Brightness contrast inhibits color induction: evidence for a new kind of color theory

A gray region can be made to look colored by a colored surround. This phenomenon, chromatic induction, depends on color differences around the boundary of the region. We performed experiments on chromatic induction with small, initially achromatic, targets on nine different colored surrounds ranging in color from blue to red. Using scaling of saturation as our measure of perceived color strength, we found that chromatic induction is at its maximum when the brightness contrast at the boundary between target and surroundings is minimal. This implies that the neural mechanism in the cerebral cortex that mediates the appearance of brightness at a boundary inhibits the activity of chromatic mechanisms at that same boundary. Observers matched the apparent brightness and luminance of each of the colored surrounds. For surround colors where brightness and luminance matches differ, brightness contrast, not luminance contrast, controls chromatic induction. These new findings, taken together with other evidence, require a new theory of color appearance that includes mutually inhibitory interactions between color and brightness mechanisms that are sensing color and brightness contrast at visual boundaries.     

Lightness, illumination, and gradients

The illumination interpretation approach claims that lightness illusions can be explained as misapplications of lightness constancy mechanisms, processes which usually enable veridical extraction of surface reflectance from luminance distributions by discounting illumination. In particular, luminance gradients are thought to provide cues about the interactions of light and surfaces. Several examples of strong lightness illusions are discussed for which explanations based on illumination interpretation can be proposed. In criticisms of this approach, a variety of demonstrations of similarly structured control displays are presented, which involve equivalent lightness effects that cannot readily be accounted for by illumination interpretation mechanisms. Furthermore, a number of known and novel displays are presented that demonstrate effects of gradients on the qualitative appearance of uniform regions. Finally, some simple simulations of neural effects of luminance distributions are discussed.     

The dynamic range of human lightness perception

Natural viewing challenges the visual system with images that have a dynamic range of light intensity (luminance) that can approach 1,000,000:1 and that often exceeds 10,000:1 [1, 2]. The range of perceived surface reflectance (lightness), however, can be well approximated by the Munsell matte neutral scale (N 2.0/ to N 9.5/), consisting of surfaces whose reflectance varies by about 30:1. Thus, the visual system must map a large range of surface luminance onto a much smaller range of surface lightness. We measured this mapping in images with a dynamic range close to that of natural images. We studied simple images that lacked segmentation cues that would indicate multiple regions of illumination. We found a remarkable degree of compression: at a single image location, a stimulus luminance range of 5,905:1 can be mapped onto an extended lightness scale that has a reflectance range of 100:1. We characterized how the luminance-to-lightness mapping changes with stimulus context. Our data rule out theories that predict perceived lightness from luminance ratios or Weber contrast. A mechanistic model connects our data to theories of adaptation and provides insight about how the underlying visual response varies with context.     

Adelson's tile and snake illusions: a Helmholtzian type of simultaneous lightness contrast

Adelson's tile, snake, and some other lightness illusions of the same type were measured with the Munsell neutral scale for twenty observers. It was shown that theories based on low-level luminance contrast processing could hardly explain these illusions. Neither can those based on luminance X-junctions. On the other hand, Helmholtz's idea, that simultaneous lightness contrast originates from an error in judgement of apparent illumination, has been elaborated so as to account for the tile and snake illusions as well as other demonstrations presented in this report.     

Color and luminance contrasts attract independent attention

Paying attention can improve vision in many ways, including some very basic functions such as contrast discrimination, a task that probably reflects very early levels of visual processing. Electrophysiological, psychophysical, and imaging studies on humans as well as single recordings in monkey show that attention can modulate the neuronal response at an early stage of visual processing, probably by acting on the response gain. Here, we measure incremental contrast thresholds for luminance and color stimuli to derive the contrast response of early neural mechanisms and their modulation by attention. We show that, for both cases, attention improves contrast discrimination, probably by multiplicatively increasing the gain of the neuronal response to contrast. However, the effects of attention are highly specific to the visual modality: concurrent attention to a competing luminance, but not chromatic pattern, greatly impedes luminance contrast discrimination; and attending to a competing chromatic, but not luminance, task impedes color contrast discrimination. Thus, the effects of attention are highly modality specific, implying separate attentional resources for different fundamental visual attributes at early stages of visual processing.     

EFFECT OF CHROMATIC LIGHTS ON PHYCOBILIN FORMATION IN A BLUE-GREEN ALGA, TOLYPOTHRIX TENUIS

1. The formation of phycobilin pigments in a blue-green alga Tolypothrixtenuis was investigated with special reference to the effectsof preillumination with colored lights.
2. It was discoveredthat the algal cells are capable of formingphycobilin pigmentsin the dark, if they have been previouslyilluminated for severalhours in the presence of CO₂.
3. The color of light applied inthe later period of preillumination(chromatic illumination)was found to affect the ratio of phycoerythrinto phycocyaninformed in the subsequent dark period. A greenlight acceleratesthe dark-formation of phycoerythrin, a redlight that of phycocyanin,and the two lights counteractingwith each other in their effects.
4. These directive effects of the "chromatic illumination" canbe accomplished within a very short period, for instance, in3 minutes if it is preceded by sufficient "preillumination"with an incandescent or day light fluorescent light. The reactionsoccurring during the period of chromatic illumination does notrequire the presence of CO₂ and the aerobic condition.
5. Thealga can be grown heterotrophically when supplied with casaminoacids and glucose. Under such a condition the alga forms phycocyanintogether with chlorophyll and carotenoids, but not phycoerythrin.
6. On the basis of the results obtained, a tentative scheme forthe biosynthesis of phycobilin pigments in the alga was proposed,assuming the light-induced formation of unknown precursors whichare converted into phycocyanin and phycoerythrin in the subsequentdark period.

(Received July 4, 1960; )     

Responses to lightness variations in early human visual cortex

Lightness is the apparent reflectance of a surface, and it depends not only on the actual luminance of the surface but also on the context in which the surface is viewed [1-10]. The cortical mechanisms of lightness processing are largely unknown, and the role of early cortical areas is still a matter of debate [11-17]. We studied the cortical responses to lightness variations in early stages of the human visual system with functional magnetic resonance imaging (fMRI) while observers were performing a demanding fixation task. The set of dynamically presented visual stimuli included the rectangular version of the classic Craik-O'Brien stimulus [3, 18, 19] and a variant that led to a weaker lightness effect, as well as a pattern with actual luminance variations. We found that the cortical activity in retinotopic areas, including the primary visual cortex (V1), is correlated with context-dependent lightness variations.     

视动震颤(OKN)眼动控制系统中的颜色通道

用亮度相等的不同颜色构成的等亮度彩色运动条纹(Isoluminant chromatic moving gratings)来进行OKN眼动跟踪实验,探讨它是否与由亮度差别构成的黑白运动条纹图象一样引起OKN反应。实验结果表明在等亮度彩色运动条纹图象(没有亮度差别只有颜色差别)刺激下,视动系统可产生与黑白运动条纹刺激下同样的OKN反应,并且与各单原色运动条纹刺激下的OKN反应也一致。说明0KN眼动跟踪中的运动检测存在颜色通道。本文并提出了一种基于颜色的运动检测模型。     

Spatial information capacity of eyes: Roles of Na+ channels and photon noise in photoreceptor

The spatial information capacity of the human eye for photopic vision has been determined taking into account the intensity response function of the photoreceptor. It has been found that spatial information capacity increases with the mean luminance upto a certain value of mean luminance and after that it starts decreasing. The decrement occurs below the damage threshold. These results are in agreement with the reported experimental observations. It has been concluded that the limited number of Na⁺ channels in the photoreceptor outer segment and the photon noise are responsible for the fall in the information capacity below the damage threshold.Associated with the Biochemistry Cell     

Effect of pictorial depth cues, binocular disparity cues and motion parallax depth cues on lightness perception in three-dimensional virtual scenes

Background

Surface lightness perception is affected by scene interpretation. There is some experimental evidence that perceived lightness under bi-ocular viewing conditions is different from perceived lightness in actual scenes but there are also reports that viewing conditions have little or no effect on perceived color. We investigated how mixes of depth cues affect perception of lightness in three-dimensional rendered scenes containing strong gradients of illumination in depth.

Methodology/Principal Findings

Observers viewed a virtual room (4 m width×5 m height×17.5 m depth) with checkerboard walls and floor. In four conditions, the room was presented with or without binocular disparity (BD) depth cues and with or without motion parallax (MP) depth cues. In all conditions, observers were asked to adjust the luminance of a comparison surface to match the lightness of test surfaces placed at seven different depths (8.5–17.5 m) in the scene. We estimated lightness versus depth profiles in all four depth cue conditions. Even when observers had only pictorial depth cues (no MP, no BD), they partially but significantly discounted the illumination gradient in judging lightness. Adding either MP or BD led to significantly greater discounting and both cues together produced the greatest discounting. The effects of MP and BD were approximately additive. BD had greater influence at near distances than far.

Conclusions/Significance

These results suggest the surface lightness perception is modulated by three-dimensional perception/interpretation using pictorial, binocular-disparity, and motion-parallax cues additively. We propose a two-stage (2D and 3D) processing model for lightness perception.     

Natural image statistics mediate brightness 'filling in'

Although the human visual system can accurately estimate the reflectance (or lightness) of surfaces under enormous variations in illumination, two equiluminant grey regions can be induced to appear quite different simply by placing a light-dark luminance transition between them. This illusion, the Craik-Cornsweet-O'Brien (CCOB) effect, has been taken as evidence for a low-level 'filling-in' mechanism subserving lightness perception. Here, we present evidence that the mechanism responsible for the CCOB effect operates not via propagation of a neural signal across space but by amplification of the low spatial frequency (SF) structure of the image. We develop a simple computational model that relies on the statistics of natural scenes actively to reconstruct the image that is most likely to have caused an observed series of responses across SF channels. This principle is tested psychophysically by deriving classification images (CIs) for subjects' discrimination of the contrast polarity of CCOB stimuli masked with noise. CIs resemble 'filled-in' stimuli; i.e. observers rely on portions of the stimuli that contain no information per se but that correspond closely to the reported perceptual completion. As predicted by the model, the filling-in process is contingent on the presence of appropriate low SF structure.     

An account of responses of spectrally opponent neurons in macaque lateral geniculate nucleus to successive contrast

Coloured surfaces in the normal environment may be brighter or dimmer than the mean adaptation level. Changes in the firing rate of cells of the parvocellular layers of macaque lateral geniculate nucleus were studied with such stimuli; chromatic mixtures briefly replaced a white adaptation field. This paradigm is therefore one of successive contrast. Families of intensity-response curves for different wavelengths were measured. When taking sections at different luminance ratios through these families of curves, strongly opponent cells displayed spectrally selective responses at low luminance ratios, while weakly opponent cells had higher chromatic thresholds and responded well to stimuli at higher luminance ratios, brighter than the adaptation field. Strength of cone opponency, defined as the weight of the inhibitory cone mechanism relative to the excitatory one, was thus related to the range of intensity in which cells appeared to operate most effectively. S-cone inputs, as tested with lights lying along tritanopic confusion lines, could either be excitatory or inhibitory. Families of curves for different wavelengths can be simulated mathematically for a given cell by a simple model by using known cone absorption spectra. Hyperbolic response functions relate cone absorption to the output signals of the three cone mechanisms, which are assumed to interact linearly. Parameters from the simulation provided estimates of strength of cone opponency and cone sensitivity which were shown to be continuously distributed. Cell activity can be related to cone excitation in a trichromatic colour space with the help of the model, to give an indication of suprathreshold coding of colour and lightness.     

Shading Beats Binocular Disparity in Depth from Luminance Gradients: Evidence against a Maximum Likelihood Principle for Cue Combination

Perceived depth is conveyed by multiple cues, including binocular disparity and luminance shading. Depth perception from luminance shading information depends on the perceptual assumption for the incident light, which has been shown to default to a diffuse illumination assumption. We focus on the case of sinusoidally corrugated surfaces to ask how shading and disparity cues combine defined by the joint luminance gradients and intrinsic disparity modulation that would occur in viewing the physical corrugation of a uniform surface under diffuse illumination. Such surfaces were simulated with a sinusoidal luminance modulation (0.26 or 1.8 cy/deg, contrast 20%-80%) modulated either in-phase or in opposite phase with a sinusoidal disparity of the same corrugation frequency, with disparity amplitudes ranging from 0’-20’. The observers’ task was to adjust the binocular disparity of a comparison random-dot stereogram surface to match the perceived depth of the joint luminance/disparity-modulated corrugation target. Regardless of target spatial frequency, the perceived target depth increased with the luminance contrast and depended on luminance phase but was largely unaffected by the luminance disparity modulation. These results validate the idea that human observers can use the diffuse illumination assumption to perceive depth from luminance gradients alone without making an assumption of light direction. For depth judgments with combined cues, the observers gave much greater weighting to the luminance shading than to the disparity modulation of the targets. The results were not well-fit by a Bayesian cue-combination model weighted in proportion to the variance of the measurements for each cue in isolation. Instead, they suggest that the visual system uses disjunctive mechanisms to process these two types of information rather than combining them according to their likelihood ratios.     

Chromatic and achromatic visual evoked potentials in Parkinson's disease

Chromatic and achromatic visual evoked potentials (VEP) were evaluated in 39 patients with idiopathic Parkinson's disease (PD) (age 64.0 ± 8.6 years) and 43 healthy controls (age 62.8 ± 8.7 years). The following pattern-reversal checkerboard stimuli were performed: (1) achromatic with luminance contrast 86% (achr.hk.); (2) achromatic with luminance contrast 20% (achr.lk.); (3) chromatic isoluminant blue-yellow (by.); (4) chromatic isoluminant red-green (rg.). The mean latencies N70, P100, and N135 of chromatic and achromatic VEP were significantly delayed in patients with PD as compared to controls. The highest rate (41.0%) of pathological findings could be demonstrated by achromatic stimulation (luminance contrast 86%). Isolated abnormalities of chromatic VEP (in combination with normal achromatic VEP) were found in 5 (12.8%) patients. The delay of VEP-latencies was significantly correlated with the severity of motor symptoms in PD patients. We conclude that VEP are valuable tools to demonstrate a dysfunction of the visual system in PD. Although chromatic VEP are less sensitive than achromatic VEP, the combination of both will increase the diagnostic yield. Therefore, there seems to exist a variety of individual characters of visual impairment in PD.     

Chromatic signals control proboscis movements during hovering flight in the hummingbird hawkmoth Macroglossum stellatarum

Most visual systems are more sensitive to luminance than to colour signals. Animals resolve finer spatial detail and temporal changes through achromatic signals than through chromatic ones. Probably, this explains that detection of small, distant, or moving objects is typically mediated through achromatic signals. Macroglossum stellatarum are fast flying nectarivorous hawkmoths that inspect flowers with their long proboscis while hovering. They can visually control this behaviour using floral markings known as nectar guides. Here, we investigate whether this is mediated by chromatic or achromatic cues. We evaluated proboscis placement, foraging efficiency, and inspection learning of naïve moths foraging on flower models with coloured markings that offered either chromatic, achromatic or both contrasts. Hummingbird hawkmoths could use either achromatic or chromatic signals to inspect models while hovering. We identified three, apparently independent, components controlling proboscis placement: After initial contact, 1) moths directed their probing towards the yellow colour irrespectively of luminance signals, suggesting a dominant role of chromatic signals; and 2) moths tended to probe mainly on the brighter areas of models that offered only achromatic signals. 3) During the establishment of the first contact, naïve moths showed a tendency to direct their proboscis towards the small floral marks independent of their colour or luminance. Moths learned to find nectar faster, but their foraging efficiency depended on the flower model they foraged on. Our results imply that M. stellatarum can perceive small patterns through colour vision. We discuss how the different informational contents of chromatic and luminance signals can be significant for the control of flower inspection, and visually guided behaviours in general.     

Development of contrast sensitivity and acuity of the infant colour system

We have monitored the development of infant colour vision by measuring chromatic contrast sensitivity and acuity in eight young infants over a period of 6 months. Steady-state visual evoked potentials (VEPS) were recorded in response to both chromatic (red-green) and luminance (red-black or green-black) patterns that were reversed in contrast over time. For most infants, no response could be obtained to chromatic stimuli of any size or contrast before 5 weeks of age, although luminance stimuli of 20% contrast gave reliable responses at that age. When responses to chromatic stimuli first appeared, they could be obtained only with stimuli of very low spatial frequency, 20 times lower than the acuity for luminance stimuli. Both contrast sensitivity and acuity for chromatic stimuli increased steadily, more rapidly than for luminance stimuli. As the spectral selectivities of infant cones are similar to those of adults, the difference in rate of development of luminance and chromatic contrast sensitivity and acuity stimuli probably reflects neural development of the infant colour system.     

Chromatic cues to trap the oriental fruit fly, Bactrocera dorsalis

Various colors have been used as visual cues to trap insect pests. For example, yellow traps for monitoring and control of the oriental fruit fly (Bactrocera dorsalis) have been in use for a very long time. However, the chromatic cue of using color traps has never been meticulously investigated. In this study, the spectral sensitivities of the photoreceptors in the compound eyes of B. dorsalis were measured intracellularly, and the theory of receptor quantum catch was applied to study the chromatic cue of fly attracting. Responses to five wavelength categories with peak wavelengths of 370, 380, 490, and 510 nm, and one with dual peaks at 350 and 490 nm were recorded. Based on spectral sensitivities, six colored papers were chosen to test the color preference of the fly, and an additional UV preference test was done to confirm the effect of the UV stimuli. It was concluded that UV and green stimuli (spectra: 300-380 nm and 500-570 nm) would enhance the attractiveness of a colored paper to the oriental fruit fly, and blue stimuli (380-500 nm) would diminish the attractiveness.     

The angular range of achromatic target detection by honey bees

Honeybees Apis mellifera were trained to enter a Y-maze and choose the arm with a rewarded disc presented against a grey background. The alternative arm displayed the unrewarded grey background alone. Training and testing were performed with the rewarding disc subtending different visual angles. The training disc was either achromatic and provided green contrast, or chromatic and provided the same amount of green contrast as the achromatic one. The bee-achromatic disc could be learned and detected by the bees whenever it subtended 5° or 10°, but not if it subtended 30°. The chromatic disc was learned well and detected at all three visual angles. However, at 5° the maximum level of correct choices was ca. 75% with the achromatic disc whilst it was ca. 90% with the chromatic one. Thus, the presence of chromatic contrast enhances considerably the level of correct choices for the same amount of green contrast. The lower threshold of achromatic target detection lies between 3.7° and 5°; the upper threshold between 15° and 10°. At the upper threshold, detection switches from chromatic-based to achromatic-based. Thus, in the context of target detection, the achromatic green contrast channel specialises in the detection of objects of reduced angular size, whilst the chromatic channels are specialised for objects of large angular size. We suggest that achromatic detectors with a centre-surround organisation are involved in the task of detecting achromatic targets. Accepted: 23 February 1998     

Spatiotemporal frequency responses of cat retinal ganglion cells

Spatiotemporal frequency responses were measured at different levels of light adaptation for cat X and Y retinal ganglion cells. Stationary sinusoidal luminance gratings whose contrast was modulated sinusoidally in time or drifting gratings were used as stimuli. Under photopic illumination, when the spatial frequency was held constant at or above its optimum value, an X cell's responsivity was essentially constant as the temporal frequency was changed from 1.5 to 30 Hz. At lower temporal frequencies, responsivity rolled off gradually, and at higher ones it rolled off rapidly. In contrast, when the spatial frequency was held constant at a low value, an X cell's responsivity increased continuously with temporal frequency from a very low value at 0.1 Hz to substantial values at temporal frequencies higher than 30 Hz, from which responsivity rolled off again. Thus, 0 cycles X deg-1 became the optimal spatial frequency above 30 Hz. For Y cells under photopic illumination, the spatiotemporal interaction was even more complex. When the spatial frequency was held constant at or above its optimal value, the temporal frequency range over which responsivity was constant was shorter than that of X cells. At lower spatial frequencies, this range was not appreciably different. As for X cells, 0 cycles X deg-1 was the optimal spatial frequency above 30 Hz. Temporal resolution (defined as the high temporal frequency at which responsivity had fallen to 10 impulses X s-1) for a uniform field was approximately 95 Hz for X cells and approximately 120 Hz for Y cells under photopic illumination. Temporal resolution was lower at lower adaptation levels. The results were interpreted in terms of a Gaussian center-surround model. For X cells, the surround and center strengths were nearly equal at low and moderate temporal frequencies, but the surround strength exceeded the center strength above 30 Hz. Thus, the response to a spatially uniform stimulus at high temporal frequencies was dominated by the surround. In addition, at temporal frequencies above 30 Hz, the center radius increased.     

Spatiochromatic properties of natural images and human vision

The human visual system shows a relatively greater response to low spatial frequencies of chromatic spatial modulation than to luminance spatial modulation. However, previous work has shown that this differential sensitivity to low spatial frequencies is not reflected in any differential luminance and chromatic content of general natural scenes. This is contrary to the prevailing assumption that the spatial properties of human vision ought to reflect the structure of natural scenes. Now, colorimetric measures of scenes suggest that red-green (and perhaps blue-yellow) color discrimination in primates is particularly suited to the encoding of specific scenes: reddish or yellowish objects on a background of leaves. We therefore ask whether the spatial, as well as chromatic, properties of such scenes are matched to the different spatial-encoding properties of color and luminance modulation in human vision. We show that the spatiochromatic properties of a wide class of scenes, which contain reddish objects (e.g., fruit) on a background of leaves, correspond well to the properties of the red-green (but not blue-yellow) systems in human vision, at viewing distances commensurate with typical grasping distance. This implies that the red-green system is particularly suited to encoding both the spatial and the chromatic structure of such scenes.