How the Human Eye Focuses

As people age, their ability to focus on nearby objects

gradually declines. Several probable causes have been identified,
including changes in the eye's geometry and biochemistry

by Jane F. Koretz and George H. Handelman

I
t happens to almost everyone: by focus, the light must be bent so that point closer than 20 feet away, the
about the age of 45 it becomes the rays converge at the fovea, the ciliary muscle contracts, reducing the
impossible to read without the center of the retina. The nearer some­ diameter of its opening and also caus­
help of glasses. How does the healthy thing is to the eye, the more the light ing the muscle to move slightly for­
young eye focus on a nearby object? must be bent if the object is to be seen ward. Both changes reduce the stress
Why does near vision fade? The an­ clearly. The cornea, aqueous humor on the zonules and thereby lessen the
swer to the first question has long and vitreous body each have a fixed stress exerted by the zonules on the
been incomplete, and the answer to refractive power, or ability to bend lens. The lens thereupon undergoes
the second remains a matter of con­ light, but the lens can accommodate: it elastic recovery: much as a foam-rub­
jecture. By means of photographic can sharpen the curvature of its front ber ball expands after it has been
studies of the lens of the eye and and back surfaces, thereby increasing compressed, the lens rebounds to a
mathematical modeling, we have re­ its focusing power. more relaxed state. As the lens focus­
cently gained new insight into both es on progressively closer objects, it

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problems. We have shown that several hat, then, enables the lens to becomes thicker from front to back,
processes conspire to progressively accommodate? A broad expla­ its surfaces become more sharply
limit one's ability to focus on close nation, which our work and curved and the diameter of the equa­
objects; other processes counteract that of others generally supports, was tor shrinks. This relaxation process is
the decline for a while, but these ulti­ put forward in the mid-19th century precisely controlled to provide the ex­
mately fail-typically in the fifth dec­ by the German physicist Hermann von act degree of extra refractive power
ade of life. Helmholtz in his Treatise on PhYSiolog­ needed for focusing on objects closer
When a person looks at something, ical Optics. Helmholtz noted that the than 20 feet away.
light reflected from the object pass­ lens is suspended by filaments that The lens, then, is unaccommodat­
es through the cornea (a transparent project from the so-called ciliary mus­ ed-flattest and the least refractive­
sheath across the front of the eye) and cle, which encircles the equator, or when it is under maximum stress:
a fluid known as the aqueous humor, rim, of the lens like a collar but does when the eye focuses on infinity and
on through the pupil of the iris and not come in direct contact with it. the ciliary muscle is totally relaxed.
into the lens, which is normally clear These nonelastic filaments, called the The lens is maximally accommodat­
and is shaped and oriented something zonules of Zinn, are now known to ed-most sharply curved and most
like the lens of a camera. From there form three rings of hairlike "spokes" refractive-when it is under the least
the light travels through the gel-like around the lens; one ring attaches to stress: when the eye focuses on the
vitreous body to the retina, the part of the equator and the other two attach closest discernible object and the cili­
the eye that converts light into electri­ somewhat in front and in back of it. ary muscle is fully contracted.
cal signals that are transmitted to the Helmholtz proposed that when the Helmholtz' accommodative model
brain for interpretation. eye is focused on infinity (which for is widely accepted today, but it leaves
In order for the image to come into human beings begins about 20 feet many questions unanswered. For in­
away), the sphincterlike ciliary muscle stance, what effects do small changes
relaxes and therefore expands; the di­ in the pattern of stress exerted by the
JANE F. KORETZ AND GEORGE H. HAN­ ameter of the circular muscle reaches zonules have on lens shape? How
DELMAN are colleagues at the Rensse­ a maximum. As the muscle expands it much relaxation of the force exerted
laer Polytechnic Institute. Koretz, who pulls the zonules taut, causing them by the zonules is needed to produce
earned her PhD. in biophysics from the
in turn to pull on the lens. The pull­ enough lens curvature for, say, read­
University of Chicago, is associate pro­
ing flattens the front and back of the ing, and at what angle must the zon­
fessor of biology and a member of the
Center for Biophysics. Handelman holds lens and increases the diameter of its ules meet the lens? Does the vitreous,
the Amos Eaton Chair at Rensselaer. He equator. In this condition-called the to which Helmholtz paid little atten­
received his doctorate in applied math­ unaccommodated state-the ability of tion, have a role in accommodation?
ematics from Brown University and the lens to bend light is at a minimum. Moreover, Helmholtz conceived of
served as chairman of the department The combined refractivity of the cor­ the lens as a readily deformable bag of
of mathematics and dean of the School
nea, aqueous humor, unaccommodat­ fluid. Actually the material within the
of Science at Rensselaer for 18 years.
ed lens and vitreous is just right for "bag" consists of long, ribbonlike fi­
The authors dedicate this article to the
memory of Henry N. Fukui of the Nation­ focusing an image of a distant object bers that interlock and also nest with­
al Eye Institute. on the fovea: in one another like the layers of an
When the eye attempts to focus on a onion. The bag itself, the lens capsule,

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LENS OF HUMAN EYE is suspended by filaments known as the zonules exert major control over the ability of the lens to
zonules of Zinn (purple "threads") from the ciliary muscle, a accommodate: to change shape so as to increase its focusing
ring of tissue essentially concentric with the lens. In this power. The accommodative facility usually is lost by the age of
frontal view the muscle is not visible; it is embedded in the about 45. Patricia N. Farnsworth of the University of Medicine
ciliary body (dark, ruffled region). The ciliary muscle and the and Dentistry of New Jersey in Newark made the photograph.

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 is composed of fibers of a different
kind, oriented parallel to the lens sur­
face. Both kinds of fiber, which are rich
in protein, resist stretching but not
bending. These findings raise an ad­
ditional question: What effect does
the structural organization of the lens
have on accommodation?
Someone searching for answers to
such questions would ideally examine
the zonules and the vitreous directly
in the living eye, measuring the mag­
nitude and direction of the forces
they exert on the lens capsule and
the lens body. The investigator would
also measure the distribution of for­
ces throughout the lens to determine
the effect of the internal structure on
the response to external stresses. In
reality, of course, it is impossible to
make such direct measurements.
It is possible, however, to describe
changes in the shape of the living lens
as it accommodates and to calculate
the magnitude and direction of the
forces that would have to act on the
lens in order to produce the described
changes. This information can then be
correlated with what is known about
the architecture of the eye to deter­
mine which structures are capable of
exerting the calculated forces.

T
hat is the approach we adopted.
We began by creating a mathe­
matical model of the lens body­
the lens minus the capsule. For sim­
plicity we concentrated on the lens of'
a young person about 10 years old.
Such a lens can be represented quite
accurately as a somewhat distorted
sphere composed of two hemispheres
of differing radii, one representing the
front of the lens and the other the
back. We then made certain assump­
tions in our model about the elasticity
of the lens body, for instance that it
responds differently to stress exerted
parallel to and at a right angle to the
optical axis of the eye. The model also

STRUCTURE of the lens is shown in this

drawing. The zonules attach to the cap­
sule, or outer membrane, in three plac­
es-at the equator, or rim, and at points
somewhat anterior and posterior to it.
The inner "lens body" has two main re­
gions: the nucleus (the original fetal
lens) and the cortex, consisting of the
fibers that have been laid down since
birth. The fibers, which extend from the
anterior to the posterior pole, originate
as epithelial cells (detail) at the outer
boundary of the lens body. With time the
cells elongate into ribbons, lose their
nuclei and are covered by newer cells; as
a consequence the lens grows thicker.
The lens fibers nest within one another
much the way the layers of an onion do.

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assumed that, because they are inter­
locking, the lens fibers cannot slide
past one another: that the only way
the lens can change shape is for the
fibers to alter their curvature.
Our calculations indicated that all
the forces acting on the surface of the
accommodating lens body are approx­
imately equal and are in a direction
approximately perpendicular to the
lens surface. This finding suggests
that the capsule, which is subject to
discretely applied stresses from the
zonules and is the only material in
direct contact with the lens body (the
zonules do not penetrate the body),
transforms these discrete stresses
into a uniform compressive force
against the entire surface of the lens.
When the zonular stress on the cap­
sule is released, the compressive force
acting on the lens body is also re­
leased; the compressive force within
the lens is reduced as well, and the
lens undergoes elastic recovery. The
finding that the force exerted by the
capsule is perpendicular to the sur­
face of the lens body is not entirely
surprising. The zonules exert a force
that has both a parallel (stretching) LENS FIBERS, here enlarged about 8,000 times, interlock along their long edges, as
component and a perpendicular (com­ is seen in this electron micrograph made by Richard G. Kessel of the University of
pressive) one. The capsular fibers, Iowa. Each fiber is also linked to those above and below it, as is shown in the illus­
however, resist stretching, and so only tration on the opposite page. The interlocking of the fibers affects the distribution
the perpendicular force is transmitted of forces within the lens and constrains the shape it adopts during accommodation.
to the lens.
Another important implication is
CILIARY BODY
that the shape assumed by the lens at
a given accommodation state is in­ ZONULES
deed different from the shape it would
assume if it were merely a bag of fluid. CORNEA
Under uniform pressure on its surface
a bag of fluid could not achieve the
known curvatures of the accommo­
dated lens. The observed changes in
shape must therefore be influenced by
the internal structure of the lens.
We were now ready to determine the
forces exerted on the lens by other
structures in the eye. Relaxation of the
zonules can account for most but not
all of the shape change seen in the
lens during elastic recovery. This sug­
gests that the vitreous humor may
take part in the focusing process-by
lending support to the posterior of the
lens. Interestingly, at about the age LENS
when most people need reading glas­
ses, the gel-like vitreous begins to liq­
VITREOUS BODY
uefy, losing its ultrastructure. Exactly
how such liquefaction would contrib­
EYE FOCUSES on an object by refracting, or bending, the light (cone) reflected from
ute to the loss of accommodative abil­
the object so that the light rays converge on the retina. Nerve cells in the retina
ity is not clear, but it may have some
transform the light into electrical signals that are transmitted to the brain for
role to play.
interpretation. Light is bent at the front and back surfaces of both the cornea and
The only way the zonules could pro­ the lens, but only the lens can accommodate. Excessive refraction causes light
duce the calculated change in forces '
to converge in front of the retina and thus impairs distance vision, whereas insuffi-
during accommodation would be to cient refraction causes light to converge behind the retina and impairs near vi­
lose tension themselves or to change sion. A progressive decrease in the refractive nature of the lens and in the ability
the angle at which they apply stress to of the lens to accommodate is thought to undermine near vision in middle age.

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the lens in such a way that the com­ ever ; if they did, there would be no clines from about 14 diopters at the
pressive component of the stress is way to hold the lens in position. age of lO-when it is possible to focus
decreased. Our data suggest that they on the tip of one's nose-to about nine

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do both. In addition to becoming less he model based on the young diopters at 20, four diopters in the
taut, the zonules, which are attached eye enabled us to understand mid-thirties, one or two diopters in
to the capsule, move slightly, becom­ many of the details of accommo­ the mid-forties and close to zero by
ing more parallel to the surface of the dation, but it did not explain why the the age of 70. A zero-diopter eye can­
capsule; the more parallel the zonules refractive power of the eye, and hence not focus on anything nearer than
are, the less compression they cause the ability to focus on nearby objects, infinity. The change from four diop­
in the lens. Experimental evidence declines with age. The eye's refractive ters (roughly 10 inches) to two (rough­
from other investigators confirms that power is measured in diopters, the ly 20 inches) or less is the one people
in at least one individual (who has a reciprocal of the distance in meters notice because it affects reading; most
rare disorder of t he iris that makes between the eye and an object. For people hold books from 12 to 16 inch­
the normally ob�cured zonules visi­ instance, an eye with a refractive pow­ es away from their eyes.
ble through a microscope) the zonules er of 10 diopters can bend light suffi­ As a first step toward understanding
hang quite luosely when the lens is ciently to focus on an object a tenth of the cause of this gradual age-related
maximally accommodated. Our calcu­ a meter, or roughly four inches, away. decline in near vision, we decided to
lations suggest that the forces acting In human beings who start off with gather as much information as possi­
on the lens never decline to zero, how- normal vision the refractive power de- ble about the way the lens changes
shape with age and accommodative
state. Having done thiS, we would at­
tempt to determine both the effect of
such changes on refractive power and
the likely causes of the changes. We
began by examining a series of cross­
sectional photographs of the lens
made by Nicholas Phelps Brown of the
Institute of Ophthalmology in London
in the early 1970's and by preparing
100 similar sets of photographs of our
own. Brown, who gave us technical
advice, provided four photographic
sets from subjects 11, 19, 29 and 45
years old, who focused on objects at
varying distances from the eye; our
study included subjects ranging in age
from 18 to 69 who had healthy eyes
and normal distance vision. All the
cross sections are vertical "slices" run­
ning from the front to the back of the
lens and were made with the aid of
a slit lamp, which shines a narrow
("slit") beam of light into the eye.
A quick look at the photographs
confirmed the well-known fact that
the size of the lens increases as a
function of age. The unaccommodated
lens of the infant has been found to be
about 3.3 millimeters thick from front
to back. As time passes, the cells that
constitute the outer layer of the lens
body (the epithelial cells lying just
inside the capsule) grow and are trans­
formed into the kind of ribbonlike
fibers that constitute the bulk of the
lens. As new epithelial cells are laid
down over the older fibers, they un­
dergo the same growth process as
their predecessors, and so the lens
LENS INCREASES its focusing power by becoming more sharply curved. When the
thickens. By the time a person is 70
initially imaccommodated, or flattened, lens is viewed from the side (a), it looks fairly
the unaccommodated lens can be as
thin from front to back. It thickens as it accommodates (b); the front surface moves
much as five millimeters thick.
closer to the cornea but the back surface remains in place. The change in shape is
effected primarily by the contraction of the ciliary muscle. A front view shows that The photographs also revealed a se­
the lens adopts the unaccommodated state (c) when the muscle expands so that its ries of bands in the interior of the lens
diameter is at a maximum. The expansion of the muscle pulls the zonules taut, and known as zones of discontinuity. The
they pull on the lens and flatten it. When the muscle contracts (d), the zonules relax, bands in the front of the lens exhibit
and the lens rebounds into a rounded state, much as a foam-rubber ball rebounds roughly the same curvature as the
after compression. The illustrations of accommodation are exaggerated for clarity. front surface, and the bands in the

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back are similar to the posterior sur­ creasingly curved shape as it grows in er lens had to move more than those
face, although the curves become pro­ size throughout a life span. Moreover on a younger lens.
gressively sharper as they approach (as Brown had discovered with sur­ A critical finding was that the total
the nucleus, or core, of the lens. In the prise in his subjects), in people young­ range of movement declines gradual­
young lens the bands are few and er than about 45 the older the lens ly with age. Indeed, in subjects older
faint. As the lens ages, the number was, the more sharply curved it was than about 45 the front of the lens was
and clarity of the zones increase until for any given accommodation state. unable to change shape and so could
sometime in or soon after the fifth For example, a 33-year-old lens curved not accommodate: it appeared to be
decade, when the zones merge. more than a 19-year-old lens focus­ locked in the unaccommodated state.
ing at the same close distance. Brown This suggests either that the front

O
n the basis of the photographs called the phenomenon of increas­ zonules become progressively less
and other sources of informa­ ing curvature with age the lens para­ able to relax when the ciliary muscle
tion, we created another mod­ dox, because one would expect more contracts in the attempt to accommo­
el. This time we described the surface sharply curved lenses to produce a date, or that the zonules relax but
and interior curves of each lens in greater degree of refractive power become less able to exert much influ­
mathematical terms. (They were pa­ than less curved ones. ence on the lens, or both. The zonules
rabolas and hence could be re-created When we traced the movements of might be unable to relax if the front
with simple equations.) Then we ex­ selected points on the lens surface of the enlarged lens is so far from
trapolated from the cross sections to and in the interior as the eye accom­ the ciliary muscle that the lens pulls
determine the shape of the entire lens modated, we found a parallel effect: the zonules taut. Moreover, just as
in all accommodative states as a func­ equal amounts of movement during the zonules meet the lens at a differ­
tion of age. This enabled us to make accommodation produced less change ent angle when the lens is accommo­
extensive comparisons and to track in focus in the older eyes than in the dated than when it is unaccommo­
the movement of selected points in younger ones. Considered in another dated, so they meet the lens at a differ­
the lenses during focusing. way, the data showed that in order to ent angle when the lens is thick from
The comparisons revealed that the achieve, say, a one-diopter increase in aging than when it is younger and
unaccommodated lens adopts an in- focusing power, the points on an old- thinner. Ultimately the filaments may

SOME AGE-RElATED CHANGES are readily apparent in slit-lamp discontinuity. As the lens ages,the bands multiply and become
photographs of unaccommodated (top) and maximally accom­ more prominent; when the lens is more than about 45 years
modated (bottom) lenses from subjects (left to right) 19,33,45 old, the zones merge. The authors suggest that the increas­
and 69 years old. (The subjects' maximum focusing power is ing thickness of the lens and a rise in the fraction of insol­
respectively 9,4.5, 1 and .25 diopters; lower numbers indicate uble protein in the zones contribute to an age-related decline
less power.) The photographs show vertical, front-to-back in the refractive nature of the lens. For a time the increasing
cross sections; the front of the lens is to the left. The changes curvature of the lens may help to compensate for the de­
include increased growth and curvature with time and, in the cline; so might the extra refractive surfaces provided by the
45- and 69-year-old lenses, an almost complete failure to ac­ zones. Eventually, however, compensatory mechanisms fail­
commodate. Also apparent are dark bands called zones of probably at about the time the lens ceases to accommodate.

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come to exert a force that is tangen­ light's trajectory, such as the curva­ include each zone of discontinuity as
tial, or nearly so, to the surface of the ture of each lens in accommodated yet another refractive surface.
lens. Once this stage is reached, relax­ (well-focused) states, the curvature of We ran another set of simulations in
ation of the zonules would have little the cornea and the spacing between which we kept the overall refractive
effect on the shape of the lens, which the cornea and the lens and between index of the lens constant but allowed
in turn would exhibit little or no elas­ the front and the back of the lens. the indexes of the zones to vary. This
tic recovery. (Along with curvature, the distance time the simulation allowed the eye of
In other words, presbyopia-the between refractive surfaces influences a 45 -year-old subject to focus, but the
name given to the age-related loss of the path of the light; closely spaced simulations for younger subjects re­
near vision-appears to be a geomet­ surfaces bend light more than widely mained underfocused. Moreover, the
ric disorder, stemming to a large ex­ spaced ones.) We then assigned to younger the subject was, the more the
tent from a change in the size and each part of the eye a refractive in­ simulated eye was underfocused. The
angular relations between the lens dex-one that is generally accepted results suggest that the overall index
and the zonules. in the literature-and instructed the of refraction for young eyes is actually
computer to bend the light at the somewhat higher than the values in

W
e think geometry explains boundaries between materials having the literature and that the younger the
why the lenses of people old­ different indexes of refraction. If the subject is, the more the index varies
er than 45 or so can no long­ overall index of refraction of the lens from the accepted value.
er accommodate, but what explains remained constant throughout life, Looked at in another way, the re­
Brown's paradox? Why is it that an the simulations would have indicated sults showed that the index of refrac­
older lens has to be more curved than that the light passing through each tion of the lens decreases with age.
a younger one to focus on the same eye was bent in a way that focused an This finding fits with the age-related
object? One possibility is that the na­ image on the retina. changes in the lens discussed above. If
ture of the cytoplasm in the lens fibers In the simplest simulation we had the refractive index of the lens materi­
changes in a way that decreases the the light bend at the front and back of al decreases with age, the only way the
lens's refractive index-a measure ofa both the cornea and the lens. In anoth­ reduction could be lessened would be
material's ability to refract light. If the er simulation we treated the bound­ by increasing the sharpness of curva­
refractive index declines with age, aries between the nucleus and the ture of the surfaces of the lens or the
then the decline would solve the para­ outer "cortex" of the lens as addition­ zones of discontinuity within the lens,
dox: the increased curvature would al refractive surfaces and assigned to or by increasing the number of re­
not increase the focusing power of the the two regions different, but again fractive surfaces. Clearly all these
lens; instead it would simply compen­ accepted, indexes of refraction. mechanisms operate. In addition to
sate somewhat for the decline in the Both simulations failed abysmally. increasing the curvature of the lens
refractive nature of the medium. For every accommodative state and surfaces, the aging eye also develops
To explore this possibility, we un­ age at which data were available, the more-and more sharply curved­
dertook computerized "ray tracing" focal point of the simulation was be­ zones of discontinuity. Indeed, the
experiments that simulated the pas­ hind the retina, as if the lenses of all contribution of the zones to the eye's
sage of light through every eye we the subjects had too little refractive overall refractive power becomes in­
had photographed. We did this by de­ power. Something had to be wrong creasingly important with age.
scribing the factors that influence the with the model. Perhaps we needed to One remaining mystery is the nature

AITACHMENT SITES of zonules on the front surface of the lens lens ultimately loses its ability to accommodate in part be­
change as the lens ages and grows. In a 17 ·year-old (left) the cause the zonules are almost parallel to the face of the lens. A
zonules are close to the equator,but they move progressively release of tension on such zonules can have little effect on the
onto the face of the lens in a 46- and an 85-year-old (middle and lens. The photographs were made by Farnsworth,who was the
right). As the zonules shift, so must the angle formed by the first to propose that changes in the lens-zonule geometry
lens surface and the fiJament. The authors propose that the could contribute to the failure of aged eyes to accommodate.

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of the zones of discontinuity that are
so apparent in slit-lamp images. Ul­
trastructural examination of the lens
provides no evidence that they exist.
Furthermore, if protein concentration
is measured as a function of distance
from the lens surface, one finds a
small but smooth increase rather than
the alternating increases and decreas­
es that might be expected if the dark
bands resulted from differences in
protein concentration. Why, then, are
these distinct regions clearly visible
in photographs?
The answer, we concluded, had to
lie in the nature of the photograph­
ic technique, which transmits the slit
beam into the eye and produces an
image from the light that is reflected
into a camera located to the side of the
original light source. (The film in the
camera is tilted to compensate for the
ALPHA-CRYSTALLIN,the major protein in the lens, is usually present as flat sheets
angular distortion.) We reasoned that (larger shapes), but these can break apart to form small, insoluble spheres (smaller
in order for the zones of discontinuity spots). The spheres in turn can join to form rodlike bodies that can adhere to one
to appear in the photographs, the ma­ another. If the aggregates grew very large-as it is thought they may-they would
terial in these regions would have to scatter a great deal of light. Such scattering could account for the presence of the
interact with light differently from the zones of discontinuity in the photographs on page 97. The insoluble nature of the
material in adjacent regions. In partic­ particles could also account for the apparent age-related decline in the refractivity
ular, one would expect to find dark of the lens. The spheres shown here are about 11 billionths of a meter in diameter.
bands in the photographs if the pro­
tein in the corresponding regions scat­
tered light to a great extent. Is such Our data suggest that the degree of ses are somewhat counteracted-at
scattering possible? scattering is minimal in the first few least for a time-by the development
At first glance the answer appears to decades of life but slowly increases. of new refractive surfaces (the zones
be no. In the eye alpha-crystallin, the The increasing presence of insolu­ of discontinuity), by the overall sharp­
major protein component of the lens, ble protein particles could also ex­ ening of lens curvature and by fur­
is generally thought to assemble into plain why the index of refraction of ther sharpening during accommoda­
aggregates that are smaller than the the lens declines with age. The index tion. Eventually, at about the time the
minimum size that would cause scat­ of refraction of a solution (such as the vitreous liquefies, such compensato­
tering. Yet the size of such particles is cytoplasm in lens fibers) depends on ry mechanisms fail and the lens loses
known to increase in response to in­ the nature and concentration of its its ability to accommodate.
creases in temperature, and it is solutes, or dissolved materials. Add­ Perhaps someday investigators will
thought to change also in response to ing soluble protein to an aqueous me­ learn enough to reverse or prevent the
such environmental factors as small dium increases the medium's refrac­ natural, age-related decline of near
alterations in pH level or calcium con­ tive index. On the other hand, if a large vision. For now, however, the need
centration. Many vision scientists have fraction of the protein is converted for reading glasses is-like death and
also shown that although the protein into large, insoluble particles, the re­ taxes-inevitable.
concentration in the eye remains con­ fractive index will decline appreciably.
stant over the course of time, the frac­ Such may be the case in the lens.
FURTHER READING
tion of protein that is insoluble-and
ANALYSIS OF HUMAN CRYSTALLINE LENS

B
therefore particulate-increases. oth microscopic and macro­ CURVATURE AS A FUNCTION OF ACCOM­
Such findings raise the possibility scopic factors must be invoked MODATIVE STATE AND AGE. J. F. Koretz,
that, with time, alpha-crystallin may to explain why the nearest point G. H. Handelman and N. P. Brown in
form larger aggregates than has been one can see with clarity moves pro­ Vision Research, Vol. 24, No. 10, pages
suspected. If this is the case, the pres­ gressively farther away as time pass­ 1 14 1- 1 15 1; 1984.
ence of a significant number of large, es. In our conception, the increasing ON THE HYDRAULIC SUSPENSION THEORY
OF ACCOMMODATION. D. jackson Cole­
insoluble particles of alpha-crystallin amount of insoluble lens protein, the
man in Transactions of the Ameri­
would help to explain not only the growing size of the lens (and hence
can Ophthalmological Society, Vol. 84,
appearance of the zones of disconti­ the increased spaCing between the pages 846-868; 1986.
nuity in our photographs but also a front and the back of the lens) and the MODELING AGE-RElATED ACCOMMODA­
phenomenon known as glare, in which concomitant reduction in the index of TIVE Loss IN THE HUMAN EYE. jane F.
intense light whites out the entire vis­ refraction tend to interfere with near Koretz and George H. Handelman in
ual field. The phenomenon is particu­ vision. Closeup viewing is also in­ Mathematical Modelling, Vol. 7, pages
1003- 10 14; 1986.
larly common in people in their forties creasingly hampered by the declining
A POSSIBLE STRUCTURE FOR ()(-CRYSTAL­
and older, who are most bothered by it ability of the lens to accommodate,
LIN. Robert C. Augusteyn and jane F.
on a sunny day or when they have to probably because of gradual changes Koretz in FEES Letters, Vol. 2 2 2, No. 1,
contend with the headlights of on­ in the geometry of the lens-zonule­ pages 1-5; September, 1987.
coming traffic while driving at night. ciliary-muscle complex. These proces-

SCIENTIFIC AMERICAN July 1988 99

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