Cf. Stromeyer et al., COLOR ADAPTATION MODIFIES THE LONG-WAVE VERSUS MIDDLE-WAVE CONE WEIGHTS AND TEMPORAL PHASES IN HUMAN LUMINANCE (BUT NOT RED-GREEN) MECHANISM, Journal of physiology, 499(1), 1997, pp. 227-254
The human luminance (LUM) mechanism detects rapid flicker and motion,
responding to a linear sum of contrast signals, L' and M', from the lo
ng-wave (L) and middle-wave (M) cones. The red-green mechanism detects
hue variations, responding to a linear difference of L' and M' contra
st signals. 2. The two detection mechanisms were isolated to assess ho
w chromatic adaptation affects summation of L' and M' signals in each
mechanism. On coloured background (from blue to red), we measured, as
a function of temporal frequency: both the relative temporal phase of
the L' and M' signals producing optimal summation and the relative L'
and M' contrast weights of the signals (at the optimal phase for summa
tion). 3. Within the red-green mechanism at 6 Hz, the phase shift betw
een the L' and M' signals was negligible on each coloured field, and t
he L' and M' contrast weights were equal and of opposite sign. 4. Rela
tive phase shifts between the L' and M' signals in the LUM mechanism w
ere markedly affected by adapting field colour. For stimuli of 1 cycle
deg(-1) and 9 Hz, the temporal phase shift was zero on a green-yellow
field (similar to 570 nm). On an orange held, the L' signal lagged M'
by as much as 70 deg phase while on a green field M' lagged L' by as
much as 70 deg. The asymmetric phase shift about yellow adaptation rev
eals a spectrally opponent process which controls the phase shift. The
phase shift occurs at an early site, for colour adaptation of the oth
er eye had no effect, and the phase shift measured monocularly was ide
ntical for flicker and motion, thus occurring before the motion signal
is extracted (this requires an extra delay). 5. The L' versus M' phas
e shift in the LUM mechanism was generally greatest at intermediate te
mporal frequencies (4-12 Hz) and was small at high frequencies (20-25
Hz). The phase shift was greatest at low spatial frequencies and stron
gly reduced at high spatial frequencies (5 cycle deg(-1)), indicating
that the receptive field surround of neurones is important for the pha
se shift. 6. These temporal phase shifts were confirmed by measuring m
otion contrast thresholds for drifting L cone and M cone gratings summ
ed in different spatial phases. Owing to the large phase shifts on gre
en or orange fields, the L and M components were detected about equall
y well by the LUM mechanism (at 1 cycle deg(-1) and 9 Hz) when summed
spatially in phase or in antiphase. Antiphase summation is typically t
hought to produce an equiluminant red-green grating. 7. At low spatial
frequency, the relative L' and M' contrast weights in the LUM mechani
sm (assessed at the optimal phase for summation) changed strongly with
field colour and temporal frequency. 8. The phase shifts and changing
contrast weights were modelled with phasic retinal ganglion cells, wi
th chromatic adaptation strongly modifying the receptive field surroun
d. The cells summate L' and M' in their centre, while the surround L'
and M' signals are both antagonistic to the centre for similar to 570
nm yellow adaptation. Green or orange adaptation is assumed to modify
the L and M surround inputs, causing them to be opponent with respect
to each other, but with reversed polarity on the green versus orange f
ield (to explain the chromatic reversal of the phase shift). Large cha
nges in the relative L' and M' weights on green versus orange fields i
ndicate the clear presence of the spectrally opponent surround even at
20 Hz. The spectrally opponent surround appears sluggish, with a long
delay (similar to 20 ms) relative to the centre.