Chapter |2|

Physiology of the eye

CHAPTER CONTENTS Alignment of the eyes

Alignment of the eyes 13
Looking straight ahead (fixation) 14
In human beings the two eyes work as though they were
Locking images (fusion) 14
one, both projecting to the same point in space and fus-
Eye movements 15
ing their images so that a single mental impression is
Looking toward a close object 16 obtained by this collaboration. Without this delicate bal-
Seeing in depth 16 ance we would “see double” because two images would
Focusing at near (accommodation) 18 be formed by the independent action of each eye. In other
Transparent pathway for light 19 words, stereopsis would be lost because this faculty is
Retinal images 22 totally dependent on the eyes seeing in unison. The ability
Intraocular pressure 22 of the eyes to fuse two images into a single one is called
Tears 23 binocular vision.
Color vision 24 Binocular vision depends on an exquisite balance of
motor and sensory function. The eyes must be parallel
when looking straight ahead and they must be able to
Physiology of the eye deals with the function of the eye, maintain this alignment when gazing in other positions.
its capacities, and its limitations. The actual perception of Each impulse that directs an eye to move in one direc-
light takes place in a well-delineated area called the field of tion must be equally received by the other eye. Further,
vision. What is not seen beyond these boundaries is cata- the contraction of an eye muscle pulling the eye in one
loged and stored in our visual memory center, so that we direction must be accompanied by an equivalent amount
are not uncomfortable or handicapped by this imposition. of relaxation of its opponent muscle. Without perfectly
Most eyes cannot form a sharp image on the retina without harmonious eye movement, binocular vision would be
an internal adjustment made by focusing or by some exter- impossible because eyes that do not move together do not
nal appliance, such as lenses placed before them. There is see together.
a limit to how much detail the eye can resolve, its magnify- Each eye must have good vision because a clear image
ing abilities being only 15×, considerably less than most and a fuzzy image cannot be fused. The brain usually
microscopes. The spectrum of light to which our retinal ignores the fuzzy image (suppression). Each macula must
receptors are sensitive is confined to specific wavelengths have its projection straight ahead, so that the line of vision
of light; the world of ultraviolet and infrared is invisible to from each eye intersects at one point in space. Also, the field
ordinary perception. of vision from each eye must overlap (Fig. 2.1). Although
Despite these limitations, the human eye is an extremely we can see more with two eyes than with one, this differ-
versatile instrument capable of seeing both in daylight and ence is not great (~35 degrees) because most of the field of
in dim light, registering colors, appreciating depth, and vision from one eye overlaps the field from the other eye.
exercising rapid focusing adjustments. This chapter deals Overlapping visual fields act as a locking device, forging
with the mechanisms that enable the eye to carry out these our peripheral vision in place and thereby ensuring central
tasks. fusion.

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 Section |1| Basic Sciences

120°

Binocular field

Monocular Monocular
field 35° 35° field

Blind area 170° A Macular pathway

True

Fig. 2.1 Field of vision. Binocular field of vision (120 degrees)

represents the overlapping field of vision from each eye.

Looking straight ahead (fixation)

Fixation involves the simple task of looking straight ahead B False

toward an object in space. Fixation requires stability of the
eyes and good monocular function. If the eyes are con- Fig. 2.2 (A) Binocular vision (both eyes looking at the same
figure). (B) One eye is turned in, resulting in double vision. In
stantly moving, such as occurs with congenital nystagmus
this case the figure is received by the macula of one eye and a
(shaking of the eyes), the eyes can make only scanning point nasal to the macula of the turned eye. The projection of
motions around an object and never adequately see it in this nasal point results in the person seeing two images instead
detail. Needless to say, if the ability to fixate becomes com- of one of the same figure. This is an example of uncrossed
promised by constant eye movements, then the visual acu- diplopia, as seen in esodeviations.
ity of the affected eyes is reduced. If the macula is damaged,
then fixation is difficult because anything viewed directly
ahead becomes enshrouded in relative darkness. easily disrupted by covering one eye. The eye under cover
Fixation can be reduced without organic changes in the eye. drifts to its fusion-free position. The amount of movement
Children with strabismus often are found to have poor vision that the eye makes is a measure of the latent muscular
in the turned eye. If a child has crossed eyes, we would think imbalance kept in check by fusion, or the amount of heter-
that double vision would occur because the two eyes would ophoria. Heterophoria, then, may be defined as the position
not be directed to the same point in space (Fig. 2.2). Children, the eyes assume when fusion is disrupted. The eye under
however, have a wonderful faculty for completely ignoring cover may drift in, called esophoria, or drift out, called exo-
the image in the turned eye to avoid confusion. It is this con- phoria. The eye also may drift up and down; this position
stant habit of actively suppressing the image in the turned eye is called hyperphoria. Fusion also may be disrupted by plac-
that eventually leads to loss of vision or amblyopia. In some ing a Maddox rod before one eye. The Maddox rod changes
of these children, in whom the suppression mechanism has the size and shape of the image presented to the eye under
become profound and the resultant vision very poor, foveal cover so that fusion becomes impossible.
function becomes so depressed that a new point just outside The power of fusion is measurable by prisms (see Ch. 3).
the fovea is used. Such an eye can no longer see straight ahead For example, a four-diopter prism is placed with the base
and the fixation pattern is described as eccentric. toward the nose of an observer looking at a small letter
placed 16 inches (40 cm) from the eye. The prism will dis-
place the image before that eye in a direction toward its
Locking images (fusion) apex and the eye moves outward to follow it because of the
power exerted by the fusional reflex (Fig. 2.3A). Now the
prism is removed and the uncovered eye returns to its origi-
Fusion is the power exerted by both eyes to keep the posi- nal position in response to the fusional reflex (Fig. 2.3B).
tion of the eyes aligned so that both foveae project to the Normally, 20- to 40-prism diopters can be exercised by
same point in space. Because fusion is a binocular act, it is fusional convergence. The amount of fusion exercised with

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 Physiology of the eye Chapter |2|

eye is either covered or patched. The movement of

two eyes in the same direction is called a version
(dextro-, levo-, sursum-, and deorsumversion)
(Fig. 2.4).
• Eyes right: dextroversion
A • Eyes left: levoversion
• Eyes up: sursumversion
• Eyes down: deorsumversion
An outline of the functions of the extraocular muscles
is given in Table 2.1. The medial and lateral rectus mus-
B cles have only one action: to move the eye horizontally.
Fig. 2.3 (A) The prism displaces the image toward its apex
The other four muscles of the eye have auxiliary functions.
and the eye moves outward because of the fusional reflex.
(B) When the prism is removed, the eye returns to its original
position because of the fusional reflex.
Table 2.1 Actions of extraocular muscles
respect to divergence is less, being only 10- to 20-prism
diopters. This is measured by using base-out prisms. Vertical Secondary
imbalances are difficult to overcome because our eyes can Muscle Prime action action
overcome only about 2- to 4-prism diopters.
Medial Turns eye inward toward nose None
rectus or adducts eye

Eye movements Lateral Turns eye outward toward None

rectus temples or abducts eye

The primary position of the eyes is the straight-ahead posi- Superior Elevates eye Intorsion
rectus Adduction
tion as they look at a point just below the horizon with
the head held erect. Movement of the eye from the primary Inferior Depresses eye Extorsion
position to a secondary position occurs when the eyes rectus Adduction
are moved either horizontally or vertically. If the eyes are
Superior Intorts eye Depression
directed in an oblique position (up and in or down and
oblique Abduction
in), they are said to be in a tertiary position.
• The movement of one eye from one position to another Inferior Extorts eye Elevation
in one direction is called duction. In duction, the fellow oblique Abduction

Sursumversion

Dextroversion Levoversion

Deorsumversion
Fig. 2.4 Version movements of the eyes or movements formed by both eyes working together.

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 Section |1| Basic Sciences

Right eye Left eye

Superior rectus Inferior Inferior Superior

oblique oblique rectus

Lateral rectus Medial rectus Medial rectus Lateral rectus

Inferior rectus Superior Superior Inferior

oblique oblique rectus

Fig. 2.5 Action of the extraocular muscles. The arrows reveal that the superior and inferior rectus muscles function best
as an elevator and a depressor, respectively, when the eye is abducted. The inferior and superior oblique muscles function best as
an elevator and depressor, respectively, when the eye is adducted.

When these secondary roles are used, assisting the lateral or exactly. With greater speeds, following movement becomes
medial rectus muscles to abduct or adduct, these muscles difficult and the smooth, gliding movement is replaced with
are called synergists (Fig. 2.5). an irregular, jerky movement. Pursuit movements are con-
The main function of the oblique muscles is to rotate trolled from centers in the occipital lobe of the brain.
the globe either inward (intorsion) or outward (extorsion).
Intorsion occurs when the eye rotates on its long axis so
that the 12 o’clock position on the cornea moves toward the Looking toward a close object
nose. For example, if a point on the cornea of the right eye
moves inward from 12 to 1 o’clock, then intorsion is said
to occur because of the primary action of the right superior Vergence is the term applied to simultaneous ocular move-
oblique muscle or secondary action of the right superior rec- ments in which the eyes are directed to an object in the
tus muscle. Similarly, if the point on the right cornea moves midline in front of the face. The term is usually applied to
outward from 12 to 11 o’clock, then extorsion is said to convergence, in which the eyes rotate inward toward each
occur because of the primary action of the right inferior other, or to divergence, in which they rotate outward simul-
oblique muscle or secondary action of the right inferior rec- taneously (Fig. 2.6).
tus muscle. Convergence is invariably accompanied by narrowing,
or constriction, of the pupils and by accommodation. The
triad of convergence, pupillary constriction, and accommo-
Control centers for eye movements dation is often called the accommodative reflex, although in
The eyes move in response to our own volition or in a pas- the true sense these movements are merely associated reac-
sive manner, such as in following a slow-moving target. tions (synkinesis) rather than a true reflex. Each component
Volitional eye movements usually are rapid, starting at high of the triad facilitates fixation at near. The constriction of
speeds and ending just as abruptly. Such movements occur the pupil is the attempt by the eye to form a pinhole cam-
with reading, when words or phrases are quickly scanned, era device so that a clearer image is seen. Accommodation
with an abrupt halt coming at the end of a section or a line. enables the object to be focused on the retina; convergence
These voluntary eye movements are controlled from centers brings the eye inward toward the object of regard.
in the frontal lobe of the brain.
Whereas voluntary eye movements tend to be short and
choppy, following or pursuit eye movements are rather slow, Seeing in depth
smooth, and gliding. The velocity of a following movement
depends entirely on the speed of the object the eye is tracking.
If the fovea is fixed on a moving target with an angular veloc- The ability to see in depth enables us to travel comfort-
ity (<30 degrees per second), the eye follows the target almost ably in space. Without it, we could not judge distances,

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 Physiology of the eye Chapter |2|

Convergence
A
A

B Divergence

Fig. 2.6 (A) Convergence. The eye is turned in toward the

midline plane. (B) Divergence. The eye is turned out, away B
from the midline plane.
Fig. 2.7 (A) Artist has drawn the picture with proper depth
perspective. Monocular clues include decrease in size of dogs
and confluence of lines toward a point. (B) Artist has ignored
the usual monocular clues so that our appreciation of depth
estimate the size of objects beyond us, or avoid bumping and size is erroneous. The second dog appears larger than the
into things. Without depth perception, even the simplest first, although both are the same size.
of tasks would be difficult. We would be unable to reach
accurately for our morning coffee, and passing a car on the
highway would be tantamount to suicide. Fortunately, eve-
ryone has some depth perception, whether the person has • Parallax: if two objects situated at different points
one eye or two. Those with only one eye learn to estimate in space are aligned and the head of the observer is
depth with monocular clues (Figs. 2.7 and 2.8). They know moved in one direction, the nearer object will appear
that the speck in the distance that becomes a huge train to move in the opposite direction
standing beside them in the station has not grown larger A monocular person, however, if removed from familiar
but has merely come closer. There are other clues in addi- surroundings, would have great difficulty in judging dis-
tion to changes in object size. The train tracks spread from tances because of a lack of any intrinsic depth-perception
a point and become parallel, the color of the train changes mechanism. For example, a one-eyed pilot would create a
from a misty blue-gray to dark green, the sound increases, hazard because of the difficulty he or she would experience
and when the train is alongside, one can feel the heat. in maneuvering in space without the normal monocular
• There are many monocular clues that facilitate depth clues.
perception, including the following: magnification: Stereopsis is a higher quality of binocular vision. Each eye
well-recognized objects, if they become larger, are views an object at a slightly different angle, so that fusion
deemed to be nearer of images occurs by combining slightly dissimilar images.
• Confluence of parallel lines to a point (e.g., railway It is the combination of these angular views that yields ste-
tracks) reopsis. The same method is used in photography in mak-
• Interposition of shadows ing three-dimensional pictures. The stereoscopic picture is
• Blue-gray mistiness of objects at a great distance taken at slightly different angles and later viewed that way.

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 Section |1| Basic Sciences

A B
Fig. 2.9 (A) Crystalline lens of the eye is thin for distant
objects. (B) Crystalline lens accommodates for near objects by
becoming thicker. This increases its effective power.

B
Fig. 2.8 (A) The scene is drawn using normal monocular clues
of distance, thereby giving it perspective. (B) The same scene is
drawn without regard to the normal impressions of distance.
Therefore the scene loses its perspective.

Fig. 2.10 Measurement of the near point of accommodation.

Focusing at near (accommodation)
accommodative target (such as a small letter) can be seen
Any object can be moved from a distance to about 20 feet clearly. It usually is equal in both eyes. The range of accom-
in front of an observer and still be seen clearly without modation is measured in centimeters, whereas the power is
accommodation. This distance is called the range of focus. As converted to diopters (Table 2.2).
the object is brought closer than 20 feet, however, the eye This stimulus for accommodation is a blurred image on
must continuously readjust to keep the image of the object the retina. As an object is moved closer to the eye, the rays
clearly focused on the retina. This readjustment requires an of light entering the pupil must be continuously converged.
increase in the power of the eye and is brought about by an This change in focusing power of the eyes is brought about
automatic change in the shape of the lens in response to a by active contraction of the ciliary muscle. The contrac-
blurred image (Fig. 2.9). This zoom-lens mechanism in the tion of this muscle causes the zonular fibers of the lens to
eye is very active in children; they are able to see a small let- relax, which in turn allows the lens of the eye to change
ter in clear focus only 7 cm from the eye, whereas an adult its shape (Fig. 2.11). In the child and the young adult, the
of 55 years can focus no closer than 55 cm. The range of lens can be molded, and it increases its power by becoming
accommodation is the distance in which an object can be thicker and increasing the curvature of its anterior space.
carried toward an eye and be kept in focus. The power of In an adult, the ability of the ciliary muscle to effectively
accommodation of an eye is the dioptric equivalent of this contract declines with age and the lens becomes harder and
distance. By age 75 years, this power is zero. less malleable with advancing years.
Both the range and the power of accommodation are The decline in accommodation with age, called presbyo-
measured quite easily (Fig. 2.10). When the full spectacle pia, is remedied with reading glasses or bifocals. It usually
correction is worn, it is merely the closest point at which an becomes apparent by the age of 45 years.

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