The biological basis of a universal constraint on color naming: cone contrasts and the two-way categorization of colors

Many studies have provided evidence for the existence of universal constraints on color categorization or naming in various languages, but the biological basis of these constraints is unknown. A recent study of the pattern of color categorization across numerous languages has suggested that these patterns tend to avoid straddling a region in color space at or near the border between the English composite categories of "warm" and "cool". This fault line in color space represents a fundamental constraint on color naming. Here we report that the two-way categorization along the fault line is correlated with the sign of the L- versus M-cone contrast of a stimulus color. Moreover, we found that the sign of the L-M cone contrast also accounted for the two-way clustering of the spatially distributed neural responses in small regions of the macaque primary visual cortex, visualized with optical imaging. These small regions correspond to the hue maps, where our previous study found a spatially organized representation of stimulus hue. Altogether, these results establish a direct link between a universal constraint on color naming and the cone-specific information that is represented in the primate early visual system.     

Platyrrhine color signals: New horizons to pursue

Like catarrhines, some platyrrhines show exposed and reddish skin, raising the possibility that reddish signals have evolved convergently. This variation in skin exposure and color combined with sex‐linked polymorphic color vision in platyrrhines presents a unique, and yet underexplored, opportunity to investigate the relative importance of chromatic versus achromatic signals, the influence of color perception on signal evolution, and to understand primate communication broadly. By coding the facial skin exposure and color of 96 platyrrhines, 28 catarrhines, 7 strepsirrhines, 1 tarsiiform, and 13 nonprimates, and by simulating the ancestral character states for these traits, we provide the first analysis of the distribution and evolution of facial skin exposure and color in platyrrhini. We highlight ways in which studying the presence and use of color signals by platyrrhines and other primates will enhance our understanding of the evolution of color signals, and the forces shaping color vision.     

Discovery of androecium color polymorphism in Epimedium pubescens with habitat preference of anther/pollen color in the genus

Aims Interspecific and intraspecific variation in flower color in natural populations provides an opportunity for us to understand the evolution and maintenance of diversity of floral traits. Compared to corolla color, little is known about the color polymorphism of sexual organs in flowering plants. To explore evolutionary transitions of androecium color and polymorphism within species, interspecific and intraspecific variation in androecium (anther and pollen) color in the genus Epimedium (Berberidaceae) was investigated.Methods To explore the geographical patterns of anther/pollen color variation in Epimedium species, data of 45 species were collected and their phylogeny was constructed based on available DNA sequences. To investigate whether intraspecific variation in androecium color relates to habitat preference, three environmental factors were measured in the field population of Epimedium pubescens in northeastern Sichuan, China, which plants had green or yellow androecia. Vegetative and reproductive traits of this species were compared between the two color morphs.Important findings Androecium (anther and pollen) color polymorphism in field populations of Epimedium pubescens is reported here where nine populations are monomorphic with a green androecium but three populations are dimorphic with individuals having either a green or a yellow androecium. Inflorescence stalk height, stalk diameter, leaf number, flower number and spur length (as well as spur and nectar volume) were not significantly different between two morphs. Compared to the yellow morph, the green morph had relatively larger leaves and anthers, but smaller sepals. The green morph produced more pollen and larger seeds, but the same number of ovules. Seed set was not significantly different between green and yellow morph. Investigations of environmental factors in the color dimorphic populations of E. pubescens indicated that the green morph was more likely to occur in habitats with relatively lower light intensity. The distribution survey of 45 Epimedium species showed that species with a green androecium tended to appear at lower elevations. Comparative phylogenetic analysis showed that transitions from yellow to a green androecium or to androecial color dimorphism occurred at least seven times. This genus, characterized by anther color diversity and containing some species with anther color polymorphism, provides a model system in which to study the evolution and maintenance of colorful sexual organs in flowering plants.     

Signatures of selection and gene conversion associated with human color vision variation

Trichromatic color vision in humans results from the combination of red, green, and blue photopigment opsins. Although color vision genes have been the targets of active molecular and psychophysical research on color vision abnormalities, little is known about patterns of normal genetic variation in these genes among global human populations. The current study presents nucleotide sequence analyses and tests of neutrality for a 5.5-kb region of the X-linked long-wave "red" opsin gene (OPN1LW) in 236 individuals from ethnically diverse human populations. Our analysis of the recombination landscape across OPN1LW reveals an unusual haplotype structure associated with amino acid replacement variation in exon 3 that is consistent with gene conversion. Compared with the absence of OPN1LW amino acid replacement fixation since divergence from chimpanzee, the human population exhibits a significant excess of high-frequency OPN1LW replacements. Our results suggest that subtle changes in L-cone opsin wavelength absorption may have been adaptive during human evolution.     

Hemispheric difference in human skin color

Previous studies of human skin color have shown a strong relationship between skin color and distance from the equator, which has been interpreted as a link between skin color, latitude, and the intensity of ultraviolet radiation. The underlying assumptions are that UV radiation is greatest at the equator and that it diminishes with increasing latitude to the same extent in both the Northern and Southern Hemispheres. The standard analysis of human skin color is based on these assumptions, such that skin color is assumed to be darkest at the equator, and the decrease of skin color with latitude is assumed to be the same in both hemispheres. A nonlinear piecewise regression model was developed to test these assumptions and applied to mean skin reflectance data from 102 male samples and 65 female samples from across the Old World. For both males and females, skin reflectance (%) is lowest at the equator (darkest skin). Among males, skin reflectance increases roughly 8.2% for every 10 degrees of latitude in the Northern Hemisphere but only 3.3% for every 10 degrees of latitude in the Southern Hemisphere. Among females, the corresponding numbers are 8.1% in the Northern Hemisphere and 4.7% in the Southern Hemisphere. These results indicate that human skin color is darker in the Southern Hemisphere than in the Northern Hemisphere at equivalent latitude. Recent research shows that UV radiation is higher in the Southern Hemisphere than in the Northern Hemisphere at similar latitude. This difference, relating to astronomical and climatic conditions, may have existed in the past at different times and perhaps influenced the evolution of human skin color. Am J Phys Anthropol 104:449–457, 1997. © 1997 Wiley-Liss, Inc.     

How to reach linguistic consensus: a proof of convergence for the naming game

In this paper we introduce a mathematical model of naming games. Naming games have been widely used within research on the origins and evolution of language. Despite the many interesting empirical results these studies have produced, most of this research lacks a formal elucidating theory. In this paper we show how a population of agents can reach linguistic consensus, i.e. learn to use one common language to communicate with one another. Our approach differs from existing formal work in two important ways: one, we relax the too strong assumption that an agent samples infinitely often during each time interval. This assumption is usually made to guarantee convergence of an empirical learning process to a deterministic dynamical system. Two, we provide a proof that under these new realistic conditions, our model converges to a common language for the entire population of agents. Finally the model is experimentally validated.     

Group-size diversity in public goods games

Public goods games are models of social dilemmas where cooperators pay a cost for the production of a public good while defectors free ride on the contributions of cooperators. In the traditional framework of evolutionary game theory, the payoffs of cooperators and defectors result from interactions in groups formed by binomial sampling from an infinite population. Despite empirical evidence showing that group-size distributions in nature are highly heterogeneous, most models of social evolution assume that the group size is constant. In this article, I remove this assumption and explore the effects of having random group sizes on the evolutionary dynamics of public goods games. By a straightforward application of Jensen's inequality, I show that the outcome of general nonlinear public goods games depends not only on the average group size but also on the variance of the group-size distribution. This general result is illustrated with two nonlinear public goods games (the public goods game with discounting or synergy and the N-person volunteer's dilemma) and three different group-size distributions (Poisson, geometric, and Waring). The results suggest that failing to acknowledge the natural variation of group sizes can lead to an underestimation of the actual level of cooperation exhibited in evolving populations.     

Effects of substrate color on intraspecific body color variation in the toad‐headed lizard,Phrynocephalus versicolor

Diversity in animal coloration is generally associated with adaptation to their living habitats, ranging from territorial display and sexual selection to predation or predation avoidance, and thermoregulation. However, the mechanism underlying color variation in toad‐headed Phrynocephalus lizards remains poorly understood. In this study, we investigated the population color variation of Phrynocephalus versicolor. We found that lizards distributed in dark substrate have darker dorsal coloration (melanic lizards) than populations living in light substrates. This characteristic may improve their camouflage effectiveness. A reciprocal substrate translocation experiment was conducted to clarify the potential role of morphological adaptation and physiological plasticity of this variation. Spectrometry technology and digital photography were used to quantify the color variation of the above‐mentioned melanic and nonmelanic P. versicolor populations and their native substrate. Additionally, substrate color preference in both populations was investigated with choice experiments. Our results indicate that the melanic and nonmelanic populations with remarkable habitat color difference were significantly different on measured reflectance, luminance, and RGB values. Twenty‐four hours, 30 days, and 60 days of substrate translocation treatment had little effects on dorsal color change. We also found that melanic lizards choose to live in dark substrate, while nonmelanic lizards have no preference for substrate color. In conclusion, our results support that the dorsal coloration of P. versicolor, associated with substrate color, is likely a morphological adaptation rather than phenotypic plasticity.     

Color categorization and color constancy in a neural network model of V4

We develop a neural network model that instantiates color constancy and color categorization in a single unified framework. Previous models achieve similar effects but ignore important biological constraints. Color constancy in this model is achieved by a new application of the double opponent cells found in the blobs of the visual cortex. Color categorization emerges naturally, as a consequence of processing chromatic stimuli as vectors in a four-dimensional color space. A computer simulation of this model is subjected to the classic psychophysical tests that first uncovered these phenomena, and its response matches psychophysical results very closely.     

Using Delaunay triangulation to sample whole‐specimen color from digital images

1. Color variation is one of the most obvious examples of variation in nature, but biologically meaningful quantification and interpretation of variation in color and complex patterns are challenging. Many current methods for assessing variation in color patterns classify color patterns using categorical measures and provide aggregate measures that ignore spatial pattern, or both, losing potentially important aspects of color pattern.
2. Here, we present Colormesh, a novel method for analyzing complex color patterns that offers unique capabilities. Our approach is based on unsupervised color quantification combined with geometric morphometrics to identify regions of putative spatial homology across samples, from histology sections to whole organisms. Colormesh quantifies color at individual sampling points across the whole sample.
3. We demonstrate the utility of Colormesh using digital images of Trinidadian guppies (Poecilia reticulata), for which the evolution of color has been frequently studied. Guppies have repeatedly evolved in response to ecological differences between up‐ and downstream locations in Trinidadian rivers, resulting in extensive parallel evolution of many phenotypes. Previous studies have, for example, compared the area and quantity of discrete color (e.g., area of orange, number of black spots) between these up‐ and downstream locations neglecting spatial placement of these areas. Using the Colormesh pipeline, we show that patterns of whole‐animal color variation do not match expectations suggested by previous work.
4. Colormesh can be deployed to address a much wider range of questions about color pattern variation than previous approaches. Colormesh is thus especially suited for analyses that seek to identify the biologically important aspects of color pattern when there are multiple competing hypotheses or even no a priori hypotheses at all.

     

Evolutionary dynamics in structured populations

Evolutionary dynamics shape the living world around us. At the centre of every evolutionary process is a population of reproducing individuals. The structure of that population affects evolutionary dynamics. The individuals can be molecules, cells, viruses, multicellular organisms or humans. Whenever the fitness of individuals depends on the relative abundance of phenotypes in the population, we are in the realm of evolutionary game theory. Evolutionary game theory is a general approach that can describe the competition of species in an ecosystem, the interaction between hosts and parasites, between viruses and cells, and also the spread of ideas and behaviours in the human population. In this perspective, we review the recent advances in evolutionary game dynamics with a particular emphasis on stochastic approaches in finite sized and structured populations. We give simple, fundamental laws that determine how natural selection chooses between competing strategies. We study the well-mixed population, evolutionary graph theory, games in phenotype space and evolutionary set theory. We apply these results to the evolution of cooperation. The mechanism that leads to the evolution of cooperation in these settings could be called ‘spatial selection’: cooperators prevail against defectors by clustering in physical or other spaces.     

Egg color variation,but not egg rejection behavior,changes in a cuckoo host breeding in the absence of brood parasitism

Interactions between parasitic cuckoos and their songbird hosts form a classical reciprocal “arms race,” and are an excellent model for understanding the process of coevolution. Changes in host egg coloration via the evolution of interclutch variation in egg color or intraclutch consistency in egg color are hypothesized counter adaptations that facilitate egg recognition and thus limit brood parasitism. Whether these antiparasitism strategies are maintained when the selective pressure of parasitism is relaxed remains debated. However, introduced species provide unique opportunities for testing the direction and extent of natural selection on phenotypic trait maintenance and variation. Here, we investigated egg rejection behavior and egg color polymorphism in the red‐billed leiothrix (Leiothrix lutea), a common cuckoo (Cuculus canorus) host, in a population introduced to Hawaii 100 years ago (breeding without cuckoos) and a native population in China (breeding with cuckoos). We found that egg rejection ability was equally strong in both the native and the introduced populations, but levels of interclutch variation and intraclutch consistency in egg color in the native population were higher than in the introduced population. This suggests that egg rejection behavior in hosts can be maintained in the absence of brood parasitism and that egg appearance is maintained by natural selection as a counter adaptation to brood parasitism. This study provides rare evidence that host antiparasitism strategies can change under parasite‐relaxed conditions and reduced selection pressure.     

Participation costs dismiss the advantage of heterogeneous networks in evolution of cooperation

Real social interactions occur on networks in which each individual is connected to some, but not all, of others. In social dilemma games with a fixed population size, heterogeneity in the number of contacts per player is known to promote evolution of cooperation. Under a common assumption of positively biased pay-off structure, well-connected players earn much by playing frequently, and cooperation once adopted by well-connected players is unbeatable and spreads to others. However, maintaining a social contact can be costly, which would prevent local pay-offs from being positively biased. In replicator-type evolutionary dynamics, it is shown that even a relatively small participation cost extinguishes the merit of heterogeneous networks in terms of cooperation. In this situation, more connected players earn less so that they are no longer spreaders of cooperation. Instead, those with fewer contacts win and guide the evolution. The participation cost, or the baseline pay-off, is irrelevant in homogeneous populations, but is essential for evolutionary games on heterogeneous networks.     

Sexual selection and trichromatic color vision in primates: statistical support for the preexisting-bias hypothesis

The evolution of trichromatic color vision in primates may improve foraging performance as well as intraspecific communication; however, the context in which color vision initially evolved is unknown. We statistically examined the hypothesis that trichromatic color vision in primates represents a preexisting bias for the evolution of red coloration (pelage and/or skin) through sexual selection. Our analyses show that trichromatic color vision evolved before red pelage and red skin, as well as before gregarious mating systems that would promote sexual selection for visual traits and other forms of intraspecific communication via red traits. We also determined that both red pelage and red skin were more likely to evolve in the presence of color vision and mating systems that promote sexual selection. These results provide statistical support for the hypothesis that trichromatic color vision in primates evolved in a context other than intraspecific communication with red traits, most likely foraging performance, but, once evolved, represented a preexisting bias that promoted the evolution of red traits through sexual selection.     

Mating status correlates with dorsal brightness in some but not all poison frog populations

Sexual signals are important for intraspecific communication and mate selection, but their evolution may be driven by both natural and sexual selection, and stochastic processes. Strawberry poison frogs (Oophaga pumilio) show strong color divergence among populations, but coloration also varies among individuals of the same population. The importance of coloration for female mate choice has been studied intensely, and sexual selection seems to affect color divergence in strawberry poison frogs. However, the effect of coloration on mating success under field conditions has received very little attention. Furthermore, few studies examined how phenotypic variation among individuals of the same color morph affects mate selection under natural conditions. We measured the spectral reflectance of courting and noncourting individuals and their background substrates in three geographically separated populations. In one population (Sarapiquí, Costa Rica), we found that naturally occurring courting pairs of males and females had significantly brighter dorsal coloration than individual males and females not engaged in courtship interactions. Our field observations suggest that, in the wild, females prefer brighter males while the reason for the higher courtship activity of brighter females remains unclear. Overall our results imply that brightness differences among individuals of the same color morph may actually affect reproductive success in some populations of strawberry poison frogs.     

Reduced color polymorphism in a population of Bunodosoma granulifera

The proportions of carotenoids responsible for column coloration were determined for a population of the sea anemone Bunodosoma granulifera from Grand Cayman using TLC. This population exhibited a very low degree of polymorphism for these proportions, compared to previously sampled populations from Puerto Rico and Curaçao. In addition to the carotenoid pigment system a second system possessing pigments similar to bilins was found for the first time in this genus. This population was also assayed electrophoretically for enzymes encoded by 12 loci. Analysis of these data suggested that the reduced polymorphism for column color may be the result of selection. Although the proportions of carotenoids varied little, the amount of carotenoid per unit of epidermal tissue varied almost four-fold from 0.05 to 0.19 μg/mg dry wt.     

Different selective pressures shape the molecular evolution of color vision in chimpanzee and human populations

A population genetic analysis of the long-wavelength opsin (OPN1LW, "red") color vision gene in a global sample of 236 human nucleotide sequences had previously discovered nine amino acid replacement single nucleotide polymorphisms, which were found at high frequencies in both African and non-African populations and associated with an unusual haplotype diversity. Although this pattern of nucleotide diversity is consistent with balancing selection, it has been argued that a recombination "hot spot" or gene conversion within and between X-linked color vision genes alone may explain these patterns. The current analysis investigates a closely related primate with trichromatism to determine whether color vision gene amino acid polymorphism and signatures of adaptive evolution are characteristic of humans alone. Our population sample of 56 chimpanzee (Pan troglodytes) OPN1LW sequences shows three singleton amino acid polymorphisms and no unusual recombination or linkage disequilibrium patterns across the approximately 5.5-kb region analyzed. Our comparative population genetic approach shows that the patterns of OPN1LW variation in humans and chimpanzees are consistent with positive and purifying selection within the two lineages, respectively. Although the complex role of color vision has been greatly documented in primate evolution in general, it is surprising that trichromatism has followed very different selective trajectories even between humans and our closest relatives.     

Single cell heterogeneity

Multi-level heterogeneity is a fundamental but underappreciated feature of cancer. Most technical and analytical methods either completely ignore heterogeneity or do not fully account for it, as heterogeneity has been considered noise that needs to be eliminated. We have used single-cell and population-based assays to describe an instability-mediated mechanism where genome heterogeneity drastically affects cell growth and cannot be accurately measured using conventional averages. First, we show that most unstable cancer cell populations exhibit high levels of karyotype heterogeneity, where it is difficult, if not impossible, to karyotypically clone cells. Second, by comparing stable and unstable cell populations, we show that instability-mediated karyotype heterogeneity leads to growth heterogeneity, where outliers dominantly contribute to population growth and exhibit shorter cell cycles. Predictability of population growth is more difficult for heterogeneous cell populations than for homogenous cell populations. Since “outliers” play an important role in cancer evolution, where genome instability is the key feature, averaging methods used to characterize cell populations are misleading. Variances quantify heterogeneity; means (averages) smooth heterogeneity, invariably hiding it. Cell populations of pathological conditions with high genome instability, like cancer, behave differently than karyotypically homogeneous cell populations. Single-cell analysis is thus needed when cells are not genomically identical. Despite increased attention given to single-cell variation mediated heterogeneity of cancer cells, continued use of average-based methods is not only inaccurate but deceptive, as the “average” cancer cell clearly does not exist. Genome-level heterogeneity also may explain population heterogeneity, drug resistance, and cancer evolution.