Asymmetries between achromatic increments and decrements: Perceptual scales and discrimination thresholds.

The perceptual response to achromatic incremental (A+) and decremental (A-) visual stimuli is known to be asymmetrical, due most likely to differences between ON and OFF channels. In the current study, we further investigated this asymmetry psychophysically. In Experiment 1, maximum likelihood difference scaling (MLDS) was used to estimate separately observers' perceptual scales for A+ and A-. In Experiment 2, observers performed two spatial alternative forced choice (2SAFC) pedestal discrimination on multiple pedestal contrast levels, using all combinations of A+ and A- pedestals and tests. Both experiments showed the well-known asymmetry. The perceptual scale curves of A+ follow a modified Naka-Rushton equation, whereas those of A- follow a cubic function. Correspondingly, the discrimination thresholds for the A+ pedestal increased monotonically with pedestal contrast, whereas the thresholds of the A- pedestal first increased as the pedestal contrast increased, then decreased as the contrast became higher. We propose a model that links the results of the two experiments, in which the pedestal discrimination threshold is inversely related to the derivative of the perceptual scale curve. Our findings generally agree with Whittle's previous findings (Whittle, 1986, 1992), which also included strong asymmetry between A+ and A-. We suggest that the perception of achromatic balanced incremental and decremental (bipolar) stimuli, such as gratings or flicker, might be dominated by one polarity due to this asymmetry under some conditions.

Poster Session: Threshold versus intensity curves measured with a new high-brightness display system.

Classical threshold vs. intensity (tvi) curves were measured using optical systems and were generally limited to increment test stimuli and relatively simple spatial patterns. Modern displays provide more flexibility in terms of stimuli spatial profiles but are usually dim enough that there may be rod intrusion when measuring cone responses. Here we describe a high-brightness display system and present tvi's for increment and decrement achromatic tests. The system consists of a PROPixx three-chip DLP LED color projector (VPixx Technologies, Saint-Bruno, Canada) controlled via a Datapixx display driver, with 12-bit digital to analog conversion per RGB channel. Light from the projector is collected in a large diameter lens and focused on high gain rear projection screen. Retinal illuminance of the background may be varied in three ways: (a) varying the mean current supplied to the LEDs from the controller (adjustable in software); (b) using calibrated neutral density filters mounted near the eye; and (c) changing the midpoint of the RGB channels in software (e.g., making the white background as R=G=B=0.1 instead of 0.5). Method (c) is made easier by the fact that the PROPixx "gamma curve" is linear, which also means that no RGB bits are lost to gamma correction. We will show thresholds for achromatic tests on a white background varying from 0.56 to 4.03 log trolands, with preliminary results suggesting differences in the tvi curves between the increment and decrement tests.

Visual psychophysics: Luminance and color.

Human visual psychophysics is a mature field of research that employs specialized methods and has generated a large body of established findings, a few of which are summarized in this overview. The methods reviewed include those known as classical psychophysical methods, signal detection theory, and the efficient modern Bayesian adaptive methods. The covered results emphasize well-established findings in both rod and cone vision, including some effects of light adaptation, luminous efficiency and spectral sensitivity, color detection, and spatial and temporal contrast sensitivities.

Modeling individual variations in equiluminance settings.

Recently, we reported measurements of heterochromatic flicker photometry (HFP) in 22 young observers, with stimuli that (nominally) modulated only L- and M-cones and were kept at (approximately) a constant multiple of detection threshold. These equiluminance settings were represented as the angle in the (L, M) cone contrast plane, with the greenish peak of the flicker in quadrant II and the reddish peak in quadrant IV; equiluminance settings were reported as the greenish angle. The mean equiluminance angle was 116.3° (an M:L cone contrast ratio of -2 at equiluminance), but individual differences in the settings were substantial, with the variation across individuals almost five times larger than the within-subject precision in the settings. In the present study we sought to determine the degree to which we could account for our observers' HFP settings by plausible variations in the macular pigment optical density (MPOD), the lens pigment optical density (LPOD), the cone photopigment optical densities (PPOD), and serine/alanine polymorphism in L-cone opsin (λmax shift). Most of the range of our measured equiluminance angles could be accounted for by these factors, although the largest two angles (smallest |ΔM/M: ΔL/L| ratio at equiluminance) could not. Individual differences in HFP have sometimes been taken to indicate variations in the ratio of L:M cone number; our results suggest that most of the individual differences in HFP might be equally well ascribed to physiological factors other than cone number. Simple linear models allow predictions of equiluminance angle, cone adapting level, and artifactual S-cone contrast from the values of the four factors considered here.

Signals for defocus arise from longitudinal chromatic aberration in chick.

Chicks respond to two signals from longitudinal chromatic aberration (LCA): a wavelength defocus signal and a chromatic signal. Wavelength defocus predicts reduced axial eye growth in monochromatic short-wavelength light, compared to monochromatic long-wavelength light. Wavelength defocus may also influence growth in broadband light. In contrast, a chromatic signal predicts increased growth when short-wavelength contrast > long-wavelength contrast, but only when light is broadband. We aimed to investigate the influence of blue light, temporal frequency and contrast on these signals under broadband conditions. Starting at 12 to 13 days-old, 587 chicks were exposed to the experimental illumination conditions for three days for 8h/day and spent the remainder of their day in the dark. The stimuli were flickering lights, with a temporal frequency of 0.2 or 10 Hz, low (30%) or high contrast (80%), and a variety of ratios of cone contrast simulating the effects of defocus with LCA. There were two color conditions, with blue contrast (bPlus) and without (bMinus). Stimuli in the "bPlus" condition varied the amounts of long- (L), middle- (M_) and double (D-) cone contrast, relative to short- (S-) and (UV-) cone contrast, to simulate defocus. Stimuli in the "bMinus" condition only varied the relative modulations of the L + D vs. M cones. In all cases, the average of the stimuli was white, with an illuminance of 777 lux, with cone contrast created through temporal modulation. A Lenstar LS 900 and a Hartinger refractometer were used to measure ocular components and refraction. Wavelength defocus signals with relatively high S-cone contrast resulted in reduced axial growth, and more hyperopic refractions, under low-frequency conditions (p = 0.002), in response to the myopic defocus of blue light. Chromatic signals with relatively high S-cone contrast resulted in increased axial growth and more myopic refractions, under high frequency, low contrast, conditions (p < 0.001). We conclude that the chromatic signals from LCA are dependent on the temporal frequency, phase, and relative contrast of S-cone temporal modulation, and recommend broadband spectral and temporal environments, such as the outdoor environment, to optimize the signals-for-defocus in chick.

Methods for determining equiluminance in terms of L/M cone ratios.

Heterochromatic flicker photometry (HFP), minimum motion (MM), and minimally distinct border (MDB) settings have often been used to determine equiluminance, a relative intensity setting for two chromaticities that, in theory, eliminates the responses of a luminance or achromatic psychophysical mechanism. These settings have been taken to reflect the relative contribution of the long (L) and medium (M) wavelength cones to luminance, which varies widely across individuals. The present study compares HFP, MM, and MDB using stimuli that do not modulate the short (S) wavelength cones, in both practiced and naïve observers. MDB was performed with both flashed and steadily viewed stimuli. Results are represented in the (∆L/L, ∆M/M) plane of cone contrast space. Considering both practiced and naïve observers, both MM and HFP had excellent within-subject precision and high test-retest reliability, whereas HFP also had low between-subject variability. The MDB tasks were less reliable and less precise. The mean L:M contrast ratios at equiluminance were lower for the two temporal tasks (HFP and MM) compared to the spatial tasks (MDB), perhaps consistent with the existence of multiple luminance mechanisms. Overall, the results suggest that the best method for determining equiluminance is HFP, with MM being a close second.

Sensitivity to S-Cone Stimuli and the Development of Myopia.

Purpose: Longitudinal chromatic aberration (LCA) is a color signal available to the emmetropization process that causes greater myopic defocus of short wavelengths than long wavelengths. We measured individual differences in chromatic sensitivity to explore the role LCA may play in the development of refractive error. Methods: Forty-four observers were tested psychophysically after passing color screening tests and a questionnaire for visual defects. Refraction was measured and only subjects with myopia or hyperopia without severe astigmatism participated. Psychophysical detection thresholds for 3 cyc/deg achromatic, L-, M-, and S-cone-isolating Gabor patches and low-frequency S-cone increment (S+) and decrement (S-) blobs were measured. Parametric Pearson correlations for refractive error versus threshold were calculated and nonparametric bootstrap 95% percentage confidence intervals (BCIs) for r were computed. Results: S-cone Gabor detection thresholds were higher than achromatic, L-, and M-cone Gabors. S-cone Gabor thresholds were higher than either S+ or S- blobs. These results are consistent with studies using smaller samples of practiced observers. None of the thresholds for the Gabor stimuli were correlated with refractive error (RE). A negative correlation with RE was observed for both S+ (r = -0.28; P = 0.06; BCI: r = -0.5, -0.04) and S- (r = -0.23; P = 0.13; BCI = -0.46, 0.01) blobs, although this relationship did not reach conventional statistical significance. Conclusions: Thresholds for S+ and S- stimuli were negatively related to RE, indicating that myopes may have reduced sensitivity to low spatial frequency S-cone stimuli. This reduced S-cone sensitivity might have played a role in their failure to emmetropize normally.

A tour of contemporary color vision research.

The study of color vision encompasses many disciplines, including art, biochemistry, biophysics, brain imaging, cognitive neuroscience, color preferences, colorimetry, computer modelling, design, electrophysiology, language and cognition, molecular genetics, neuroscience, physiological optics, psychophysics and physiological optics. Coupled with the elusive nature of the subjective experience of color, this wide range of disciplines makes the study of color as challenging as it is fascinating. This overview of the special issue Color: Cone Opponency and Beyond outlines the state of the science of color, and points to some of the many questions that remain to be answered in this exciting field.

Harmonics added to a flickering light can upset the balance between ON and OFF pathways to produce illusory colors.

The neural signals generated by the light-sensitive photoreceptors in the human eye are substantially processed and recoded in the retina before being transmitted to the brain via the optic nerve. A key aspect of this recoding is the splitting of the signals within the two major cone-driven visual pathways into distinct ON and OFF branches that transmit information about increases and decreases in the neural signal around its mean level. While this separation is clearly important physiologically, its effect on perception is unclear. We have developed a model of the ON and OFF pathways in early color processing. Using this model as a guide, we can produce imbalances in the ON and OFF pathways by changing the shapes of time-varying stimulus waveforms and thus make reliable and predictable alterations to the perceived average color of the stimulus-although the physical mean of the waveforms does not change. The key components in the model are the early half-wave rectifying synapses that split retinal photoreceptor outputs into the ON and OFF pathways and later sigmoidal nonlinearities in each pathway. The ability to systematically vary the waveforms to change a perceptual quality by changing the balance of signals between the ON and OFF visual pathways provides a powerful psychophysical tool for disentangling and investigating the neural workings of human vision.

Asymmetric high-contrast masking in S cone increment and decrement pathways.

Physiological, anatomical, and psychophysical evidence points to important differences between visual processing of short-wave cone increments and decrement (S+ and S-) stimuli. The present study uses the pedestal discrimination paradigm to investigate potential differences, using S+ and S- tests presented on (L)ong-wave, (M)edium-wave, S, L+M, L-M, and achromatic pedestals, of both contrast polarities. Results show that high contrast 'purplish' (S+ or -(L+M)) pedestals produce substantially more masking of both S+ and S- tests than 'yellowish' (S- or +(L+M)) pedestals do. The other pedestals produce no masking. These findings suggest greater nonlinearity - either a static nonlinearity or contrast gain control - in the mechanisms responsible for the 'purplish' polarity, likely the S ON pathway.

Conscious awareness is necessary for affective faces to influence social judgments.

A growing body of research claims that stimuli presented outside conscious awareness can influence affect, speech perception, decision-making, eating behavior, and social judgments. However, research has shown that conscious awareness is a continuous phenomenon. Using a continuous flash suppression (CFS) paradigm to suppress awareness of affective faces (smiling and scowling), we demonstrate that some awareness of suppressed stimuli is required for the stimuli to influence social judgments. We discovered this using a rigorous within-participants psychophysics method that allowed us to assess awareness at very low levels, which is difficult using traditional methods. Our findings place boundary conditions on claims (made previously by us and others) that stimuli presented completely outside conscious awareness influence judgments. This work contributes to the literature highlighting the need to study conscious awareness as a continuous phenomenon and provides a framework for researchers to ask and answer questions regarding conscious awareness and its relation to judgment and behavior.

Labeling the lines: A test of a six-mechanism model of chromatic detection.

Six linear chromatic mechanisms are sufficient to account for the pattern of threshold elevations produced by chromatic noise masking in the (L,M) plane of cone space (Shepard, Swanson, McCarthy, & Eskew, 2016). Here, we report results of asymmetric color matching of the threshold-level tests from that detection study and use those matches to test the detection model. We assume the mechanisms are univariant labeled lines (Rushton, 1972; Watson & Robson, 1981), implying that the chromaticities of physically different stimuli that are detected by a single mechanism should all be the same-they are postreceptoral metamers-but the chromaticities of two stimuli detected by different mechanisms should be different. The results show that color matches fall into six clusters in CIE (u',v') space (across all the noise conditions) and that these clusters correspond closely to the six mechanisms in the model. Most importantly, where the detection model determines that a given test angle is detected by different mechanisms under different noise conditions, the hue of that test angle changes in a consistent way. These color matches allow us to apply a color label to each of the mechanisms, confirm the six-mechanism model, and quantify the hue signaled by each mechanism.

A model of selective masking in chromatic detection.

Narrowly tuned, selective noise masking of chromatic detection has been taken as evidence for the existence of a large number of color mechanisms (i.e., higher order color mechanisms). Here we replicate earlier observations of selective masking of tests in the (L,M) plane of cone space when the noise is placed near the corners of the detection contour. We used unipolar Gaussian blob tests with three different noise color directions, and we show that there are substantial asymmetries in the detection contours-asymmetries that would have been missed with bipolar tests such as Gabor patches. We develop a new chromatic detection model, which is based on probability summation of linear cone combinations, and incorporates a linear contrast energy versus noise power relationship that predicts how the sensitivity of these mechanisms changes with noise contrast and chromaticity. With only six unipolar color mechanisms (the same number as the cardinal model), the new model accounts for the threshold contours across the different noise conditions, including the asymmetries and the selective effects of the noises. The key for producing selective noise masking in the (L,M) plane is having more than two mechanisms with opposed L- and M-cone inputs, in which case selective masking can be produced without large numbers of color mechanisms.

Noise masking of S-cone increments and decrements.

S-cone increment and decrement detection thresholds were measured in the presence of bipolar, dynamic noise masks. Noise chromaticities were the L-, M-, and S-cone directions, as well as L-M, L+M, and achromatic (L+M+S) directions. Noise contrast power was varied to measure threshold Energy versus Noise (EvN) functions. S+ and S- thresholds were similarly, and weakly, raised by achromatic noise. However, S+ thresholds were much more elevated by S, L+M, L-M, L- and M-cone noises than were S- thresholds, even though the noises consisted of two symmetric chromatic polarities of equal contrast power. A linear cone combination model accounts for the overall pattern of masking of a single test polarity well. L and M cones have opposite signs in their effects upon raising S+ and S- thresholds. The results strongly indicate that the psychophysical mechanisms responsible for S+ and S- detection, presumably based on S-ON and S-OFF pathways, are distinct, unipolar mechanisms, and that they have different spatiotemporal sampling characteristics, or contrast gains, or both.

Perceiving opponent hues in color induction displays.

According to Hering's color theory, certain hues (red vs green and blue vs yellow) are mutually exclusive as components of a single color; consequently a color cannot be perceived as reddish-green or bluish-yellow. The goal of our study is to test this key postulate of the opponent color theory. Using the method of adjustment, our observers determine the boundaries of chromatic zones in a red-green continuum. We demonstrate on two distinct stimulus sets, one formed using a chromatic grid and neon spreading and the other based on solid colored regions, that the chromatic contrast of a purple surround over a red figure results in perception of 'forbidden' reddish-green colors. The observed phenomenon can be understood as resulting from the construction of a virtual filter, a process that bypasses photoreceptor summation and permits forbidden color combinations. Showing that opponent hue combinations, previously reported only under artificial image stabilization, can be present in normal viewing conditions offers new approaches for the experimental study of the dimensionality and structure of perceptual color space.

Visual noise disrupts conceptual integration in reading.

The Effortfulness Hypothesis suggests that sensory impairment (either simulated or age-related) may decrease capacity for semantic integration in language comprehension. We directly tested this hypothesis by measuring resource allocation to different levels of processing during reading (i.e., word vs. semantic analysis). College students read three sets of passages word-by-word, one at each of three levels of dynamic visual noise. There was a reliable interaction between processing level and noise, such that visual noise increased resources allocated to word-level processing, at the cost of attention paid to semantic analysis. Recall of the most important ideas also decreased with increasing visual noise. Results suggest that sensory challenge can impair higher-level cognitive functions in learning from text, supporting the Effortfulness Hypothesis.

Quantifying the effect of natural and arbitrary sensorimotor contingencies on chromatic judgments.

When a body movement systematically co-occurs with a stimulus, a change in perception to compensate for this correlation may occur. In Experiment 1 (similar to A. Bompas & J. K. O'Regan, 2006), we induced a correlation between leftward eye saccades and a red stimulus, and rightward eye saccades and a green stimulus. In a subsequent test phase, observers compared the color of two stimuli after leftward and rightward saccades. The major result was that stimuli tended to look greener after a leftward saccade and redder after a rightward saccade (A. Bompas & J. K. O'Regan, 2006). Measured here in meaningful units for the first time, the shift in the point of subjective equality was approximately d' = 0.4, a remarkably large effect for only 40 minutes of eye movement/color exposure. A control experiment ruled out a simple reduction in initial bias as the cause of the effect. In Experiment 2, we hypothesized that the blue of the sky might cause an initial bias to judge spots seen with upward gaze as "yellower"; this expectation was not met, but the basic effect was replicated and extended to other chromaticities and eye movement directions. Experiment 3 substituted listening to tones for the eye movements (a sensorisensory correlation) to explore differences between sensorimotor adaptation and sensory integration; neither effect was found for our task. Sensorimotor adaptation can be a remarkably powerful influence on perception; because it operates in a compensatory direction, it may oppose the effects of sensory integration, depending upon task demands.

Higher order color mechanisms: a critical review.

A large number of studies, using a wide variety of experimental techniques, have investigated the "higher-order" color mechanisms proposed by Krauskopf and colleagues in 1986. Results reviewed here come from studies of chromatic discrimination at threshold, habituation, classification images, spatial alignment and orientation effects, and noise masking. The bulk of the evidence has been taken to support the existence of multiple, linear color mechanisms in addition to (or after) the three putative low-level cardinal mechanisms. But there remain disconcerting inconsistencies in the results of noise masking experiments, and the results of chromatic discrimination experiments clearly show that there are a very limited number of labeled-line mechanisms near threshold. No consensus on higher order mechanisms has been reached even after more than 20 years of study.

Theory of chromatic noise masking applied to testing linearity of S-cone detection mechanisms.

A method for testing the linearity of cone combination of chromatic detection mechanisms is applied to S-cone detection. This approach uses the concept of mechanism noise, the noise as seen by a postreceptoral neural mechanism, to represent the effects of superposing chromatic noise components in elevating thresholds and leads to a parameter-free prediction for a linear mechanism. The method also provides a test for the presence of multiple linear detectors and off-axis looking. No evidence for multiple linear mechanisms was found when using either S-cone increment or decrement tests. The results for both S-cone test polarities demonstrate that these mechanisms combine their cone inputs nonlinearly.