Eye Anatomy

Introductory article
Article Contents

Thomas C Litzinger, Miami University, Oxford, Ohio, USA

Katia Del Rio-Tsonis, Miami University, Oxford, Ohio, USA

. Introduction

The eye is a small yet multifaceted unit of anatomical machinery in which each structure
works in accord with the next, refracting, constricting, dilating and chemically reacting to
convert patterns of light into discernible images. Eyes can be divided into two broad
categories: the simple eye of vertebrates and the compound eye of invertebrates.

. An Overview of the Basic Structures and Functions of

the Simple Eye: Human as Primary Model
. The Retina as a Part of the Central Nervous System
. Metabolic Support for Photoreceptor Cells from the
Retinal Pigment Epithelium
. Focusing of Light onto the Fovea in Primates
. The Compound Eye

Introduction

. Evolutionary Trends of Eye Structures

. Summary

The eye has been described by Charles Darwin as both

perfect and complex. There are several structural and
functional variations that exist between organisms, yet it
would be incorrect to say that one is superior to the next.
This is the perfection that the eye beholds; each eye has
evolved to suit precisely the necessities of its possessor.
Though numerous and intricate, the many eyes of the
world can be placed into two very general categories:
simple and compound.
Though dierent in appearance, these two models of the
eye are actually quite similar in their most elementary
functional components. One particularly well-conserved
molecule between organisms is the light-absorbing protein
opsin, which essentially initiates the sequence of events
leading to image formation. Though the most basic visual
molecules such as opsin have not been selectively altered in
a drastic way by environmental pressures, the anatomical
morphology of the eye has. This divergent evolution has
led to the formation of such dissimilar eyes as that of the
human (simple eye), the y (compound eye), and many in
between.

An Overview of the Basic Structures and

Functions of the Simple Eye: Human as
Primary Model
Along lights journey through the eye it is slowed down,
bent, absorbed, and converted by various structures
(Figure 1a). As light approaches the eye it rst comes in
contact with the cornea. The cornea refracts the light,
causing the image to converge on its way to the iris and
pupil. Depending on the intensity and availability of the
light, the iris will contract or expand adjusting the pupil
size. In situations of low light, the pupil will be larger,
allowing for the passage of enough light to form a
discernible image. The opposite is true in situations of
abundant light, for an excess of light results in poor
imaging as well. Once through the gate of the pupil, the
light is received by the lens. With the aid of auxiliary

muscles, the lens possesses the ability to change shape.

Depending on its form, objects at various distances can be
brought into focus. The lens also slightly improves the
already rened image from the cornea, and projects it onto
the retina. The retina, which literally means net, catches
the light via its photoreceptor and pigment epithelial cells.
The photoreceptor cells photopigment molecules absorb
the light, causing a change in the photoreceptors
membrane potential. This initiates a series of signals that
travel through the neurons of the retina, and into the optic
nerve leading to the brain. This signal is then received and
processed by the brain as an interpretable image.

A closer look at the structures involved in the

entry of light into the eye
The cornea
As mentioned, along the path of light into the eye, it will
rst encounter the cornea, which is a transparent body
consisting of an epithelium, a thick brous structure made
up of connective tissue and extracellular matrix, a
homogeneous elastic lamina and a single layer of
endothelial cells. The cornea is the primary contributor
in the focusing of light on the retina. Following the basic
laws of refraction, as incident light encounters a medium
possessing a greater refractive index than that of air,
propagation slows down, thus bending the beams path.
The cornea would be an example of such a medium,
possessing refracting capabilities. When light hits the
surface of the cornea, it slows down and converges towards
the centre of the eye, thus reducing the image that has been
reected to the eye. Though the cornea bends light, its
transmission is very characteristic of the transparent media
that it is, the main characteristic of transparency being the
minimal scattering of light, and the continuing transmission of light in its original direction, both of which
contribute to discernible image formation. These intrinsic
properties of the cornea are made possible by the spatial
uniformity of its cells, which contribute to the acuity of

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Eye Anatomy

Figure 1 (a) Three-dimensional representation of the structures of the human eye. (b) Cross-section of the human eye, and an enlarged view of the
various layers of the retina.

light transmission. With these elements present, the cornea

makes up the rst of many critical components of the
functioning eye.
The pupil and iris
The light must cross through the aqueous humour, the
body of uid that lls the anterior chamber between the
cornea and the lens, so that it can reach the next group of
2

structures: the iris and pupil. The two structures work as

the regulators of the amount of light passing through the
system. The iris is a pigmented sheet of tissue that lies
directly in front of the lens, and has the ability to restrict
and dilate with the aid of sphincter and dilator muscles,
respectively. This contraction and dilation regulates the
aperture of the eye, the pupil. In cases of abundant light,
the iris lessens the pupillary aperture with the aid of the
sphincter muscles, trying to avoid the admittance of too

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Eye Anatomy

much light which would eventually result in the processing

of a muddled blur. The opposite is true when light is
lacking. The pupil becomes greatly dilated in an attempt to
gather as many photons as possible for imaging.
The lens
Once the correct amount of light has entered the eye
through the pupil, it encounters the lens. The lens,
composed of a lens epithelium layer covering a mass of
lens bres, is primarily made up of proteins called
crystallins, which further rene the image from the cornea.
Like the cornea, the molecules of the lens are densely
packed and uniformly spaced. This is necessary for its
transparency. The lens has an inherently greater index of
refraction than that of the cornea, based on its environment needs. Since the lens is surrounded by the uid of the
aqueous humour and the vitreous humour, which have a
relatively high index of refraction, the index of the lens
must be higher still if it is to focus the image further and
contribute to the optical system.
Though the lens has an inherent refractive index, it
actually has the ability to change its degree of refraction
with the aid of ciliary muscles. When discussing the process
of accommodation, the active altering of the shape of the
lens to bring close objects into focus, it would be
appropriate to start with the ciliary zonule. The ciliary
zonule consists of a series of thin, peripheral ligaments that
suspend and hold the lens in place (also known as
suspensory ligaments). These ligaments, or bres, are
attached to the area of the ciliary muscle called the ciliary
body. The ciliary body and the zonule bres work in
conjunction to alter the focal point of the eye. When the eye
is in its most relaxed state, it is focusing at distances beyond
6 metres (20 feet). In this state, the ciliary muscle is relaxed,
and the zonular bres are taut, thus pulling outward on the
lens forcing it to assume a rather attened shape. When the
eye focuses on an object within 6 metres, the ciliary muscle
must contract or close, as the tension in the zonular bres is
reduced. This results in a thickening and bulging of the
lens, in turn increasing optical power, bringing the focal
point closer and creating a clear image of an object within
6 metres of the viewer.

The Retina as a Part of the Central

Nervous System
The viewer would never perceive this image if it were not
for the retina. The retina is the light-processing centre of
the eye, where light signals are transformed into neural
signals that can be perceived and processed by the brain.
The neural cells involved in this process are remarkably
similar to those of the brain, which supports the common
assertion that the visual system is an outgrowth of the
central nervous system. The retina is in fact the only part of

the human central nervous system that is exposed to stimuli

from the outside environment.

Organization of the retina into the different

cell and synaptic layers
The retina can be divided into many distinguishable layers
(Figure 1b). The rst layer to interact with light coming from
the lens is the retinal pigment epithelium (RPE) layer. The
RPE cells do not contribute directly to the transformation
and transduction of information in the retina, but do
provide supportive functions to the photoreceptor cells,
which lie just above this layer. The next set of cells, making
up the photoreceptor layer, are the rst of three neural cell
types (photoreceptor, bipolar cells and ganglion cells) that
contribute to the vertical transferring of signals in the
retina. This photoreceptor layer consists of the outer and
inner segments of the rods and cones, which receive and
transform photons of light. The nuclei of these photoreceptor cells reside in the outer nuclear layer and their
axons and cell terminals in the outer plexiform layer and
the outer synaptic layer, respectively. The outer synaptic
layer represents the site where the photoreceptors rst
interact with the bipolar cells and other retinal neurons and
marks the transition between the outer and inner layers of
the retina. Like the outer layers, the inner layers can be
divided into nuclear and plexiform layers. The inner
nuclear layer contains the nuclei of bipolar cells, horizontal
cells, and the majority of the amacrine cells. The inner
nuclear layer is followed by the inner plexiform layer,
where vertical communication between the bipolar cells
and the ganglion cells takes place, thus making up the
second synaptic contact layer. The next layer, the ganglion
cell layer, contains the cell bodies of the ganglion cells. The
dendrites of the ganglion cells actually extend into the inner
plexiform layer, whereas their axons extend in the opposite
direction towards the nerve bre layer. It is through this
layer that all of the ganglion cells axons travel in the
direction of the optic nerve.

Five different types of neurons

Now that the groundwork of the retina has been laid out,
the cells already mentioned can be discussed further,
starting with the ve dierent kinds of neurons in the
retina. The rst three neurons are involved in the vertical
transmission of information through the retina, beginning
with the photoreceptor cells. These cells are responsible for
initiating the cascade of events that takes an image
projected onto a layer of tissue (retina), and converting it
from photons to an electrochemical signal capable of being
read by the brain. This conversion of light energy to
informative chemical energy is called phototransduction.
The two types of cells involved in this process are the
photoreceptor cells: rods and cones.

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Eye Anatomy

Rods and cones

Of the 130 million photoreceptors, about 120 million are
long cylindrical structures known as rods. Rods are
extremely sensitive to light, and only send shades of grey
to the brain. Cones are a thicker, usually shorter version of
rods that register ne detail and colour, provided they
receive enough light. The most critical element in this
process of phototransduction is the photopigments contained within the rods and cones. Both cells contain the
light sensitive protein opsin, as mentioned previously. In
rods this protein binds to a straight chain of vitamin A,
assuming a bent position. When in this conformation, the
complex is called rhodopsin. When as much as a single
photon of light strikes this compound, the energy absorbed
causes the bent vitamin A chain to snap back into its
original straightened form. This occurrence consequently
disrupts the electrical eld within the photoreceptor,
initiating an electrical impulse that begins its journey to
the brain. The cones possess three dierent forms of opsins
capable of binding to vitamin A. Each compound
eventually is responsible for the creation of one of the
three primary colours (red, blue or yellow), as interpreted
by the brain. However, as mentioned earlier, cones are
much less sensitive to low intensities of light, and therefore
require a very specic wavelength of light to initiate the
electrical impulse. This is why our daylight environment is
full of brilliant colours, whereas our rod-dominated night
vision produces various shades of grey. Essentially, the
world is colourless. Colours are merely biochemical
interpretations of wavelengths of light, whose identity is
dependent on the biochemical make-up of the particular
organism in question.
Bipolar cells
The next set of neurons that propagate the vertical, or
direct, communication pathway are the bipolar cells. As
stated earlier, their cell bodies reside in the inner nuclear
layer while their dendrites receive signals from the
photoreceptors at the rst synaptic junction. On the
opposite end of the cell body the signal travels through
the bipolar cells axon to synapse with the next vertical
neuron, the ganglion cell.
Lateral neurons: horizontal and amacrine cells
The electrical impulses running through the vertical
neurons are not completely independent of one another,
because most are linked by lateral neurons. One type of
lateral neuron is the horizontal cell. Horizontal cells are
found in the inner nuclear layer of the retina. These cells are
commonly linked to more than one photoreceptor, meaning that the subsequent bipolar cells receive signals from
more than one photoreceptor. This pathway would
intuitively seem to lessen visual acuity, but in most cases
serves a useful purpose, as it increases the perception of
contrast. The nal type of neuron in the retina is another
4

lateral body, the amacrine cell. These cells form links

between vertical pathway neurons in the inner layers, and
sometimes the ganglion layer of the retina. Their eects are
not entirely clear, but they are thought to contribute to the
eect of contrast.
Retinal ganglion and output from the retina
The last neurons of the network to receive the signals are
the retinal ganglion cells. When activated by an incoming
signal, the ganglion cells produce an action potential that
begins its journey down the cells axons. The axons of the
ganglion cells of the retina converge, forming the optic
nerve. The optic nerve represents a highway for electrical
signals en route to the brain.

Metabolic Support for Photoreceptor

Cells from the Retinal Pigment
Epithelium
The RPE is located underneath the neural retina and it is
characterized by having tight junctions, forming the
bloodretinal barrier. The RPE regulates and transports
ions, water, growth factors and nutrients such as glucose
and amino acids to the outer portions of photoreceptors.
The RPE is also involved in the maintenance of retinal cell
adhesion by supporting the interphotoreceptor matrix
(IPM). This extracellular matrix is bound to the outer
limiting membrane and the apical membrane of the RPE
(membrane facing photoreceptors). The IPM is critical for
the metabolic exchanges between the photoreceptors and
the RPE. Its bonding properties and viscosity are regulated
by the RPE, which tightly controls the ionic environment
in that region. The RPE cells are essential for the
regeneration of photopigments, since they uptake, store
and reisomerize vitamin A, which is necessary for the
future synthesis of rhodopsin used by photoreceptors. The
RPE also phagocytoses the tips of the outer segments of
photoreceptors on a regular basis, then it digests the
absorbed material to nally recycle it. Melanin, the visual
pigment present in the RPE, reduces scatter to the
photoreceptors and shields them from excessive light
exposure.

Focusing of Light onto the Fovea in

Primates
As mentioned earlier, the cones of the eye are responsible
for discerning minute details. The highest concentration of
cone photoreceptors is found at the centre of an area of the
retina called the fovea (Figure 1a). The fovea is about
1500 mm in diameter, a third of which comprises cone
photoreceptors. This area contains the highest frequency

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Eye Anatomy

of cones per unit area in the entire retina. However, this

structural oddity goes beyond its compositional homogeneity; it actually lacks many of the common retinal layers
as well. The only stratications present are the pigment
epithelium cells, photoreceptor layer, the outer nuclear
layer, and a bit of the outer plexiform. Owing to its
compositional nature and resolving capabilities, the fovea
is an obvious target for light as it enters the eye. The cornea
and lens make it possible to focus light onto this small area
in order to produce images possessing the nest details we
are capable of visualizing.

The Compound Eye

The human and other vertebrate eyes are considered to be
simple not because of any restraints of function, but
because they consist of a singular optical system (primarily
the cornea and lens). On the other hand, the eyes of most
insects and a few crustaceans are considered compound,
with each eye possessing multiple components or optical
systems. The surface of the compound eye is divided into
separate circular or hexagonal facets that act as individual
refractive units, called ommatidia. Each ommatidium does
not receive an entire image, but rather a small part of the
whole. Each parcel of the perceived image travels through
the optical system of the organism, and is eventually fused
to some degree, creating the overall image (Figure 2).

Many structures of the ommatidium are analogous to

those of the simple eye. On the surface of an ommatidium is
the cornea, followed by a conical lens. The two structures
may be physically separated, or fused depending on the
organism. Either way, the lens cannot be adjusted;
therefore the compound eye is a xed-focus eye. Only the
position of the organism can determine what objects are in
focus. Just below the cornea and lens there are nerve cells
called retinula cells, which contain photoreceptors called
rhabdomes. There are anywhere between one and eight
rhabdomes in each retinula cell, which are encased by a
periphery of pigment cells that absorb any excessive light.
However, it is in the rhabdomes that an image forms. As
in most organisms, this image is sent through a series of
neural bres to the brain.

The two types of compound eyes

There are two types of compound eyes: the apposition and
superposition eye. The apposition eye is characterized by
the optical system being continuous with the photoreceptor cells, and is usually a trait of insects adapted to a well-lit
environment. The lens and rhabdomes are constantly
surrounded by pigment, not allowing transmittance into
adjacent ommatidia. This results in many isolated imaging
systems, whose photoreceptors only receive the light that
enters their respective cornea and lens.

Figure 2 Cross-section of a compound eye illustrating a group of ommatidia, and an enlarged view of a single ommatidium.

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Eye Anatomy

The superposition eye is characterized by a separation

between the optical system and the photoreceptor cells,
and is commonly found in nocturnal organisms. Being
creatures of the night, they must gather as much light as
possible, so the ommatidia are often not completely
surrounded by pigment cells. The intended function of
this is to enable the ommatidia to share incoming light in an
attempt to form an image. The eort is often aided by the
convergence of light from many optical systems onto a
single rhabdome.

Evolutionary Trends of Eye Structures

It is believed that eyes have evolved over 40 times,
independently, during the course of their evolution. It is

no wonder that there is such a diversity of eye structures

found in nature. The planarian or atworm eye represents
the most primitive invertebrate eye, made up of visual cells
within a pigmented mantle. In contrast, arthropods have
complex eyes consisting of up to 800 ommatidia, each
containing all the basic eye components. These basic
components of an ommatidium include the cornea and/or
lens for focusing light, pigment cells with absorbing and/or
reecting properties, and retinula cells essential for light
processing.
It is believed that the vertebrate eye evolved independently from its invertebrate counterpart, keeping the basic
eye function but increasing its complexity to accommodate
the needs of the organisms. This adjustment resulted in the
production of a complex eye that contained elaborated
focusing equipment including a cornea, lens, pupil and iris.

Figure 3 (a) Top: cross-section of the newt (vertebrate) eye. Bottom: scanning electron micrograph of a cross-section of the newt eye;  35
magnification. Evident structures include the retina (R), iris (I), cornea (C), and lens (L).
(b) Top: cross-section of a small area of the Drosophila compound eye. Bottom: scanning electron micrograph of a cross-section of the Drosophila eye;
 648 magnification. Evident structures include the cornea (C), and ommatidium (O). The thin, hair-like structures are setae (S), and are believed to reduce
glare.
(c) Top: cross-section of the squid (invertebrate) eye, illustrating the striking resemblances to the vertebrate eye. Bottom: scanning electron microscope
composite image of a cross-section of the squid eye;  5 magnification. Evident structures include the retina (R) and lens (L).

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Eye Anatomy

These eyes also contained an intrinsic light-processing

machinery made up of neural retina cells and the important
supportive cells, the RPE cells. It is amazing, then, how the
cephalopod eye (an invertebrate eye) developed so
similarly to the vertebrate eye (see Figure 3). Overall, the
basic eye function of detecting and transforming light
signals into neuronal signals has been conserved regardless
of the origin of the eye.

Each structure works in accord with the next, refracting,

constricting, dilating, and chemically reacting to convert
patterns of light into interpretable images. Not only are the
mechanisms numerous, but they occur involuntarily and
with extremely high frequency. The functions of the eye
represent a symphony of activity that has been perfected
over millions of years, resulting in each organisms detector
of light, their sculptor of subjective reality, their own
respective evolutionary masterpiece.

Summary
Following the light path through the vertebrate and
invertebrate eye, we have compared the light-focusing
structures as well as the light-transforming cells in both the
vertebrate eye and the compound eye of invertebrates. The
dierent evolutionary trends that shaped the eyes of the
world have also been discussed.
Charles Darwin asserted that the eye is both perfect and
complex. The eye is a small yet multifaceted unit of
anatomical machinery, with intricate design and function.

Further Reading
Dawkins R (1996) The forty-fold path to enlightenment. Climbing
Mount Improbable, pp. 138197. New York: WW Norton.
Kessel D and Kardon RH (1979) Tissues and Organs: A Text-atlas of
Scanning Electron Microscopy. San Francisco: WH Freeman.
Marmor MF and Wolfensberger TJ (1998) The Retinal Pigment
Epithelium. New York: Oxford University Press.
Oyster CD (1999) The Human Eye. Sunderland, MA: Sinauer
Associates.
Sinclair S (1985) How Animals See. New York: Facts On File.

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