J. Phyaiol. (1971), 213, pp.

157-174 157
With 9
text-figureM
Printed in Great Britain
THE ORIENTATION SPECIFICITY OF TWO
VISUAL AFTER-EFFECTS
By COLIN BLAKEMORE AND JACOB NACHMIAS*
From the Physiological Laboratory, University of Cambridge,
Cambridge CB2 3EG
(Received 21 August 1970)
SUMMARY
1. Inspection of a high-contrast adapting grating produces two visual
after-effects: (a) the contrast threshold is raised for test gratings of similar
spatial frequency to that of the adapting pattern and (b) the apparent
spatial frequency of test gratings shifts away from that of the adapting
grating-higher frequencies seem higher and lower ones lower than they
really are.
2. Both after-effects are orientation-specific. A horizontal adapting
grating influences neither the threshold nor the apparent spatial fre-
quency of vertical test gratings.
3. The magnitude of the two after-effects was measured with vertical
test gratings as a function of (a) tilt of a high-contrast adapting grating
and (b) contrast of a vertical adapting grating.
4. At all frequencies of the test grating, the decline of both after-effects
produced by an increase in tilt of approximately
61'
could be matched by
a reduction in contrast by a factor of 2.
5. We take this as evidence for a common neural origin for these two
visual phenomena.
INTRODUCTION
If one stares at a high contrast pattern of bright and dark bars for a
minute or so, there are two striking consequences. First, it is much harder
afterwards to detect gratings of similar bar width: the contrast threshold
is elevated by about five times (Blakemore & Campbell, 1969a, b). There
is, however, no such threshold elevation for gratings of a very different
spatial frequency (the number of cycles of the pattern per degree of visual
angle). The other outcome of adaptation to a grating is that the apparent
size of the bars of another grating can seem changed. Gratings of a lower
spatial frequency than the adapting pattern appear to be of even lower
*
On leave from Department of Psychology, University of Pennsylvania, 3815
Walnut Street, Philadelphia, Pennsylvania 19104, U.S.A. This address should be
used for correspondence.
158 COLIN BLAKEMORE AND JACOB NACHMIAS
frequency and higher frequency ones seem higher still (Blakemore &
Sutton, 1969; Blakemore, Nachmias & Sutton, 1970).
The threshold elevation phenomenon implies that there are neural
elements in the human visual system that are selectively sensitive to the
spatial dimensions of retinal images. The perceived spatial-frequency shift
is directly predictable on the basis of the hypothesis that adaptation to a
grating depresses the sensitivity of a limited group of 'tuned' frequency-
selective neurones amongst the whole population of such cells, each
optimally sensitive to slightly different spatial frequencies.
The visual cortices of the cat and the monkey do indeed contain neurones
that are selectively sensitive to target size (Hubel & Wiesel, 1962, 1968)
and Campbell, Cooper & Enroth-Cugell (1969) have described the response
of cortical neurones in the cat to moving gratings of different spatial fre-
quencies. These cells are also critically sensitive to the orientation of the
image and will respond to linear targets, even of the optimal spatial
dimensions, only over a limited range of orientation. It is of particular
interest, then, that the perceptual phenomena described above are also orien-
tation-specific. Adaptation to a horizontal grating will influence neither the
threshold (Gilinsky, 1968; Mayo, Gilinsky & Jochnowitz, 1968; Sekuler,
Rubin & Cushman, 1968; Blakemore & Campbell, 1969a) nor the apparent
spatial frequency (Blakemore et al. 1970) of a vertical grating. All of these
findings suggest that the visual system of man is organized somewhat
similarly to that of the cat and monkey.
We have made some effort to emphasize the probable common neural
origin of the threshold elevation and perceived frequency-shift phenomena,
and in a previous paper we drew comparisons between many of their pro-
perties (Blakemore et al. 1970). This notion would be further strengthened
if the two after-effects were found to have very similar orientation
selectivity. This possibility, and the current interest in orientation
detection in the visual system, motivated us to look more closely at this
property of the two perceptual after-effects.
Definitions. Spatial frequency: the number of cycles of a repetitive
grating pattern per degree of visual angle.
Spatial period: the visual angle subtended by a single cycle of the
grating. Period =
1/spatial frequency.
Octave: a change in spatial frequency by a factor of 2.
Imax: luminance of the brightest part of the grating.
Imin: luminance of the dimmest part of the grating.
Contrast:
Imax
-
Imin
Imax + Imin
Mean luminance:
Imax +
Imin
2
ORIENTATION SPECIFICITY IN HUMAN VISION 159
METHODS
From a distance of 114 in. a subject (the first
author, C. B.) looked at two oscillo-
scope tubes with green-yellow (P-31) phosphors. Each screen was masked down to a
rectangular area subtending 1-50 in width, 1-25 in height. One screen was directly
above the other with a gap subtending 25 min between them. In the middle of the
gap was a self-luminous horizontal fixation bar, about 30 min by 4 min.
Schade's method (1956) with the modifications of Green & Campbell (1965) was
used to display vertical gratings with sinusoidal luminance profile on the faces of
the oscilloscopes. The gratings could be turned on and off and their spatial frequen-
cies and contrasts altered by the experimenter without appreciable change in the
mean luminance of the screen, which was 2-2 cd/M2. In the frequency matching
experiments, the subject had access to a potentiometer in series with the time-base
velocity control of the lower oscilloscope, and he could therefore change the spatial
frequency of gratings on that screen. He displayed the voltage across the potentio-
meter on a digital voltmeter and a printer by pressing a button. The period of the
lower grating varied linearly with this voltage. Alternatively, for the experiments on
threshold elevation, the subject controlled, by means of a high precision logarithmic
potentiometer, the contrast of the pattern on the upper screen. He pressed a button
to display the voltage across a ganged linear potentiometer on the voltmeter and
the printer. This gave a direct measure of the logarithm of the contrast.
Above the two oscilloscopes was a simple apparatus to display a high contrast
sinusoidal grating at any orientation. A sheet of opal Perspex (Lucite) was illuminated
from behind by two projectors. Masks on the Perspex exposed two rectangular
transilluminated areas, one above the other, and a fixation bar in between, of exactly
the same dimensions and spatial arrangement as the two oscilloscope displays. In a
rotatable frame over the upper aperture we fixed a negative of a photograph of a
high-contrast sinusoidal grating generated on an oscilloscope.
By means of suitable colour filters and adjustment of the voltages powering the
projectors, we made the mean luminance of the blank lower aperture identical to
that of the upper one containing the grating. Their mean luminance was, in fact,
2-2 cd/m2, exactly the same as that of the two oscilloscope tubes. Under these con-
ditions the contrast of the rotatable grating was found, by matching it with a grating
on an oscilloscope, to be about 047 and its spatial frequency was 84 c/deg. The
luminance of the dim background was about 0-07 cd/M2 throughout.
RESULTS
If the reader wishes to see the two phenomena and their orientation
specificity demonstrated, Blakemore & Campbell (1969a) show an illu-
stration of the threshold-elevation effect and Blakemore et al. (1970) the
perceived-frequency shift.
The concept of equivalent contrast
Blakemore et al. (1970) made the point that changing the orientation of
the adapting grating has an effect on the perceived spatial-frequency shift
just like reducing the contrast of an
adapting grating of the same orienta-
tion as the test grating. It occurred to us that an
appropriate way to
compare the orientation selectivity of the threshold elevation and the
6 P H Y
21I3
160 COLIN BLAKEMORE AND JACOB NACHMIAS
perceived-frequency effects was to express the decline of both effects
with increasing discrepancy between adapting and testing orientations
as an equivalent reduction in contrast of an adapting grating of the
same orientation as the test pattern. First, we justify this operation and
then we use it to make the comparison.
PART I
The perceived spatial-frequency shift
Orientation of the adapting grating. The subject C. B. adapted by fixating
the horizontal bar between the two apertures in the opal Perspex sheet.
The lower of the two fields was always blank and the upper one contained
a sine-wave grating of 8-4 c/deg at a contrast of approximately 0 7. He
moved his gaze backwards and forwards along the horizontal bar to avoid
the formation of a conventional negative after-image of the grating.
After 3 min initial adaptation he quickly transferred his fixation to the
horizontal bar between the two oscilloscope screens directly below the
Perspex sheet. He found vertical sinusoidal gratings, of some spatial
frequency set by the experimenter and always at a contrast of 0x32, on the
face of both oscilloscopes. He quickly tried to match the spatial frequency
of the lower grating, by turning the potentiometer, to the apparent fre-
quency of the upper one, which fell upon the 'adapted' area of retina. He
took about 2 see to make the match, pressed the button, and returned his
gaze to the adapting display for a further 13 see before looking down to
make another setting. This was repeated again and again, the subject
offsetting the potentiometer and re-adapting each time. The experimenter
changed the spatial frequency of the test pattern in half octave steps,
taking two readings at each test frequency on the way up through the
frequency spectrum, and two more on the way down. For test frequencies
lower than 6 c/deg the experimenter also changed the phase of the grating
with respect to the lateral boundaries of the rectangular mask from trial
to trial so as to make it easier for the subject to ignore features other
than 'apparent frequency' in making his judgements.
At least 30 min was allowed to pass between the experimental sessions.
Furthermore, before adapting in the manner described, the subject always
made a series of unadapted frequency matches, with four observations at
each test frequency, in exactly the same fashion, but simply looking between
the two blank screens between readings. These unadapted settings served
two purposes: they acted as base line settings from which to estimate the
effect of adaptation, and also they were a check that there was no residual
after-effect from the preceding experiment. This latter point is important
because Blakemore et al. (1970) have shown that following a long period of
ORIENTATION SPECIFICITY IN HUMAN VISION 161
adaptation, the after-effect lasts for a very long time. The value of taking
base line readings was that they enabled us to subtract out any slight drift
in subjective equality of spatial frequency, possibly caused by a change in
the subject's criterion.
We changed the orientation of the adapting grating in 50
steps between
vertical (00) and 350 off vertical, in both clockwise and anticlockwise
directions. The results are plotted in Fig. 1 for three orientations of the
adapting grating. The means of the base line settings (N = 4) are shown as
IA4 1:4
1-3 -*-1.3
1.2
00
150 300 1.2
.210ro >l
0
0.9 _ _0.9
0.8 0.8
0*7 _
1
X
1
U1 I I I
"
ill I I I Ilf
il l
0.7
3 4 5 10 20 30 3 4 5 10 20 30 3 4 5 10 20 30
Spatial frequency of upper grating (c/deg)
Fig. 1. The effect of the orientation of the adapting grating upon the per-
ceived spatial-frequency shift for vertical test gratings. The logarithmic
abscissa is the spatial frequency of the test grating on the upper oscillo-
scope screen, and the ordinate is the ratio of the period of the lower grating
to that of the upper. The open circles show the means (N
=
4) of settings
made before adapting and the solid circles are means of settings made after
adapting to the rotatable grating of 8-4 c/deg. The results for three different
orientations of the adapting grating are shown: vertical, 150, and 300
clockwise.
open circles and the means of the settings made during adaptation (N
=
4)
as filled circles. If
Po
is the period of the grating on the upper screen, Pi
is the matching grating on the lower screen without prior adaptation, and
P2
is the
matching
lower
grating during adaptation,
then the
period
ratio
plotted on the ordinate of Fig. 1 is
Pl/Po
for the open circles and P2/Po for
the filled circles. Thus, for a perfect spatial frequency match, the period
ratio has a value of 1-0. An over-estimate of spatial frequency (an
under-
estimate of the period) produces
a value of less than
1.0,
and an under-
estimate of frequency, a value greater than 1.0.
Fig. 1 illustrates the effects of adapting to the
grating
at
vertical,
150
6-2
162 COLIN BLAKEMORE AND JACOB
NACHMIAS
clockwise and 300 clockwise respectively. The spatial frequency of the
adapting grating, 8-4 c/deg, is marked on each abscissa with a solid arrow.
Contrast of the adapting grating. In this experiment we used the oscil-
loscope tubes for both adaptation and testing. During adaptation, the
subject fixated the bar between the two screens, the upper one of which
bore a grating. Its spatial frequency was 8-4 c/deg and its contrast was
varied, up to a maximum of 0-72, in steps of
77%
(0-25 log units). The
luminance of the lower screen was uniform.
1 3 1*3
12
- 0072
0*128
0023
-12
.0 2a
~0.9 0*9
0t8
-
- 0.8
0.7
0.7
! |1 ll 1 I Ill 111 1ll
4 5 10 20 4 5 10 20 4 5 10 20
Spatial frequency of upper grating (c/deg)
Fig. 2. The effect of the contrast of the vertical adapting grating upon
the perceived spatial frequency shift for vertical test gratings. The results
are plotted exactly as in Fig. 1 for three contrast levels of adaptation, 0-72,
0-128, and 0023.
After the initial 3 min adaptation, and periodically after re-adaptation
as described above, a test grating was suddenly substituted on both
screens. Without taking his eyes off the fixation bar, the subject quickly
matched in spatial frequency the lower grating to the upper one, and
pressed the button to display the result, whereupon the adaptation display
re-appeared. Again, four readings were taken for each frequency. 30 min
were allowed between sessions and an unadapted series of settings was
made before each one, to be sure that there was no residual after-effect.
The results are plotted in Fig. 2 in much the same fashion as those in
Fig. 1. They show the after-effect at five spatial frequencies spanning two
octaves around the adapting frequency. There are three different contrasts
of the adapting pattern, 0-72, 0-128, and 0-023.
Compare Figs. 1 and 2 to see just how similar is the
change
in the
shape
of the function for a tilt of the adapting pattern or a reduction of its
contrast.
ORIENTATION SPECIFICITY IN HUMAN VISION 163
The equivalent-contrast transformation. The results from the experiments
on adapting contrast and orientation are combined in Fig. 3. On the left-
hand side, the magnitude of the after-effect is plotted against the orienta-
tion of the adapting grating, 00 being vertical. The filled circles represent
the means (N = 4) of readings taken with the adapting grating tilted
clockwise from the vertical, and the open circles are for anti-clockwise
rotation. The results for test frequencies of 4-2, 5.9, 11P9 and 16-6 c/deg
are shown separately, these being frequencies on each side of the adapting
frequency at which considerable after-effects were found.
The magnitude of the after-effect plotted on the ordinate in this case is
the ratio of the period of the matching grating during adaptation to that
of the mean unadapted setting made in the immediately preceding base
line session:
P2/P1.
Therefore for each test frequency, the ordinate value of
1*0 represents no after-effect.
This side of Fig. 3 shows that there is no consistent difference between
the decline of the effect for clockwise and anticlockwise rotation of the
adapting pattern, and that for this level of adapting luminance and con-
trast, the after-effect is hardly detectable for a relative tilt of 350, what-
ever the test frequency.
On the right-hand side of Fig. 3, the magnitude of the effect, expressed
in exactly the same way, is plotted against the logarithm of the contrast
of a vertical adapting grating on the abscissa. The after-effect at the same
four test frequencies is shown with exactly the same ordinates that were
used on the left-hand side of the figure. Again, for each frequency, an
ordinate value of 1P0 represents no after-effect. Two different determina-
tions (N = 4 for each) at each contrast of the adapting grating are plotted
as open and solid circles.
An open arrow on the contrast abscissa marks the approximate con-
trast threshold for the subject just to detect the presence of the adapting
grating while fixating the bar. For each test frequency the after-effect falls
to zero at about this contrast level. Fortunately all the sets of points on
both sides of Fig. 3 are fitted fairly well on these co-ordinates by straight
lines. Regression lines, derived by the method of least squares, are drawn
through them.
Now the equivalent-contrast transformation can be performed. For any
test frequency, one can look up the magnitude of the effect for a parti-
cular orientation of the adapting grating. For example, a grating tilted
by 100 produces a period of ratio about 1-26, at 5-9 c/deg. Now one can
look across to exactly the same position on the ordinate on the right-hand
side and one finds from the regression line that this magnitude of after-
effect at 5-9 c/deg is also caused by a vertical adapting grating with a
contrast of about 0 3.
164 COLIN BLAKEMORE AND JACOB NACHMIAS
It is a simple matter to construct a graph, for each test
frequency,
of the
adapting contrast equivalent to a tilt of the fixed-contrast
adapting
grating. Since both original functions are straight lines, the transformation
will, of course, also be a straight line, on the same co-ordinates.
Fig. 4 shows the equivalent contrast transformation functions for the
four test frequencies. It will be remembered that the contrast of the
1*3
1*2
1X1
Orientation (deg)
0 10 20 30 40
1
11
I I I I
l;I
0
* 0
0 *
0 e
Contrast (log units below 0.72)
-2*0 -1*5 -1.0 -0*5 0
I I I I I I I I I
4X2
c/deg
0 o
10
0
C
00
o0
1*3
1-2
1.1
Vu - -___a1U
1*3 F-
1*2
1*1
0
o 1-0
0
09
0*8
0
0
5*9 c/deg
0
0
0
00 0
ID
0 080
11.9cfdeg
.0
0
0.7
L
1*4
1-2
1*1
1.0
Is
10o
0
00
0*8
0*7
0 10 20 30 40- 0.01 0.1
Orientation (deg) Contrast
Fig. 3. For legend see facing page.
1'
,
s.n L
10
ORIENTATION SPECIFICITY IN HUMAN VISION 165
rotatable adapting grating was about 0 7. Therefore, when it was vertical,
it should have had exactly the same effect as a grating of that contrast
displayed on the upper oscilloscope screen. It is obvious, then, that all the
transformation functions should pass quite close to this equivalent contrast
level, marked with an open arrow on the ordinate, for an adapting orienta-
tion of 00. They do, and, what is more, they all have approximately the same
slope. Here is the justification for this equivalent-contrast manoeuvre:
it produces nearly the same function, whatever the test frequency.
Look back at Fig. 3 to see another way of making this point. Again
follow through the transformation to find the equivalent contrast for an
adapting grating tilted by 100. But this time do it for all four test fre-
quencies by following across horizontally to the equivalent position on the
contrast function on the right-hand side and you will find that it always
produces approximately the same result, 0-3.
Clearly, so far as the perceived-frequency shift at all test frequencies
is concerned, changing the orientation of the adapting grating has the
same effect as reducing its contrast by a fixed amount. Next we derive a
similar transformation for the threshold-elevation effect, and compare the
two transformations.
PART II
The threshold-elevation phenomenon
Orientation of the adapting grating. For this experiment, the subject
adapted by looking directly at a fixation mark in the centre of the rotatable
grating, and moved his gaze around the mark to prevent the generation
of an after-image.
Legend to Fig. 3.
The equivalent-contrast transformation for the perceived spatial-frequency
shift phenomenon. The magnitude of the after-effect is plotted, on the
left, against the orientation of an adapting grating of 0-7 contrast, and,
on the right, against the contrast of a vertical adapting grating.
Results are shown for four test frequencies, 4-2, 5-9, 11-9 and 16-6 c/deg.
Each ordinate is the ratio of the period of the matching grating during
adaptation to that in the immediately preceding unadapted 'base line'
session. A value of 1-0 on each ordinate marks the level of zero after-effect
for that frequency. On the left, the filled circles are for clockwise rotation
and the open circles for anticlockwise. Split symbols are used for the
results with a vertical adapting grating. On the right the filled and open
symbols are results for two separate sessions. The logarithmic
abscissa is
the contrast of the adapting grating and the open arrow marks the subject's
approximate contrast threshold for a grating of 8-4 c/deg. Regression lines
are fitted through each set of data points so that for any orientation of the
adapting grating it is possible to determine the contrast of a vertical
grating that would cause the same magnitude of after-effect.
166 COLIN BLAKEMORE AND JACOB NACHMIAS
After periods of initial adaptation and successive readaptation, he looked
down to a fixation point in the middle of the upper oscilloscope screen.
(The lower part of the display was not needed for this experiment.) Here
he found a vertical test grating turned on and off 1-5 times per second,
without any change in mean luminance. He used the logarithmic
potentiometer to attenuate the modulating signal to the Z-axis of the
1*0
co
es 0-5
(U
to 0-4
.X 0-3
0.
.w
la 0-2
co
.'U
W 001
0
0-
:3
0-04
0
~.. 0-03
1v
0-02
001
I I I I I I I I
0 10 20 30 40
Orientation of adapting grating (deg)
Fig. 4. The equivalent-contrast transformation functions for the per-
ceived spatial-frequency shift at the four test frequencies analysed in
Fig. 3. Each line on this graph plots, for the test frequency indicated, the
contrast of a vertical adapting grating of 8-4 c/deg that will produce the
same change in apparent spatial frequency as an adapting grating, also
of 8-4 c/deg, and at a contrast of 0 7, tilted at various angles to the vertical.
The orientation of the adapting grating is plotted on the abscissa, zero
being vertical, and the equivalent adapting contrast on the logarithmic
ordinate. The contrast of the rotatable grating, 0-7, is marked with an open
arrow on the ordinate. Clearly all the lines should have this value at zero
on the abscissa.
oscilloscope, until the grating was at threshold, whereupon he pressed the
button to print out the contrast level. All this he could do in a few seconds
and then he looked back to the adapting grating.
The spatial frequency of the test grating was varied by the experimenter
ORIENTATION SPECIFICITY IN HUMAN VISION 167
and four estimates of the threshold taken for each value, just as in the
previous experiments. At least 30 min recovery was allowed between each
session and a set of unadapted estimates of the contrast threshold taken
before each adaptation run. Again these unadapted estimates served the
0.1
04
0=
0 05
L
0
004
W 003
4A0
0
0*01
4
~~~~~"_
II
g1_
-I ElIlt I IX 11 1111 I I I I I I1111 I I
-
3 4 5 7 10 20 303 4 5 7 10 20 303 4 5 7 10 20 30
Spatial frequency (c/deg)
0.1
005
004
003
0*02
0*01
Fig. 5. The effect of the orientation of the adapting grating upon the ele-
vation of threshold for vertical test gratings. The logarithmic abscissa is the
spatial frequency of the test grating and the logarithmic ordinate is the
threshold contrast. The open circles are the means (N
=
4) of threshold
estimates made before adaptation and the filled circles means (N = 4) of
settings made after adaptation to the rotatable grating of 8-4 c/deg
(marked with a solid arrow). Results are plotted for three adapting orien-
tations, vertical, 15 and 300 clockwise.
0.1
o 0*05..
a 0-04
W 0-03
= 0*02
C
0
U
0*01
4*
0-023
0-1
005
0-04
0-03
0-02
0*01
I I II I I I I I 111 111 I L
4 5 678910 20 4 5 678910 20 4
Spatial frequency (c/deg)
I I I 11 1 1
5678910 20
Fig. 6. The effect of the contrast of the vertical adapting grating upon the
elevation of threshold for vertical test gratings. The results are plotted in
exactly the same way as in Fig. 5 for three contrast levels of adaptation,
0-72, 0-128 and 0-023.
double function of acting as local base lines from which to judge the rise
in threshold and as a check on whether there was any residual after-effect.
The results are plotted in Fig. 5. The logarithmic ordinate is contrast
168 COLIN BLAKEMORE AND JACOB NACHMIAS
Orientation (deg)
0 1.0 20 30 40
I
I
I
I'I-
11
0
0
rhI 0'
Contrast(Itg
units below 0.72)
-2-0 -1-5 -1-0 -0-5 0
1 - A I I
o o _ to0
4-2
c/deg
0
*
0
To
0 0
-0
0
0 8
- a_
S-9 c/deg
0
*
,
8.4-c/deg
0
10 20 30 40 0-01 0.1
Orientation (deg) Contrast
Fig. 7. The equivalent-contrast transformation for the threshold-elevation
phenomenon. As in Fig. 3 the magnitude of the after-effect is plotted, on
the left, against the orientation of an adapting grating of 0-7 contrast,
and, on the right, against the contrast of a vertical adapting grating.
Results are shown for four test frequencies, 4-2, 5-9, 8-4 and 11-9 c/deg.
Each ordinate is the logarithm of the ratio of the mean adapted contrast
threshold to the mean contrast threshold in the immediately preceding
unadapted base line session. Zero on each ordinate, then, indicates the
level at which there is no after-effect. The symbols and abscissa are exactly
as in Fig. 3 and again the open arrow on the right shows the subject's
approximate contrast threshold for a grating of 8-4 c/deg, the adapting
frequency. Regression lines are fitted through each set of data points.
'.3 0-3
0-2
0-1
U-s
o-s
04
0-3
?0s2
6.
to
0 04
-; 0-7
1.0.61
05
2
0-4
%
0-3
04
0
0-2
0-1
O
0-6
0-5
0-4
'0-3
0-2
0-1
0
a
91%
-
ORIENTATION SPECIFICITY IN HUMAN VISION 169
threshold and the abscissa is the spatial frequency
of the test
pattern.
The
open and filled circles are, respectively,
determinations of threshold before
and during adaptation to the grating of 8-4
c/deg,
marked with a solid
arrow. Results for adapting orientations of
vertical, 15 and 300 clockwise
are plotted in this Figure. As reported by Blakemore &
Campbell (1969b)
the elevation of threshold is greatest in the
region of the
adapting spatial
frequency, and the whole after-effect declines as the
adapting grating
is
tilted.
1.0
bO
C4 0*5
bO
0-3
ba
C
CL
.0-2
co
0-
0
0.05
0.04
0~
LU
0119
I I I I I I I I
I
0 10 20 30' 40
Orientation of adapting grating (deg)
Fig. 8. The equivalent-contrast transformation functions for the eleva-
tion of threshold at the four test frequencies analysed in Fig. 7. The curves
were derived from the regression lines of Fig. 7 in exactly the same way as
those of Fig. 4 were produced from Fig. 3. The axes are identical to those of
Fig. 4 and the four lines refer to the four test frequencies indicated.
Contrast of the adapting grating. The subject looked at the single oscillo-
scope screen throughout this
experiment. The
adapting grating of 8-4
c/deg and variable contrast was periodically replaced by a low contrast test
grating, flashing on and off 1-5 times per see, which the subject adjusted to
threshold.
Fig. 6 shows some of the results. Contrast thresholds before and
during
adaptation are plotted as open and filled
circles, respectively,
for
adapting contrasts of 0-72, 0-128 and 0-023.
170 COLIN BLAKEMORE AND JACOB NACHMIAS
The equivalent-contrast transformation. Fig. 7 is analogous to Fig. 3. In
Fig. 7, the magnitude of the threshold elevation effect (expressed as the
log of the ratio of the adapted to the unadapted contrast threshold) is
plotted as a function of the orientation of a high-contrast adapting grating
and as a function of the contrast of a vertical adapting grating. Results are
shown for the four test frequencies exhibiting the largest threshold eleva-
tions; 4-2, 5 9, 11-9 and the adapting frequency itself, 8-4 c/deg. For each
test frequency, the same ordinates are used on the two sides of the figure.
Zero on the ordinate represents no elevation of threshold.
1.0 00
0.
10 20 30 4 0
0
20051
0. ~ rinato
300Nraizdeuvletcnrs
0.4
4J(~O
> 0*02
O
Ui0~0
0 10 20 30 40 0 0.5 1.C
Orientation of Normalized
equivalent
contrast
adapting grating (deg)
Fig. 9. The orientation specificity of the two after-effects, expressed in
terms of equivalent contrast.
A. The results of Figs. 4 and 8 are reproduced on the same co-ordinates
but only the mean of the four regression lines is reproduced in each case.
The lines are marked, one for the perceived-frequency shift and the other
for threshold elevation.
B. The mean regression line of Fig. 9A, for the two after-effects, is
plotted here on polar co-ordinates and a linear rather than a logarithmic
scale. The function is reproduced symmetrically on each side of vertical
(00). The equivalent contrast of the vertical adapting grating is, of course,
approximately the same as its actual contrast, 0 7, and it has been arbi-
trarily assigned a value of 1.0 on the ordinate. The width of the function
at half amplitude (0-5 on the equivalent-contrast scale) is about
131'
(half-width, 610).
On the right-hand abscissa the open arrow
again
marks the
approxi-
mate psychophysical threshold for detection of the adapting pattern. It is
evident that at each test frequency, there is virtually no elevation of
threshold if the adapting grating is tilted
by
350 or reduced in contrast to
its own threshold.
ORIENTATION SPECIFICITY IN HUMAN VISION 171
Regression lines, determined by the method of least squares, are fitted
to each set of points in Fig. 7. Fig. 8 shows the equivalent contrast trans-
formation functions derived as in Fig. 4. Again the equivalent-contrast
transformation appears to be legitimate, for the four regression lines are
very similar in both slope and intercept. Moreover, a comparison of Figs.
4 and 8 should convince the reader that the decline of equivalent contrast
with rotation of the adapting pattern is practically identical for the two
after-effects. In Fig. 9A the mean regression line from Fig. 4 and the one
from Fig. 8 are plotted on the same graph to show their similarity.
DISCUSSION
In this paper we have introduced the idea of equivalent adapting con-
trast and have shown that it is a reasonable way of expressing the orienta-
tion specificity of both the perceived-frequency shift and the threshold-
elevation phenomenon.
The real value of this transformation is that it makes the exact nature of
the measure for the after-effect irrelevant, as long as the same metric is
used to express the results of both the orientation and contrast experi-
ments. A direct comparison can then be made, in exactly the same terms,
of the properties of the two visual phenomena. And it turns out that the
orientation sensitivity, expressed in this way, is practically identical for
the two (Fig. 9A).
This is not, of course, conclusive proof that the neural origin of the two
effects is the same but it is certainly a strong indication. All the more so
when one remembers that the perceived-frequency shift was predicted,
a priori, from frequency-specific threshold elevation (Blakemore & Sutton,
1969; Blakemore et al. 1970).
This kind of data reduction is analogous to the so-called 'Crawford
transformation' (Crawford, 1947). Under a variety of conditions, the
dark adaptation curve, plotting the threshold for a flash of light against
time after an intense bleach of the retina, varies in the same way as the
increment threshold curve, plotting the amount of extra light necessary
to see a flash on a background of variable intensity. It is possible, therefore,
to compute, for any given amount of time in the dark, the luminance of an
'equivalent background' that would raise threshold by the same amount.
Just as the concept of equivalent background, or 'dark light' has impor-
tant theoretical implications for students of dark adaptation (Barlow &
Sparrock,. 1964; Rushton, 1965) the idea of equivalent adapting contrast
may have theoretical consequences for any model of how spatial adapta-
tion works.
It is natural to suppose that the site of the adaptation whose after-
172 COLIN BLAKEMORE AND JACOB NACHMIAS
effects we have been investigating psychophysically is a population of
frequency and orientationally selective neurones in the human visual
cortex, similar to those already revealed in cat and monkey by means of
single-unit recording (Hubel & Wiesel, 1968; Campbell et al. 1969). It
should be possible to demonstrate in such cells some electrophysiological
after-effect of prolonged stimulation by a high-contrast grating. In that
case, several questions spring to mind, among them the following: in a
single cell, does the magnitude of the after-effect, regardless of the test
frequency at which it is measured, vary in a similar manner with the con-
trast and orientation of an adapting grating? Is the frequency of the most
effective adapting grating independent of its orientation and vice versa?
Our results suggest affirmative answers to both questions. For example,
recall that the threshold-elevation effect remains centred at 8-4 c/deg
whatever the orientation of the adapting grating (Fig. 7).
The curves of Fig. 9A show that the decline of equivalent adapting
contrast with orientation is approximately exponential (since the ordinate
is logarithmic) and that it has dropped by a factor of two (0.3 logarithmic
units) after about 6 ' of rotation. The mean function for the two after-
effects is replotted in Fig. 9B with a linear scale, on polar co-ordinates, and
this shows more conveniently that the half-width of the function at half
amplitude is about 63', the full width
131o.
Other investigators of the threshold elevation phenomenon have
measured its orientation specificity (Mayo et al. 1968; Sekuler et al. 1968)
or its dependence on adapting contrast (Pantle & Sekuler, 1969; Blake-
more & Campbell, 1969b). Qualitatively our results, displayed in Fig. 7,
agree with theirs; there are some quantitative discrepancies, which may
well be due to differences in procedure and stimulus conditions. The
conditions employed by Blakemore & Campbell (1969b, Fig. 4) and
Sekuler et al. (1968, Fig. 3) were most similar to ours and their results
are in excellent agreement with our own (Fig. 7, 8x4 c/deg).
Campbell & Kulikowski (1966) measured the threshold for a vertical
grating in the presence of a 'masking' grating, as a function of the contrast
and orientation ofthe latter. They found that the threshold elevation caused
by simultaneous masking also decreases exponentially as the masking
pattern is rotated but that the effect does not fall by a factor of two until
it is tilted by 120. We have no explanation for this big difference in
orientation specificity ofsimultaneous and successive masking phenomena,
but it must be pointed out that Campbell & Kulikowski were working at
considerably higher luminance levels than we were, and this may account
for the discrepancy.
Campbell, Cleland, Cooper & Enroth-Cugell (1968) found that the fre-
quency of nerve impulses generated by cat cortical cells falls off linearly
ORIENTATION SPECIFICITY IN HUMAN VISION 173
with the orientation of a moving grating, expressed in degrees. It would be
most interesting to know the manner in which such cells respond to varia-
tion in contrast of a grating of the preferred orientation, so that their
orientation selectivity could also be expressed in terms of the 'equivalent
contrast'.
One cannot help wondering why orientation-selectivity
is such a uni-
versal property of visual systems. Cats and monkeys are not the only
species to have orientational units. They have, for example, also been
found at various stages in the visual systems of the rabbit (Levick, 1967),
ground squirrel (Michael, 1967), pigeon (Maturana & Frenk, 1963) and
goldfish (Cronly-Dillon, 1964). One obvious function for such neurones is
that they actually encode the orientations of parts of retinal images and
thus contribute directly to form perception. But an alternative, or addi-
tional idea has been suggested by Blakemore & Campbell (1969b). The
orientation-selective neurones, by virtue of their spatial-frequency selec-
tivity, may be acting as sets of spatial-frequency filters, each set per-
forming a crude Fourier analysis of the luminance profile of the retinal
image along a different meridian. In this manner, the harmonic content of
a pattern would be simultaneously analysed along many meridia. In
either case the orientation-selective and frequency-selective neural
mechanisms at the root of the threshold-elevation phenomenon and the
perceived spatial-frequency shift may have a vital role to play in the whole
business of image recognition.
J.N. was sponsored, during his visit to Cambridge, by a research grant (NB
06050) from the U.S. Public Health Service. C. B. holds a grant (No. G968/190/B)
from the Medical Research Council, London. The Wellcome Trust provided a travel
grant for C. B. to visit the University of Pennsylvania.
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