Temporal asymmetry in motion masking: a shortening of the temporal impulse response function.

Morgan and Chubb observed a striking temporal asymmetry in motion masking (Vis. Res. 39 (1999) 4217). Motion was produced with a two-frame sequence of gratings presented in spatial quadrature phase; the second grating (100 ms) was presented immediately after the first grating (100 ms), with no temporal overlap. The contrast threshold for detecting the direction of motion of the stimulus pair was facilitated when the first grating was of low-contrast and the second grating was of high-contrast, but strong masking occurred when the order was reversed, so the high-contrast grating came first. We replicated this result, but showed that the masking mostly disappeared when the two gratings temporally overlapped only slightly. The high sensitivity to the precise temporal pattern of the stimulus can be explained by a small temporal 'shortening' of the temporal impulse response function (IRF) as stimulus contrast is increased. The IRF is biphasic with a negative inhibitory lobe. When the first grating has high-contrast, its flash response (owing to the shortening of the IRF) may be in a fairly strong negative phase by the time that the positive response to the second, lower-contrast grating has reached appreciable strength--this reduces the magnitude of the motion signal generated by the two flashes and can account for the masking. A shortening of the IRF with increased contrast (a nonlinearity) is supported by psychophysical studies in humans and by recordings of magnocellular retinal ganglion cells in macaque, and the present results bolster this concept.

Human temporal impulse response speeds up with increased stimulus contrast.

It is well known that raising mean luminance speeds-up the visual response to temporal change. At higher mean luminance, the temporal impulse response function (IRF) becomes more transient or biphasic. An analogous effect is observed physiologically when stimulus contrast is increased, at constant mean luminance. As stimulus contrast is raised, the temporal response to flicker advances in phase and becomes more transient (bandpass). The MC (magnocellular) retinal ganglion cells manifest this temporal contrast gain control, but the PC (parvocellular) cells do not. We show psychophysically that the temporal response in humans speeds-up in an analogous manner as stimulus contrast is raised. Low spatial-frequency gratings, of suprathreshold contrast, were presented as pairs of pulses, separated by brief delays. Responses became more transient with increasing contrast in both our motion task (direction discrimination) and in our flicker task ('agitation' discrimination), mimicking the temporal contrast gain control seen in the physiological studies. Results could be modeled with a nonlinearity, in which the IRF shortens with increasing contrast.

Colour adaptation modifies the temporal properties of the long- and middle-wave cone signals in the human luminance mechanism.

The human luminance mechanism (LUM) detects rapid flicker and motion, summating the neurally integrated L' and M' 'contrast' signals from the long- and middle-wave cones, respectively. We previously observed large temporal phase shifts between the L' and M' signals in LUM, which were maximal and of reversed sign on green versus orange background fields and which were accompanied by large variations in the relative L' and M' contrast weights. The effects were modelled with phasic magnocellar retinal ganglion cells. The changing L' versus M' contrast weights in the model predict that the temporal dynamics of the L' and M' luminance signals will differ on green and orange fields. This is assessed with several protocols. Motion thresholds for 1 cycle deg(-1) drifting gratings or static pulsed gratings on the orange field show that the M' signal is more temporally bandpass than the L' signal; this reverses on the green field. Strong motion due to the different dynamics of the L' and M' signals is even seen with a pair of L' and M' gratings pulsed simultaneously. Impulse response functions were measured with gratings pulsed spatially in phase or antiphase. The impulse response was clearly biphasic for the M' signal on the orange field and L' signal on the green field, while the other signals were more sustained. The impulse responses predicted the motion seen with gratings pulsed in spatial quadrature.

Spatial masking does not reveal mechanisms selective to combined luminance and red-green color.

Detection thresholds plotted in the L and M cone-contrast plane have shown that there are two primary detection mechanisms, a red-green hue mechanism and a light-dark luminance mechanism. However, previous masking results suggest there may be additional mechanisms, responsive to combined features like bright and red or dark and green. We measured detection thresholds for a 1.2 c deg-1 sine-wave grating in the presence of a spatially matched mask grating which was either stationary, dynamically jittered or flickered. The stimuli could be set to any direction in the L,M plane. The appearance of selectivity for combined hue and luminance arose only in conditions where adding the test to the mask modified the spatial phase offset between the luminance and red-green stimulus components. Sensitivity was very high for detecting this spatial phase offset. When this extra cue was eliminated, masking contours in the L,M plane could be largely described by the classical red-green and luminance mechanisms.

Second-site adaptation in the red-green detection pathway: only elicited by low-spatial-frequency test stimuli.

The red-green (RG) detection mechanism was revealed by measuring threshold detection contours in the L and M cone contrast plane for sine-wave test gratings of 0.8-6 c deg-1 on bright adapting fields of yellow or red. The slope of the RG detection contours was unity, indicating that the L and M contrast signals contribute equally (with opposite signs) on both the yellow and the red fields: this reflects first-site, cone-selective adaptation. Second-site adaptation, which may reflect saturation at a color-opponent site, was evidenced by the RG detection contours being further out from the origin of the cone contrast plane on the red field than on the yellow field. Second-site adaptation was strong (3-fold) for low spatial frequency test gratings but greatly diminished by 6 c deg-1. The disappearance of second-site adaptation with increasing spatial frequency can be explained by spatial frequency channels. The most sensitive detectors may comprise a low spatial frequency channel which is susceptible to masking by the chromatic, spatial DC component of the red field. The 6 c deg-1 patterns may be detected by a less sensitive, higher frequency channel which is less affected by the uniform red field. The RG spatial frequency channels likely arise in the cortex, implicating a partially central site for the second-site effect.

Detection of flickering edges: absence of a red-green edge detector.

Kelly ((1975) Science, 188, 371-372) showed that a centrally-fixated, contrast-reversing edge has a very different effect on the detection of luminance and red-green flicker. Red-green flicker sensitivity was approximately 3-fold greater for a uniform field than for a 'split' field with the two sides flickering out-of-phase. Just the opposite effects were observed for luminance flicker--the split field yielded a 7-fold advantage over the uniform field at 2 or 4 Hz and a 3-fold advantage at 12 Hz. Contrary to Kelly, we find that the split field offers only a very small advantage of 40% for luminance flicker at 2 Hz and virtually no advantage at 4 Hz and above. Kelly's chromatic results are surprising since one might expect that the larger color difference (or step) across the central edge would aid chromatic discrimination rather than strongly suppressing sensitivity. We show that the central chromatic edge only weakly impairs detection. Further results show that the two sides of the chromatic split field are detected essentially independently by red or green 'blob' detectors, which do not take advantage of the color difference across the edge. This has a remarkable implication: when wavelength discrimination is measured with a bipartite field whose two side are slowly modulated in opposite directions, then one side may be deleted with little adverse effect.

Facilitation between the luminance and red-green detection mechanisms: enhancing contrast differences across edges.

Previous studies have shown that detection of a red-green test pattern, such as a spot or grating, may be facilitated two to three times by a suprathreshold luminance pedestal of the same shape. We measured facilitation between the red-green (RG) and luminance (LUM) detection mechanisms using sine and square-wave gratings. Facilitation of RG by luminance pedestals was 3-fold for in phase sine-wave gratings of 0.8 cpd and a remarkable 7-fold for square-wave gratings. The latter facilitation was greatly reduced at intermediate relative phases and was generally reduced at higher spatial frequencies. We show that on a uniform field, the red or green regions of low spatial frequency test patterns are detected approximately independently, but in the presence of the LUM pedestal RG becomes sensitive to the red-green difference across the luminance edges. Under optimal conditions (with the low-frequency, square-wave luminance pedestal) this increased red-green sensitivity corresponds to a wavelength discrimination threshold as small as approximately 0.04 nm. This conversion of RG into an 'edge detector' may explain why facilitation is twice as large for square-wave gratings (bipolar patterns) than spots (unipolar patterns). The reverse facilitation, that of LUM by the red-green pedestal, is weaker and the results suggest that this is because LUM is initially sensitive to the light-dark difference across luminance edges even in the absence of the red-green pedestal.

Motion detection on flashed, stationary pedestal gratings: evidence for an opponent-motion mechanism.

Contrast thresholds were measured for discriminating left vs right motion of a vertical, 1 c/deg luminance grating lasting for one cycle of motion. This test was presented on a 1 c/deg stationary grating (pedestal) of twice-threshold, flashed for the duration of the test motion. Lu and Sperling [(1995). Vision Research, 35, 2697-2722] argue that the visual system detects the underlying, first-order motion of the test and is immune to the presence of the stationary pedestal (and the 'feature wobble' which it induces). On the contrary, we observe that the stationary pedestal has large effects on motion detection at 7 and 15 Hz, and smaller effects at 0.9-3.7 Hz, evidenced by a spatial phase dependency between the stationary pedestal and moving test. At 15 Hz the motion threshold drops as much as five-fold, with the stationary pedestal in the optimal spatial phase (i.e., pedestal and test spatially in phase at middle of motion), and the perceived direction of the test motion reverses with the pedestal in the opposite phase. Phase dependency was also explored using a very brief (approximately 1 msec) static pedestal presented with the moving test. The pedestal of Lu and Sperling (flashed for the duration of the test) has a broad spectrum of left and right moving components which interact with the moving test. The pedestal effects can be explained by the visual system's much higher sensitivity to the difference of the contrast of right vs left moving components than to either component alone.

Short-wave cone signal in the red-green detection mechanism.

Previous work shows that the red-green (RG) detection mechanism is highly sensitive, responding to equal and opposite long-wave (L) and middle-wave (M) cone contrast signals. This mechanism mediates red-green hue judgements under many conditions. We show that the RG detection mechanism also receives a weak input from the short-wave (S) cones that supports the L signal and equally opposes M. This was demonstrated with a pedestal paradigm, in which weak S cone flicker facilitates discrimination and detection of red-green flicker. Also, a near-threshold +S cone flash facilitates detection of red flashes and inhibits green flashes, and a near-threshold -S cone flash facilitates detection of green flashes and inhibits red flashes. The S contrast weight in RG is small relative to the L and M contrast weights. However, a comparison of our results with other studies suggests that the strength of the absolute S cone contrast contribution to the RG detection mechanism is 1/4 to 1/3 the strength of the S contribution to the blue-yellow (BY) detection mechanism. Thus, the S weight in RG is a significant fraction of the S weight in BY. This has important implications for the 'cardinal' color mechanisms, for it predicts that for detection or discrimination, the mechanisms limiting performance do not lie on orthogonal M-L and S axes within the equiluminant color plane.

Colour adaptation modifies the long-wave versus middle-wave cone weights and temporal phases in human luminance (but not red-green) mechanism.

1. The human luminance (LUM) mechanism detects rapid flicker and motion, responding to a linear sum of contrast signals, L' and M', from the long-wave (L) and middle-wave (M) cones. The red-green mechanism detects hue variations, responding to a linear difference of L' and M' contrast signals. 2. The two detection mechanisms were isolated to assess how 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 summation). 3. Within the red-green mechanism at 6 Hz, the phase shift between the L' and M' signals was negligible on each coloured field, and the L' and M' contrast weights were equal and of opposite sign. 4. Relative phase shifts between the L' and M' signals in the LUM mechanism were 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 (approximately 570 nm). On an orange field, 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 reveals a spectrally opponent process which controls the phase shift. The phase shift occurs at an early site, for colour adaptation of the other eye had no effect, and the phase shift measured monocularly was identical for flicker and motion, thus occurring before the motion signal is extracted (this requires an extra delay). 5. The L' versus M' phase shift in the LUM mechanism was generally greatest at intermediate temporal frequencies (4-12 Hz) and was small at high frequencies (20-25 Hz). The phase shift was greatest at low spatial frequencies and strongly reduced at high spatial frequencies (5 cycle deg-1), indicating that the receptive field surround of neurones is important for the phase shift. 6. These temporal phase shifts were confirmed by measuring motion contrast thresholds for drifting L cone and M cone gratings summed in different spatial phases. Owing to the large phase shifts on green or orange fields, the L and M components were detected about equally 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 thought to produce an equiluminant red-green grating. 7. At low spatial frequency, the relative L' and M' contrast weights in the LUM mechanism (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, with chromatic adaptation strongly modifying the receptive field surround. The cells summate L' and M' in their centre, while the surround L' and M' signals are both antagonistic to the centre for approximately 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 field (to explain the chromatic reversal of the phase shift). Large changes in the relative L' and M' weights on green versus orange fields indicate the clear presence of the spectrally opponent surround even at 20 Hz. The spectrally opponent surround appears sluggish, with a long delay (approximately 20 ms) relative to the centre.

Human cones appear to adapt at low light levels: measurements on the red-green detection mechanism.

Recent physiological evidence suggests that cones do not light adapt at low light levels. To assess whether adaptation is cone-selective at low light levels, the red-green detection mechanism was isolated. Thresholds were measured with a large test flash, which stimulated the L and M cones in different fixed amplitude ratios, on different colored adapting fields. Thresholds were plotted in L and M cone contrast coordinates. The red-green mechanism responded to an equally-weighted difference of L and M cone contrast on each colored field, demonstrating equivalent, Weberian adaptation of the L and M cone signals. The L and M cone signals independently adapted for illuminance levels as low as 60 effective trolands (e.g. M-cone trolands). Since this adaptation is entirely selective to cone type, it suggests that the cones themselves light-adapt. The red-green detection contour on reddish fields was displaced further out from the origin of the cone contrast coordinates, revealing an additional sensitivity loss at a subsequent, spectrally-opponent site. This second-site effect may arise from a net "red" or "green" signal that represents the degree to which the L and M cones are differently hyperpolarized by the steady, colored adapting field. Such differential hyperpolarization is compatible with equivalent, Weberian adaptation of the L and M cones.

Contributions of human long-wave and middle-wave cones to motion detection.

1. It has been suggested that motion may be best detected by the luminance mechanism. If this is the most sensitive mechanism, motion thresholds may be used to isolate the luminance mechanism and study its properties. 2. A moving (1 cycle deg-1), vertical, heterochromatic (red-plus-green), foveal grating was presented on a bright yellow (577 nm wavelength) field. Detection and motion (direction identification: left versus right) thresholds were measured for different amplitude ratios of the red and green components spatially summed in phase or in antiphase. Threshold contours plotted in cone-contrast co-ordinates (L',M') for the long-wave (L) and middle-wave (M) cones, revealed two motion mechanisms: a luminance mechanism that responds to a weighted sum of L and M contrasts, and a spectrally opponent mechanism that responds to a weighted difference. 3. Detection and motion thresholds, measured at 1-4 Hz, were identical for luminance gratings, having equal cone contrasts, L' and M', of the same sign. For chromatic gratings, with L' and M' of opposite sign, motion thresholds were higher than detection thresholds. A red-green hue mechanism may mediate chromatic detection, and a separate spectrally opponent motion mechanism may mediate motion. 4. The red-green hue mechanism was assessed from 1 to 15 Hz with an explicit hue criterion. The detection contour had a constant slope of one, implying equal L' and M' contributions of opposite sign. For motion identification, L' and M' contributed equally at 1 Hz, but the M' contribution was attenuated at higher velocities. 5. The cone-contrast metric provides a physiologically relevant comparison of sensitivities of the two motion mechanisms. At 1 Hz, the spectrally opponent motion mechanism is approximately 4 times more sensitive than the luminance mechanism. As temporal frequency is increased, the relative sensitivities change so that the luminance mechanism is more sensitive above 9 Hz. 6. The less sensitive motion mechanism was isolated with a quadrature phase protocol, using a pair of heterochromatic red-plus-green gratings, counterphase flickering in spatial and temporal quadrature phase with respect to each other. One grating was set slightly suprathreshold and oriented in cone contrast (L',M') so as to potentiate a single motion mechanism, the sensitivity of which was probed with the second grating, which was varied in (L',M'). This allowed us to measure the motion detection contour of the less sensitive luminance mechanism at low velocities.(ABSTRACT TRUNCATED AT 400 WORDS)

Temporal properties of the red-green chromatic mechanism.

The temporal properties of the red-green chromatic mechanism were studied with red and green equiluminant flashes of 1 deg diameter presented in the center of a bright (800-3000 td) yellow adapting field. A subthreshold 200 msec red or green flash makes an immediately subsequent, suprathreshold yellow luminance flash appear tinged with the complementary color. The chromatic flash also makes a subsequent chromatic flash of the same hue harder to detect and identify, and makes a flash of the opposite hue easier to detect and identify. These results indicate that the response of the red-green mechanism changes polarity during its time-course, suggesting that the chromatic temporal impulse-response function has a negative lobe. Pairs of chromatic pulses were used to estimate the shape of the chromatic impulse-response. The estimated impulse-response function has a zero crossing near 90 msec, followed by a long, shallow negative lobe. We also measured threshold-duration functions; the critical duration for the chromatic and luminance flashes is about 95 and 45 msec, respectively. Chromatic sensitivity (measured in cone contrast units) is 10 times greater than luminance sensitivity for long durations, and is 3 times greater for all durations less than 45 msec.

The time-course of chromatic facilitation by luminance contours.

The threshold for detecting an equiluminant chromatic spot is approximately halved by the presentation of a coincident, suprathreshold luminance pedestal flash. The dynamics of this facilitation were studied by varying the duration and temporal asynchrony of the chromatic test flash and luminance pedestal. Facilitation occurs in a narrow temporal window near the chromatic test presentation; masking may occur when the pedestal is temporally displaced from the test by longer times. The mechanism producing facilitation lags behind the chromatic signal by at least 20 msec.

The foveal color-match-area effect.

Rayleigh matches for foveal, temporally alternating fields showed only a small increase in the log green/red matching ratio (0.03--average of 10 observers) as the field was decreased from 116 to 19 min arc. This is consistent with only a small, 10%, change in photopigment density or lengthening of the cone outer segments in the central fovea. The change in matches with field size is considerably less than reported in several previous studies.

Separable red-green and luminance detectors for small flashes.

Detection contours were measured in L and M cone contrast coordinates for foveal flashes of 200 msec duration and 2.3, 5, 10 and 15 min arc diameter on a bright yellow field. The test flash consisted of simultaneous incremental and decremental red and green lights in various amplitude ratios. At all sizes, the most sensitive detection mechanism was not a luminance mechanism, but rather a red-green mechanism that responds to the linear difference of equally weighted L and M cone contrasts, and signals red or green sensations at the detection threshold. Both temporal and spatial integration were greater for red-green detection than luminance detection. A coincident, subthreshold, yellow flash (a luminance pedestal) did not affect the threshold of the red-green mechanism. Such a pedestal is a sum of equal L and M cone contrast--it represents a vector parallel to the red-green detection contour and thus is expected not to stimulate directly the red-green mechanism. When suprathreshold, the coincident pedestal facilitated chromatic detection by approximately 2x at all tested sizes; intense pedestals did not mask chromatic detection. This insensitivity to intense luminance pedestals further indicates that the red-green mechanism has fixed spectral tuning with balanced opponent L and M contrast inputs. This view of fixed spectral weights contrasts with the "variable tuning hypothesis", which postulates that the weights change with spatial-temporal variations in the test stimulus.(ABSTRACT TRUNCATED AT 250 WORDS)

Colour is what the eye sees best.

It has been argued by Watson, Barlow and Robson that the visual stimulus that humans detect best specifies the spatial-temporal structure of the receptive field of the most sensitive visual neurons. To investigate 'what the eye sees best' they used stimuli that varied in luminance alone. Because the most abundant primate retinal ganglion cells, the P cells, are colour-opponent, we might expect that a coloured pattern would also be detected well. We generalized Watson et al.'s study to include variations in colour as well as luminance. We report here that our best detected coloured stimulus was seen 5-9-fold better than our best luminance spot and 3-8-fold better than Watson's best luminance stimulus. The high sensitivity to colour is consistent with the prevalence and high colour contrast-gain of retinal P cells, and may compensate for the low chromatic contrasts typically found in natural scenes.

Peripheral chromatic sensitivity for flashes: a post-receptoral red-green asymmetry.

Thresholds of luminance and red-green chromatic flashes (200 msec) were measured on a yellow adapting field in the fovea and periphery (up to 12 degrees eccentricity for 1 degree flashes and 21 degrees eccentricity for 2 degrees flashes). Chromatic sensitivity (in cone contrast coordinates) is about 7 times higher than luminance sensitivity in the fovea but falls faster with eccentricity, so that luminance and chromatic sensitivities are similar at eccentricities of 20 degrees or less. At eccentricities greater than about 14 degrees, there is a clear asymmetry wherein green chromatic flashes are considerably less detectable than red ones. By measuring complete detection contours for many ratios of incremental and decremental red and green flashes, we isolated the red and green chromatic detection mechanisms, and demonstrated that the red-green asymmetry is not a property of the L- or M-cone response per se, but rather is a property of the post-receptoral, chromatic mechanisms. The peripheral luminance and chromatic mechanisms could be further separated with a suprathreshold luminance flash (a pedestal), since an intense pedestal masks coincident luminance test flashes but facilitates the chromatic flashes. The luminance pedestal approximately linearizes the chromatic detection function (the psychometric function).

Detection uncertainty and the facilitation of chromatic detection by luminance contours.

A suprathreshold luminance flash (1 degree, 200 msec) on a large uniform yellow field facilitates detection of a coincident (1 degree, 200 msec) red or green equiluminant flash and approximately linearizes the psychometric function for detecting the chromatic flash. The facilitation is produced by the suprathreshold contour created by the luminance flash. We tested whether the contour facilitates detection by reducing spatiotemporal uncertainty in detecting the chromatic flash. Uncertainty increase false alarms, and this effect can be factored out by correcting yes-no psychometric functions for guessing. Uncertainty also alters the shape of the receiver operating characteristic. Measurements of yes-no psychometric functions and receiver operating characteristics do not support the uncertainty reduction hypothesis.

Temporal phase response of the short-wave cone signal for color and luminance.

A chromatic discrimination paradigm was used to measure the temporal phase of the S (short-wave cone) signal relative to the L--M (long-wave cone minus middle-wave cone) signal. Suprathreshold equiluminant red-green flicker that stimulates the L--M mechanism was presented on a steady, intense yellow-green adapting field. Violet flicker that stimulates the S cones was added to the red-green flicker at different temporal phase angles, and the violet modulation depth was varied to achieve a chromatic discrimination threshold. A template was fitted to the data relating thresholds to phase: the location of the template symmetry axis showed that the S signal lagged L--M by about 75-90 degrees at 10 Hz. This is about one half the phase lag obtained for luminance or motion discrimination. The phase discrepancy shows that there are separate luminance and chromatic mechanisms receiving S cone inputs. The hue of the flicker in the present study varied strongly with phase angle, with the positive and negative excursions of the S cone signal producing a reddish-blue and greenish-yellow, respectively, and these colors combined with the reddish and greenish hues produced by the L--M signal. The observed phase shift, and measured color appearance of the combined flicker, account for the colors seen on a radially segmented disk of Munsell hues when rotated: the colors differ strikingly depending on the direction of rotation.