Eleetroencephalography and clinical Neuropt~vsiologv, 1984, 58: 107-119 107

Elsevier Scientific Publishers Ireland, Ltd.

C O L O U R A N D B R I G H T N E S S C O M P O N E N T S OF FOVEAL VISUAL EVOKED P O T E N T I A L S IN

MAN

W.M. PAULUS ,.i, V. HOMBERG *, K. CUNNINGHAM **, A.M. HALLIDAY ** and N. ROHDE *
• Neurologische UniversitiJtsl~linik Di~sseldorf, Moorenstr. 5. 4000 Di~sseldorf (F.R.G.) and ** National Hospital for Ner~ous Diseases,
Queen Square, l+ondon WC1N 3BG (England)

(Accepted for publication: January 3+ 1984)

While much of the literature on colour evoked
potentials deals with steady-state potentials (e.g.,
Regan 1970, 1972) or measurement of spectral
luminosity functions (e.g., Siegfried 1971; Est6vez
et al. 1975; Zrenner 1983), there have been rela-
tively few parametric studies of the different ef-
fects of colour and brightness changes on transient
visual evoked responses. One reason for this may
be the difficulty of defining colour and brightness
parameters psychophysically. VEPs have fre-
quently been recorded to colour onset, but these
responses inevitably have both colour and bright-
ness components (for definitions of luminance,
brightness and additivity failure, see Appendix).
Experiments in which coloured patterns wer& used
as the stimuli have indicated that colour-related
effects may be masked by dominating pattern
responses (Shipley et al. 1965; Spekreijse et al.
1977; Est6vez and Dijkhuis 1983).
The present study addresses the problem of
discriminating colour and brightness effects by
using unstructured colour substitution stimuli de-
fined both psychophysically and photometrically.
Equal brightness matches were performed by het-
erochromatic brightness matching carried out by a
trained observer between a yellow reference colour
and red, orange and green test colours. Replace-
ment of the yellow reference by an equally bright
colour should result in a pure colour evoked re-
sponse. To discriminate any additional effects of
brightness changes, a range of colour stimuli with
1 Present address: Neurologische Klinik. Alfried Krupp Krhs,
43 Essen, F.R.G. Supported by Deutsche Forschungs-
gemeinschaft, Grant No. Pa 267.
luminance levels above and below the equal
brightness level were introduced. Uncontrolled
adaptation effects to colour or luminance are to be
avoided in colour experiments and care was taken
in the present study to guard against this.
The results obtained from a group of 10 sub-
jects are presented and the chosen method of
heterochromatic brightness matching is discussed.
A preliminary account of this work has appeared
elsewhere (Paulus et al. 1984).
Methods
Stimulating equipment
The stimulator used provided a circular 4 ° di-
ameter centrally fixated test field surrounded by a
rectangular background field with a diameter of
60 °. The fixation spot was a small black dot of 0.5
m m diameter. The test field alternated between a
yellow and either a red, orange or green stimulus.
This was provided by an intermingled matrix of
about 100 red and 100 green LEDs whose inten-
sity could be independently controlled. By simul-
taneous illumination of the red and green LEDs at
various intensities it was possible to produce a
stimulus of yellow, or of differently saturated reds
or greens. The matrix of diodes and the lamps
illuminating the background were placed about 10
cm behind a translucent screen, so that a homoge-
neous field could be produced. Because of this
simple construction, no lens system or other opti-
cal devices were needed. The test field and the
background were separated by an opaque partition
mounted behind the screen so that they would not

0013-4649/84/$03.00 ,-~ 1984 Elsevier Scientific Publishers Ireland, Ltd.

108
influence each other. The subject was seated with
his eyes 40 cm in front of the screen. The luminos-
ity of the surrounding background, derived from 4
tungsten lamps, was kept constant at 25 c d / m 2.
Recording system
Silver-silver chloride disc electrodes were located
at FZ, PL, O, and half-way between P~ and O~ (POx)
and at both mastoids, serving as a linked refer-
ence. Electrode-skin impedance was kept below 5
kg2 measured at a frequency of 400 Hz. Electrical
activity was recorded by 4 T6nnies amplifiers with
time constants of 0.3 sec and an upper frequency
response 3 dB down at 30 Hz. The 26 different
stimuli were presented randomly under the control
of a Nicolet MED-80 computer, which stored ev-
ery trial with a sampling rate of 1 kHz, a pre-
stimulus time of 50 msec and a total sweep dura-
tion of 512 msec. The data were digitized with
12-bit resolution. An artifact rejection facility ex-
cluded single trials with significant eye movement
artifacts by discarding trials with amplitudes ex-
ceeding the input range of the AD convertor.
Psychophysicalprocedures
VEPs were measured to substitution of the ref-
erence yellow (wavelength 567 nm, luminance 21.5
c d / m 2) by a red, orange or green stimulus of
equal brightness, as well as by an identical yellow
control stimulus, i.e., without any test field change.
The colour loci of the red, orange and green
stimuli were spectral colours of 595, 578 and 552
nm wavelength 2. In Fig. 1 their colour loci are
plotted on the CIE u',v' chromaticity diagram, in
which equal distances are supposed to represent
equal colour differences (Wyszecki and Stiles 1982).
Equal brightness of red, orange and green, com-
pared with the reference yellow, had been de-
termined psychophysically at a flicker rate of 2 Hz
by one of the authors (W.P.), who had experience
2 Although in physical terms the stimuli are not monochro-
matic, their colour loci are identical with spectral colours. This
is due to the involvement of only two receptor systems, the
'green' and 'red' receptors. For example a monochromatic
yellow colour of 567 n m is indistinguishable from an adequate
additive mixture of green and red, which in fact was used in
our equipment.
W.M. P A U L U S ET AL.
0.6
552
567
• ¥ 578
O 595 n m
Ve • R
21.9 21.5 ¢d
m2
0,5 I
15.3
01 0.2 0.3 0./.
UI
Fig. 1. Colour loci (circles) of the colour stimuli within a part
of the CIE u',v' diagram. In this type of chromaticity diagram,
equal spatial distances are supposed to represent equal percep-
tual steps. One row (triangles) of the Uniform Colour Scales
(MacAdam 1974), representing equal perceptual distances, has
been added for comparison and shows a similar decline in
luminance for red colours of equal subjective brightness.
with heterochromatic brightness matching (Paulus
1978/79).
The resulting luminance values for red, orange
and green, equated in brightness to the reference
yellow, were: 13.0, 16.6 and 23.1 c d / m z respec-
tively. To demonstrate the perceptual colour dif-
ferences of our stimuli a comparable row (L = - 2,
j = 6, - 1 0 less than or = g less than or = 4) of
colours of the Uniform Colour Scales (McAdam
1974, 1978) has been added (triangles). The col-
ours are separated by equal perceptual steps differ-
ing only in red-green hues whilst the blue-yellow
and brightness, j and L, are kept constant. Their
luminance value can be seen to decline towards
the red colours with a similar slope. (For an over-
view of 'colour brightness' perception, see
Wyszecki and Stiles 1982.)
The Uniform Colour Scales data represent the
most modern estimation of subjective scaling of
saturation and brightness changes in colour per-
ception. They provide a 'grand average' reference
for the subjective procedure of heterochromatic
brightness matching, in which there is a great deal
of inter-individual variability between different
observers. The estimations of our trained observer
(W.P.) correspond closely to the UCS data (Fig. 1)
and on this basis the same stimuli were employed
for all the subjects in the group.
The colour loci were derived by measurement
with a Photo Research Spectra Spotmeter photom-
eter, which provides CIE tristimulus values X, Y,
Z for each colour measured. These values are
transformed in the chromaticity diagram coordi-
C O L O U R A N D BRIGHTNESS COMPONENTS OF VEPs
nates x, y by:
x = X / ( X + Y + Z)
and
y = Y / ( X + Y + Z)
The appropriate wavelength has then been de-
picted from the CIE 1931 Standard Colorimetric
System, which is printed in Wyszecki and Stiles
(1982), in 1 nm intervals. Our stimuli were spectral
colours with maximum possible saturation, since a
blue cone intrusion was not present. This was
reflected in zero values for Z.
VEP procedure
To study the influence of additional brightness
increments or decrements on the different colours,
we introduced 26 different stimuli. In addition to
the equally bright stimuli, brighter and darker
stimuli with luminance increments and decrements
of 15~/~ and 50% for each colour were provided.
Some further stimuli of +100% and 75% were
included for red, yellow and green stimuli. A yel-
low to no-stimulus control was also included and
is plotted at the bottom of the 'orange' column in
Fig. 2 (off response). Photometric calibration was
performed with a Photo Research Spectra Spotme-
ter photometer.
The stimuli were presented in random order to
avoid adaptation to brighter or darker luminance
levels. The stimulus sequence alternated between
the reference yellow, presented with a randomized
time interval between 300 and 500 msec, and a
following 300 msec test stimulus. Thereafter
another randomized 300 500 msec yellow stimu-
lus occurred, followed by a compensating stimulus
with the complementary ~ colour to the test stimu-
lus, again of 300 msec duration, before the se-
quence started off again. The responses to the
complementary colour stimuli were not analysed.
The use of yellow instead of white as a refer-
ence stimulus should not substantially alter re-
"Complementary' is used in terms of colour opponent theory.
The intention was to avoid adaptation in the red green oppo-
nent channel. Since neither the red nor the green colours
stimulated the 'blue' receptor, the blue-yellow channel was
constantly unbalanced towards yellow.
109
sponse wave forms. Yellow is the most desaturated
spectral colour and, due probably to adaption to
yellow during the course of the experiment, the
tungsten lamp illuminated background was, in fact,
almost indistinguishable from the reference yellow.
Fourteen subjects (11 male, 3 female), aged
between 25 and 35 years, served as experimental
subjects. Their visual acuity was normal and all
the subjects could read the Ishihara plates without
difficulty. Data from 4 subjects had to be excluded
from further analysis because of high noise con-
tamination of their records, probably due to photic
alpha-driving in spite of the random stimulus pre-
sentation. One hundred responses were averaged
from each subject for each colour. The data for
each subject were collected in two 1 h sessions,
presented on the same day with a 30 rain break
between them.
Results
To demonstrate the cardinal wave form fea-
tures, Fig. 2 presents grand averages of 10 subjects
for all colour and luminance levels at the 4 differ-
ent electrode sites. Inspection of the wave forms
reveals an inconstant, very small, positive-going
peak at about 55 msec followed by a prominent
negative component with onset at around 60 msec
and peaking at around 87 msec, showing a clear
occipital maximum and a gradual decrease to-
wards more anterior leads. This N87 component is
most pronounced for red, somewhat smaller for
orange, even smaller for green and virtually absent
for the purely luminance changes of the yellow
stimuli.
For a given colour, N87 is more accentuated for
stimuli with additional luminance increments. For
a saturated red stimulus, the N87 peak is followed
by a long-lasting negative wave which obscures the
later positive components. This negative after-wave
appears most prolonged for the brightest red
stimuli.
For the colour substitution stimuli of equal
brightness, the N87 component shows a very simi-
lar configuration for red, orange and green stimuli,
although it is somewhat smaller in amplitude for
the latter. This component is still seen with the
colour stimuli with a reduced luminance of 15%,
110 W.M. PAULUS ET AL

FZ--LM

Y R Y Y 0 Y Y Y Y Y G Y

J [ _ l J L__ _J l__

• 100 . ~ A ~ _ _

•15 ~ V p~ -15
~ %/A .50
*15
15

/ -

m2 I 16,6

-15

-50 J"

-75

5M¥I
' ' I ' P ' I ' 1 ' ' ' I I ' I ' I ' ' ' I ' I ' I ' I ' ' I ' I ' f ' '
100 200 300 LO0 ms 100 200 300 400 ms 100 200 300 l.O0 ms 100 200 300 l.O0 ms

PZ--LM

Y Y Y 0 Y Y Y Y Y O Y

I__ __J [__ J I__ _ L _ _

• 100 !
*75

*50 *50 ~

*15 -15 *15 ~ .15 . /

c~
13,0 / 16,6 21.5 ~ 23,1
m2

-15 -15 -IS../~/~ A /-15"~

-50 / -50 ~

-75 - - off ~" -75 "-- -75

SM¥I
I ' I ' I ' I ' I ' ' ' I ' I ' I ' I ' ' ' f ' [ ' t ' I '
100 200 300 /.C~3ms 0 1130 200 300 z,O0 ms 100 200 300 l.O0ms 100 200 300 400 ms
Fig. 2. For legend see facing page.
('OLOUR AND BRIGHTNESS COMPONENTS O F VEPs 111

POZ--LM

Y R Y Y 0 Y Y Y Y Y G Y
_ i j [__ J L__ _ I

1 ~'J .15 .15 .J"

] - 5 -15 -15

i I ~ I i I ' I '
0 100 200 300 t.O0 ms 100 200 300 z.O0 ms 100 200 300 L.O0ms 100 200 300 /.00 ms

O Z -- L M

O Y Y Y Y Y G Y
Y R Y Y
!
-
t__ / F _

1.,,'
• 100 ~ *75

• 50 "~ *50 ~ ' V ~ /r~ /.~ /~ I *50

.15

C~ 16,6 21.5 23,1

13.0 z~"
m2

-15 -'~ -15

-50 /J" _ 50 ...~ - "*v ~ -50 -'~

-75 off -75 ~ -75

SPYI

r ' I ' l ' ! ' I ' ' ' 1 i I ' I ' I ' l i ' I i ,

0 100 200 300 /,00 ms 100 200 300 /*00 ms 100 200 300 /.00 ms 100 200 300 400 ms

l:ig. 2. ( ; r a n d averages of V E R s from 10 subjects at 4 different electrode sites. F,. P~. PO, and O,. The mMdlc ro,.', of traces (marked
on the left by tile actual test sth+;ulus intensity hi c d / m 2 ) represents the responses to subjecfivel~ equally bright colour stimuli. C o l o u r
tcspollSe>, with a d d i t i o n a l JtlFnill;.mce illl..'felllelltS and decrenlents (marked ill percentages at tile left of e a c h trace) arc sJlo~.~l~ D.bove and
below respecti,,elx. A l l c o l o u r stimuli are preceded and follo,,~ed b', the s a m e , , e l l o ~ reference stimulus, as indicated at the top of each
trace {for further e x p l a n a t i o n , see texl). Negati',h~ plotted up'~ards.
112 W.M. P A U L U S ET AL.

but is not evident for the transition to the two
dimmer colour stimuli.
In contrast to the colour-brightness dependent
N87, the most pronounced feature in the wave
form correlating with pure luminance changes is a
positive deflection starting at around 75 msec and
peaking at around 120 msec (P120). It increases in
amplitude, both for luminance decrements and
increments, i.e., for any change in luminance. For
the green stimuli, P120 follows the preceding N87
potentials. For bright orange and red stimuli, P120
seems to be superimposed on the longer-lasting
negative wave and is only visible as a small deflec-
tion for the brightest stimuli, although it reappears
with decreasing brightness and dominates the wave
forms for the darkest orange and red stimuli.
Topographically, N87 and P120 have an occipital
maximum with a steep decline in amplitude in the
parietal leads. P120 has an obvious polarity inver-
sion at F~ (although this may be partly contributed
by the linked mastoid reference).
At longer latency a non-specific P250 compo-

red orange yellow green

'I/I~'A S pV J~ SpV s yv

/~ vvw
I /h ,, ~ / I ,
, A I
/ ; 'w.,< '\U~/

z,,~.-.~I ..,~ ~1 ~ ~ '~ ~"%'~/

4...~-~.¸~,

6~, V
',j

C) " 100' 200 3(~0' Z~O mS () ' 100'2~' 300' ~0 'mS {~ ' 100 200' 3~0' /~0 ms {~ ' 1C}0' 2~' 300' ~}0 'ms
Fig. 3. Individual wave forms of the average responses to 100 bright red, orange, ~ellow and green stimuli, recorded from O, in 10
different subjects.
C O L O U R A N D B R I G H T N E S S C O M P O N E N T S OF VEPs 113

nent, with a predominantly frontal maximum, N 87 - AMPLITUDE

shows somewhat increased amplitudes for more
-8- red
saturated colour stimuli.
-7--
In the late part of the analysis epoch a clear pV_6_
sequence of colour-offset potentials is elicited by
-5--
the transition from the test stimulus to the con- -4--
stant yellow reference stimulus. Only the first 162
'\ -3~
msec of this 'off response' were included in the
analysis epoch. A positive peak at 85 msec (P85
-2--
'
\ i

-1 --
off), a negative peak at around 95 msec (N95 off) O-
and a second positive peak at around 125 msec b d b d
(P125 off) can be seen. Whereas P85 off and N95
Fig. 4. Mean amplitude+_standard error of component N87 in
off are best developed for transitions from bright all 10 subjects, measured at O, for all red and orange stimuli
red or green to yellow, P125 off increases in ampli- (excluding the off stimulus). The responses to the brightest red
tude with decreasing luminance of the test stimu- or orange are labelled b, the darkest d. Responses to red and
lus. i.e.. with transitions in which the return to the orange stimuli of equal brightness to the yellow reference are
labelled with an asterisk.
reference yellow represents a luminance increment.
P125 off, like the earlier P120 to the test stimulus,
shows a clear occipital maximum with a polarity
inversion at the frontal electrode. N95 off is seen for red and orange test stimuli. As can be seen, the
chiefly in occipital and (to a lesser extent) parietal amplitude for both colours steadily declines as
leads. The following negative-going slope, only luminance decreases, red stimuli yielding larger
partially represented in the analysis epoch, ap- amplitudes than orange. Asterisks indicate the am-
pears most pronounced for transitions from low plitude of potentials elicited by stimuli matched in
luminance test stimuli to the standard yellow, brightness to the reference yellow. In Fig. 5,
maximal for the yellow to yellow transition. latencies to the onset and peak of N87 are plotted
A basic problem to be faced, especially with for all red stimuli excluding the darkest. Only a
foveal and unstructured test fields, is the variabil- small trend towards latency reduction with brighter
ity of the representation of the visual cortex colours is observed.
(Brindley 1970) which is supposed to cause a great PI20 amplitudes are plotted in Fig. 6 for the
deal of inter-individual variability of evoked visual
responses (Halliday 1982). For this reason the
main features of the derived potentials have been N 8 7 - LATENCY
demonstrated in a grand average from 10 subjects Onset Peak
(Fig. 2). To illustrate inter-individual consistency ms red
of the observed wave forms, Fig. 3 depicts individ- ms red
ual O, averages for the brightest red, orange, yel-
low and green stimuli. The variability between
records seems to fit the model of a more or less
continuous gradation in the distinctness of the
component configuration rather than any bimodal
distribution into colour coders and non-coders, as
suggested by earlier work (Shipley et al. 1968).
55- 80-
1 I ~ I I ~ q - q
To quantify the results, the component latencies d b d
and amplitudes were measured from the individual
Fig. 5. Mean latency+ standard error of the onset and peak of
wave forms by means of an interactive cursor component N87 for all red stimuli. The values for dark red
display. Fig. 4 illustrates the group mean ampli- ha,~e been omitted, as N87 was too small for reliable latency
tudes and standard errors of component N87 at O, t11c;LMirelllelllS (see Fig. 4).
114 W.M. PAULUS ET AL.

~
P120- AMPLITUDE Oz POz Pz Fz
-100

0-- I
b yellow
I ,T,
' \
I I
d
I O~--
1
~ red d
%

P 120
1- I1 ~'.
2-
0
,,' uV 3--
3-
/.,- ' L 2--
5-
red
5-
6- 6-
%
7-
blV +100
Fig. 6. Mean amplitude in microvolts of component P120 for Fig. 8. Relative amplitudes of components N87 and P120
all yellow and red stimuli, labelled as in Fig. 4. across all 4 electrodes.

yellow and red stimuli. In the case of the red Discussion

stimuli, this component is masked by the predomi-
nant N87 component until a luminance decrement
of at least 50% from the equally bright red is
presented. For the yellow stimuli, luminance incre-
ments of 15% and 50% yield the same amplitude of
P120 as the corresponding decrements. No really
systematic changes in the latency of P120 for the
yellow stimuli could be demonstrated (Fig. 7).
Topographically, both N87 and P120 show a
clear-cut occipital maximum with a comparable
decrease in amplitude in more anterior leads. P120
even shows a definite polarity inversion at F, (Fig.
8).
P 1 2 0 - LATENCY
yellow
ms
125-
115--
I 1 I 1
b "
Fig. 7. Latency of PI20 for all yellow stimuli, labelled as in Fig.
7.
These experiments distinguish the different ef-
fects of colour change, achromatic brightness
change and colour plus brightness change on the
VEP in man. When a coloured stimulus is viewed,
both achromatic (luminance) and chromatic sys-
tems contribute to the subjective sensation of
brightness (Guth et al. 1969). When stimuli of
different colours are compared, those which are
equal in luminance may not evoke equal sensa-
tions of brightness 4. In an attempt to produce
pure colour changes, the colour stimuli, compris-
ing orange and green colours of equal saturation
and a red with a saturation about 2.5 times higher,
were matched with the reference yellow by hetero-
chromatic brightness matching to provide stimuli
of equal subjective brightness. In the authors'
opinion this is the optimum method of brightness
matching in recording transient evoked potentials
to colour, as compared to flicker photometry,
which depends in a major way on the temporal
behaviour of retinal colour opponent cells (Gouras
and Zrenner 1979) and tests mainly achromatic
properties because of the relatively high flicker
rate (see also footnote 5). The inclusion of stimuli
with additional luminance increments and decre-
ments made it possible to differentiate between
I 1
d
4 See note on additivity failure in Appendix.
C O L O U R A N D B R I G H T N E S S C O M P O N E N T S OF VEPs
predominantly colour-evoked components and
luminance-evoked components.
Large inter-individual differences (up to 100%
according to Wagner and Boynton 1972) com-
plicate heterochromatic brightness matching. Our
approach was to apply a single set of stimuli,
determined by the match of the trained observer
W.P,, to the whole group of subjects. The hetero-
chromatic brightness match of W.P. stands in close
relation to the Uniform Colour Scale data (Fig. 1)
which provide the best currently available estima-
tion of equal brightness of different colours.
In the present results the first small component,
P55, is followed by a colour-dominated compo-
nent, N87, which is present for red, orange and
green stimuli and is strongly accentuated, at least
for red and orange, if luminance increments are
incorporated.
For saturated red stimuli with large brightness
increase N87 predominates, so that all other com-
ponents up to 200 msec are obliterated in the
response. P120 seems to be a predominantly lumi-
nance-mediated component, as may be seen from
Fig. 2, where the yellow increment as well as the
decrement stimuli produce a response dominated
almost exclusively by this component. P120 is also
present for luminance changes in the colour stimuli,
but more evident for green than for red, where it is
progressively masked by the negative prolongation
of N87, as already described.
The frontally dominant component, P250, is
probably related to further cortical processing. Its
amplitude seems to depend in part on stimulus
discriminability, since it shows increasing ampli-
tude with increasing brightness steps or colour
saturation. It may represent the P3a component of
the task-related cognitive ERPs since the random-
ized presentation of 26 different stimuli means
that any particular one of them is relatively un-
expected.
As compared with the colour onset potentials
the early offset potentials are smaller but appear
to show a similar colour-related negative peak,
N95 off, and a distinct luminance-related positive
peak, P125 off. Both have an occipital maximum.
The components P55 and N87 are among the
shortest latency components described in human
VEP literature (except those components gener-
115
ated in the retina or lateral geniculate) and can be
correlated with the early processing in the visual
cortex. This is well seen for bright red responses
which are closely similar in shape to those re-
corded in monkeys by Padmos and Van Norren
(1975). Their small component P1 (50-65 msec)
could be shown in an earlier single cell investiga-
tion by Gouras and Padmos (1974) to represent
the arrival of the lateral geniculate signals at the
cortex 5. Padmos and Van Norren's (1975) N1
(65-100 msec), is almost identical to our N87,
although the former shows a more rapidly declin-
ing trailing edge and is about 10 times larger,
presumably due to the absence of skull attenua-
tion. Their P2 (90-130 msec) corresponds closely
to our P120.
An important observation, illustrated in Fig. 2,
is that VEPs to red and green colours have com-
parable wave forms and latencies when the stimuli
are equal in brightness to the reference yellow,
although the amplitude is somewhat larger for red.
However, with the addition of brightness incre-
ments to the tegt stimuli the colour-related N87
component behaves differently for red compared
to green colours, becoming much enhanced for the
former. With the brightest stimuli N87 has a com-
pletely different time-course for red and green
responses up to 150 msec, although in photometric
terms the + 100% red stimulus (26 c d / m 2) differs
less in luminance from the reference yellow (21.5
c d / m 2) than the red of matched brightness does
(13.0 cd/m2). It appears therefore that past re-
ports of different specific red and green compo-
nents are probably due to brightness changes and
not to pure colour stimulation (Wilson et al. 1981).
Larger components for red stimuli in comparison
to other colours seem to be a consistent finding by
many authors (Padmos and Van Norren 1975;
s The onset latency of the firing of red/green colour opponent
retinal ganglion cells following stimulation of the centre of their
receptive fields lies between 20 and 45 msec for low flicker
rates, decreasing with additional stimulus intensity (Gouras
and Zrenner 1979). The findings by these authors, that cells
showing clear green/red opponent properties with stimulus
durations of 200 msec lose their opponent properties for flashes
of 10 msec duration, may explain why methods based on colour
flashes have encountered more difficulty in getting reliable
results (e.g., White et al. 1977).
116
White et al. 1977; Novikova et al. 1980).
Comparable components to N87 have been re-
ported by several workers in addition to Padmos
and Van Norren as already discussed. White et al.
(1977, 1979) and Novikova et al. (1980) recorded a
similar peak for bright onset flashes. Kellermann
and Adachi-Usami (1972/73) (using red, green
and blue colour onset flashes) recorded a negative
component at around 80 msec and a positive
component at around 105 msec but the negative
wave was much less prominent than in the present
study. As they used a strongly adapting chromatic
background, for the purposes of isolating cone
mechanisms, it may be that N87 is better seen for
a transition from an unsaturated colour or white.
The greater part of the experimental work on
colour VEPs is, however, difficult to correlate be-
cause of widely differing experimental aims and
designs, as summarized in a recent review by
Zrenner (1983). Such, for example, are the experi-
ments aimed at the detection of signals from dif-
ferent cones, investigation of spectral sensitivity
functions, examination of congenital and acquired
colour vision deficiencies, investigation of the
peculiarities of the blue sensitive mechanism in
VERs and other aspects. The importance of the
particular methodology employed is evident in
Padmos and Van Norren's (1975) statement that
the mere withdrawal of the achromatic back-
ground causes a delay of 20 msec without altering
the shape of the VEP. The lack of a background,
together with the prolonged ganglion cell response
to short flashes (Zrenner 1983) may, at least in
part, be responsible for the later components found
by previous authors (Perry et al. 1969; greater than
215 msec). Shipley et al. (1968) found configura-
tion shifts between bright red and green colours
similar to those in Fig. 2, starting, however, at
around 175 msec.
Unstructured stimuli may result in weaker re-
sponses in comparison with checkerboard fields,
but they help to avoid complications due to chro-
matic aberration (Regan 1972). White et al. (1979)
also used unstructured test fields after their own
and previous findings (Shipley et al. 1965) had
shown that the VEP responses following check-
erboard stimulation persisted regardless of its col-
our. This has been supported by Spekreijse et al.
W . M . P A U L U S ET A L .
(1977) and recently by Estdvez and Dijkhuis (1983).
Recent electrophysiological data suggest that, un-
der conditions of constant light adaptation, the
luminance of a brief stimulus is more reliably
coded by cells without orientation specificity
(Maguire and Baizer 1982). Their cortical response
latencies also shortened with increasing luminance,
as suggested by Fig. 5.
It is interesting to speculate on the electrophysi-
ological correlates of the colour-related compo-
nents in the light of recent work. The stronger N87
component, which can be elicited with bright red
as compared to pure colour or luminance stimuli,
may be due to inherent properties of the receptive
fields of opponent colour cells (Paulus and
KrOger-Paulus 1983). Pure unstructured luminance
stimuli produce almost no response from an oppo-
nent X-type unit (Zrenner 1983), which in contrast
reacts much better either to luminance contrast or
to the unbalancing of the receptive fields centre
and surround with chromatic stimuli. An addi-
tional luminance component for coloured stimuli
will cause an even greater imbalance.
Recent electrophysiological work by Michael
(1978a, b, 1981), indicates that geniculate fibres
end on double opponent colour-coded units 6
organized in colour-coded columns in the visual
cortex. At the next stage of cortical colour process-
ing, simple colour-coded cells with an orientation-
specific receptive field structure can be found.
According to Michael (1978a, b), however, double
opponent cells are the first processing stage for
chromatic information, which is bypassed by
achromatic signals. The earlier colour-related N87
may in some way reflect these early stages of
colour processing in the visual cortex. There re-
mains, however, the difficulty of explaining the
difference in the VEPs to bright red and bright
green.
In psychophysical terms it may be seen from
the results of the heterochromatic brightness
matching that there is a greater contribution to the
perceived brightness from the chromatic systems
for the red and orange stimuli than for the green
stimulus, in which the conditions for brightness
6 T h e s e are u n i t s in w h i c h b o t h the c e n t r e a n d s u r r o u n d of the
receptive field r e s p o n d r e c i p r o c a l l y to c o m p l e m c n l a r y colours.
C O L O U R A N D B R I G H T N E S S C O M P O N E N T S OF VEPs
matching with the reference yellow were at almost
equal luminance. This correlates well with the
observation that the main effects of additional
brightness increments are seen in the colour-re-
lated N87 component for the red colour stimuli,
while for the green stimuli they are mainly seen in
the luminance-related P120 component. The EP
method appears to offer a means of measuring the
relative magnitude of colour brightness and lumi-
nance components in different test stimuli. It re-
mains to be seen how well such measurements will
correlate with the subjective sensation of bright-
ness.
Summa~
The different effects of colour and brightness
on the transient visual evoked potentials to a
foveal stimulus have been investigated in a psy-
cho-physically controlled stimulus set-up, in which
equally bright red, orange and green stimuli were
substituted for a standard yellow stimulus. These
colour-evoked responses were compared with the
responses to additional brightness increments and
decrements of each of the colour stimuli. An initial
component of small amplitude, P55, was followed
by a colour-dominated component, N87, and a
luminance-dominated component, P120, with a
maximum at the occipital electrode. Both N87 and
P120 showed a decline in amplitude at the parietal
electrode and P120 had a reversed polarity at Fz.
These results indicate that the responses to equally
bright green and red stimuli have closely similar
wave forms, but that this changes rapidly with
additional brightness differences. Comparison with
the reports of subdural recording of colour-evoked
potentials in the macaque striate cortex suggests
that P55 corresponds with the primary excitation
via geniculo-cortical fibres and that N87 and P120
represent later stages of cortical processing.
117
Resume
Composantes correspondant gtla brillance et h la
couleur clans les potentiels kvoqubs visuels fovbaux
chez l'homme
Les diff6rents effets de la couleur et de la
brillance sur les potentiels 6voqu6s visuels tran-
sitoires ~ un stimulus fov6al ont 6t6 6tudi6s dans
un dispositif de stimulations contr616s psycho-
physiquement et dans laquelle des stimulus rouges,
oranges et verts de mEme luminance furent sub-
stitu6s au stimulus jaune standard. Ces r6ponses
6voqu6es aux couleurs furent compar6es aux
r6ponses a des augmentations et diminutions de
brillance, de chaque stimulus color& Une com-
posante initiale de faible amplitude, P55, 6tait
suivie par une composante principalement li6e ~ la
couleur, N87, et une composante principalement
li6e ~ la luminance, P120, avec un maximum sous
l'61ectrode occipitale. Les deux, N87 et P120,
pr6sent6rent une baisse d ' a m p l i t u d e sous
l'61ectrode pari6tate et P120 une inversion de
polarit6 sous F~. Ces r6sultats indiquent que les
r6ponses aux stimulus rouge et vert de brillance
6gale ont des ondes tr6s similaires mais que celles-ci
changent tr6s vite lorsque des modifications ad-
ditionnelles de brillance interviennent. La com-
paraison avec des rapports d'enregistrements sub-
duraux de potentiels 6voqu6s "h la couleur dans le
cortex stri6 du macaque sugg6re que P55 corre-
spond h l'excitation primaire par les fibres
g6niculo-corticales et que N87 et P120 repr6sentent
des 6tapes plus tardives de l'int6gration corticale.
Appendix
Definitions according to Wyszecki and Stiles
(1982):
LUMINANCE in c d / m 2 is the areal density of
luminous intensity and is defined in photometric
terms. It depends on the spectral sensitivity func-
tion V (?~) adopted in the Standard Normal Ob-
server CIE 1931 (quantitative data in Wyszecki
and Stiles 1982).
118
B R I G H T N E S S is defined as the attribute of a
visual sensation according to which a given visual
stimulus appears to be more or less intense. Lumi-
nance and brightness may be related by a power
law:
B = a Lp - - BO
B denotes a measure of the brightness estimated
by an observer presented with a stimulus of lumi-
nance L under specified conditions of observation.
The exponent p has a value of approximately 1/3.
The values of a and BO depend on the observing
conditions. This relation, however, is only valid for
achromatic stimuli because of:
A D D I T I V I T Y F A I L U R E : if in a bipartite field,
a chromatic stimulus C1 is equated in brightness
with a white stimulus W and this procedure is
repeated for a second stimulus C2 with an oppo-
nent hue, the additive mixture of C1 and C2 is
found to be less bright than the white reference of
twice its original radiance. Additivity failure is
involved in the derivation of luminance, which
depends on the luminosity function V (X), which
in turn has been derived predominantly by flicker
photometry and not by heterochromatic brightness
matching. If flicker photometry is done with low
flicker rates (about 1 2 Hz) the resulting spectral
sensitivity is much broader than the one derived
with high flicker rates and has 3 peaks instead of
one (Zrenner 1983). (For a critical review of the
CIE luminosity data see Est6vez 1979.)
Additivity failure is most pronounced for col-
ours at the two ends of the spectrum, namely blue
and red (Guth et al. 1969). It causes declining
luminance values for the equally bright red and
orange stimuli to the reference yellow in Fig. 2,
while the luminance for equally bright green is
almost unaffected. Colour spaces such as the Uni-
form Colour Scales (MacAdam 1974, one row of
which is incorporated in Fig. 1) try to quantify this
relation. Other authors use the term CHRO-
MATIC BRIGHTNESS for A D D 1 T I V 1 T Y
F A I L U R E (e.g., Ingling et al. 1977). For a com-
parison of 4 methods of heterochromatic photome-
try see Wagner and Boynton (1972).
SATURATION is the attribute of a visual
W.M. PAULUS ET AL.
sensation which permits a judgement to be made
of the degree to which a chromatic stimulus differs
from an achromatic stimulus regardless of their
brightness (Wyszecki and Stiles 1982). A satura-
tion scale is one in which colours vary in paleness
or their white content. The spectral hues are called
saturated colours, whereas white is called an un-
saturated colour. The spectral colours are not
themselves seen as being equal in saturation: spec-
tral blues and reds appear more saturated than
spectral greens and yellows. An unsaturated colour
may be specified as a mixture of a spectral colour
and white. The term colorimetric purity, p, has
been defined for such mixtures as the ratio p =
Lx/(L~, + Lwx), where L x is the luminance of the
spectral component and Lw x is the luminance of
the white component. Saturation discrimination
usually refers to the measurement of least colori-
metric purity (Ap) or the minimal amount of
spectral wavelength that allows a mixture of col-
orimetric purity p to be distinguished from a spec-
tral white (Pokorny et al. 1979).
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