The colour hexagon: a chromaticity diagram based on photoreceptor excitations as a generalized representation of colour opponency

Summary A chromaticity diagram which plots the 3 photoreceptor excitations of trichromatic colour vision systems at an angle of 120° is presented. It takes into acount the nonlinear transduction process in the receptors. The resulting diagram has the outline of an equilateral hexagon. It is demonstrated by geometrical means that excitation values for any type of spectrally opponent mechanism can be read from this diagram if the weighting factors of this mechanism add up to zero. Thus, it may also be regarded as a general representation of colour opponent relations, linking graphically the Young-Helmholtz theory of trichromacy and Hering's concept of opponent colours. It is shown on a geometrical. basis that chromaticity can be coded unequivocally by any two combined spectrally opponent mechanisms, the main difference between particular mechanisms being the extension and compression of certain spectral areas. This type of graphical representation can qualitatively explain the Bezold-Brücke phenomenon. Furthermore, colour hexagon distances may be taken as standardized perceptual colour distance values for trichromatic insects, as is demonstrated by comparison with behavioural colour discrimination data of 3 hymenopteran species.     

Behavioural evidence for body colour signaling in the fiddler crab Uca perplexa (Brachyura: Ocypodidae)

In the reproductive season, mature females of the fiddler crab Uca perplexa leave their burrows and wander about their habitat for mating. To clarify whether the fiddler crabs respond to colour or luminosity, I examined the behavioural responses of the males to the wandering females before and after the females were painted white, red, black or blue. The behaviours of the males were categorized into three types: lateral–circular wave and lateral–straight wave for courting, and repelling. Before painting, almost all of the males courted the females. After painting, significantly fewer males courted the red-, black- and blue-painted females than courted the white-painted females. These results mean that the fiddler crabs can discriminate colours or luminosity. The role of body colour as a visual signal in crab society is discussed.     

Floral colour variation of Nicotiana glauca in native and non-native ranges: Testing the role of pollinators' perception and abiotic factors

• Invasive plants displaying disparate pollination environments and abiotic conditions in native and non-native ranges provide ideal systems to test the role of different ecological factors driving flower colour variation.
• We quantified corolla reflectance of the ornithophilous South American Nicotiana glauca in native populations, where plants are pollinated by hummingbirds, and in populations from two invaded regions: South Africa, where plants are pollinated by sunbirds, and the Balearic island of Mallorca, where plants reproduce by selfing. Using visual modelling we examined how corolla reflectance could be perceived by floral visitors present in each region. Through Mantel tests we assessed a possible association between flower colour and different abiotic factors.
• Corolla reflectance variation (mainly along medium to long wavelengths, i.e. human green-yellow to red colours) was greater among studied regions than within them. Flower colour was more similar between South America and South Africa, which share birds as pollinators. Within invaded regions, corolla reflectance variation was lower in South Africa, where populations could not be distinguished from each other by sunbirds, than in Spain, where populations could be distinguished from each other by their occasional visitors. Differences in corolla colour among populations were partially associated with differences in temperature.
• Our findings suggest that shifts in flower colour of N. glauca across native and invaded ranges could be shaped by changes in both pollination environment and climatic factors. This is the first study on plant invasions considering visual perception of different pollinators and abiotic drivers of flower colour variation.

     

Defective colour vision and taste sensitivity to PTC in the Maharashtrian population

Three caste groups of the Maharashtrian population, namely Brahmin (N=58), Maratha (N=989) and Scheduled caste (N=1073), were studied for defective colour vision and for the ability to taste phenylthiocarbamide (PTC). Comparisons of defective colour vision and PTC taste sensitivity were made with other Maharashtrian populations studied previously by various authors. The incidence of defective colour vision is lowest among the Brahmins (3.44%) and highest in the Scheduled caste people (4.28%). Taste sensitivity to PTC is highest among the Brahmins (72.5%) with a value which is close to the Vednagar Brahmins (73.3%) who show the highest frequency of the ‘T’ gene recorded so far in Maharashtra.     

Natural selection in a postglacial range expansion: the case of the colour cline in the European barn owl

Gradients of variation—or clines—have always intrigued biologists. Classically, they have been interpreted as the outcomes of antagonistic interactions between selection and gene flow. Alternatively, clines may also establish neutrally with isolation by distance (IBD) or secondary contact between previously isolated populations. The relative importance of natural selection and these two neutral processes in the establishment of clinal variation can be tested by comparing genetic differentiation at neutral genetic markers and at the studied trait. A third neutral process, surfing of a newly arisen mutation during the colonization of a new habitat, is more difficult to test. Here, we designed a spatially explicit approximate Bayesian computation (ABC) simulation framework to evaluate whether the strong cline in the genetically based reddish coloration observed in the European barn owl (Tyto alba) arose as a by‐product of a range expansion or whether selection has to be invoked to explain this colour cline, for which we have previously ruled out the actions of IBD or secondary contact. Using ABC simulations and genetic data on 390 individuals from 20 locations genotyped at 22 microsatellites loci, we first determined how barn owls colonized Europe after the last glaciation. Using these results in new simulations on the evolution of the colour phenotype, and assuming various genetic architectures for the colour trait, we demonstrate that the observed colour cline cannot be due to the surfing of a neutral mutation. Taking advantage of spatially explicit ABC, which proved to be a powerful method to disentangle the respective roles of selection and drift in range expansions, we conclude that the formation of the colour cline observed in the barn owl must be due to natural selection.     

Turn the temperature to turquoise: cues for colour change in the male chameleon grasshopper (Kosciuscola tristis) (Orthoptera: Acrididae)

Rapid, reversible colour change is unusual in animals, but is a feature of male chameleon grasshoppers (Kosciuscola tristis). Understanding what triggers this colour change is paramount to developing hypotheses explaining its evolutionary significance. In a series of manipulative experiments the author quantified the effects of temperature, and time of day, as well as internal body temperature, on the colour of male K. tristis. The results suggest that male chameleon grasshoppers change colour primarily in response to temperature and that the rate of colour change varies considerably, with the change from black to turquoise occurring up to 10 times faster than the reverse. Body temperature changed quickly (within 10 min) in response to changes in ambient temperature, but colour change did not match this speed and thus colour is decoupled from internal temperature. This indicates that male colour change is driven primarily by ambient temperature but that their colour does not necessarily reflect current internal temperature. I propose several functional hypotheses for male colour change in K. tristis.     

High sensitivity to short wavelengths in a lizard and implications for understanding the evolution of visual systems in lizards

Progress in developing animal communication theory is frequently constrained by a poor understanding of sensory systems. For example, while lizards have been the focus of numerous studies in visual signalling, we only have data on the spectral sensitivities of a few species clustered in two major clades (Iguania and Gekkota). Using electroretinography and microspectrophotometry, we studied the visual system of the cordylid lizard Platysaurus broadleyi because it represents an unstudied clade (Scinciformata) with respect to visual systems and because UV signals feature prominently in its social behaviour. The retina possessed four classes of single and one class of double cones. Sensitivity in the ultraviolet region (UV) was approximately three times higher than previously reported for other lizards. We found more colourless oil droplets (associated with UV-sensitive (UVS) and short wavelength-sensitive (SWS) photoreceptors), suggesting that the increased sensitivity was owing to the presence of more UVS photoreceptors. Using the Vorobyev-Osorio colour discrimination model, we demonstrated that an increase in the number of UVS photoreceptors significantly enhances a lizard's ability to discriminate conspecific male throat colours. Visual systems in diurnal lizards appear to be broadly conserved, but data from additional clades are needed to confirm this.     

Sometimes slower is better: slow-exploring birds are more sensitive to changes in a vocal discrimination task

Animal personality, defined as consistent individual differences across context and time, has attracted much recent research interest in the study of animal behaviour. More recently, this field has begun to examine how such variation arose and is maintained within populations. The habitat-dependent selection hypothesis, which posits that animals with differing personality types may fare better (i.e. have a fitness advantage) in different habitats, suggests one possible mechanism. In the current experiment, we tested whether slow- and fast-exploring black-capped chickadees (Poecile atricapillus), determined by performance in a novel environment exploration task, perform differentially when the demands of an acoustic operant discrimination (cognitive) task were altered following successful task acquisition. We found that slow-exploring birds learn to reverse previously learned natural category rules more quickly than faster exploring conspecifics. In accordance with the habitat-dependent selection hypothesis, and previous work with great tits (Parus major), a close relative of the black-capped chickadee, our results suggest that fast-exploring birds may perform better in stable, predictable environments where forming a routine is advantageous, while slow-exploring birds are favoured in unstable, unpredictable environments, where task demands often change. Our results also support a hypothesis derived from previous work with great tits; slow-exploring birds may be generally more flexible (i.e. able to modify their behaviour in accordance with changes in environmental stimuli) in some learning tasks.     

From silk to satellite: half a century of ocean colour anomalies in the Northeast Atlantic

Changes in phytoplankton dynamics influence marine biogeochemical cycles, climate processes, and food webs, with substantial social and economic consequences. Large‐scale estimation of phytoplankton biomass was possible via ocean colour measurements from two remote sensing satellites – the Coastal Zone Colour Scanner (CZCS, 1979–1986) and the Sea‐viewing Wide Field‐of‐view Sensor (SeaWiFS, 1998–2010). Due to the large gap between the two satellite eras and differences in sensor characteristics, comparison of the absolute values retrieved from the two instruments remains challenging. Using a unique in situ ocean colour dataset that spans more than half a century, the two satellite‐derived chlorophyll‐a (Chl‐a) eras are linked to assess concurrent changes in phytoplankton variability and bloom timing over the Northeast Atlantic Ocean and North Sea. Results from this unique re‐analysis reflect a clear increasing pattern of Chl‐a, a merging of the two seasonal phytoplankton blooms producing a longer growing season and higher seasonal biomass, since the mid‐1980s. The broader climate plays a key role in Chl‐a variability as the ocean colour anomalies parallel the oscillations of the Northern Hemisphere Temperature (NHT) since 1948.     

Host discrimination and evidence for a host marking pheromone in the almond seed wasp, Eurytoma amygdali

Laboratory experiments and observations on the oviposition behaviour of the almond seed wasp Eurytoma amygdali Enderlein (Hymenoptera, Eurytomidae) revealed that the females of this species deposit a host-marking pheromone, immediately after an oviposition, by dragging the tip of their abdomen on the fruit surface. This pheromone enables them to discriminate between the infested and uninfested fruit and to select for oviposition the latter. Its primary function is apparently the prevention of repeated ovipositions in already infested fruit, thus contributing to the optimal utilisation of the available resources for larval development. The responses of individual females to different treatments of almonds, in a series of two-choice tests, revealed that the pheromone can be perceived by the females on direct contact and, when at high concentrations, also olfactorily from a short distance. The pheromone was present inside the abdomen and thorax of females but not of males, and, although water soluble, could not be entirely removed from heavily infested almonds when rinsed with water. Direct observations revealed that after an average of 3.7 successive visits to pheromone-bearing almonds, females were induced to walk or, most often, fly away from the experimental set-up. This suggests that the pheromone may also contribute to the dispersion of the wasps.

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Résumé Des expériences de laboratoire et des observations concernant le comportement de ponte de Eurytoma amygdali Enderlein (Hymenoptera, Eurytomidae) ont montré que, immédiatement après la ponte, les femelles déposent une phéromone de marquage de l'hôte en trainant le bout de leur abdomen sur la surface de l'amande. Cette phéromone les rends capable de distinguer les fruits infectés des non-infectés dt de sélectionner pour la ponte les derniers. Apparemment, la fonction principale de cette phéromone est la prévention de la répétition des ovipositions dans les fruits déjà infectés et la répartition uniforme des oeufs dans les amandes, contribuant ainsi à la meilleure utilisation des ressources disponsibles pour le développement des larves. Des expériences de deux choix entre des fruits de différents traitements ont montré que la phéromone pouvait être perçue par les femelles par le direct contact et, quand elle était à hautes concentrations, par olfaction d'une courte distance. La phéromone était présente dans l'abdomen et dans le thorax des femelles, et bien qu'elle soit soluble à l'eau, elle ne pouvait pas s'éloigner entièrement par lavage des amandes sérieusement infectées à l'eau. Les observations ont démontré qu'après un numéro de 3.7 visites successives sur des amandes portant de la phéromone les femelles s'éloignaient du lieu de ponte en marchant ou, le plus souvant, en s'envolant, ce qui suggère que la phéromone contribue à la dispersion des femelles.

     

Flower colour variation in the montane plant Gentiana lutea L. (Gentianaceae) is unrelated to abiotic factors

Background: Flower colour variation among populations may result from the spatial variation of selective agents. The structure of phenotypic variation informs on the ecological processes related to this variation. Variation in floral traits is mainly attributed to variation in the pollinator fauna, while variation in vegetative traits is usually linked to abiotic factors or herbivores.

Aims: We investigated the geographical variation of flower colour (and correlated traits) in Gentiana lutea and the relationship with the variation in abiotic factors.

Methods: Phenotypic variation (flower colour, petal length, petal width, stalk length, leaf length, flower number, petal number and number of basal leaves) was assessed in 429 plants of 12 populations located at north-west Iberia. Additionally, we obtained data on the geographical coordinates, elevation, temperature, rainfall and radiation for each population.

Results: Populations mostly differed in flower colour, from orange to yellow from west to the east. Abiotic factors were unrelated to variation in either floral or vegetative traits.

Conclusions: Phenotypic variation among G. lutea populations does not result from adaptation to environmental factors. Other factors, such as historical events or selective pressure exerted by biotic interactions, might explain the flower colour variation in G. lutea along the Cantabrian Range.     

Beauty in the eyes of the beholders: colour vision is tuned to mate preference in the Trinidadian guppy (Poecilia reticulata)

A broad range of animals use visual signals to assess potential mates, and the theory of sensory exploitation suggests variation in visual systems drives mate preference variation due to sensory bias. Trinidadian guppies (Poecilia reticulata), a classic system for studies of the evolution of female mate choice, provide a unique opportunity to test this theory by looking for covariation in visual tuning, light environment and mate preferences. Female preference co‐evolves with male coloration, such that guppy females from ‘low‐predation’ environments have stronger preferences for males with more orange/red coloration than do females from ‘high‐predation’ environments. Here, we show that colour vision also varies across populations, with ‘low’‐predation guppies investing more of their colour vision to detect red/orange coloration. In independently colonized watersheds, guppies expressed higher levels of both LWS‐1 and LWS‐3 (the most abundant LWS opsins) in ‘low‐predation’ populations than ‘high‐predation’ populations at a time that corresponds to differences in cone cell abundance. We also observed that the frequency of a coding polymorphism differed between high‐ and low‐predation populations. Together, this shows that the variation underlying preference could be explained by simple changes in expression and coding of opsins, providing important candidate genes to investigate the genetic basis of female preference variation in this model system.     

Field observations and feeding experiments on the responses of rufous-tailed jacamars (Galbula ruficauda) to free-flying butterflies in a tropical rainforest

Wild rufous-tailed jacamars (Galbula ruficauda) were shown to prey frequently, but selectively, upon butterflies in a Costa Rican rainforest. Two individually caged birds (a male and a female) were further tested with over 1000 butterflies of 114 morphs. Both wild jacamars and the two captive individuals were able to capture and handle all kinds and sizes of local butterflies. These butterflies (and other winged insects) were recognized by the jacamars as prey only through their movement. The captive birds discriminated between an unacceptable group of butterflies, which generally fly slowly or regularly, are warningly coloured and mimetic, with transparent, or white, orange, red, and/or black coloration, and an acceptable group that generally fly fast or erratically, are cryptic (on one or both sides), and have yellow, orange, green, blue, and/or brown coloration. These different morphological and behavioural characteristics of butterflies presumably helped the jacamars to assess their palatability. Most individuals of unacceptable butterflies (e.g. Battus and Parides (Papilionidae), some Pieridae, Diaethria and Callicore (Nymphalinae), Heliconiinae, Acraeinae, Ithomiidae, and Danaidae) were sight-rejected by the male jacamar (Jacamar 2), and many of the same were also sight-rejected by the female (Jacamar 1). In cases when the above butterflies were attacked, they were quickly released and usually unharmed. The captive female bird, after long periods without food, consumed many pierid and heliconiine butterflies that were consistently rejected by the male for their distasteful and dangerous qualities. In contrast, palatable butterflies (e.g. Papilio, Charaxinae, most Nymphalinae, Morpho, Brassolinae, and Satyrinae) were usually quickly attacked and consumed. The captive jacamars were able to discriminate between the very similar colour patterns of some Batesian mimics and their models, and could memorize the palatability of a large variety of butterflies. The discriminatory abilities of specialized insectivorous birds such as jacamars are likely to play a major role in the evolution of neotropical butterfly mimicry.     

Early male production is not linked to a reproductive strategy in the Japanese paper wasp,<Emphasis Type="Italic"> Polistes chinensis antennalis</Emphasis> (Hymenoptera: Vespidae)

Diploid males in haplo-diploid insects are sterile, because they produce diploid sperm. Our previous report revealed that early males of the Japanese paper wasp Polistes chinensis antennalis are diploid but did not reveal how often this occurs. We analyzed the genotypes of early males using six microsatellite markers. Two of the 41 early males (5%) from six colonies were haploid, but the other males were diploid. This evidence suggests that we can ignore the reproductive success of the early males of P. chinensis antennalis.     

Enhancing Nanos expression via the bacterial TomO protein is a conserved strategy used by the symbiont Wolbachia to fuel germ stem cell maintenance in infected Drosophila females

The toxic manipulator of oogenesis (TomO) protein has been identified in the wMel strain of Wolbachia that symbioses with the vinegar fly Drosophila melanogaster, as a protein that affects host reproduction. TomO protects germ stem cells (GSCs) from degeneration, which otherwise occurs in ovaries of host females that are mutant for the gene Sex‐lethal (Sxl). We isolated the TomO homologs from wPip, a Wolbachia strain from the mosquito Culex quinquefasciatus. One of the homologs, TomO_wPip1, exerted the GSC rescue activity in fly Sxl mutants when lacking its hydrophobic stretches. The GSC‐rescuing action of the TomO_wPip1 variant was ascribable to its abilities to associate with Nanos (nos) mRNA and to enhance Nos protein expression. The analysis of structure–activity relationships with TomO homologs and TomO deletion variants revealed distinct modules in the protein that are each dedicated to different functions, i.e., subcellular localization, nos mRNA binding or Nos expression enhancement. We propose that modular reshuffling is the basis for structural and functional diversification of TomO protein members.     

The population demography of Betula maximowicziana,a cool‐temperate tree species in Japan,in relation to the last glacial period: its admixture‐like genetic structure is the result of simple population splitting not admixing

Conservation of the local genetic variation and evolutionary integrity of economically and ecologically important trees is a key aspect of studies involving forest genetics, and a population demographic history of the target species provides valuable information for this purpose. Here, the genetic structure of 48 populations of Betula maximowicziana was assessed using 12 expressed sequence tag–simple sequence repeat (EST‐SSR) markers. Genetic diversity was lower in northern populations than southern ones and structure analysis revealed three groups: northern and southern clusters and an admixed group. Eleven more genomic‐SSR loci were added and the demographic history of these three groups was inferred by approximate Bayesian computation (ABC). The ABC revealed that a simple split scenario was much more likely than isolation with admixture, suggesting that the admixture‐like structure detected in this species was due to ancestral polymorphisms. The ABC analysis suggested that the population growth and divergence of the three groups occurred 96 800 (95% CI, 20 500–599 000) and 28 300 (95% CI, 8700–98 400) years ago, respectively. We need to be aware of several sources of uncertainty in the inference such as assumptions about the generation time, overlapping of generations, confidence intervals of the estimated parameters and the assumed model in the ABC. However, the results of the ABC together with the model‐based maps of reconstructed past species distribution and palaeoecological data suggested that the modern genetic structure of B. maximowicziana originated prior to the last glacial maximum (LGM) and that some populations survived in the northern range even during the LGM.