condition, the amount of exertion
needed is minimized. Incidentally,
this system is also related to the
deterioration of eyesight with
advancing age. Older lenses are
stiffer and less deformable, and
more effort is needed. So close
vision becomes more difficult with
each year. Humans get around
this by equipping themselves with
artificial lenses; animals don't
worry about it. Most wild animals
don't live long enough in any
event for accommodation
problems to become serious; but
the situation does arise in

domestic animals whose protected

environments make it possible for
them to live lives much longer
than "normal" for their species.
Another age related degenerative
change in the lens is the
development of cataracts,
opacities that impair the passage
of light and cause loss of vision.
There are many kinds of
cataracts, and many causes for
them, but they are a fairly
common consequence of living
beyond the "normal" life span.
They're quite common in older
domestic animal, especially dogs,
because pets tend to live much
longer lives than their feral
counterparts.
Accessory Structures

Accessory structures of the eye

include the eyelids and the
lachrymal glands.
Eyelids
The eyelid (and the that part of
the eye socket which faces it) is
lined with an interesting and
unusual conjunctival epithelium.
The part lining the inside of the
lid is the tarsal conjunctiva, which
continues onto the outer surface
of the eyeball as the bulbar
conjunctiva. The conjunctival
epithelium is unusual in that it's
classified as stratified columnar,
and it also contains goblet cells.
The tarsal conjunctiva forms a
mucocutaneous junction with the
integument at the edge of the
eyelid, and the bulbar conjunctiva
transitions to the stratified

squamous epithelium of the

cornea.
Slide 266 is a vertical section
through an eyelid, showing a
typical stratified squamous
epithelium and adnexal structures
of skin on the outside surface, and
a transition region of smooth
conjunctiva on the bulbar side.
The center of the eyelid is filled
with bundles of skeletal muscle,
blood vessels, nerves, glands, etc.
most of which are visible on slide
266.
To see the conjunctiva, click here.
To see the eyelid, click here.
The large tarsal glands (also
called Meibomian glands, after
Hendrick Meibom, 1638-1700, a
German anatomist) are quite

obvious. These are modified

sebaceous glands, opening into a
long central duct which isn't
visible on this slide. (If your eyes
have ever been gummed shut after
a hard night at the local
Brauhaus, that crud that held
them closed came from these
glands.)
There are also modified sweat
glands present, the glands of Moll
(Jacob A. Moll, 1832-1914, a
Dutch oculist) whose secretions
are released into the follicles of
the eyelashes.
Lachrymal Glands
It's necessary to keep the surface
of the eye moist, to prevent
abrasion and damage to the
cornea. The eye produces a fluid

secretion, the tears, in glands

located nearby. These secretions
are conducted via ducts to the
surface of the eyeball. None of the
slides in the set have lachrymal
glands, but a demonstration slide
(which also shows all of the other
features of the eye) is available on
which you can see these.
Lachrymal glands are very similar
to serous type salivary glands in
structure and operation. The cells
closely resemble those in salivary
glands, too. Tears, in addition to
providing lubrication, have an
antibacterial effect because of the
inclusion of lysozyme in their
composition.
To see an example of the
lachrymal gland, click here.

Lab Exercise List