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AbstractIntraspecific variation in camouflage is common in animals. Sexual dimorphism in camouflage is less common and, where observed, attributed to trade-offs between natural selection for predator avoidance and sexual selection for conspicuous mating signals. Here we report on variation in putatively cryptic ventral hindwing patterns in the American snout butterfly, Libytheana carinenta. We use field surveys and crowdsourced data to characterize three morphs. One is found in both sexes, one is male specific, and one is female specific. The sex-specific morphs constitute a sexually dimorphic set whose frequencies change together in time. Field surveys indicate that butterflies in southern Arizona transition from midsummer dominance of the sexually monomorphic pattern to early-fall dominance of the sexually dimorphic set. Crowdsourced data indicate that the sexually dimorphic set dominates in early spring, transitioning later into a mixture of morphs dominated by the monomorphic pattern, with the dimorphic set rising in frequency again in late fall. We discuss this unique pattern of camouflage variation with respect to contemporary theory on animal coloration.

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Structural color is an optical phenomenon resulting from light interacting with nanostructured materials. Although structural color (SC) is widespread in the tree of life, the underlying genetics and genomics are not well understood. Here, we collected and sequenced a set of 87 structurally colored bacterial isolates and 30 related strains lacking SC. Optical analysis of colonies indicated that diverse bacteria from at least two different phyla (Bacteroidetes and Proteobacteria) can create two-dimensional packing of cells capable of producing SC. A pan-genome-wide association approach was used to identify genes associated with SC. The biosynthesis of uroporphyrin and pterins, as well as carbohydrate utilization and metabolism, was found to be involved. Using this information, we constructed a classifier to predict SC directly from bacterial genome sequences and validated it by cultivating and scoring 100 strains that were not part of the training set. We predicted that SCr is widely distributed within gram-negative bacteria. Analysis of over 13,000 assembled metagenomes suggested that SC is nearly absent from most habitats associated with multicellular organisms except macroalgae and is abundant in marine waters and surface/air interfaces. This work provides a large-scale ecogenomics view of SC in bacteria and identifies microbial pathways and evolutionary relationships that underlie this optical phenomenon.

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Colors are well studied in bird plumage but not in other integumentary structures. In particular, iridescent colors from structures other than plumage are undescribed in birds. Here, we show that a multilayer of keratin and lipids is sufficient to produce the iridescent bill of Spermophaga haematina. Furthermore, that the male bill is presented to the female under different angles during display provides support for the hypothesis that iridescence evolved in response to sexual selection. This is the first report of an iridescent bill, and only the second instance of iridescence in birds in which melanosomes are not involved. Furthermore, an investigation of museum specimens of an additional 98 species, showed that this evolved once, possibly twice. These results are promising, as they suggest that birds utilize a wider array of physical phenomena to produce coloration and should further stimulate research on nonplumage integumentary colors.

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BACKGROUND AND AIMS: Structural colour is responsible for the remarkable metallic blue colour seen in the leaves of several plants. Species belonging to only ten genera have been investigated to date, revealing four photonic structures responsible for structurally coloured leaves. One of these is the helicoidal cell wall, known to create structural colour in the leaf cells of five taxa. Here we investigate a broad selection of land plants to understand the phylogenetic distribution of this photonic structure in leaves. METHODS: We identified helicoidal structures in the leaf epidermal cells of 19 species using transmission electron microscopy. Pitch measurements of the helicoids were compared with the reflectance spectra of circularly polarized light from the cells to confirm the structure-colour relationship. RESULTS: By incorporating species examined with a polarizing filter, our results increase the number of taxa with photonic helicoidal cell walls to species belonging to at least 35 genera. These include 19 monocot genera, from the orders Asparagales (Orchidaceae) and Poales (Cyperaceae, Eriocaulaceae, Rapateaceae) and 16 fern genera, from the orders Marattiales (Marattiaceae), Schizaeales (Anemiaceae) and Polypodiales (Blechnaceae, Dryopteridaceae, Lomariopsidaceae, Polypodiaceae, Pteridaceae, Tectariaceae). CONCLUSIONS: Our investigation adds considerably to the recorded diversity of plants with structurally coloured leaves. The iterative evolution of photonic helicoidal walls has resulted in a broad phylogenetic distribution, centred on ferns and monocots. We speculate that the primary function of the helicoidal wall is to provide strength and support, so structural colour could have evolved as a potentially beneficial chance function of this structure.

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This study used scanning electron microscopy via 3D dissection coupled with synchrotron radiation with microfocal beams of both small-angle X-ray scattering and wide-angle X-ray diffraction to analyze the periodic crystal aggregates of unusual poly(p-dioxanone) (PPDO) dendritic cactus-arm-like ring bands upon crystallization with a diluent poly(vinyl alcohol) (PVA) that is capable of hydrogen bonding interactions with PPDO. Three-dimensional microscopy interior dissection clearly expounds that the banded periodic architectures are packed by alternately normal-oriented flat-on crystals underneath the valley, periodically interfaced/branched with horizontal-oriented edge-on fibrils underneath the ridge. The oblique angles between the valley's flat-on crystals with the branches are ca. 25-45° (depending on gradient inclines and bending), which is also proved by the azimuthal angle in microbeam X-ray diffraction. The grating-like strut-rib assembly in the PPDO cactus-arm-like ring bands is further proved by novel iridescence tests.

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Using sodium alginate (Alg) aqueous solution containing indigo carmine (IdC) at various concentrations we characterized the rippled surface pattern with micro-spacing on a flexible film as intriguing bluish Alg-IdC iridescence. The characterization was performed using Fourier-transform infrared spectroscopy, ultraviolet-visible spectroscopy, field emission scanning electron microscopy, atomic force microscopy, electron microscopy, differential scanning calorimetry, thermogravimetric analysis, X-ray diffraction analysis, and photoluminescence detection. The edge pattern on the film had a maximum depth of 825 nm, a peak-to-peak distance of 63.0 nm, and an average distance of 2.34 nm. The center of the pattern had a maximum depth of 343 nm and a peak-to-peak distance of 162 nm. The pattern spacing rippled irregularly, widening toward the center and narrowing toward the edges. The rippled nano-patterned areas effectively generated iridescence. The ultraviolet absorption spectra of the mixture in the 270 and 615 nm ranges were the same for both the iridescent and non-iridescent film surfaces. By adding Ag+ ions to Alg-IdC, self-assembled microspheres were formed, and conductivity was improved. Cross-linked bluish materials were immediately formed by the addition of Ca2+ ions, and the film was prepared by controlling their concentration. This flexible film can be used in applications such as eco-friendly camouflage, anti-counterfeiting, QR code materials for imaging/sensing, and smart hybrid displays.

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Hibiscus trionum, commonly known as the 'Flower of an Hour', is an easily cultivated plant in the Malvaceae family that is widespread in tropical and temperate regions, including drylands. The purple base part of its petal exhibits structural colour due to the fine ridges on the epidermal cell surface, and the molecular mechanism of ridge formation has been actively investigated. We performed genome sequencing of H. trionum using a long-read sequencing technology with transcriptome and pathway analyses to identify candidate genes for fine structure formation. The ortholog of AtSHINE1, which is involved in the biosynthesis of cuticular wax in Arabidopsis thaliana, was significantly overexpressed in the iridescent tissue. In addition, orthologs of AtCUS2 and AtCYP77A, which contribute to cutin synthesis, were also overexpressed. Our results provide important insights into the formation of fine ridges on epidermal cells in plants using H. trionum as a model.

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Iridescence is a taxonomically widespread form of structural coloration that produces often intense hues that change with the angle of viewing. Its role as a signal has been investigated in multiple species, but recently, and counter-intuitively, it has been shown that it can function as camouflage. However, the property of iridescence that reduces detectability is, as yet, unclear. As viewing angle changes, iridescent objects change not only in hue but also in intensity, and many iridescent animals are also shiny or glossy; these "specular reflections," both from the target and background, have been implicated in crypsis. Here, we present a field experiment with natural avian predators that separate the relative contributions of color and gloss to the "survival" of iridescent and non-iridescent beetle-like targets. Consistent with previous research, we found that iridescent coloration, and high gloss of the leaves on which targets were placed, enhance survival. However, glossy targets survived less well than matt. We interpret the results in terms of signal-to-noise ratio: specular reflections from the background reduce detectability by increasing visual noise. While a specular reflection from the target attracts attention, a changeable color reduces the signal because, we suggest, normally, the color of an object is a stable feature for detection and identification.

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Meat color is an important aspect for the meat industry since it strongly determines the consumers' perception of product quality and thereby significantly influences the purchase decision. Emergence of new vegan meat analogs has renewed interest in the fundamental aspects of meat color in order to replicate it. The appearance of meat is based on a complex interplay between the pigment-based meat color from myoglobin and its chemical forms and light scattering from the muscle's microstructure. While myoglobin biochemistry and pigment-based meat color have been extensively studied, research on the physicochemical contribution of light scattering to meat color and the special case of structural colors causing meat iridescence has received only little attention. Former review articles focused mostly on the biochemical or physical mechanisms rather than the interplay between them, in particular the role that structural colors play. While from an economic point of view, meat iridescence might be considered negligible, an enhanced understanding of the underlying mechanisms and the interactions of light with meat microstructures can improve our overall understanding of meat color. Therefore, this review discusses both biochemical and physicochemical aspects of meat color including the origin of structural colors, highlights new color measurement methodologies suitable to investigate color phenomena such as meat iridescence, and finally presents approaches to modulate meat color in terms of base composition, additives, and processing.

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Patterned photonic crystals (PCs) have great application potential in the textile field owing to their attractive high-saturation iridescent effect. Herein, based on the idea of resist printing, a novel approach to constructing patterned photonic crystals via screen printing was designed and achieved. A colorless pattern with hydrophilic and hydrophobic difference was firstly prepared by screen printing using a hydrophilic polymer paste printed on a hydrophobic fabric, and then the PC structurally colored pattern was obtained through scrapping liquid photonic crystals (LPCs) on the fabric because the LPCs were spread and assembled in the hydrophilic pattern but resisted in the hydrophobic areas, so that to realize the rapid preparation of patterned PCs on the fabric surface. Once the contact angle difference (ΔCA) between the hydrophilic and hydrophobic areas exceeded 80, the "color paste" (that is, LPCs) did not stain the hydrophobic area at all after scrapping, and the assembled PCs pattern showed good contour sharpness and high-saturation iridescence effect. The complex multistructural color patterns on the fabrics were achieved by adjusting the size of nanospheres and using multistep printing and scrapping. The preparation of the protective layer on the PC surface effectively improved the structural stability of the patterned PCs while retaining the optical properties of the pattern. This patterned PCs preparation method was combined with a conventional responsive substance (rhodamine B) to obtain double anti-counterfeiting patterned PCs with the iridescence effect. The results suggested a promising future in both the highly efficient preparation of patterned PCs and the application of PCs in the anti-counterfeiting field.

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Structural colorations have been recognized as a significant way to replace conventional organic dyes for paints, inks, packaging, and cosmetics because of brilliant colors, high stability, and eco-friendliness. However, most current structural-color pigments present an iridescent appearance, and it remains difficult to mitigate a trade-off between lowering the iridescence effect and maintaining the color saturation and brightness. Here, we demonstrate a universal yet economical approach to prepare cellulose structural-color pigments with different sizes. A combined ultrasonication and grinding treatment is explored to adjust the pigment colors as well as control the iridescence-to-non-iridescence transition that depends on the pigment size. The cellulose pigments can be applied on irregular and curved surfaces, having high water-, chemical-, and mechanical-resistances. With humidity-sensing behaviors, the pigments can be further integrated into monitoring systems for environmental management. Such a preparation strategy overcomes the limitation of controlling iridescent and non-iridescent structural colors without sacrificing color properties, which may bring more opportunities to develop new eco-friendly pigments for wide applications.

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Hybridization is a known source of morphological, functional and communicative signal novelty in many organisms. Although diverse mechanisms of established novel ornamentation have been identified in natural populations, we lack an understanding of hybridization effects across levels of biological scales and upon phylogenies. Hummingbirds display diverse structural colours resulting from coherent light scattering by feather nanostructures. Given the complex relationship between feather nanostructures and the colours they produce, intermediate coloration does not necessarily imply intermediate nanostructures. Here, we characterize nanostructural, ecological and genetic inputs in a distinctive Heliodoxa hummingbird from the foothills of eastern Peru. Genetically, this individual is closely allied with Heliodoxa branickii and Heliodoxa gularis, but it is not identical to either when nuclear data are assessed. Elevated interspecific heterozygosity further suggests it is a hybrid backcross to H. branickii. Electron microscopy and spectrophotometry of this unique individual reveal key nanostructural differences underlying its distinct gorget colour, confirmed by optical modelling. Phylogenetic comparative analysis suggests that the observed gorget coloration divergence from both parentals to this individual would take 6.6-10 My to evolve at the current rate within a single hummingbird lineage. These results emphasize the mosaic nature of hybridization and suggest that hybridization may contribute to the structural colour diversity found across hummingbirds.

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Despite the elaborate varieties of iridescent colors in biological species, most of them are reflective. Here we show the rainbow-like structural colors found in the ghost catfish (Kryptopterus vitreolus), which exist only in transmission. The fish shows flickering iridescence throughout the transparent body. The iridescence originates from the collective diffraction of light after passing through the periodic band structures of the sarcomeres inside the tightly stacked myofibril sheets, and the muscle fibers thus work as transmission gratings. The length of the sarcomeres varies from ~1 µm from the body neutral plane near the skeleton to ~2 µm next to the skin, and the iridescence of a live fish mainly results from the longer sarcomeres. The length of the sarcomere changes by ~80 nm as it relaxes and contracts, and the fish shows a quickly blinking dynamic diffraction pattern as it swims. While similar diffraction colors are also observed in thin slices of muscles from non-transparent species such as the white crucian carps, a transparent skin is required indeed to have such iridescence in live species. The ghost catfish skin is of a plywood structure of collagen fibrils, which allows more than 90% of the incident light to pass directly into the muscles and the diffracted light to exit the body. Our findings could also potentially explain the iridescence in other transparent aquatic species, including the eel larvae (Leptocephalus) and the icefishes (Salangidae).

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The bright, saturated iridescent colours of feathers are commonly produced by single and multi-layers of nanostructured melanin granules (melanosomes), air and keratin matrices, surrounded by an outer keratin cortex of varying thicknesses. The role of the keratin cortex in colour production remains unclear, despite its potential to act as a thin film or absorbing layer. We use electron microscopy, optical simulations and oxygen plasma-mediated experimental cortex removal to show that differences in keratin cortex thickness play a significant role in producing colours. The results indicate that keratin cortex thickness determines the position of the major reflectance peak (hue) from nanostructured melanosomes of common pheasant (Phasianus colchicus) feathers. Specifically, the common pheasant has appropriate keratin cortex thickness to produce blue and green structural colours. This finding identifies a general principle of structural colour production and sheds light on the processes that shaped the evolution of brilliant iridescent colours in the common pheasant.

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The effect of Hofmeister salts (NaCl, NaSCN, Na2SO4, KCl, LiCl, CaCl2) on surface iridescence in cooked pork was investigated. Strongest iridescence occurred in samples treated with NaSCN, NaCl and KCl. Control samples and LiCl, CaCl2 and Na2SO4 treatments showed weaker iridescence. However, differences between KCl and LiCl, CaCl2 and Na2SO4 were not statistically significant (p > 0.05). Nevertheless, a tendency of chaotropic salts (NaSCN, NaCl, KCl) to cause stronger iridescence was noted that might be explained with a more effective solubilization of myofibrillar proteins (MPs), reducing incoherent scattering from the myofibrils and thus enhancing multilayer interference.

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Polarized optical microscopy (POM), scanning electron microscopy (SEM), and synchrotron microbeam wide-angle X-ray diffraction (WAXD) were used to investigate the mechanisms of periodic assemblies leading to ring-banded crystal aggregates with light-grating capacity for iridescence in poly (1,4-butylene adipate) (PBA) modulated with poly (ethylene oxide) (PEO). A critical finding is that the PBA crystal assembly on the top surface and in the interior constitutes a grating architecture, with a cross-bar pitch equaling the inter-band spacing. The inner lamellae are arranged perpendicularly to the substrate under the ridge region, where they scroll, bend, and twist 90° to branch out newly spawned lamellae to form the parallel lamellae under the valley region. The cross-hatch grating with a fixed inter-spacing in the PBA aggregated crystals is proved in this work to perfectly act as light-interference entities capable of performing iridescence functions, which can be compared to those widely seen in many of nature's organic bio-species or inorganic minerals such as opals. This is a novel breakthrough finding for PBA or similar polymers, such as photonic crystals, especially when the crystalline morphology could be custom-made and modulated with a second constituent.

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Structural colours, produced by the reflection of light from ultrastructures, have evolved multiple times in butterflies. Unlike pigmentary colours and patterns, little is known about the genetic basis of these colours. Reflective structures on wing-scale ridges are responsible for iridescent structural colour in many butterflies, including the Müllerian mimics Heliconius erato and Heliconius melpomene. Here, we quantify aspects of scale ultrastructure variation and colour in crosses between iridescent and non-iridescent subspecies of both of these species and perform quantitative trait locus (QTL) mapping. We show that iridescent structural colour has a complex genetic basis in both species, with offspring from crosses having a wide variation in blue colour (both hue and brightness) and scale structure measurements. We detect two different genomic regions in each species that explain modest amounts of this variation, with a sex-linked QTL in H. erato but not H. melpomene. We also find differences between species in the relationships between structure and colour, overall suggesting that these species have followed different evolutionary trajectories in their evolution of structural colour. We then identify genes within the QTL intervals that are differentially expressed between subspecies and/or wing regions, revealing likely candidates for genes controlling structural colour formation. This article is part of the theme issue 'Genetic basis of adaptation and speciation: from loci to causative mutations'.

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The members of the Bacteroidetes phylum move on surfaces by gliding motility in the absence of external motility appendages, leading to the formation of spreading colonies. Here, the structural features of the spreading colony were assessed in a uranium-tolerant Bacteroidetes bacterium, Chryseobacterium sp. strain PMSZPI, by using correlative light and scanning electron microscopy (CLSEM). We developed a simple and convenient workflow for CLSEM using a shuttle and find software module and a correlative sample holding slide designed to transport samples between the light/fluorescence microscope (LM/FM) and the scanning electron microscope (SEM) to image spreading colony edges. The datasets from the CLSEM studies allowed convenient examination of the colonial organization by LM/FM followed by ultrastructural analysis by SEM. The regions of interest (ROIs) of the spreading colony edges that were observed in LM/FM in the absence and presence of uranium could be re-identified in the SEM quickly without prolonged searching. Perfect correlation between LM and SEM could be achieved with minimum preparation steps. Subsequently, imaging of the correlated regions was done at higher resolution in SEM to obtain more comprehensive information. We further showed the association of uranium with the gliding PMSZPI cells by energy-dispersive X-ray spectroscopy (EDS) attached to SEM.

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OBJECTIVE: The objective of this study was to investigate the effect of pH change of cooked cured pork M. longissimus thoracis et lumborum on iridescence intensity and extent (= percentage of iridescent area) since interaction with light may be related to pH-induced alterations in microstructure. Muscles were injected with brines of different pH values, cooked, sliced perpendicular to muscle fiber direction, and visually evaluated by a panel of 20 experienced panelists. RESULTS: Muscles with lowest pH (5.38) showed the lowest iridescence score of 4.63 (p < 0.05). Iridescence was greatest in muscles with normal (5.78) and high pH (6.03, respectively 6.59), but did not differ significantly (p > 0.05). Iridescence was positively correlated (p < 0.01) with pH and water content, and negatively correlated (p < 0.01) with cooking loss. Hence, hydration state and light scattering from microstructure may be important factors that determine the degree of iridescence in cooked meat products.

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Mating cues evolve rapidly and can contribute to species formation and maintenance. However, little is known about how sexual signals diverge and how this variation integrates with other barrier loci to shape the genomic landscape of reproductive isolation. Here, we elucidate the genetic basis of ultraviolet (UV) iridescence, a courtship signal that differentiates the males of Colias eurytheme butterflies from a sister species, allowing females to avoid costly heterospecific matings. Anthropogenic range expansion of the two incipient species established a large zone of secondary contact across the eastern United States with strong signatures of genomic admixtures spanning all autosomes. In contrast, Z chromosomes are highly differentiated between the two species, supporting a disproportionate role of sex chromosomes in speciation known as the large-X (or large-Z) effect. Within this chromosome-wide reproductive barrier, linkage mapping indicates that cis-regulatory variation of bric a brac (bab) underlies the male UV-iridescence polymorphism between the two species. Bab is expressed in all non-UV scales, and butterflies of either species or sex acquire widespread ectopic iridescence following its CRISPR knockout, demonstrating that Bab functions as a suppressor of UV-scale differentiation that potentiates mating cue divergence. These results highlight how a genetic switch can regulate a premating signal and integrate with other reproductive barriers during intermediate phases of speciation.

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