Experimental verification of the opponent theory of human color vision

The colors observed by the human eye after a short flash of light of different spectral compositions were studied experimentally. The successive images and changes in their color with time confirm the opponent theory of human color vision.     

In the eye of the beholder: Is color classification consistent among human observers?

Colorful displays have evolved in multiple plant and animal species as signals to mutualists, antagonists, competitors, mates, and other potential receivers. Studies of color have long relied on subjective classifications of color by human observers. However, humans have a limited ability to perceive color compared to other animals, and human biological, cultural, and environmental variables can influence color perception. Here, we test the consistency of human color classification using fruit color as a model system. We used reflectance data of 67 tropical fruits and surveyed 786 participants to assess the degree to which (a) participants of different cultural and linguistic backgrounds agree on color classification of fruits; and (b) human classification to a discrete set of commonly used colors (e.g., red, blue, green) corresponds to natural clusters based on light reflectance measures processed through visual systems of other animals. We find that individual humans tend to agree on the colors they attribute to fruits across language groups. However, these colors do not correspond to clearly discernible clusters in di‐ or tetrachromatic visual systems. These results indicate that subjective color categorizations tend to be consistent among observers and can be used for large synthetic studies, but also that they do not fully reflect natural categories that are relevant to animal observers.     

New World Monkeys and Color

The visual worlds of most primates are rich with potential color signals, and many representatives of the order have evolved the biological mechanisms that allow them to exploit these sources of information. Unlike the catarrhines, platyrrhines typically have sex-linked polymorphic color vision that provides individuals with any of several distinct types of color vision, including both trichromatic and dichromatic variants. In recent years, this polymorphism has been the target of an expanding range of research efforts. As a result, researchers now reasonably understand the proximate biology underlying the polymorphisms, and a number of ideas have emerged as to their evolution. Progress has also been made in illuminating how color vision capacities may be related to the particular visual tasks that New World monkeys face.     

Perceptual considerations in the use of colored photographic and video stimuli to study nonhuman primate behavior

The use of photographs, slides, computerized images, and video to study behavior is increasingly being employed in nonhuman primates. However, since these mediums have been designed to simulate natural coloration for normal trichromatic human vision, they can fail to reproduce color in meaningful and accurate ways for viewers with different visual systems. Given the range of color perception that exists both across and within different species, it is necessary to consider this variation in order to discern the suitability of these mediums for experimental use. Because of the high degree of visual similarity among humans, Old World monkeys, and apes, the use of photographic and video stimuli should be acceptable in terms of replicating naturalistic coloration and making noticeable color manipulations. However, among New World primates and prosimians, there exists a considerable degree of variation in color perceptual abilities depending on the species, sex, and allelic combination of the animals involved. Therefore, the use of these mediums to study behavior is problematic for these species, and should be done with caution.     

基于H分量旋转的色盲矫正方法

为提高色盲患者的色彩分辨力,提出一种基于H分量旋转的色盲矫正方法。在颜色的HSI空间,利用H分量的连续性和周期性特点,在保持S、I分量不变的情况下,通过旋转H分量得到矫正图像,该图像以降低低频颜色的分辨率来换取高频颜色的分辨率提高。实验表明：在色盲类型给定且图像颜色是给定色盲易混淆的情况下,对H分量旋转120度能得到色彩分辨效果很好的矫正图像。     

The Causes and Consequences of Color Vision

The ability to see colors is not universal in the animal kingdom. Those animals that can detect differences in the wavelengths of the electromagnetic spectrum glean valuable sensory information about their environment. They use color vision to forage, avoid predators, and find high-quality mates. In the past, the colors that humans could see clouded scientists’ study of animals’ color perception. Leaving that bias behind has led to new insights about how and why the color vision of animals evolved. This paper provides a brief introduction to color vision, the genetics of color vision in humans, what colors other animals see, and how scientists study color vision. We examine the consequences of having color vision, including speciation, loss of olfactory capabilities, and sexual selection.     

Avian vision and the evolution of egg color mimicry in the common cuckoo

Coevolutionary arms races are a potent force in evolution, and brood parasite-host dynamics provide classical examples. Different host-races of the common cuckoo, Cuculus canorus, lay eggs in the nests of other species, leaving all parental care to hosts. Cuckoo eggs often (but not always) appear to match remarkably the color and pattern of host eggs, thus reducing detection by hosts. However, most studies of egg mimicry focus on human assessments or reflectance spectra, which fail to account for avian vision. Here, we use discrimination and tetrachromatic color space modeling of bird vision to quantify egg background and spot color mimicry in the common cuckoo and 11 of its principal hosts, and we relate this to egg rejection by different hosts. Egg background color and luminance are strongly mimicked by most cuckoo host-races, and mimicry is better when hosts show strong rejection. We introduce a novel measure of color mimicry-"color overlap"-and show that cuckoo and host background colors increasingly overlap in avian color space as hosts exhibit stronger rejection. Finally, cuckoos with better background color mimicry also have better pattern mimicry. Our findings reveal new information about egg mimicry that would be impossible to derive by the human eye.     

视觉通道中颜色与形状特征的识别

本文利用计算机设计新的心理物理实验,研究人类视觉系统的颜色、形状通道和对颜色的识别反应,证实视觉系统的颜色通道和形状通道是独立并行的.对颜色反应还进行视觉诱发电位测试,结果与心理物理实验基本一致.最后,提出颜色通道与形状通道间信息相互统一的假说模型.     

Color matching on natural substrates in cuttlefish, <Emphasis Type="Italic">Sepia officinalis</Emphasis>

The camouflaging abilities of cuttlefish (Sepia officinalis) are remarkable and well known. It is commonly believed that cuttlefish-although color blind-actively match various colors of their immediate surroundings, yet no quantitative data support this notion. We assembled several natural substrates chosen to evoke the three basic types of camouflaged body patterns that cuttlefish express (uniform/stipple, mottle, and disruptive) and measured the spectral reflectance of the camouflaged pattern and the respective background using a fiber optic spectrometer. We demonstrate that the reflectance spectra of cuttlefish skin patterns correlate closely with the spectra of these natural substrates. Since pigmented chromatophores play a key role in cephalopod color change, we also measured the spectral reflectance of individual cuttlefish chromatophores under the microscope, and confirm the results from a previous publication reporting three distinct colors of chromatophores (yellow, orange, and dark brown) on the animals' dorsal side. Taken together, our results show that the color variations in substrate and animal skin can be very similar and that this may facilitate color match on natural substrates in the absence of color vision.     

Photopigments underlying color vision in ringtail lemurs (Lemur catta) and brown lemurs (Eulemur fulvus)

A recent examination of color vision in the ringtail lemur produced evidence that these prosimians could make color discriminations consistent with a diagnosis of trichromatic color vision. However, it was unclear if this behavior reflected the presence of three classes of cone or whether lemurs might be able to utilize signals from rods in conjunction with those from only two classes of cone. To resolve that issue, spectral sensitivity functions were obtained from ringtail lemurs (Lemur catta) and brown lemurs (Eulemur fulvus) using a noninvasive electrophysiological procedure, electroretinographic flicker photometry. Results from experiments involving chromatic adaptation indicate that these lemurs routinely have only a single class of cone photopigment in the middle to long wavelengths (peak sensitivity of about 545 nm); they also have a short-wavelengthsensitive cone pigment with peak of about 437 nm. The earlier behavioral results are suggested to have resulted from the ability of lemurs to jointly utilize signals from rods and cones. The cone pigment complements of these lemurs differ distinctly from those seen among the anthropoids. © 1993 Wiley-Liss, Inc.     

视网膜周边视野人工成像研究

目的 对于中心视力受损的人群，新型人工视觉系统可以将简化后的图像投射到视网膜上黄斑区以外的区域，从而帮助他们利用周边视觉感知信息。本文探究周边视野的感知特征，为植入式光学人工视觉系统的图形编码设计提供依据。方法 设计了探索周边视野感知特征的实验环境，向被试施加符号、数字、汉字的图案刺激，并控制刺激的大小、颜色组合、偏离角度、运动情况。用图形化的方法分析感知能力与各变量的关系。结果 周边视野的感知能力随偏离角度增大而下降，其趋势分为两个阶段，且受颜色组合、大小的影响明显。结论 研究结果提供了感知识别率较高的变量组合，为人工视觉系统的光学投影、眼内光学植入装置、特殊通信彩色符号编码开发等“人机结合”新技术提供重要的实验依据。     

Dichromatic and Trichromatic Callithrix geoffroyi Differ in Relative Foraging Ability for Red-Green Color-Camouflaged and Non-camouflaged food

An advantage for trichromatic color vision in primates is shown by its presence in many lineages, but little attention has been paid to the potential disadvantages of trichromacy. Most New World monkey species are polymorphic for color vision, with both dichromats and trichromats present within a single population. We tested the foraging ability of trichromatic and dichromatic Geoffroy's marmosets (Callithrix geoffroyi) for colored cereal balls (Kix®) under conditions of red-green color camouflage (orange/green Kix® against an orange/green background) or lack of camouflage (Kix® same color as background) in a naturalized captive setting. In separate experiments designed to test foraging ability at long distances (<6 m) and short distances (<0.5 m), trichromats found significantly fewer Kix® under the camouflage condition than in the non-camouflage condition. In contrast, there is no difference in the ability of dichromats to detect color-camouflaged versus non-camouflaged Kix®. There is no significant difference between dichromats and trichromats for either camouflaged or non-camouflaged Kix®, though the power in the tests is low because of high individual variation. The results have clear implications for the foraging strategies of trichromatic marmosets. Differences in intensity of competition between trichromats and dichromats for items of food of different colors in relation to background may also have consequences for the foraging behavior of dichromats.     

Colors of primate pelage and skin: objective assessment of conspicuousness

We present a quantitative means of assessing the conspicuousness of animal coats or other objects in terms of the color vision of each possible observer. We measured reflectance spectra from the fur and skin of many primate species in order to provide an objective survey of the possibilities of pelage coloration found in extant primates. We show that the orange coloration displayed by many platyrrhine and some strepsirhine primates, while being conspicuous to humans, would be cryptic amongst foliage to all males and many females of their own species. In relation to this finding, we briefly review what is known of the color vision of birds that prey on primates, and assess how conspicuous the orange pelage would be to these predators.     

Molecular Evolution of Arthropod Color Vision Deduced from Multiple Opsin Genes of Jumping Spiders

Among terrestrial animals, only vertebrates and arthropods possess wavelength-discrimination ability, so-called “color vision”. For color vision to exist, multiple opsins which encode visual pigments sensitive to different wavelengths of light are required. While the molecular evolution of opsins in vertebrates has been well investigated, that in arthropods remains to be elucidated. This is mainly due to poor information about the opsin genes of non-insect arthropods. To obtain an overview of the evolution of color vision in Arthropoda, we isolated three kinds of opsins, Rh1, Rh2, and Rh3, from two jumping spider species, Hasarius adansoni and Plexippus paykulli. These spiders belong to Chelicerata, one of the most distant groups from Hexapoda (insects), and have color vision as do insects. Phylogenetic analyses of jumping spider opsins revealed a birth and death process of color vision evolution in the arthropod lineage. Phylogenetic positions of jumping spider opsins revealed that at least three opsins had already existed before the Chelicerata-Pancrustacea split. In addition, sequence comparison between jumping spider Rh3 and the shorter wavelength-sensitive opsins of insects predicted that an opsin of the ancestral arthropod had the lysine residue responsible for UV sensitivity. These results strongly suggest that the ancestral arthropod had at least trichromatic vision with a UV pigment and two visible pigments. Thereafter, in each pancrustacean and chelicerate lineage, the opsin repertoire was reconstructed by gene losses, gene duplications, and function-altering amino acid substitutions, leading to evolution of color vision. Mitsumasa Koyanagi and Takashi Nagata contributed equally to this work. Sequence data from this article have been deposited with the DDBJ under accession nos. AB251846–AB251851.     

Color vision models: Some simulations,a general n‐dimensional model,and the colourvision R package

The development of color vision models has allowed the appraisal of color vision independent of the human experience. These models are now widely used in ecology and evolution studies. However, in common scenarios of color measurement, color vision models may generate spurious results. Here I present a guide to color vision modeling (Chittka (1992, Journal of Comparative Physiology A, 170, 545) color hexagon, Endler & Mielke (2005, Journal Of The Linnean Society, 86, 405) model, and the linear and log‐linear receptor noise limited models (Vorobyev & Osorio 1998, Proceedings of the Royal Society B, 265, 351; Vorobyev et al. 1998, Journal of Comparative Physiology A, 183, 621)) using a series of simulations, present a unified framework that extends and generalize current models, and provide an R package to facilitate the use of color vision models. When the specific requirements of each model are met, between‐model results are qualitatively and quantitatively similar. However, under many common scenarios of color measurements, models may generate spurious values. For instance, models that log‐transform data and use relative photoreceptor outputs are prone to generate spurious outputs when the stimulus photon catch is smaller than the background photon catch; and models may generate unrealistic predictions when the background is chromatic (e.g. leaf reflectance) and the stimulus is an achromatic low reflectance spectrum. Nonetheless, despite differences, all three models are founded on a similar set of assumptions. Based on that, I provide a new formulation that accommodates and extends models to any number of photoreceptor types, offers flexibility to build user‐defined models, and allows users to easily adjust chromaticity diagram sizes to account for changes when using different number of photoreceptors.     

Evolution of color and vision of butterflies

Butterfly eyes consist of three types of ommatidia, which are more or less randomly arranged in a spatially regular lattice. The corneal nipple array and the tapetum, optical structures that many but not all butterflies share with moths, suggest that moths are ancestral to butterflies, in agreement with molecular phylogeny. A basic set of ultraviolet-, blue- and green-sensitive receptors, encountered among nymphalid butterflies, forms the basis for trichromatic vision. Screening pigments surrounding the light-receiving rhabdoms can modify the spectral sensitivity of the photoreceptors so that the sensitivity peak is in the violet, yellow, red, or even deep-red, specifically in swallowtails (Papilionidae) and whites (Pieridae), thus enhancing color discriminability. The photoreceptor sensitivity spectra are presumably tuned to the wing colors of conspecific butterflies.     

Cross-modal associations between odors and colors

In the present study, we investigated the nature of any cross-modal associations between colors and odors. In Experiment 1, we show that participants consistently match certain odors to specific colors when asked to explicitly select from among different colors the one that best matched a given odor. In Experiment 2, we investigated the robustness of these cross-modal associations using a cross-modal variant of the implicit association test (IAT). Participants made speeded discrimination responses to a random sequence of odors (strawberry vs. spearmint) and color patches (pink vs. turquoise). On the basis of the results of Experiment 1, the assignment of these targets onto the two response keys was manipulated in order to generate compatible (e.g., responding to the pink color and to the strawberry odor with the same response key) and incompatible (e.g., responding to the pink color and to the spearmint odor with the same response key) blocks of trials. The results showed that participants responded more rapidly and accurately to odor-color pairings having a stronger association than to those having a weaker (or no) association. These results suggest that odor-color associations can be both systematic and robust. The paradigm developed here provides a novel cross-modal extension of the IAT to probe the nature of color-odor associations.     

The normalized segment classification model: A new tool to compare spectral reflectance curves

1. Color patterns are complex traits under selective pressures from conspecifics, mutualists, and antagonists. To evaluate the salience of a pattern or the similarity between colors, several visual models are available. Color discrimination models estimate the perceptual difference between any two colors. Their application to a diversity of taxonomic groups has become common in the literature to answer behavioral, ecological, and evolutionary questions. To use these models, we need information about the visual system of our beholder species. However, many color patterns are simultaneously subject to selective pressures from different species, often from different taxonomic groups, with different visual systems. Furthermore, we lack information about the visual system of many species, leading ecologists to use surrogate values or theoretical estimates for model parameters.
2. Here, we present a modification of the segment classification method proposed by Endler (Biological Journal of the Linnean Society, 1990 41, 315–352): the normalized segment classification model (NSC). We explain its logic and use, exploring how NSC differs from other visual models. We also compare its predictions with available experimental data.
3. Even though the NSC model includes no information about the visual system of the receiver species, it performed better than traditional color discrimination models when predicting the output of some behavioral tasks. Although vision scientists define color as independent of stimulus brightness, a likely explanation for the goodness of fit of the NSC model is that its distance measure depends on brightness differences, and achromatic information can influence the decision‐making process of animals when chromatic information is missing.
4. Species‐specific models may be insufficient for the study of color patterns in a community context. The NSC model offers a species‐independent solution for color analyses, allowing us to calculate color differences when we ignore the intended viewer of a signal or when different species impose selective pressures on the signal.

     

Mutational changes in S-cone opsin genes common to both nocturnal and cathemeral Aotus monkeys

Aotus is a platyrrhine primate that has been classically considered to be nocturnal. Earlier research revealed that this animal lacks a color vision capacity because, unlike all other platyrrhine monkeys, Aotus has a defect in the opsin gene that is required to produce short-wavelength sensitive (S) cone photopigment. Consequently, Aotus retains only a single type of cone photopigment. Other mammals have since been found to show similar losses and it has often been speculated that such change is in some fashion tied to nocturnality. Although most species of Aotus are indeed nocturnal, recent observations show that Aotus azarai, an owl monkey species native to portions of Argentina and Paraguay, displays a cathemeral activity pattern being active during daylight hours as frequently as during nighttime hours. We have sequenced portions of the S-cone opsin gene in A. azarai and Aotus nancymaae, the latter a typically nocturnal species. The S-cone opsin genes in both species contain the same fatal defects earlier detected for Aotus trivirgatus. On the basis of the phylogenetic relationships of these three species these results imply that Aotus must have lost a capacity for color vision early in its history and they also suggest that the absence of color vision is not compulsively linked to a nocturnal lifestyle.