Quarterly Journal of Experimental Physiology (1985) 70, 249-269

Printed in Great Britain

ASSESSMENT OF THE OPTICAL CONTRIBUTIONS TO

THE AGE-RELATED DETERIORATION IN VISION
J. D. MORRISON AND C. McGRATH
Institute of Physiology, University of Glasgow, Glasgow, G12 8QQ
(RECEIVED FOR PUBLICATION 14 SEPTEMBER 1984)

SUMMARY

Contrast sensitivities were measured in human observers of different ages in response to

sinusoidal grating patterns generated by laser interferometer and cathode ray tube (c.r.t.). The
former method bypasses the effects of the optical media to assess directly the contrast sensitivity
of the retina/visual system, while the over-all assessment including optical media is made to
the c.r.t. display. The ratio of the two sets of values gives the contrast ratio of the optical media.
The latter assessment does not include the effects of wide-angle light scattering which were
measured by comparing the responses of eyes with a Perspex lens implant or without any lens
at all (i.e. without the major source of scattering) with natural eyes. With increasing age, c.r.t.
contrast sensitivities remained steady until the sixth decade when they declined while laser
interference fringe contrast sensitivities declined continuously, apart from some abnormally low
results for teenage observers. In contrast, the optical contrast did not vary systematically with
age nor did its rate of change with spatial frequency vary. The latter was always less than the
rate for laser interference fringe contrast sensitivities indicating that the retina/visual system
always sets the limit to visual resolution. Wide-angle light scattering was not found to contribute significantly to the changes in ageing nor did reduced retinal illumination, as
reproduced by reduced pupil diameter or viewing through a neutral density filter. We therefore
conclude that the age-related deterioration is primarily caused by changes within the central
nervous system rather than the optical media, the transmission quality of which remains
unaffected.
INTRODUCTION

Advancing age inevitably causes changes in vision even should disease be avoided. In old
people, pupil diameter is reduced (senile miosis) while the power of focus (accommodation)
decreases gradually due to a combination of the decline of the elasticity modulus of the
lens capsule and the increase in elasticity modulus of the lens substance (Fisher, 1973). The
lens is unusual in that it continues to grow throughout life due to the addition of new
lenticular cells to its cortex (Niesel, 1982; Tripathi & Tripathi, 1983). From birth to
adolescence, the axial length of the eye increases and is followed by a compensatory change
in the refractive power of the eye (Sorsby, Benjamin, Sheridan, Stone & Leary, 1961). Then,
the eye progressively becomes more far-sighted (hypermetropia) up to about 64 years after
which it tends towards short-sightedness (myopia), partly as a result of the formation of
senile cataract in the lens (Slataper, 1950). The proportion of water insoluble protein within
the lens increases with age (Harding & Dilley, 1976) though this need not be deleterious
unless there is an associated change in refractive index. In contrast, the increase in molecular
diameter which occurs in ac-crystallin aggregates (Hoenders & Bloemendal, 1983) may be
of consequence through an effect on wide-angle scattering of incident light within the eye.
As the lens matures, its optical density increases and the resultant increase in the absorption
of blue light leads to the characteristic yellow colour of the ageing lens (Said & Weale, 1959).
In the retina, loss and displacement of photoreceptor cell nuclei occur (Gartner &

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250

J. D. MORRISON AND C. McGRATH

Henkind, 1981) together with convolution of the outer portion of rod photoreceptor outer
segments (Marshall, Grindle, Ansell & Borwein, 1979). Morphological changes in cone
photoreceptor cells which subserve high acuity vision do not appear to have been studied
except for dark-adaptation thresholds of cones which rise threefold from ages 20 to 80 years
(Gunkel & Gouras, 1963). The pigment epithelium also undergoes marked changes which
have been correlated with the corresponding patients' visual acuity before death (Sarks,
1976). Basal linear deposits located under the pigment epithelium constituted the most
consistent indicator of ageing. In severe cases involving a marked fall in visual acuity, drusen
had accumulated under the pigment epithelium which also showed signs of degeneration
and formation of new blood vessels. When such changes arise in the fovea, they constitute
senile macular degeneration. Nor are higher centres immune from ageing. A profound loss
of neurones occurs from 46 millions per gram wet tissue at age 20 years to 24 millions at
80 years in the foveal representation of the visual cortex (Devaney & Johnson, 1980).
Furthermore, many of those neurones still present may have lost their apical and basal
dendrites as shown by the Golgi studies of Scheibel, Tomiyasu & Scheibel (1977) on the
Betz cell of the human motor cortex.
Functional assessments of the effects of ageing on vision show that visual acuity remains
steady up to about 50 years and declines thereafter (Slataper, 1950; Weale, 1975; McGrath
& Morrison, 1981). A more complete assessment of ageing effects comes from contrast
sensitivity measurements. With sinusoidal gratings printed on card, Arden (1978) reported
an absence of change with age though a decline was subsequently described (Arden &
Jacobson, 1978; Skalka, 1980). With grating patterns generated by cathode ray tube (c.r.t.)
which allow higher spatial frequencies to be studied, contrast sensitivity was found to decline
significantly with age (Derefeldt, Lennerstrand & Lund, 1979; McGrath & Morrison, 1981;
Kline, Schieber, Abusamra & Coyne, 1983; Owsley, Sekuler & Siemsen, 1983). The earliest
study which used this method (Arundale, 1978) included only five observers (aged between
45 and 66 years) as its sample of old observers while the study of Sekuler & Hutman (1980)
in which they omitted to correct the refraction of their older observers seems to have been
superseded by their later report (Owsley et al. 1983). The general view seems that at
moderate to high spatial frequencies (3-40 cycles/deg), contrast sensitivity declines with age
(Derefeldt et al. 1979; McGrath & Morrison, 1981; Kline et al. 1983; Owsley et al. 1983)
while at low spatial frequencies the consensus seems to be that little change occurs though
a decrease may be recorded when a subjective assessment of contrast threshold is used rather
than a forced choice procedure (Vaegan & Halliday, 1982).
The above methods provide an over-all assessment of visual performance without
specifying where within the visual system the changes arise. An assessment of the
contribution of the optical media to the ageing process is, however, practicable with a
psychophysical method which involves the comparison of the contrast sensitivities to c.r.t.
grating patterns which are refracted by the eye and those to interference fringes observed
in the Maxwellian view which are not refracted (Campbell & Green, 1965). The ratio of
these contrast sensitivities which represents the optical transmission quality of the eye may
then be related to the neural transmission quality obtained in response to the interference
fringes. The method, however, does not assess the possible effects of wide-angle light
scattering which would affect both types of stimulus displays. An important source of light
scattering appears to be the lens (Allen & Vos, 1967). Therefore, measurements were made
on observers with either an aphakic eye, i.e. minus its lens or one containing a replacement
Perspex lens implant, in which wide-angle light scattering would be expected to be minimal,
as well as on normal observers.

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OPTICAL CHANGES IN AGEING

251

Preliminary communications of some of the results of this study have been made
(McGrath & Morrison, 1983; Morrison & McGrath, 1983).
METHODS

Contrast sensitivity was measured in response to a vertical sinusoidal grating display generated by
both c.r.t. and laser interferometer. Before the experiments, visual acuity was measured with the
Snellen test. Additional lenses were worn if necessary for optimum acuity when viewing the c.r.t.
display but were not necessary for the laser display. Our normal observers reported no visual
abnormalities and often had Snellen acuity of 6/5 except in some older observers where it was 6/6
or 6/9. In many of these older observers and in all eyes in which an operation had been performed,
an ophthalmologist's report was to hand. For both stimulus displays, the observer viewed monocularly
with the same eye.
C.r.t. display
A uniform green raster was produced on a Telequipment DM53S oscilloscope (P31 phosphor, peak
wave-length 520 nm), with time base at 1 ms/div, by feeding a 12 V peak-to-peak triangular wave of
frequency 220 kHz into both vertical amplifiers the beams of which had been superimposed. The
grating pattern was produced by modulation of the raster by a sine wave of variable voltage and
frequency fed into the z modulation of the oscilloscope. The display was viewed through a window
which excluded the distortions at the margins of the c.r.t. This window subtended 15 deg when viewed
from 3 m and was surrounded by green card. The luminance of the raster was 0 4 foot-lambert
(1-4 cd/M2) measured with a Pentax spotmeter V, being equivalent to about 3 83 log units above foveal
threshold measured in darkness. The z modulation signal was obtained from a sine-wave oscillator
the output of which could be amplified to a maximum of 70 V peak-to-peak by a control unit. The
amplitude of this sine wave was varied by adjustment of either the oscillator output or the ten-turn
potentiometer which attenuated the output of the control unit, and was monitored on an oscilloscope.
The relation between z modulation voltage and contrast was determined by the psychophysical method
described by Campbell & Green (1965). One cycle of the grating display located in a narrow window
was viewed at 1 m. A calibrated neutral density filter was placed in the window over the brighter half
of the cycle and the z modulation voltage was increased until the two half-cycles were judged to be
of equal luminance. At this point, the z-modulation voltage was numerically equivalent to the
transmissivity of the neutral density filter. This was repeated at cycle widths corresponding to 5, 10
and 20 cycles/deg at 3 m by two observers. Contrast was linearly related to z modulation voltage with
a slope of 0 022 contrast units/V peak-to-peak up to 25 V, attaining a maximum contrast of 0 85 at
70 V. This calibration in fact remained stable over several years. An objective method employing a
photodiode with amplifier was also used but was least reliable at high contrasts just when the
psychophysical method was most reliable, due to the low luminance of the dark half-cycles.
Laser interference fringe display
This method was basically that of Westheimer (1960) and Campbell & Green (1965) in which
sinusoidal fringes generated by the interference of two lasers beams consisting of polarized
monochromatic light are observed in the Maxwellian view (Fig. 1). Instead of the incoming rays of
light being focused by the optical media of the eye, they are focused by a microscope objective onto
a plane passing through the posterior nodal point, located approximately 17 mm in front of the retina,
through which rays of light pass unrefracted. Thus, the effects of the standard geometrical aberrations
associated with any optical system, but not those of wide-angle scattering of light by small particles,
are avoided. Vertically orientated interference fringes of sinusoidal luminance profile and of variable
spatial frequency and contrast were generated by a Mach-Zehnder interferometer as described in Born
& Wolf (1980) and shown in Fig. 2. Light from a He-Ne laser (wave-length 632 nm) was attenuated
by 2 0 log units, spatially filtered by a Scie-Mechs spatial filter s.f. in which was mounted a Vickers
x 10 microplan objective focusing the laser beam onto a 50 ,tm pinhole, and collimated by a 15 cm
focal length lens L. The resultant beam was divided by a 35 mm cube beam-splitter b.s. 1 (H. W. English
Ltd) and the two beams reflected from A/10 and A/20 surface aluminized mirrors Ml and M2
(Ealing-Beck Ltd) onto a 3 in. plate beam-splitter b.s.2 (N.V. Optische Industrie). The two emerging
beams were adjusted to equal intensity and reflected from another A/20 surface objective the field of

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252

J. D. MORRISON AND C. McGRATH

Beam2I

Interferenc
Eye

Posterior
nodal

Objective lens

plane
Fig. 1. Schematic representation of the formation of laser interference fringes on the retina.
N.d. S.f.

B.s. I
Ml

.,~~~~~~~Nd
B.s.22N.d.

M3

Rotation

IMovement

C.P.:

xLooX2

LL
c >bx 1()(

Image plane
Fig. 2. Schematic diagram of laser interferometer: further details are given in the Methods. B.s., beam splitter;
c.p., calcite polariser; I, interference filter; L, lens; M, front-silvered mirror; n.d., neutral density filter;
P. Polaroid; s.f., spatial filter.

view of which was limited to 7 deg by an aperture. The plate beam-splitter b.s.2 also transmitted
another pair of beams (of unequal intensity) which was viewed through a combination of a concave
simple lens and a Vickers x 100 microplan phase-objective to obtain a magnified view of the
interference fringes. The spatial frequency of the fringes, which is directly related to the path-length
difference between the two laser beams, was set by translation of mirror M2 by a 63 mm Microcontrole
stage. This carried a rotation stage (A. J. Beck Ltd) the purpose of which was to bring the beam back
into alignment after translation. The range of spatial frequencies employed was 8-55 cycles/deg
though the apparatus was capable of a much wider range from very low up to the diffraction limit.
The spatial frequency of the interference fringes was obtained by counting the number of cycles within
the phase ring of the x 100 objective, and multiplying by the correction factor 0-79. The latter value
had been obtained by relating the number of cycles in the x 100 objective to those in the x 2 objective
obtained by photography over the range of spatial frequencies studied.
Variable contrast of the test beam was obtained by superimposing the laser beams on a diffuse beam
of equal intensity but polarized at 90 deg. The latter was generated from a tungsten source (Vickers
microscope lamp) and passed through a heat-absorbing filter, a sheet of high extinction Polaroid film

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OPTICAL CHANGES IN AGEING

253

P (H. W. English Ltd) and an interference filter I with peak transmission 630 nm and band width
5 nm (Barr & Stroud Ltd) before combining with the laser beams at the 35 mm cube beam-splitter
b.s.3. The laser beams and diffuse beam were passed through a 15 mm aperture Glan-Thompson
calcite polarizer c.p. with nominal extinction 10-5 (J. Phillips Ltd) mounted in a Beck rotation stage
whose precision of rotation was to within 5-10 minutes of arc. Thus, by rotation of the calcite polarizer
from 0 to 90 deg, the contrast of the observed interference fringes was changed from 0 to 1 0, according
to the sine-squared law (Westheimer, 1960). The intensity of the combined beams which remained
constant upon rotation of the calcite polarizer was determined psychophysically to be about 2 55 log
units above foveal threshold and allowed prolonged viewing without discomfort.

Experimental procedure
C.r.t. display. Some observers had taken part in the earlier study 3 years ago (McGrath & Morrison,
1981) and thus had seen the c.r.t. grating display previously. The measurements of contrast sensitivity
of the c.r.t. display were made from the distance of 3 m under subdued illumination of about 2 mcd.
The procedure was for the observer to say when the grating pattern was just visible and no more,
as the experimenter gradually increased the contrast from 0 (i.e. uniform screen). Account was taken
of possible differences in reaction times between individuals. After practice runs at three to five spatial
frequencies, six to ten measurements of contrast threshold were made for ten to fifteen different spatial
frequencies in the range 8-40 cycles/deg, arranged in pseudorandom order involving passing up and
down the spatial frequency range twice. Sometimes, on changing the spatial frequency, the first few
contrast thresholds showed a downward trend, indicating that the observer was becoming familiar
with that spatial frequency; these results were disregarded. A steady upward trend, sometimes
apparent at the end of a recording session, indicated fatigue and a rest was called for. Each c.r.t.
recording session lasted 1-21 h.
Laser display. For all observers, except the authors, viewing the interference fringes was a new
experience. A similar procedure was adopted with measurements in response to the laser interference
fringes, though the need for familiarization was appreciably greater. Again six to ten contrast
thresholds were recorded at each of ten to fifteen spatial frequencies over a period of 2-31 h. Problems
similar to those outlined in the previous section were also encountered. Sometimes, the laser
interferometer measurements were done first and the c.r.t. measurements second, and vice versa. The
two experiments were usually done on the same day. Particular attention was paid throughout to the
comfort of the observer and breaks were made regularly to guard against fatigue. The most usual
report was that the observer did not feel tired during the experiment but did once it was all over.
Generally, the older observers performed very well. Sometimes, difficulty in maintaining attention
was evident in some teenage observers.

RESULTS

Observers
The main normal group consisted of forty-five observers in whom contrast sensitivity was
measured in one eye. In each person, wearing spectacles if necessary, Snellen acuity was
normal. This group also contained four older observers whose eyes were described as having
minimal ageing changes and three observers for whom further description is necessary. An
80-year-old woman with Snellen acuity 6/9 possessed a cataract which was described as 'very
mild indeed' while a 78-year-old man with the same acuity had some corneal scarring which
interfered with the laser interference fringes but evidently not with the c.r.t. display. Finally,
a 15-year-old girl with severe but not complete red-green colour blindness (protanomalous
trichromat, with the Pickford-Nicolson anomaloscope) was also included since her Snellen
acuity was 6/5 and photopic thresholds to the c.r.t. and laser displays were normal.
Contrast sensitivity measurements were made in another group of eight observers who
had undergone an operation in one eye. One had had an intracapsular extraction involving
the removal of the lens complete with capsule from one eye and was thus aphakic. Six had

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J. D. MORRISON AND C. McGRATH

254

W.H.B. 61 years

1-0
-0*5

C.M. 24 years

1 0

-05
m

0.3

-03 O
0
-0-2

-0 005

-0-2

-0008

0.1

-01
0~~~~~~~~~~~~~~

1000
8

00

0.

0-05

0-05

-P

.*C.r.t. display

100

Laser display

I
m
r

m =

-0-048

r-098
%8e

O0-51
->1

0-097

10

00

m =-0-060
r

-099

01

r= -0986

m=-005.18

-1.0

J.D.E. 86 years

1-0

J.B. de V.W. 75 years

-0-5

-0-5

Cu

m-0020

n +0004

~~~~~~~~-0-2
-

1000

0-1

0-02

'01

0.

0-005

0-005

100

VC

o 1st session
A2nd session

01
-0

=-0-032

-0\92
96r =

r =

-0-064
-0-99

10

10

=-0-96

-0-06

t
lines
(m
frequency
(cycles/deg)C
C
~~~~~Spatial
Fig. 3. Contrast sensitivities for c.r.t. and laser-generated displays for different ages, showing best-fit regression
r

0 10

m =

20

30

40

60

10

20

30

40

50

60

lines (m is slope and r is correlation coefficient). In D, the regression line for laser display was calculated from
the mean values at each spatial frequency to obviate the skewed distribution of points. Contrast ratios were
calculated from the best-fit regression lines.

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OPTICAL CHANGES IN AGEING

255

had the lens removed but leaving the posterior capsular membrane intact (extracapsular
extraction) and a Pearce Perspex lens implant placed into the posterior chamber. This
procedure had been done in both eyes of the eighth observer. Of the seven of these observers
in whom the fellow eye retained the natural crystalline lens, four of these were normal and
were placed within the group of forty-five observers. Of the remainder, a 46-year-old woman
with Snellen acuity 6/9 had a mild cataract which could be bypassed by the laser fringes,
while in a 61-year-old man with Snellen acuity 6/18, the cataract was so severe that it could
not be satisfactorily bypassed. Thirdly, a 72-year-old woman with Snellen acuity 6/12 had
possible macular damage. The age distribution of the observers, with the number of
normal/abnormal eyes in parentheses, was: 15-19 (5/0), 20-29 (10/0), 30-39 (5/0), 40-49
(3/2), 50-59 (5/2), 60-69 (5/4), 70-79 (10/3), 80-89 (2/1).
The contrast sensitivity curve
Since this was obtained with stationary grating patterns of spatial frequencies 850 cycles/deg, it may be taken as representing an assessment of foveal function. The usual
relation was that logarithm contrast sensitivity was inversely linearly related to spatial
frequency, with the correlation coefficient usually much better than r = -0 90 (e.g. Fig.
3 A). In eight observers uniformly distributed over the age range, a better fit was obtained
between logarithm contrast sensitivity and logarithm spatial frequency. In the analysis of
these eight results, the rates of decline of contrast sensitivity with age were treated separately
from the main body of data.
The normal scatter of points at any given spatial frequency was usually within 0-2 log
units and did not vary systematically with age (Fig. 3 A-D). Measurements were also quite
reproducible when remeasured for both c.r.t. grating patterns and laser interference fringes.
This is illustrated in Fig. 3 C in which close agreement between contrast sensitivities to laser
interference fringes measured 1 month apart was obtained in a 75-year-old man (neither
the slopes nor intercepts of the regression lines for the two sets of data were significantly
different). Systematic analysis of contrast sensitivities for age decades (see later) showed no
difference between male and female observers in any given age range.
In a young observer, contrast sensitivities to the c.r.t. display were always less than to
the laser fringes due to degradation of image contrast by the optical media (Fig. 3 A). The
ratio of c.r.t. and laser fringe contrast sensitivities (the difference on a logarithmic scale)
is the contrast ratio determined psychophysically and, when expressed over the range of
spatial frequencies studied, represents the quality of the optical transmission. The more level
this is, i.e. all spatial frequencies are passed equally well, the more nearly neutral is the
transmission quality. This may then be compared with the neural transmission data shown
by contrast sensitivities to laser fringes. Invariably, the optical transmission which was
indeed almost level declined at a much slower rate with spatial frequency than neural
transmission which itself set the limit to visual resolution (Fig. 3 A). This confirms the results
of Campbell & Green (1965) whose experiments were performed only on themselves.

Optical and neural transmission in ageing

With increasing age, contrast sensitivity to both c.r.t. and laser-generated displays
decreased as illustrated in Fig. 3 A-D which shows typical results for observers of different
ages. The optical transmission calculated from the regression lines for c.r.t. and laser
contrast sensitivities, however, showed no such trend with age. Comparison of the rate of
decline of optical and neural transmission shows that neural transmission quality declined
faster with spatial frequency and formed the limit to resolution in even our oldest observers.
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J. D. MORRISON AND C. McGRATH

256

-0-100-

0 Laser fringes (natural eye)

fringes (implant or aphakic eye)
* Contrast ratio (natural eye)
A Contrast ratio (implant or aphakic eye)

A Laser

0r

-0-50
o

^~h0~~

BQO-

S1

I--:

00
-0---0
--20

r~ +0-020
~

-0

=-0-004
5'

0-

= -006

L0

-0004

co

--I

0+0020

~B

Cl

m~~~~~

*.A

-0-100

20

CBr.t. display
40 ~0

80

10004

00~

---

-0-050

r-0*150

0*000~
0

20

40

60

80

100

Age (years)

Fig. 4. Rates of decline of A, contrast sensitivities to laser interference fringes together with contrast ratios and
B, contrast sensitivities to c.r.t. displays. Dashed lines show + 1 S.D. of y values. Points for operated eyes
(triangles) were not included in the regression analysis.

The collected data for all observers are shown in Fig. 4A. In no case did the rate of fall
in optical transmission quality ever exceed that ofneural transmission quality. Nor was there
any significant change in the rate constant for neural transmission or optical transmission
with age (Fig. 4A). The rate constants for the c.r.t. display contrast sensitivities also
remained unaffected by age (Fig. 4B). The difference between the two sets of points in
Fig. 4A is all the more notable when it is borne in mind that these rate constants are the
indices of exponential values. Sometimes, the contrast ratio had a small positive slope
against spatial frequency as shown in Fig. 3 D. In the eight observers whose results showed a
positive slope, the latter differed significantly from a constant level value (P < 005) in only
three cases. This serves to emphasize the high quality of the optical transmission in our
observers. Visual resolution was limited by optical transmission quality only in abnormal

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257

OPTICAL CHANGES IN AGEING

Table 1. Mean correlation coefficients over 10-40 cycles/deg

Measurement (AM)

M vs. age

log M vs. age

M vs. log age

log M vs. log age

C.s.-c.r.t. display
C.s.-laser display
Contrast ratio
Contrast ratio
(values 0 1-0 5)

-0 58
-0 40
+0 044
+0 064

-0 65
-0 54
+0 055
+0 069

-0 50
-0-42
+0 055
+0 089

-0 57
-0 53
+0 073
+0 054

C.s. = contrast sensitivity.

Table 2. Correlation coefficients for logarithm measurement against age

Spatial frequency
(cycles/deg)
10
12
15
20
25
30
35
40
n=

C.s.-c.r.t

display
-0 62***
-0 66***
-0 65***
-0 70***
-0-72***
-0 63***
-0 61***

42

C.s.-laser
display
-0.63***
-0.57***
-0 56***
-0 62***
-0 62***
-0 42**
-0048**
-0 39*
37

C.s. = contrast sensitivity.

***P < 0-001; **P < 0-01; *P < 0 02;

Contrast ratio

Contrast ratio
(values 0 1-0 5)

+0 14t

+0 18t

+0 06t
+0 03t
+0 04t
0O Ot

+0-06t
+0 OSt
+ 005t
0OOt

37

30

tnot significant.

circumstances which, in the present study, were as follows: the eye with the moderately
severe cataract, but not that containing only a mild cataract; in one natural eye with
suspected pathological changes; in an implanted eye with reported haziness of the posterior
capsular membrane, and in two implanted eyes of normal appearance but of several years
standing. By contrast, all implants of recent standing had normal optical quality (see later).
The reason for the poor optical quality of the older implants remains to be determined.
A notable result was that the optical quality in eyes with reported ageing changes - yellowing
of the lens (nuclear sclerosis) and minimal cortical lens opacities - was nevertheless very
good, as if the observed defects were having no marked effect on optical quality.
Trends with increasing age
The most appropriate relation between contrast sensitivity and age or contrast ratio and
age was investigated by correlation of the possible combinations of linear or logarithmic
values at each of a range of spatial frequencies. The mean values of the correlation
coefficients at the specified spatial frequencies are compared in Table 1. The best correlation
was between logarithm contrast sensitivity against age, though the improvement over other
combinations was never significant (P > 025). Of particular note was that no combination
of optical contrast ratio and age gave a statistically significant correlation coefficient.
Generally, the mean result was representative of the individual correlation coefficients at
each spatial frequency. The full list of correlation coefficients for the logarithmic-linear

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J. D. MORRISON AND C. McGRATH

258

C.r.t. display

100-

*^

Spatial frequency 30 cycles/deg

C.r.t. display

Spatial frequency 10 cycles/deg

.oMean values for

;^v

v
age decades
'.>.^

at *

r-0 62

10
0~~~~~~~~~r-06

.@

C
1 0000-

Laser display

Laser display

10000

2.

"Io
>01;

o Natural eye

1000-

00
0

'hoNatural eye (extrapolated)

E_

*O

0XS, O

A~

iImplant or aphakic eye

o,

000

Mean values

_
A

-0-63

0E0 0F

100

A0

oc
0

e=0_0 A0_4

*0

0
10

|
3Contrast
e 251
ratio

r = 0 140

-4
=

==+ =
Contrast ratio

1-0

o-

0A

0.3

~ "'

0.2

r= 0.14
*~~~~~~~~~~~~
A

01

.0

0mln rapa

~
~_

0*05

*Natural eye

AImplant or aphakic eye

0I0

4 0

80

Mean values
100

Age (years)
Fig. 5. Contrast sensitivities for c.r.t. display, laser interference fringes and contrast ratios at 10 and 30 cycles/deg,
with regression lines and + I S.D. Mean values for each decade are also shown (filled or open circles). Two
extrapolated values are present in D as open squares. Points for operated eyes (triangles) were not included
analysis.
in the regression
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OPTICAL CHANGES IN AGEING

259

combination is shown in Table 2 and the graphs for 10 cycles/deg and 30 cycles/deg are
shown in Fig. 5A-F. The age-related changes in the measurements are as follows.

(a) C.r.t. grating display. The contrast sensitivity values to c.r.t. grating patterns showed
a statistically significant downward trend with age for spatial frequencies 10-35 cycles/deg
(Fig. 5A and B and Table 2, column 2). Although considerable scatter of the results was
present throughout, exemplified by values of r2 rarely in excess of 5000, which indicates that
most of the variation is due to scatter of the data rather than the regression itself, the
probability that the decline with age was due to chance was generally less than 01 00 . When
the data were analysed by age group in an attempt to minimize scatter within that age group,
the same downward trend was apparent at each of the seven spatial frequencies studied.
Contrast sensitivity, however, remained relatively constant up to the sixth decade, and only
thereafter did it decline with age (Figs. 5 A, B and 6 A). A similar time course was observed
when the highest frequency detectable was plotted against age (Fig. 7 A) confirming our
earlier (McGrath & Morrison, 1981) observation. It was interesting to note how the Snellen
acuity of older observers tended to be rather poorer than would be expected from their
high-frequency cut-offs, e.g. in the 80-year-old woman, the highest spatial frequency
observed was 30 cycles/deg while Snellen acuity was only 6/9. The general validity of the
above conclusions was confirmed by changing the age groups to 15-19, 20-29 etc. and 15-24,
25-34 etc. to vitiate possible sampling biases.

(b) Neural transmission quality. The contrast sensitivities to laser fringes also declined
steadily with increasing age (Fig. 5 C and D), and again scatter of the points was present.
Fig. 5 C and D also shows values for observers with an intra-ocular implant or aphakic eye
which were not included in the regression analysis and will be commented upon later. Most
noticeable were the low values for the second decade, which in some cases were similar to
their corresponding values for the c.r.t. display. Since this implies that optical transmission
was nearly 100% which seems unlikely, these contrast sensitivity measurements for the
second decade were considered to be abnormally low and omitted from the regression
analysis. From the third decade onwards, there occurred a consistently downward trend of
logarithm contrast sensitivity with age which was statistically significant at each of the eight
spatial frequencies in the range 10-40 cycles/deg (Table 2, column 3). Analysis of the results
in terms of age decades showed a continuous decline in contrast sensitivity with age from
the third decade onwards (Figs. 5 C, D and 6 B) in contrast to the results for the
c.r.t.-generated display which were constant up to the sixth decade. When the age bands
were changed, the only difference was that the initial rise in contrast sensitivity between
second and third decades was averaged out and a linear fall occurred from the 15-24 year
group and older. The highest spatial frequency observed (taken in practice at a contrast
sensitivity value of 0 4 log units) also declined linearly with age (Fig. 7B).
(c) Optical transmission quality. The contrast ratio showed an absence of any trend with
age at the five spatial frequencies in the range 10-30 cycles/deg (Fig. 5 E and F and Table
2, column 4). Nor was there any trend in the absolute value of the contrast ratio with age
(Figs. S E, F and 6 C). A noticeable feature of the grouped data for contrast ratio plotted
against spatial frequency for each decade is the near flatness of the relation, indicative of
good optical quality (Fig. 6 C). The initial fall in contrast ratio from the second to third
decade arises as a result of the abnormally low laser fringe contrast sensitivity values in
the second decade (Fig. 5E and F). After excluding the values for the second decade,

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J. D. MORRISON AND C. McGRATH

260
A

100

C.r.t. display

10

B
N

<

me\*1000
*
X

Laser display

i&\ @

Age groups (years)

* 15-20

\\\\d~

10

10

30
1-0
Contrast ratio
20

0-5
o

0-3

40

50

31-40
A~~~~~~~~~~~
41-50

1
100

51-60

61-70

0.1

10

20

30

71-80

-x

0-2

21-30

40

50

10

10

20

30

40

50

Spatial frequency (cycles/deg)

and
contrast ratios for each age decade for A, c.r.t. display; B, laser
Fig. 6. Mean values of contrast sensitivities
display and C, contrast ratios. In A, only three regression lines are displayed due to superimposition of data.

contrast ratio values are contained within the range 0 1-0-5. Even when the regression
analysis was repeated on points within this range of contrast ratios, there was still an absence
of any correlation with age (Table 2, column 5).

Assessment of the effects of wide-angle light scattering

This was not assessed directly since both c.r.t. and laser generated stimuli would be
affected by wide-angle light scattering caused by small particles within the optical media.
An indirect method which involved the comparison of contrast sensitivities to laser fringes
for eyes containing a natural lens and those either without a lens or with a Perspex lens
implant was used. In the aphakic and implanted eyes, wide-angle light scattering would be
expected to be minimal due to either the absence of the natural lens or the substitution of
the implanted Perspex lens with its regular molecular structure. The observer with the
aphakic eye wore a contact lens during the measurements. No correction was made to take
account of a possible change in magnification which might be expected to be an increase

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261

OPTICAL CHANGES IN AGEING

Laser display

C.r.t. display

A
60

~~~~~~~~~~~~0

0.. 0

r
we

'~~
40

+0014
0.
.e--...
=

40..

*
.

o0

50+

r--9-

40

60

80

00

100

r-

Implant or
::

20

0..

.
=
0~~~~~~~~~~~~~~~~~~~~~~~~~

'D

20

sn~~~~~~~~~~~~~~~~ahakic
eye

40

60

80

100

Age years)
Fig. 7. Highest spatial frequency detectable and age, showing regression line and + I S.D. A, c.r.t. display, two
components are shown: 15-50 years and 50+ years (dotted lines). B, laser display showing values at, or
extrapolated to, 0-4 log units contrast sensitivity. Points for operated eyes (triangles) were not included in the
regression analysis.

of about 12% if the observer had been emmetropic before the operation (Ogle, Burian &
Bannon, 1958). However, this observer had normal binocular vision which suggests a
magnification difference of considerably less than 12%0. In observers with the Perspex lens
implant, image size is virtually normal.
When contrast sensitivities to laser fringes for the aphakic and natural eye of the same
observer were compared, the regression lines overlay one another (Fig. 8 A). The slope for
the neural transmission line was somewhat steeper in the natural eye than in the aphakic
eye, leading to a corresponding difference in the optical qualities. In the natural eye, optical
transmission increased with spatial frequency, being significantly different from zero slope
(P < 0-001). This particular observer, a woman of 60 years, was notable in having
particularly high contrast sensitivities for her age.
In five out of the six observers with a natural and implanted eye, laser fringe contrast
sensitivities were always appreciably higher in the normal eye than in the implanted eye
(e.g. Fig. 8 B). This occurred even despite two of these observers possessing a mild cataract
in the natural eye. In one of these two, an 80-year-old woman, contrast sensitivity did drop
abruptly to less than that in the implanted eye above 25 cycles/deg, but this was attributed
to the effects of the cataract in that eye. In the sixth observer, the natural eye had such a
severe cataract that the perception oflaser fringes was adversely affected. These comparisons
depend, of course, on the assumption that in the one person the two eyes have similar laser
fringe contrast sensitivities. While our measurements show that this is likely in young
observers, it is not so probable in older observers where one eye may differ markedly from
the other. Therefore, a further comparison of the results was made between observers with
the operated eye and normal observers (Figs. 4A, 5 C, D and 7B). Contrast sensitivities
to laser fringes for aphakic or implant eyes fell within the spread of normal results (Fig.
SC and D), as did the contrast ratios (Fig. S E and F). The slope of the optical transmission

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J. D. MORRISON AND C. McGRATH

262
A

.-1 1.0- 0 5

E.E.C. 60 years

-03
-02

10000

[0.05

C)--0

- Natural eye
A

100

-01

----Aphakic eye

0-03
-028

"
8

10

------------------

-1

T.T. 75 years

"AWL

- Natural eye
Implant eye
> "..sO o Laser display natural eye
A Laser display operated eye
* C.r.t. display natural eye
C.r.t. display operated eye

A'.~~~~~~~~~~~~~~~~

1.*

0 0
1

20

30

40

50

10

20

30

40

50

Spatial frequency (cycles/deg)

Fig. 8. Contrast sensitivities and contrast ratios for observers with A, an aphakic: and a natural eye and B. an eye
containing a Perspex lens implant and a natural eye.

line was within the normal range while that for neural transmission tended to be somewhat
lower than normal (Fig. 4A). It should be noted that this is opposite to what might be
expected if the operated eye were displaying less light scattering than the natural eye. The
spatial frequency observable at a contrast sensitivity of 0 4 log units was within the normal
spread of results for these observers (Fig. 7 B). Therefore, the conclusion is drawn that there
is no evidence to indicate that an increase in wide-angle light scattering, as assessed by our
methods, occurred in the ageing eye.

reduced pupil diameter and luminance

Effects oA
Measurements of contrast sensitivity to the c.r.t. display were made in a group of young
observers viewing through an artificial pupil or neutral density filter to investigate whether
the decrease observed in ageing arose from reduced retinal illumination, resulting from senile
miosis or reduced lenticular transmission. In Fig. 9, the mean values at 10-35 cycles/deg
for the 20 s and 70 s age groups are shown by dashed lines. No difference was observed
between contrast sensitivities for viewing with a natural 8 mm pupil or through an artificial
3 mm pupil. In general, the mean values reflected the results for each individual. Both sets
of results were very similar to the control values for that age group (Fig. 9A).
The effects of viewing through a 0-5 log unit neutral density filter was either to improve
slightly or decrease slightly or leave unchanged the contrast sensitivities in five young
observers with the result that the mean values were very similar to those for full illumination,
and both were similar to the control values for that age group (Fig. 9B).
Reduced illumination caused by a reduced pupil diameter or neutral density filter
therefore had no discernible effect on contrast sensitivity and cannot be invoked to account
for the deterioration which arises as a result of ageing.
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OPTICAL CHANGES IN AGEING

A
100

C.r.t. display

C.r.t. display

Natural (8 mm) pupil

3 mm artificial pupil

A Normal illumination
AA 0-5 log unit attenuation

Ike

10 S
a)

263

'_
~~~X
'x

..means

21-30 years

Ix--- means71-80 years

10

%~~~ A%~
1~~~~~~~~

0.

1-0

10

20

30

40

10

20

30

40

Spatial frequency (cycles/deg)

Fig. 9. Effects of A, reduced pupil diameter and B, reduced retinal illumination on contrast sensitivities to c.r.t.
display in young observers. For comparison, results for third and eight decades are shown (filled circles and
crosses respectively).
DISCUSSION

The results of the present study fall broadly into three areas:
(a) The fall in contrast sensitivities for c.r.t.-generated patterns both confirms the results
of earlier studies and extends them in that the fall did not commence until the sixth decade.
The decline in the slope of the contrast sensitivity-spatial frequency relation reported by
us previously was, however, not confirmed (Fig. 4B). We can only surmise that sampling
differences were responsible since, although the total number of observers was relatively
large, that of older observers was still quite small. The reason why the results for a small
number of observers were better described by a plot of logarithm contrast sensitivity against
logarithm spatial frequency is uncertain. A possible reason may be that as a result of the
observer's profession, experience had been gained in observing high spatial frequencies. In
some observers who may be reasonably expected to have this ability, a logarithmic-linear
plot nevertheless proved to be more appropriate.
(b) The decline in contrast sensitivities to laser interference fringes is in keeping with the
results of Campbell & Green (1965) in which the older investigator (40 years) had
consistently lower sensitivities than the younger investigator (27 years). Two recent studies
have, however, stated without documentation of their results, that no change in laser fringe
contrast sensitivity occurred with age (Dressler & Rassow, 1981; Kayazawa, Yamamoto
& Itoi, 1981). If their conclusion is taken at face value, the difference from the present results
may involve the anomalous results for teenage observers which might obscure an age-related
trend were they included indiscriminately in the analysis. Other workers have also described
an improvement in contrast sensitivity in adolescence though to c.r.t. grating patterns
(Derefeldt et al. 1979; Beazely, Illingworth, Jahn & Greer, 1980; McGrath & Morrison,
1981). Such anomalous results are likely to arise from a deficiency in attention or
inexperience in applying themselves to the task of visual discrimination rather than from
immaturity of the visual system itself.
The time course of the decline consisted of a linear fall in logarithm contrast sensitivity
with age, in contrast to the c.r.t. values which remained steady until the sixth decade.

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J. D. MORRISON AND C. MCGRATH

However, since considerable scatter is present in both sets of results, this difference must
be viewed with some caution. The decline is consistent with the linear increase in logarithm
threshold of vision which was described by Hinchcliff (1962) in his analysis of ageing
changes in the central nervous system, though it is at variance with the time course derived
by Weale (1975, 1982) from visual acuity data. The basis for the continuous decline from
the third decade onwards may possibly be in the fall in the numbers of retinal and cortical
neurones occurring over this period (Hinchcliff, 1962; Devaney & Johnson, 1980; Gartner
& Henkind, 1981).
(c) Optical transmission represented by the contrast ratio over 10-30 cycles/deg remained
essentially unchanged during ageing in terms of both the absolute value of the contrast ratio
and its rate of change with spatial frequency. Since the latter is always less than the rate
of change of neural transmission (Fig. 4 A), visual resolution will never be set by the optical
system, provided that retinal illumination does not become limiting. That the absolute value
of the contrast ratio remains invariant with age is surprising, given the well-known rise in
optical density of the lens during ageing (Said & Weale, 1959; Mellerio, 1971). Our
measurement of contrast ratio is, however, the most susceptible to criterion differences to
the c.r.t. and laser displays, which may have obscured any age-related trend.
Our results are consistent with the recent report of Weale (1983) that the resolving power
of excised lenses from cadavers of different ages remained invariant with age. Also, in
patients aged 50-80 years in whom a lens implant operation had been performed during
cataract surgery, thus eliminating the part of the optical system most susceptible to ageing,
Snellen acuity nevertheless declined with age (Dr J. L. Jay, personal communication). In
the present study, optical .transmission was usually of high quality, showing a slight decline
with increasing spatial frequency. The significance of a positive slope which occurred
occasionally (Figs. 3D and 8 A) is uncertain since the contrast ratio should decline with
spatial frequency regardless of the aberrations present (van Meeteren, 1974). A rising optical
quality is possible with defocusing (Campbell & Green, 1965) or by application of spots
of petroleum jelly to a contact lens to reproduce the effects of a cataract (Zuckerman, Miller,
Dyes & Keller, 1973), but only over a localized range of spatial frequencies before declining
again; neither situation is likely to be applicable to the present study.
Further evidence for the absence of an optical contribution to ageing comes from the
lack of effect of a reduced pupil diameter and reduced illumination on contrast sensitivity
to c.r.t.-generated displays. A reduction in pupil diameter would lead to reduced retinal
illumination and increased diffraction which would be expected to diminish contrast
sensitivity, and reduced geometrical aberrations which would be expected to improve
contrast sensitivity. The lack of effect actually observed also agrees with the study of Sekuler
& Hutman (1980) who found little loss in contrast sensitivity in one young observer viewing
through a 2 mm artificial pupil and with the observation of Woodhouse (1975) that at the
luminance of our c.r.t.-display - 0-4 foot-lambert - the resolution limit of grating patterns
of fixed contrast was the same for artificial pupils of 3 and 8 mm. Further, mydriatics have
been shown not to affect the dark-adaptation thresholds to different spectral bands of light
in older observers (Gunkel & Gouras, 1963) and neither mydriatics or miotics had an effect
on the scoring to Arden gratings in young observers (Singh, Cooper, Alder, Crawford, Terrel
& Constable, 1981). The absence of an effect in the present study of reduced retinal
illumination caused by viewing through a neutral density filter agrees with the observation
of Westheimer (1960) that an increase or decrease in the intensity of laser interference fringes
by 1 log unit was without effect on contrast sensitivity. Owsley et al. (1983) reported that
when young observers viewed through a 0 5 log unit neutral density filter, the differences

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265

OPTICAL CHANGES IN AGEING

between old and young observers were minimized. However, a significant difference
remained at their highest spatial frequency 16 cycles/deg which corresponds with the lower
part of our range of study (10 cycles/deg). Also, Sekuler & Hutman (1980) had previously
noted that viewing through a 2-0 log unit neutral density filter - a substantial reduction in
illumination - did not affect contrast sensitivity at low frequencies. Our conclusion is
therefore that a reduction in pupil diameter to 3 mm and a modest reduction in illumination
are without effect on contrast sensitivity determinations.
The present conclusion that optical quality remains of high quality during ageing and
is not responsible for the age-related loss of vision does depend critically upon the
experimental methods. Of great importance is the implicit assumption that the criteria for
determining contrast thresholds are the same for the c.r.t. display and laser interference
fringes. Certainly in viewing the c.r.t. display, one is aware of the sensation of viewing at
a distance in contrast to the apparent proximity of the laser display. Gstalder & Green
(1971) referred to the laser display as 'attention getting', but this is probably no more so
than viewing a glowing c.r.t. display in a darkened room. Providing time is allowed for
familiarization, the measurements with each technique do seem to be quite reproducible.
A more intractable problem is the comparability between observers who may choose
different criteria for contrast threshold. Apart from careful briefing, the total number of
observers was made as large as possible to obviate this problem. Another factor is the
difference in colour of the two displays. In young observers this is probably unimportant
since Campbell & Green (1965) found no differences in contrast sensitivities with orange
and blue filters in front of their c.r.t. display and our red-green colour-blind observer was
similar to other observers in her age group. In older observers, the differential loss in optical
transmission for green compared with red light amounts to only 0-08 to 0- 13 log units, from
20 to 60 years (Said & Weale, 1959; Mellerio, 1971), which is unlikely to significantly affect
the results given that 0 5 log unit attenuation is without affect on contrast sensitivity (Fig.
9 B). Also, in patients with a Perspex lens intra-ocular implant, which has a more even
spectral transmission than the natural lens (Jay, Gautam & Allan, 1982), their contrast
sensitivity and contrast ratio measurements fell within the normal range (Fig. 5 C-F).
An advantage of the subjective psychophysical method which we used to measure optical
quality over a physical method measuring the reflected image (Campbell & Gubisch, 1966)
is that, in addition to assessing neural transmission quality, it is a single-pass method not
complicated by back-scattered illumination. The laser interferometric method does not,
however, assess two important properties of the optical system, namely chromatic
aberration and wide-angle light scattering. Campbell & Gubisch (1966) assessed their effects
from a comparison of their own results with the earlier interferometric studies of Campbell
& Green (1965) and Arnulf & Dupuy (1960). Inspection of their optical transfer functions
for young eyes gives a value for the proportion of incident light dispersed by chromatic
aberration of about 3% at 20 cycles/deg. In ageing, however, chromatic aberration
decreases from -1.0 D to -0-25 D in the blue region of the spectrum and from + 0-25 D
to 0 in the red region (Millidot, 1976). The main cause of this reduction was attributed to
a rise in the refractive index of the vitreous humour so that the optical system acted like
a well-corrected achromatic doublet, rather than an effect of reduced pupil diameter
improving optical quality. While there is evidence of an increase in the density of the vitreous
humour above the age of 60 years (Wolf & Gardiner, 1965), it would nevertheless be difficult
to exclude an effect of pupil diameter since on the whole this does constrict in ageing, e.g.
McGrath & Morrison (1981). Regardless of the cause of the reduction in chromatic
aberration, its effect would be to underestimate the ageing changes reported in the present

study.
10

EPH

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70

266

J. D. MORRISON AND C. McGRATH

The proportion of wide-angle light scattering as calculated from the data of Campbell
& Gubisch (1966) is about 4 o incident illumination at 20 cycles/deg. This may be compared
with the value of 2% obtained from direct measurements of forward scattered X-rays in
the isolated lens of the calf (Delaye & Tardieu, 1983), while measurements in the intact cat
eye in vivo with a fibre-optic, light-guide system placed at the level of the retina (Robson
& Enroth-Cugell, 1978) indicate a carpet of scattered light over the retina of about 6 % peak
image intensity. Account must also be taken of light scattering by the retina itself which
at 20 cycles/deg may amount to a further reduction in contrast of 4000 (Ohzu & Enoch,
1972). Hence, appreciable degradation of image contrast occurs before detection by the
photoreceptors of the retina. The nature of the wide-angle light scattering remains uncertain
despite suggestions that it may be of the Rayleigh type in which the intensity of scattered
light is inversely proportional to A4 (Said & Weale, 1959; Mellerio, 1971). This, however,
is only an approximation of the more general Mie theory and depends upon the scattering
particles being very small (less than 1/20A) and with a refractive index (or more correctly,
a polarizability) approximately invariant with A (Kerker, 1969; Bohren & Huffman, 1983).
Experimental evidence, however, suggests this may not be the case. Scattering particles as
large as 100 nm have been described in the lens by Randall & Vaughan (1982) and the
intensity of scattered light may increase with A (Boynton, 1953; DeMott & Boynton, 1958).
In the general solution for the far field, the intensity of scattered light is actually directly
proportional to A2. Back scattering of incident light outwards from the eye also depends
upon particle size. Starting from a very small particle size, back scatter actually decreases
as particle size increases - the Mie effect. This will affect the interpretation of measurements
of back-scattered illumination taken to be an indicator of forward scattering in the ageing
lens (Allen & Vos, 1967; Sigelman, Trokel & Spector, 1974; Spector, Li & Sigelman, 1974).
The observation of Spector and co-workers that back-scattered illumination from the
nuclear region of the lens increased with age therefore suggests that particle size actually
decreases in ageing.
The origin of forward-scattered light appears to be the anterior cortex and nucleus of
the lens, with only a negligible contribution from the cornea (Allen & Vos, 1967) or vitreous
humour (Wolf & Gardiner, 1965). Psychophysical measurements of glare sensitivity
correlate well with the amount of scattered light obtained either directly in the same
observers through slit-lamp measurements (Wolf & Gardiner, 1965) or from direct physical
measurements in the cow eye (DeMott & Boynton, 1958). However, there was an absence
of correlation between measurements of back-scattered light, and the sensitivity for the
detection of the Landolt C, which was taken to indicate the absence of a role for scattered
light in the fall-off in vision with age (Allen & Vos, 1967), though the equivalence between
forward- and back-scattered light must remain in doubt. Our own studies on observers in
whom either the lens had been completely removed, or a Perspex implant substituted in
its place, also suggest an absence of a detectable change in wide-angle light scattering in
ageing. While it is never possible to rule out a deleterious effect of the operation itself, for
instance in terms of long-term changes in the implanted lens (Drews, 1982), the type of
operation involved - a posterior-chamber implant - is least likely to have complications like
corneal oedema or retinal detachment (Fagadau, Maumenee, Stark & Datiles, 1984).
Indeed no such complications which are easily detectable were observed in our patients.
It is concluded that the age-related deterioration in contrast sensitivity to medium and
high spatial frequencies observed in the present study is attributable to changes within the
retina/visual system rather than in the optical media. This complements existing evidence
that contrast sensitivities at low spatial frequencies remain unaffected with age (see

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OPTICAL CHANGES IN AGEING

267

Introduction) and lends support to the view of Weale (1975, 1982) that neural changes are
of prime importance in the loss of vision in ageing. Hence, provided retinal illumination
does not become limiting, the quality of the optical media remains uniformly good
throughout the lifespan and does not, in the absence of pathological changes, ever set the
limit to visual resolution.
To each observer who participated in the study, we give our thanks for his/her co-operation and
forebearance. We are especially grateful to Dr J. L. Jay for the opportunity to examine his patients
and for his comments on the manuscript. We acknowledge with thanks the financial support of the
Royal National Institute for the Blind and the Visual Research Trust and a research studentship to
C. McG. from the Medical Research Council.

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