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      Caution with colour calculations: spectral purity is a poor descriptor of flower colour visibility

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          Abstract

          Background

          The colours of flowers are of key interest to plant and pollination biologists. An increasing number of studies have investigated the importance of saturation of flower colours (often called ‘spectral purity’ or ‘chroma’) for visibility to pollinators, but the conceptual, physiological and behavioural foundations for these metrics as well as the calculations used rest on slender foundations.

          Methods

          We discuss the caveats of colour attributes that are derived from human perception, and in particular spectral purity and chroma, as variables in flower colour analysis. We re-analysed seven published datasets encompassing 774 measured reflectance spectra to test for correlations between colour contrast, spectral purity and chroma.

          Main findings and Conclusions

          We identify several concerns with common calculation procedures in animal colour spaces. Studies on animal colour vision provide no ground to assume that any pollinator perceives (or responds to) saturation, chroma or spectral purity in the way humans do. A re-analysis of published datasets revealed that values for colour contrast between flowers and their background are highly correlated with measures for spectral purity and chroma, which invalidates treating these factors as independent variables as is currently commonplace. Strikingly, spectral purity and chroma – both of which are metrics for saturation and are often used synonymously – are not correlated at all. We conclude that alternative, behaviourally validated metrics for the visibility of flowers to pollinators, such as colour contrast and achromatic contrast, are better in understanding the role of flower colour in plant–pollinator signalling.

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          Most cited references66

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          Evolution of avian plumage color in a tetrahedral color space: a phylogenetic analysis of new world buntings.

          We use a tetrahedral color space to describe and analyze male plumage color variation and evolution in a clade of New World buntings--Cyanocompsa and Passerina (Aves: Cardinalidae). The Goldsmith color space models the relative stimulation of the four retinal cones, using the integrals of the product of plumage reflectance spectra and cone sensitivity functions. A color is represented as a vector defined by the relative stimulation of the four cone types--ultraviolet, blue, green, and red. Color vectors are plotted in a tetrahedral, or quaternary, plot with the achromatic point at the origin and the ultraviolet/violet channel along the Z-axis. Each color vector is specified by the spherical coordinates theta, phi, and r. Hue is given by the angles theta and phi. Chroma is given by the magnitude of r, the distance from the achromatic origin. Color vectors of all distinct patches in a plumage characterize the plumage color phenotype. We describe the variation in color space occupancy of male bunting plumages, using various measures of color contrast, hue contrast and diversity, and chroma. Comparative phylogenetic analyses using linear parsimony (in MacClade) and generalized least squares (GLS) models (in CONTINUOUS) with a molecular phylogeny of the group document that plumage color evolution in the clade has been very dynamic. The single best-fit GLS evolutionary model of plumage color variation over the entire clade is a directional change model with no phylogenetic correlation among species. However, phylogenetic innovations in feather color production mechanisms--derived pheomelanin and carotenoid expression in two lineages--created new opportunities to colonize novel areas of color space and fostered the explosive differentiation in plumage color. Comparison of the tetrahedral color space of Goldsmith with that of Endler and Mielke demonstrates that both provide essentially identical results. Evolution of avian ultraviolet/violet opsin sensitivity in relation to chromatic experience is discussed.
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            Receptor noise as a determinant of colour thresholds.

            Inferences about mechanisms at one particular stage of a visual pathway may be made from psychophysical thresholds only if the noise at the stage in question dominates that in the others. Spectral sensitivities, measured under bright conditions, for di-, tri-, and tetrachromatic eyes from a range of animals can be modelled by assuming that thresholds are set by colour opponency mechanisms whose performance is limited by photoreceptor noise, the achromatic signal being disregarded. Noise in the opponency channels themselves is therefore not statistically independent, and it is not possible to infer anything more about the channels from psychophysical thresholds. As well as giving insight into mechanisms of vision, the model predicts the performance of colour vision in animals where physiological and anatomical data on the eye are available, but there are no direct measurements of perceptual thresholds. The model, therefore, is widely applicable to comparative studies of eye design and visual ecology.
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              The evolution of color vision in insects.

              We review the physiological, molecular, and neural mechanisms of insect color vision. Phylogenetic and molecular analyses reveal that the basic bauplan, UV-blue-green-trichromacy, appears to date back to the Devonian ancestor of all pterygote insects. There are variations on this theme, however. These concern the number of color receptor types, their differential expression across the retina, and their fine tuning along the wavelength scale. In a few cases (but not in many others), these differences can be linked to visual ecology. Other insects have virtually identical sets of color receptors despite strong differences in lifestyle. Instead of the adaptionism that has dominated visual ecology in the past, we propose that chance evolutionary processes, history, and constraints should be considered. In addition to phylogenetic analyses designed to explore these factors, we suggest quantifying variance between individuals and populations and using fitness measurements to test the adaptive value of traits identified in insect color vision systems.
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                Author and article information

                Contributors
                Journal
                Ann Bot
                Ann Bot
                annbot
                Annals of Botany
                Oxford University Press (US )
                0305-7364
                1095-8290
                01 July 2022
                21 June 2022
                21 June 2022
                : 130
                : 1
                : 1-9
                Affiliations
                Groningen Institute for Evolutionary Life Sciences, University of Groningen , The Netherlands
                Department of Behavioral Physiology and Sociobiology, University of Würzburg , Germany
                Author notes
                For correspondence. E-mail C.J.van.der.Kooi@ 123456rug.nl
                Author information
                https://orcid.org/0000-0003-0613-7633
                https://orcid.org/0000-0002-0734-2869
                Article
                mcac069
                10.1093/aob/mcac069
                9295922
                35726715
                cb103e9f-4559-4f3c-a0dd-d6559986904d
                © The Author(s) 2022. Published by Oxford University Press on behalf of the Annals of Botany Company.

                This is an Open Access article distributed under the terms of the Creative Commons Attribution License ( https://creativecommons.org/licenses/by/4.0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited.

                History
                : 13 January 2022
                : 09 May 2022
                : 25 May 2022
                : 26 May 2022
                : 30 June 2022
                Page count
                Pages: 9
                Funding
                Funded by: AFOSR, DOI 10.13039/100000181;
                Funded by: EOARD, DOI 10.13039/100015464;
                Award ID: FA9550-19-1-7005
                Funded by: Bavarian Research Alliance, DOI 10.13039/501100011906;
                Award ID: BayIntAn_UWUE_2020_134
                Categories
                Research in Context
                AcademicSubjects/SCI01080
                AcademicSubjects/SCI01130
                AcademicSubjects/SCI01210

                Plant science & Botany
                spectral purity,vision model,colour contrast,pollination,flower colour,saturation,chroma,plant–pollinator signalling

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