The carotenoid redshift: Physical basis and implications for visual signaling.

Carotenoid pigments are the basis for much red, orange, and yellow coloration in nature and central to visual signaling. However, as pigment concentration increases, carotenoid signals not only darken and become more saturated but they also redshift; for example, orange pigments can look red at higher concentration. This occurs because light experiences exponential attenuation, and carotenoid-based signals have spectrally asymmetric reflectance in the visible range. Adding pigment disproportionately affects the high-absorbance regions of the reflectance spectra, which redshifts the perceived hue. This carotenoid redshift is substantial and perceivable by animal observers. In addition, beyond pigment concentration, anything that increases the path length of light through pigment causes this redshift (including optical nano- and microstructures). For example, male tanagers appear redder than females, despite the same population and concentration of carotenoids, due to microstructures that enhance light-pigment interaction. This mechanism of carotenoid redshift has sensory and evolutionary consequences for honest signaling in that structures that redshift carotenoid ornaments may decrease signal honesty. More generally, nearly all colorful signals vary in hue, saturation, and brightness as light-pigment interactions change, due to spectrally asymmetrical reflectance within the visible range of the relevant species. Therefore, the three attributes of color need to be considered together in studies of honest visual signaling.