Luminance Gratings Research Articles

Automatic gain control contrast mechanisms are modulated by attention in humans: evidence from visual evoked potentials

This study investigated the effect of attention on the contrast response curves of steady-state visual evoked potentials (VEPs) to counter-phased sinusoidal gratings. The 1 cyc/deg gratings were modulated either in luminance or chromaticity (equiluminant red-green). The luminance grating counter-phased at 9 Hz (to favour activation of the magno-cellular system), and the chromatic grating at 2.5 Hz (to favour activation of the parvo-cellular system). Attention was directed towards the gratings (displayed in the left visual field) by requiring subjects to detect and respond to randomly occurring changes in contrast. In the control condition, attention towards the grating was minimised by requiring subjects to detect a target letter amongst distracters briefly flashed in the contra-lateral visual field. Attention increased VEP amplitudes for both luminance and chromatic stimuli, more so at high than at low contrasts, increasing the slope of the contrast amplitude curves (over the non-saturating range of contrasts). The estimates of contrast threshold from extrapolation of amplitudes were unaffected by attention. Attention also changed the VEP phases, but only for luminance gratings, where it acted to reduce the magnitude of phase advance with contrast. Attention had no effect on the average phases for chromatic gratings. The results are consistent with the notion that attention acts on cortical gain control mechanisms, which are known to be different for the magno- and parvo-cellular systems.

Electrophysiological Correlates of the Visual After Effect by Means of Visual Evoked Potentials

An attempt was made to determine whether changes of electrical activity could be seen in the posterior cortex during an after image of high frequency luminance gratings. Steady state visual evoked potentials were recorded (midoccipital, right and left temporo-occipital sites) immediately after a period of visual adaptation (15 min) to the stimulus, while the subjects experienced the after image. During this illusion, frequencies of the fast Fourier transform spectra linked to the stimulation differed from the noise and were larger at temporo-occipital sites than at the midoccipital on e. in view of these results, the hypothesis that the after effect represents a short term storage of the temporal characteristics of the stimulus is evoked.

Inter-attribute tilt effects and orientation analysis in the visual brain

A test grating appears to be tilted away from an inducing grating for small angular separations (repulsion), but towards the inducing grating for larger angular separations (attraction). Previous research on luminance gratings suggests that repulsion is caused by local inhibition in cortical areas V1 and/or V2, and that the attraction involves global interactions beyond V1, in extrastriate areas. Experiments reported here demonstrate attribute invariant attraction and repulsion effects for gratings specified by luminance, motion, and disparity contrasts. A frame surrounding the inducing grating abolishes only the attraction effect, but a spatial frequency difference, or a small gap between the inducer and test gratings, abolishes only the repulsion effect, irrespective of the attributes that specify the gratings. It is proposed that detectors selectively sensitive to attribute invariant orientation and size exist in early cortical sites such as V1 and/or V2.

The temporal properties of first- and second-order vision

Vision is sensitive to first-order modulations of luminance and second-order modulations of image contrast. There is now a body of evidence that the two types of modulation are detected by separate mechanisms. Some previous experiments on motion detection have suggested that the second-order system is quite sluggish compared to the first-order system. Here we derive temporal properties of first- and second-order vision at threshold from studies of temporal integration and two-pulse summation. Three types of modulation were tested: luminance gratings alone, luminance modulations added to dynamic visual noise, and contrast modulations of dynamic noise. Data from the two-pulse summation experiment were used to derive impulse response functions for the three types of stimulus. These were then used to predict performance in the temporal integration experiment. Temporal frequency response functions were obtained as the Fourier transform of impulse responses derived from data averaged across two observers. The response to noise-free luminance gratings of 2 c/deg was bi-phasic and transient in the time domain, and bandpass in the frequency domain. The addition of dynamic noise caused the response to become mono-phasic, sustained and low-pass. The response to contrast modulated noise (second-order) was also mono-phasic, sustained and low-pass, with only a slightly longer integration time than in the first-order case. The ultimate roll-off at high frequencies was about the same as for the first-order case. We conclude that second-order vision may not be as sluggish as previously thought.

The temporal course of suppression during binocular rivalry.

Orthogonally oriented sinusoidal luminance gratings were dichoptically presented to the observers' left and right eyes. During the subsequent binocular rivalry, a small target was briefly presented (4AFC) to probe the strength of interocular suppression at various temporal latencies. Both stationary and moving rivalrous patterns were investigated. The purpose of experiment 1 was to compare the temporal characteristics of stationary and motion rivalry (0 and 1.2 deg s-1), while that of experiment 2 was to examine rivalry suppression for higher speeds (2 and 4 deg s-1). In all cases, it was found that the strength of suppression remained essentially constant throughout a single phase of binocular rivalry. The results of the investigation also revealed that moving rivalrous patterns lead to greater magnitudes of interocular suppression than static patterns. Despite these differences in the strength of suppression, the results of both experiments show that the temporal characteristics of motion and static rivalry are essentially identical.

Local Disparity Not Perceived Depth Is Signaled by Binocular Neurons in Cortical Area V1 of the Macaque

Binocular neurons that are closely related to depth perception should respond selectively for stimuli eliciting an appropriate depth sensation. To separate perceived depth from local disparity within the receptive field, sinusoidal luminance gratings were presented within a circular aperture. The disparity of the aperture was coupled to that of the grating, thereby rendering unambiguous the psychophysical matching between repeating cycles of the grating. In cases in which the stimulus disparity differs by one horizontal period of the grating, the portion of the grating that locally covers a receptive field is binocularly identical, but the depth sensation is very different because of the aperture. For 117 disparity-selective V1 neurons tested in two monkeys, the overwhelming majority responded equally well to configurations that were locally identical but led to different perceptions of depth. Because the psychophysical sensation is not reflected in the firing rate of V1 neurons, the signals that make stereo matches explicit are most likely elaborated in extrastriate cortex.

Channel selection with non-white-noise masks.

It is frequently assumed that the accuracy with which luminance gratings can be detected depends solely on the signal-to-noise ratio at the output of a single linear channel. Proportionality between threshold elevation and power spectral density is implicit in this assumption. I demonstrate that this proportionality does not hold for 1-cycle/degree gratings masked by low-pass noise with a 0.5-cycle/degree cutoff frequency. This implies that different channels can mediate detection, depending on the contrast of masking stimuli.

Linear coupling between cerebral blood flow and oxygen consumption in activated human cortex.

The aim of this study was to test the hypothesis that, within a specific cortical unit, fractional changes in cerebral blood flow (CBF) and cerebral metabolic rate of oxygen consumption (CMR(O(2))) are coupled through an invariant relationship during physiological stimulation. This aim was achieved by simultaneously measuring relative changes in these quantities in human primary visual cortex (V1) during graded stimulation with patterns designed to selectively activate different populations of V1 neurons. Primary visual cortex was delineated individually in each subject by using phase-encoded retinotopic mapping. Flow-sensitive alternating inversion recovery MRI, in conjunction with blood oxygenation-sensitive MRI and hypercapnic calibration, was used to monitor CBF and CMR(O(2)). The stimuli used included (i) diffuse isoluminant chromatic displays; (ii) high spatial-frequency achromatic luminance gratings; and (iii) radial checkerboard patterns containing both color and luminance contrast modulated at different temporal rates. Perfusion responses to each pattern were graded by varying luminance and/or color modulation amplitudes. For all stimulus types, fractional changes in blood flow and oxygen uptake were found to be linearly coupled in a consistent ratio of approximately 2:1. The most potent stimulus produced CBF and CMR(O(2)) increases of 48 +/- 5% and 25 +/- 4%, respectively, with no evidence of a plateau for oxygen consumption. Estimation of aerobic ATP yields from the observed CMR(O(2)) increases and comparison with the maximum possible anaerobic ATP contribution indicate that elevated energy demands during brain activation are met largely through oxidative metabolism.

Stimulus-Dependent BOLD and Perfusion Dynamics in Human V1

Blood oxygenation level-dependent (BOLD) fMRI signals often exhibit pronounced over- or undershoot upon changes in stimulation state. Current models postulate that this is due to the delayed onset or decay of perfusion-dependent attenuating responses such as increased cerebral blood volume or oxygen consumption, which are presumed to lag behind the rapid adjustment of blood flow rate to a new steady-state level. If this view is correct, then BOLD overshoot amplitudes in a specific tissue volume should be correlated with steady-state increases in perfusion, independent of stimulus type. To test this prediction, we simultaneously recorded BOLD and relative perfusion signals in primary visual cortex while inducing graded perfusion increases with three types of visual stimulus. Two of these, a diffuse chromatic stimulus with no luminance variation and a very high spatial frequency luminance grating, did not produce detectable BOLD overshoot (or undershoot) when an equal mean luminance baseline was used. Radial checkerboard stimuli, however, caused pronounced over/undershoot of both BOLD and perfusion signals even when temporal mean luminance was held constant and stimulus contrast was adjusted to produce the same steady-state blood flow increases evoked by the other stimuli. Transient amplitudes were relatively invariant in spite of large changes in steady-state response, demonstrating nonlinear BOLD and perfusion step responses in human V1. These findings suggest that, rather than a purely tissue-specific biomechanical or metabolic phenomenon, BOLD overshoot and undershoot represent transient features in the perfusion signal whose effects may be amplified by slowly evolving blood volume changes.

Modelling spatial contrast sensitivity functions for chromatic and luminance-modulated gratings

We extended our detection model of achromatic spatial vision (Rovamo, J., Mustonen, J., & Näsänen, R. (1994a). Modelling contrast sensitivity as a function of retinal illuminance and grating area. Vision Research, 34, 1301–1314) to colour vision by taking into account the fact that due to the spatio-chromatic opponency of retinal ganglion cells and dorsal lateral geniculate nucleus (dLGN) neurons, equiluminous chromatic gratings are not affected by precortical lateral inhibition. We then tested the extended model by using Mullen’s experimental data (Mullen, K. J. (1985). The contrast sensitivity of human color vision to red–green and blue–yellow chromatic gratings. Journal of Physiology, 359, 381–400). The band-pass shape of the spatial contrast sensitivity function for luminance-modulated green and yellow gratings transformed to a low-pass shape, resembling the chromatic spatial contrast sensitivity function for red–green and blue–yellow equiluminous gratings, when the effect of precortical lateral inhibition on grating contrast was computationally removed by dividing luminance contrast sensitivities by spatial frequency (i.e. by af, where a=1°). After the removal of this direct effect of lateral inhibition, there still remained a residual shape difference between the spatial contrast sensitivity functions for chromatic and luminance gratings. It was due to indirect reduction of grating visibility by quantal noise high-pass filtered by precortical lateral inhibition. When this indirect effect of quantal noise was also removed, contrast sensitivity for luminance gratings was about twice the sensitivity for chromatic gratings at all spatial frequencies. This was evidently due to the fact that the chromatic contrast of the equiluminous grating at the opponent stage (Cole, G. R., Hine, T. & McIihagga, W. (1993). Detection mechanisms in L-, M-, and S-cone contrast space. Journal of the Optical Society of America A, 10, 38–51) was about half of the luminance contrast of either of its chromatic component. Thus, if the contrast of the equiluminous chromatic grating were not expressed as the Michelson contrast of one chromatic component grating against its own background (Mullen, K. J. (1985). The contrast sensitivity of human color vision to red–green and blue–yellow chromatic gratings. Journal of Physiology, 359, 381–400) but as chromatic contrast at the opponent stage, contrast sensitivity would be the same for chromatic and luminance gratings.

A measurement of visual extinction by contrast threshold of luminance grating

A measurement of visual extinction by contrast threshold of luminance grating

Binocular phase interactions in area 21a of the cat.

1. Binocular interactions related to retinal disparity were investigated in single neurons in area 21a of extrastriate cortex in the anaesthetized cat using sinusoidal luminance gratings. 2. The responses of approximately two-thirds of neurons were profoundly modulated by a relative phase difference between identical drifting gratings presented to each eye. This modulation included both facilitatory and inhibitory interocular interactions. The selectivity for binocular disparity was about twice as sharp as the selectivity for monocular spatial position. 3. Significant phase modulation was retained in many neurons at interocular orientation differences exceeding 45 deg. The response suppression associated with stimulation at a phase shift 180 deg from the optimum was stronger than the response suppression to an interocular orientation difference of 90 deg. 4. The proportion of phase modulated neurons and the potency of modulation in area 21a neurons exceed that reported for phase-selective complex cells in area 17. Neurons in area 21a show sharp disparity tuning that is relatively insensitive to changes in orientation and monocular position, which suggests that this extrastriate region has a role in stereoscopic depth perception.

Evidence for a mechanism sensitive to the speed of cyclopean form1Part of the results reported here were presented at the 1997 ARVO meeting. Kohly, Hajdur & Regan (1997). There is a speed-sensitive cyclopean mechanism, Investigative Ophthalmology and Visual Science, 38, S906.1

We measured Weber fractions for discriminating the speed of cyclopean gratings and Weber fractions for discriminating the speed of luminance gratings. Of our 14 observers, five were unable to see the cyclopean grating sufficiently well to discriminate its speed. One observer experienced great difficulty in discriminating the speed of cyclopean gratings, even though her threshold for detecting cyclopean gratings was low, and even though she discriminated the speed of luminance gratings on the basis of the task-relevent variable. But several observers based their speed discriminations on trial-to-trial variations of the task-relevent variable while ignoring associated trial-to-trial variations in all task-irrelevant variables (specifically: displacement; temporal frequency; spatial frequency; and presentation duration). We conclude that the visual systems of these observers contain a specialized neural mechanism for the speed of cyclopean gratings that supports acute discriminations of speed (Weber fractions were as low as 0.05-0.07).

Temporal and spatial response to second-order stimuli in cat area 18.

Temporal and spatial response to second-order stimuli in cat area 18. J. Neurophysiol. 80: 2811-2823, 1998. Approximately one-half of the neurons in cat area 18 respond to contrast envelope stimuli, consisting of a sinewave carrier whose contrast is modulated by a drifting sinewave envelope of lower spatial frequency. These stimuli should fail to elicit a response from a conventional linear neuron because they are designed to contain no spatial frequency components within the cell's luminance-defined frequency passband. We measured neurons' responses to envelope stimuli by varying both the drift rate and spatial frequency of the contrast modulation. These data were then compared with the same neurons' spatial and temporal properties obtained with luminance-defined sinewave gratings. Most neurons' responses to the envelope stimuli were spatially and temporally bandpass, with bandwidths comparable with those measured with luminance gratings. The temporal responses of these neurons (temporal frequency tuning and latency) were systematically slower when tested with envelope stimuli than with luminance gratings. The simplest kind of model that can accommodate these results is one having separate, parallel streams of bandpass processing for luminance and envelope stimuli.

Detection of chromatic and luminance contrast modulation by the visual system.

The data presented in this paper examine the ability of observers to detect a modulation in the contrast of chromatic and luminance gratings as a function of the carrier contrast, duration, and spatial frequency. The nature of the signal underlying this ability is investigated by examining both the paradigm used to make the measurement and the effect of grating masks on performance in the tasks. The results show that observers' ability to discriminate amplitude modulation from an unmodulated carrier is dependent on carrier contrast but only up to approximately 5-8 times carrier-detection threshold. Discrimination is, however, independent of spatial frequency [10-1 cycles per degree (cpd) component-frequency range], carrier color, and, most surprisingly, stimulus duration (1000-30 ms). This set of experiments compliments data from previous papers and assimilates many of the conclusions drawn from this previous data. There is absolutely no evidence for the existence of a distortion product mediating performance under any of the current conditions, and the data seriously question whether the visual system might use such a signal even if it does exist under more extreme conditions than those used here. The evidence suggests that the visual system detects variations in both chromatic and luminance contrast by means of a mechanism operating locally upon the spatial structure of the carrier.

Chromatic and luminance interactions in spatial contrast signals.

We report VEP studies which delineate interactions between chromatic and luminance contrast signals. We examined responses to sinusoidal luminance gratings undergoing 4-Hz square-wave contrast reversal, upon which standing gratings with various admixtures of luminance and chromatic contrast were alternately superimposed and withdrawn. The presence of the standing grating induced a VEP component at the fundamental frequency of the contrast-reversal grating. This VEP component appeared without any appreciable lag, and did not vary in amplitude over the 4 s during which the standing grating was present. The observed fundamental response differed from the fundamental component that would be expected from the known interaction between the luminance component of the standing grating with the modulated grating (Bodis-Wollner et al., 1972; Bobak et al., 1988), in three ways: (1) The fundamental response was not nulled for standing gratings that were isoluminant or near-isoluminant. (2) The chromatic dependence of the fundamental response implied an S-cone input to the interaction. (3) No single mechanism (driven by a linear combination of cone signals) could account quantitatively for the size of this response, particularly when the standing grating strongly modulated two cones in phase.

Effects of contrast and temporal frequency on orientation discrimination for luminance and isoluminant stimuli.

We compared the mechanisms responsible for orientation discrimination of stimuli defined by luminance and red/green isoluminant contrast. A four-alternative forced-choice (4-AFC) paradigm was used to determine thresholds for discriminating 1 cpd sinewave gratings differing in orientation, contrast, or both. When measuring orientation thresholds as a function of stimulus contrast, we found a decrease in thresholds with increasing stimulus contrast. For three temporal frequencies (0, 1, and 8 Hz) the functions relating orientation thresholds to stimulus contrast had similar shapes for luminance and isoluminant gratings, indicating similar processing mechanisms. Thresholds for stationary or slowly moving gratings were consistently lower for isoluminant than for luminance gratings, when contrast was expressed on an absolute RMS-cone-contrast scale. When contrast was defined as multiples of detection thresholds, discrimination was slightly better for luminance gratings. Thresholds for fast moving gratings were similar, irrespective of the definition of contrast. In contrast to previous work, we found a marked "oblique-effect" for both luminance and isoluminant gratings, when measuring discrimination thresholds as a function of standard orientation. Finally, we measured discrimination thresholds for gratings that varied in contrast and orientation simultaneously. The shapes of the resulting two-dimensional threshold contours were similar for luminance and isoluminant gratings, indicating again that these stimuli undergo similar neuronal processing. Performance of the observers could be described by probability summation of the orientation and contrast cues, resulting in an elliptical shape of the two-dimensional threshold contours. In conclusion, our results show similar performance for luminance and isoluminant gratings in several orientation discrimination tasks. The similarity in shape of the different threshold functions presents strong evidence that similar mechanisms underlie orientation discrimination of luminance and isoluminant stimuli.

Asymmetric Masking: Luminance Gratings Mask Second-Order Gratings, but Not Vice Versa

Human vision can detect spatiotemporal information conveyed by first-order modulations of luminance and by second-order, non-Fourier modulations of image contrast. Models for second-order motion have suggested two filtering stages separated by a rectifying nonlinearity. We explore here the encoding of stationary first-order and second-order gratings, and their interaction. Stimuli consisted of 2-D broad-band static visual noise sinusoidally modulated in luminance (first-order, LM) or contrast (second-order, CM). Modulation thresholds were measured in a two-interval forced-choice staircase procedure. With increasing noise contrast, first-order sensitivity decreased (owing to masking) but sensitivity to contrast modulation increased. Weak background gratings present in both intervals produced order-specific facilitation: LM background facilitated LM detection (the ‘dipper function’) and CM facilitated CM detection. LM did not facilitate CM, nor vice versa, and this is strong evidence that LM and CM are detected via different mechanisms. Nevertheless, suprathreshold LM gratings masked CM detection, but not vice versa. High-amplitude CM masks had little or no effect on CM or LM detection. A broadly tuned divisive gain-control mechanism applied to the first-order filtering stage has been proposed by Foley (1994 Journal of the Optical Society of America A11 1710 – 1719) to account for masking of luminance gratings, and this might also explain the masking of second-order by first-order stimuli. First-order maskers would drive down the effective contrast of the carrier, thus reducing second-order sensitivity. But for second-order maskers the mean contrast, and hence contrast gain, remained constant, independent of modulation depth. Thus second-order gratings would produce no masking effects, as observed.

What is the Transition Point between Static and Dynamic Motion Aftereffects?

The motion aftereffect (MAE) measured with a dynamic test pattern (eg a counterphase-flickering grating) is distinguishable by a number of properties from the classical MAE obtained with a static test pattern. For a dynamic MAE, however, it is not sufficient simply to introduce dynamic properties into the test pattern. In two experiments we attempted to determine the transition point in the temporal-frequency domain at which a dynamic MAE becomes distinguishable from the static MAE. First, we examined the interocular transfer (IOT) of the MAE with conventional first-order (luminance) gratings. The amount of IOT increased with temporal frequency, and was almost complete at 1 Hz and above. In addition, the IOT of a dynamic MAE shows a drastic reduction in the peripheral visual field, possibly reflecting difficulties in feature tracking or the loss of involuntary attention. Second, we examined the MAE with second-order motion as the adaptation stimulus (contrast modulation of two-dimensional static noise). Under these conditions, similar results were obtained for first-order and second-order test gratings: MAE was not observed at low temporal frequencies and a substantial MAE was observed only at 1 Hz and above. The results agree with recent findings which showed a gradual loss of spatial-frequency selectivity with increasing temporal frequency of the test pattern (Mareschal et al, 1997 Vision Research37 1755 – 1759). The present results support the idea that two mechanisms underlie the different kinds of MAE: a low-level mechanism responsible for the MAE observed at low temporal frequencies, and a high-level mechanism operating predominantly at high temporal frequencies with a transition point at about 1 Hz.

Absence of linear subthreshold summation between red-green and luminance mechanisms over a wide range of spatio-temporal conditions.

We have tested the independence of red-green chromatic and luminance mechanisms at detection threshold using a method of subthreshold summation. Stimuli were isoluminant red-green gratings and yellow-black luminance gratings that uniquely activate the red-green color and luminance mechanisms, respectively. Stimuli were Gaussian enveloped 0.25, 1 or 4 cpd sinewave gratings, counter-phase flickered at 0, 5 or 9 Hz. The threshold detection of red-green color contrast was measured in the presence of a subthreshold amount of luminance contrast, and vice versa. The results allow a model of linear summation between the color and luminance mechanisms to be rejected, but are well fitted by a model, assuming that these mechanisms are independent but combine to determine detection by probability summation, with a high summation index (median value = 4). We conclude that there are independent red-green chromatic mechanism and luminance detection mechanisms over this range of spatio-temporal conditions.