Artificial selection for food colour preferences

Colour is an important factor in food detection and acquisition by animals using visually based foraging. Colour can be used to identify the suitability of a food source or improve the efficiency of food detection, and can even be linked to mate choice. Food colour preferences are known to exist, but whether these preferences are heritable and how these preferences evolve is unknown. Using the freshwater fish Poecilia reticulata, we artificially selected for chase behaviour towards two different-coloured moving stimuli: red and blue spots. A response to selection was only seen for chase behaviours towards the red, with realized heritabilities ranging from 0.25 to 0.30. Despite intense selection, no significant chase response was recorded for the blue-selected lines. This lack of response may be due to the motion-detection mechanism in the guppy visual system and may have novel implications for the evolvability of responses to colour-related signals. The behavioural response to several colours after five generations of selection suggests that the colour opponency system of the fish may regulate the response to selection.

Flight initiation distances in relation to sexual dichromatism and body size in birds from three continents

Predators exert strong selection pressures on their prey. Prey would therefore benefit by adjusting their behaviour to the risk of predation, while predators conversely would benefit from adjusting their behaviour to that of their prey. Extravagant ornamentation has evolved to attract mates and/or successfully compete with conspecifics of the same sex to secure high mating success, even if that occurs at a cost of increased risk of predation. Thus, sexually dichromatic species may be more susceptible to predation than sexually monochromatic species, and the presence of compensation is indicative of such species being more vulnerable. If extravagant ornamentation is costly in terms of predation risk, then we should expect sexually dichromatic species to have longer flight initiation distances (FID) than sexually monochromatic species. If ornamentation is acquired as a handicap with only individuals in prime condition being able to display with the smallest viability cost, we should expect sexually dichromatic individuals to have shorter FID than sexually monochromatic individuals. Such differences among individuals should, on an evolutionary time scale, translate into differences in FID being related to differences in sexual dichromatism among species. We investigated the relationship between FID and sexual dichromatism in phylogenetic analyses, while accounting for effects of continent (Australia, North America, and Europe), body mass, the interaction between sexual dichromatism and body mass and the interaction between sexual dichromatism and continent. In an analysis of 447 species we found shorter FID in sexually dichromatic than in sexually monochromatic species (consistent with the handicap hypothesis because sexually dichromatic species took greater risks), especially so at large body size. FID differed among continents and the relative difference in FID between sexually monochromatic and sexually dichromatic species was larger in Europe than in Australia and North America. These differences among continents may be attributed to latitudinal effects of predation. These findings are important for current ideas about the evolution of secondary sexual characters because they imply covarying continental differences in predation, especially for large bodied sexually dichromatic species.

Physiological, morphological and behavioural effects of selecting zebra finches for divergent levels of corticosterone

The effects of environmental stress on the physiology and behaviour of higher vertebrates has become an important avenue of research in recent years. Evidence from recent studies has suggested that the avian stress-related hormone corticosterone (CORT) may play a role in immunocompetence and sexual selection. We tested whether CORT is immunosuppressive by studying humoral and cell-mediated immune responses in populations of captive zebra finches selected for divergent peak levels of CORT. We also investigated whether selection for peak CORT has an effect on the quality of several sexually selected regions of the male zebra finch; in addition we compared morphometric parameters and the dominance ranking in males from the different selection lines. We also tested whether different components of the immune system compete for limited resources. We found that selection for divergent levels of peak CORT had little effect on humoral immunity, male sexual signal quality or dominance ranking. However, contrary to expectations, we did find a positive relationship between CORT titre and cell-mediated immunity, as well as a greater cell-mediated response in the birds selected for high CORT titre than those selected for low CORT titre. Consistent with predictions, significant negative relationships were found between both testosterone and CORT titre on humoral immunity. Birds from the low CORT lines were significantly larger in terms of skeletal size than those from the high CORT lines. Overall, our results suggest that the cell-mediated immune response is associated with a reduction in the humoral response, but only in males, and that there is no simple relationship between peak CORT levels and immune function.

Song as an honest signal of developmental stress in the zebra finch (Taeniopygia guttata)

In a wide range of bird species, females have been shown to express active preferences for males that sing more complex songs. Current sexual selection theory predicts that for this signal to remain an honest indicator of male quality, it must be associated with an underlying cost of development or maintenance. There has been considerable debate questioning the costs associated with song production and learning. Recently, the nutritional stress hypothesis proposed that song complexity could act as an indicator of early developmental history, since the song control nuclei in the brain are laid down early in life. Here we test the nutritional stress hypothesis, by investigating the effects of dietary stress on the quality of adult song produced. In addition, we tested the effects of elevated corticosterone during development on song production to test its possible involvement in mediating the effects of developmental stress. The results demonstrate that both dietary restriction and elevated corticosterone levels significantly reduced nestling growth rates. In addition, we found that experimentally stressed birds developed songs with significantly shorter song motif duration and reduced complexity. These results provide novel experimental evidence that complex song repertoires may have evolved as honest signals of male quality, by indicating early developmental rearing conditions.

Song sharing and repertoire size in the sedge warbler, Acrocephalus schoenobaenus: changes within and between years

The complex song of the male sedge warbler functions mainly in sexual attraction and the evolution of repertoire size is driven primarily by female choice. As male song ceases upon pairing, male–male singing interactions are relatively brief and have not been studied to our knowledge. This study shows that young males in their first breeding season shared significantly more syllables with their nearest neighbour than with their fathers or more distant males. Moreover, daily recordings revealed that rapid learning and modification of syllable repertoires occurred, resulting in a progressive increase in sharing within just a few days. This does not lead to a gradual increase in repertoire size as some syllables are dropped and new ones are acquired. This turnover process allows males to share syllables with their neighbours, whilst repertoire size, known to be important in female choice, remains relatively constant in any one year. Individual males were followed for several years and also showed considerable syllable turnover between years. However, in this case, repertoire size was found to increase between years, the largest increase occurring between the first and second years. We obtained a significant positive correlation between repertoire size and age, suggesting that females choosing males with larger repertoires may gain indirect (genetic) benefits for their offspring, such as good genes for viability. Whilst these results reveal a more flexible picture of repertoire turnover than previously suspected, the relative stability of repertoire size within a season and the increase with age suggests that repertoire size remains a likely target for sexual selection by female choice.

Mate choice in zebra finches: does corticosterone play a role?

The importance of stress as a factor in influencing life history strategies has received considerable attention in recent years, because it appears to have a substantial impact on an individual's behaviour and physiology. Birds respond to environmental and social stressors by the production of corticosterone, a glucocorticoid hormone released by the adrenal gland. In this experiment, we tested whether female zebra finches preferred males selected to produce low or high peak levels of circulating plasma corticosterone. Plasma corticosterone and testosterone levels of the males were recorded, as were morphometric measurements and perch activity. Spectrophotometric measurements were also taken from several putatively sexually selected regions of the males. The females preferred the males from the low corticosterone lines to the high corticosterone males. In addition to, and consistent with this effect, females preferred males with the lowest corticosterone titres. Male activity, testosterone level, body size and mass had no effect on female preference. Leg and beak brightness were important, however, as were the brightness and chromaticity of the male cheek patch. These results are discussed in relation to contemporary hypotheses in sexual selection, particularly in the context of stress-mediated signalling.

Developmental stress selectively affects the song control nucleus HVC in the zebra finch brain

Songbirds sing complex songs as a result of evolution through sexual selection. The evolution of such sexually selected traits requires genetic control, as well as selection on their expression. Song is controlled by a discrete neural pathway in the brain, and song complexity has been shown to correlate with the volume of specific song control nuclei. As such, the development of these nuclei, in particular the high vocal centre (HVC), is thought to be the mechanism controlling signal expression indicating male quality. We tested the hypothesis that early developmental stress selectively affects adult HVC size, compared with other brain nuclei. We did this by raising cross–fostered zebra finches (Taeniopygia guttata) under stressed and controlled conditions and determining the effect on adult HVC size. Our results confirm the strong influence of environmental conditions, particularly on HVC development, and therefore on the expression of complex songs. The results also show that both environmental and genetic factors affect the development of several brain nuclei, highlighting the developmental plasticity of the songbird brain. In all, these results explain how the complex song repertoires of songbirds can evolve as honest indicators of male quality.

Song as an honest signal of past developmental stress in the European starling (Sturnus vulgaris)

Bird song is a sexually selected male trait where females select males on the basis of song quality. It has recently been suggested that the quality of the adult male song may be determined by nutritional stress during early development. Here, we test the 'nutritional–stress hypothesis' using the complex song of the European starling. Fledgling starlings were kept under experimental treatment (unpredictable short–term food deprivations) or control conditions (ad libitum food supply), for three months immediately after independence. We measured their physiological and immune responses during the treatment and recorded song production during the following spring. Birds in the experimental group showed increased mass during the treatment and also a significantly suppressed humoral response compared with birds in the control group. There was no difference between the groups in the cell–mediated response. Next spring, males in the experimental group spent less time singing, sang fewer song bouts, took longer to start singing and also sang significantly shorter song bouts. These data support the hypothesis that both the quality and quantity of song produced by individual birds reflect past developmental stress. The results also suggest the 'nutritional–stress hypothesis' is best considered as a more general 'developmental–stress hypothesis'.

Color signals in the Black Grouse (Tetrao tetrix) : signal properties and their condition dependency

Recent progress in techniques of quantifying between‐individual differences of color‐based ornaments has revealed undiscovered possibilities for research in sexual selection. We present how the color spectra data can be comprehensively used for studying the importance of sexual ornaments in the black grouse and how these ornaments are related to a male condition. For this, we used both correlative field and experimental data. Field data indicated that older males had more chromatic coloration than yearlings. Blue chroma of males was correlated with male mating success. We experimentally manipulated yearling birds with testosterone implants and found that testosterone‐implanted males had impaired expression of several sexual ornaments: 10 months after the implantation, both structural‐based blue and carotenoid‐based red eye comb coloration were diminished, as well as lyre (tail) length. However, the manipulation did not affect vital traits under natural selection (wing length or body mass). Our data indicate that structural color is an important trait in sexual selection in this lekking species. Importantly, the data also indicate that male sexual ornaments are more susceptible to environmental conditions than the other male traits, thus showing their heightened condition dependency compared with the other traits mediating the honesty of signaling.

Parasites affect song complexity and neural development in a songbird

There is now considerable evidence that female choice drives the evolution of song complexity in many songbird species. However, the underlying basis for such choice remains controversial. The developmental stress hypothesis suggests that early developmental conditions can mediate adult song complexity by perturbing investment in the underlying brain nuclei during their initial growth. Here, we show that adult male canaries (Serinus canaria), infected with malaria (Plasmodium relictum) as juveniles, develop simpler songs as adults compared to uninfected individuals, and exhibit reduced development of the high vocal centre (HVC) song nucleus in the brain. Our results show how developmental stress not only affects the expression of a sexually selected male trait, but also the structure of the underlying song control pathway in the brain, providing a direct link between brain and behaviour. This novel experimental evidence tests both proximate and ultimate reasons for the evolution of complex songs and supports the Hamilton–Zuk hypothesis of parasite-mediated sexual selection. Together, these results propose how developmental costs may help to explain the evolution of honest advertising in the complex songs of birds.

Developmental stress, social rank and song complexity in the European starling (Sturnus vulgaris)

Bird song is a sexually selected trait and females have been shown to prefer males that sing more complex songs. However, for repertoire size to be an honest signal of male quality it must be associated with some form of cost. This experiment investigates the effects of food restriction and social status during development on song complexity in the European starling (Sturnus vulgaris). Birds that experienced an unpredictable food supply early in life produced a significantly smaller repertoire of song phrases than those with a constant food supply. Social status during development was also significantly correlated with repertoire size, with dominant birds producing more phrase types. This study therefore provides novel evidence that social as well as nutritional history may be important in shaping the song signal in this species.

Developmental stress affects the attractiveness of male song and female choice in the zebra finch (Taeniopygia guttata)

Developmental stress has recently been shown to have adverse effects upon adult male song structure in birds, which may well act as an honest signal of male quality to discriminating females. However, it still remains to be shown if females can discriminate between the songs of stressed and non-stressed males. Here we use a novel experimental design using an active choice paradigm to investigate preferences in captive female zebra finches (Taeniopygia guttata). Nine females were exposed to ten pairs of songs by previously stressed and non-stressed birds that had learned their song from the same tutor. Song pairs differed significantly in terms of song complexity, with songs of stressed males exhibiting lower numbers of syllables and fewer different syllables in a phrase. Song rate and peak frequency did not differ between stressed and non-stressed males. Females showed a significant preference for non-stressed songs in terms of directed perching activity and time spent on perches. Our results therefore indicate that developmental stress affects not only the structure of male song, but that such structural differences are biologically relevant to female mate choice decisions.

Ultraviolet vision in birds: what is its function?

Although UV vision was first demonstrated in birds in the early 1970s, its function is still unknown. Here we review the evidence for UV vision in birds, discuss the special properties of UV light, lay out in detail hypotheses for the function of UV vision in birds and discuss their plausibility. The main hypotheses are that UV vision functions: (i) in orientation, (ii) in foraging and (iii) in signalling. The first receives support from studies of homing pigeons, but it would be unwise to conclude that orientation is UV's primary function in all birds. It is especially important to test the signalling hypothesis because bird plumage often reflects UV and tests of theories of sexual selection have virtually always assumed that birds perceive plumage ''colours'' as humans do. A priori this assumption is unlikely to be correct, for unlike humans, birds see in the UV, have at least four types of cones and have a system of oil droplets which filters light entering individual cones.

Ultraviolet plumage colors predict mate preferences in starlings

Avian plumage has long been used to test theories of sexual selection, with humans assessing the colors, However, many birds see in the ultraviolet (<400 nm), to which humans are blind, Consequently, it is important to know whether natural variation in UV reflectance from plumage functions in sexual signaling, We show that female starlings rank males differently when UV wavelengths are present or absent, Principal component analysis of approximate to 1300 reflectance spectra (300-700 nm) taken from sexually dimorphic plumage regions of males predicted preference under the UV+ treatment. Under UV- conditions, females ranked males in a different and nonrandom order, but plumage reflectance in the human visible spectrum did not predict choice, Natural variation in UV reflectance is thus important in avian mate assessment, and the prevailing light environment can have profound effects on observed mating preferences.

Ultraviolet vision and mate choice in zebra finches

SEXUAL selection is one of the most actively studied areas of evolutionary biology(1-3), and ever since Darwin(1) birds have been probably the most popular taxon for testing the predictions about colour variation. Humans have been used to assess 'colour', an approach which may be flawed(4,5) as many birds see in the ultraviolet (to which humans are blind), and have at least four spectral classes of retinal cone cells (humans have only three), Here we report experiments on zebra finches which test the hypothesis that the ultraviolet waveband (300-400 nm) is used in avian mate-choice decisions. We found that the ultraviolet is used, and that it probably contributes to hue perception. This finding may have,vide implications for future studies of avian sexual selection and colour, and supports one hypothesized function of avian ultraviolet vision, the role of which is largely unknown.(4,6,7)