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Three-dimensional photonic-crystal grains were found in the scales of the longhorn beetle Prosopocera lactator (Cerambycidae). The local geometric structure can be described as a face-centered-cubic array of spheres, connected by short rods, reminiscent of the "ball-and-stick" models used by solid-state chemists to visualize atomic structures. Based on scanning electron microscopy, x-ray nanotomography, optical measurements, photonic band-structure calculations, and computer simulations of the reflectance, the desaturated greenish coloration is shown to arise from the observed photonic polycrystalline structure. X-ray nanotomography is revealed as a very promising tool for photonic-crystal morphology studies. [ABSTRACT FROM AUTHOR]

The cuticle of the longhorn beetle Pseudomyagrus waterhousei shows a diffuse pattern of mixed blue and violet colors. These colorations arise from a dense layer of droplet-shaped scales covering the dorsal parts of the cuticle. In spite of their lack of iridescence, these colors are shown to be structural and produced by an aggregate of internally ordered photonic-crystal grains. Computer simulations confirm that the blue and violet colors are caused by face-centered-cubic crystallites which dominantly expose their (111) surface to illumination and viewing.

The butterfly Pierella luna (Nymphalidae) shows an intriguing rainbow iridescence effect: the forewings of the male, when illuminated along the axis from the body to the wing tip, decompose a white light beam as a diffraction grating would do. Violet light, however, emerges along a grazing angle, near the wing surface, while the other colors, from blue to red, exit respectively at angles progressively closer to the direction perpendicular to the wing plane. This sequence is the reverse of the usual decomposition of light by a grating with a periodicity parallel to the wing surface. It is shown that this effect is produced by a macroscopic deformation of the entire scale, which curls in such a way that it forms a "vertical" grating, perpendicular to the wing surface, and functions in transmission instead of reflection.

The blue scales on the cuticle of the male beetle Hoplia coerulea can absorb water, with the consequence that these scales, which have been shown to be responsible for the beetle's bright blue coloration, reversibly turn to emerald green with increasing water contents. Optical measurements are shown, by analytic photonic-crystal models, to be compatible with the full filling of the scales structures with water. The natural mechanism shows the way to produce a very efficient hygrochromic material: a medium which significantly changes color when its water contents are modified.

The African shield-backed bug Calidea panaethiopica is a very colorful insect which produces a range of iridescent yellow, green, and blue reflections. The cuticle of the dorsal side of the insect, on the shield, the prothorax and part of the head, is pricked of uniformly distributed hemispherical hollow cavities a few tens micrometers deep. Under normal illumination and viewing the insect's muffin-tin shaped surface gives rise to two distinct colors: a yellow spot arising from the bottom of the well and a blue annular cloud that appears to float around the yellow spot. This effect is explained by multiple reflections on a hemispherical Bragg mirror with a mesoscopic curvature. A multiscale computing methodology was found to be needed to evaluate the reflection spectrum for such a curved multilayer. This multiscale approach is very general and should be useful for dealing with visual effects in many natural and artificial systems.

Iridescent butterfly wing colours result from the interaction of light with sub-micrometre structures in the scales. Typically, one scale contains one such photonic structure that produces a single iridescent signal. Here, however, we show how the dorsal wings of male Lamprolenis nitida emit two independent signals from two separate photonic structures in the same scale. Multiple independent signals from separate photonic structures within the same sub-micrometre device are currently unknown in animals. However, they would serve to increase the complexity and specificity of the optical signature, enhancing the information conveyed. This could be important during intrasexual encounters, in which iridescent male wing colours are employed as threat displays. Blazed diffraction gratings, like those found in L. nitida, are asymmetric photonic structures and drive most of the incident light into one diffraction order. Similar gratings are used in spectrometers, limiting the spectral range over which the spectrometer functions. By incorporating two interchangeable gratings onto a single structure, as they are in L. nitida, the functional range of spectrometers could be extended.

The male Troides magellanus--a birdwing butterfly that lives in a restricted area of the Philippines--concentrates on its hindwings at least two distinct optical processes that contribute to its exceptional visual attraction. The first is the very bright uniform yellow coloration caused by a pigment which generates yellow-green fluorescence, and the other is a blue-green iridescence which results from light diffraction at grazing emergence under a specific illumination. Detailed optical measurements reveal that these optical effects are correlated, the fluorescence being enhanced by illuminations conditions that favor the occurrence of the iridescence. These effects are analyzed, with the conclusion that both of them depend on the same optical device: a one-dimensional microribs grating which appear on the sides of the ridges that run along the yellow scales.

The tortoise beetle Charidotella egregia is able to modify the structural color of its cuticle reversibly, when disturbed by stressful external events. After field observations, measurements of the optical properties in the two main stable color states and scanning electron microscope and transmission electron microscope investigations, a physical mechanism is proposed to explain the color switching of this insect. It is shown that the gold coloration displayed by animals at rest arises from a chirped multilayer reflector maintained in a perfect coherent state by the presence of humidity in the porous patches within each layer, while the red color displayed by disturbed animals results from the destruction of this reflector by the expulsion of the liquid from the porous patches, turning the multilayer into a translucent slab that leaves an unobstructed view of the deeper-lying, pigmented red substrate. This mechanism not only explains the change of hue but also the change of scattering mode from specular to diffuse. Quantitative modeling is developed in support of this analysis.

The three-dimensional structure that causes the coloration of the tropical weevil Pachyrrhynchus congestus pavonius was studied, using a combination of electron microscopy, optical spectroscopy, and numerical modeling. The orange scales that cover the colored rings on the animal's body were opened, to display the structure responsible for the coloration. This structure is a three-dimensional photonic polycrystal, each grain of which showing a face-centered cubic symmetry. The measured lattice parameter and the observed filling fraction of this structure explain the dominant reflected wavelength in the reddish orange. The long-range disorder introduced by the grain boundaries explains the paradoxical observation that the reflectance, although generated by a photonic crystal, is insensitive to changes in the viewing angle.

Photonic-crystal-type nanostructures occurring in the scales of the butterfly Cyanophrys remus were investigated by optical and electron microscopy (scanning and transmission electron microscopy), reflectance measurements (specular, integrated, and goniometric), by fast Fourier transform analysis of micrographs, by modeling, and by numerical simulation of the measured reflectance data. By evaluating the collected data in a cross-correlated way, we show that the metallic blue dorsal coloration originates from scales which individually are photonic single crystals of 50 x 120 microm2 , while the matt pea-green coloration of the ventral side arises from the cumulative effect of randomly arranged, bright photonic crystallites (blue, green, and yellow) with typical diameters in the 3-10-mum range. Both structures are based on a very moderate refractive index contrast between air and chitin. Using a bleached specimen in which the pigment has decayed with time, we investigated the role of pigment in photonic-crystal material in the process of color generation. The possible biologic utility of the metallic blue (single-crystal) and dull green (polycrystal) textures both achieved with photonic crystals are briefly discussed. Potential applications in the field of colorants, flat panel displays, smart textiles, and smart papers are surveyed.

Using transmission electron microscopy, analytical modeling, and detailed numerical simulations, the iridescence observed from the comb rows of the ctenophore Beroë cucumis was investigated. It is shown that the changing coloration which accompanies the beating of comb rows as the animal swims can be explained by the weakly-contrasted structure of the refractive index induced by the very coherent packing of locomotory cilia. The colors arising from the narrow band-gap reflection are shown to be highly saturated and, as a function of the incidence angle, cover a wide range of the visible and ultraviolet spectrum. The high transparency of the structure at the maximal bioluminescence wavelength is also explained.

The structural origin of the weak iridescence on some of the dark feathers of the black-billed magpie, Pica pica (Corvidae), is found in the structure of the ribbon-shaped barbules. The cortex of these barbules contains cylindrical holes distributed as the nodes of an hexagonal lattice in the hard layer cross section. The cortex optical properties are described starting from a photonic-crystal film theory. The yellowish-green coloration of the bird's tail can be explained by the appearance of a reflection band related to the photonic-crystal lowest-lying gap. The bluish reflections from the wings are produced by a more complicated mechanism, involving the presence of a cortex second gap."

The microscopic structure of the hard external parts of the body of the iridescent blue-violet chaffer beetle Hoplia coerulea is studied using scanning electron microscopy. The blue iridescence is shown to originate from the structure of the squamae within scales covering the dorsal side of the beetle. The internal structure of the scales shows a stack of planar sheets, separated by a well-organized network of spacers, a structure which belongs to the family of the layer-by-layer photonic crystals. The blue iridescence is easily explained by a planar multilayer approximation model, deduced from the observed three-dimensional structure.