Colour formation on the wings of the butterfly Hypolimnas salmacis by scale stacking.

The butterfly genus Hypolimnas features iridescent blue colouration in some areas of its dorsal wings. Here, we analyse the mechanisms responsible for such colouration on the dorsal wings of Hypolimnas salmacis and experimentally demonstrate that the lower thin lamina in the white cover scales causes the blue iridescence. This outcome contradicts other studies reporting that the radiant blue in Hypolimnas butterflies is caused by complex ridge-lamellar architectures in the upper lamina of the cover scales. Our comprehensive optical study supported by numerical calculation however shows that scale stacking primarily induces the observed colour appearance of Hypolimnas salmacis.non

Light manipulation principles in biological photonic systems

© 2013 Science Wise Publishing and DE GRUYTER.The science of light and colour manipulation continues to generate interest across a range of disciplines, from mainstream biology, across multiple physicsbased fields, to optical engineering. Furthermore, the study of light production and manipulation is of significant value to a variety of industrial processes and commercial products. Among the several key methods by which colour is produced in the biological world, this review sets out to describe, in some detail, the specifics of the method involving photonics in animal and plant systems; namely, the mechanism commonly referred to as structural colour generation. Not only has this theme been a very rapidly growing area of physics-based interest, but also it is increasingly clear that the biological world is filled with highly evolved structural designs by which light and colour strongly influence behaviours and ecological functions.We acknowledge the financial support of DARPA contract W911NF-10-C-0069 and of AFOSR grant FA9550-10-1-0020. We also wish to thank Caroline Pouya, Helen Ghiradella, Radislav Potyrailo, Roy Sambles, Shuichi Kinoshita and Doekele Stavenga for helpful discussions

Liquid – liquid phase separation morphologies in ultra-white beetle scales and a synthetic equivalent

Cyphochilus beetle scales are amongst the brightest structural whites in nature, being highly opacifying whilst extremely thin. However, the formation mechanism for the voided intra- scale structure is unknown. Here we report 3D x-ray nanotomography data for the voided chitin networks of intact white scales of Cyphochilus and Lepidiota stigma. Chitin-filling frac- tions are found to be 31 ± 2% for Cyphochilus and 34 ± 1% for Lepidiota stigma, indicating previous measurements overestimated their density. Optical simulations using finite- difference time domain for the chitin morphologies and simulated Cahn-Hilliard spinodal structures show excellent agreement. Reflectance curves spanning filling fraction of 5-95% for simulated spinodal structures, pinpoint optimal whiteness for 25% chitin filling. We make a simulacrum from a polymer undergoing a strong solvent quench, resulting in highly reflective ( 94%) white films. In-situ X-ray scattering confirms the nanostructure is formed through spinodal decomposition phase separation. We conclude that the ultra-white beetle scale nanostructure is made via liquid–liquid phase separation

Complex photonic structures in nature: from order to disorder

Structural colours arise from the interaction of visible light with nano-structured materials. The occurrence of such structures in nature has been known for over a century, but it is only in the last few decades that the study of natural photonic structures has fully matured due to the advances in imagining techniques and computational modelling. Even though a plethora of different colour-producing architectures in a variety of species has been investigated, a few significant questions are still open: how do these structures develop in living organisms? Does disorder play a functional role in biological photonics? If so, is it possible to say that the optical response of natural disordered photonics has been optimised under evolutionary pressure? And, finally, can we exploit the well-adapted photonic design principles that we observe in Nature to fabricate functional materials with optimised scattering response? In my thesis I try to answer the questions above: I microscopically investigate $\textit{in vivo}$ the growth of a cuticular multilayer, one of the most common colour-producing strategies in nature, in the green beetles $\textit{Gastrophysa viridula}$ showing how the interplay between different materials varies during the various life stages of the beetles; I further investigate two types of disordered photonic structures and their biological role, the random array of spherical air inclusions in the eggshells of the honeyguide $\textit{Prodotiscus regulus}$, a species under unique evolutionary pressure to produce blue eggs, and the anisotropic chitinous network of fibres in the white beetle $\textit{Cyphochilus}$, the whitest low-refractive index material; finally, inspired by these natural designs, I fabricate and study light transport in biocompatible highly-scattering materials.European Research Council grant awarded to Dr Silvia Vignolin

Photonics in Nature: From Order to Disorder

The most vibrant and striking colours in living organisms are often caused by a combination of pigments and nano-scale transparent architectures, which interact with light to produce so-called structural colours. These colours are the result of light interfering with the nanoscale structures that are present in the materials. Such colour-producing structures are not perfect, and irregularities in the arrangements (disorder) are present in many organisms. However, disorder in natural structures is not detrimental but functional, as it allows a broader range of optical effects. This chapter reviews and attempts to classify structurally coloured organisms, highlighting the influence that disorder has on their visual appearance. It also showcases how photonic systems, such as the blue Morpho butterfly and the white Cyphochilus beetle, are capable of obtaining optical properties (long-distance visibility and whiteness, respectively) where disorder seems to be highly optimized, indicating that disorder is important for obtaining complex visual effects in natural systems. The chapter first introduces the mathematical concepts required for analysing disordered systems, such as the Fourier transform and the structure factor. Then, ordered and disordered natural photonic systems are reviewed. This is followed by examples of completely disordered structures responsible of white appearances. Finally, we review the possibilities of hierarchical organisation and pixelated surfaces to widen the range of optical appearances

Rainbow peacock spiders inspire miniature superiridescent optics

Colour produced by wavelength-dependent light scattering is a key component of visual communication in nature and acts particularly strongly in visual signalling by structurallycoloured animals during courtship. Two miniature peacock spiders (Maratus robinsoni and M. chrysomelas) court females using tiny structured scales (~ 40 × 10 μm2) that reflect the full visual spectrum. Using TEM and optical modelling, we show that the spiders’ scales have 2D nanogratings on microscale 3D convex surfaces with at least twice the resolving power of a conventional 2D diffraction grating of the same period. Whereas the long optical path lengths required for light-dispersive components to resolve individual wavelengths constrain current spectrometers to bulky sizes, our nano-3D printed prototypes demonstrate that the design principle of the peacock spiders’ scales could inspire novel, miniature light-dispersive components

Absence of circular polarisation in reflections of butterfly wing scales with chiral gyroid structure

The single Gyroid, a triply-periodic ordered chiral network of cubic symmetry, appears as a nanostructure in the green-colored wing scales of various butterflies. In lossless and perfectly ordered single Gyroid materials, the structural chirality leads to circularly polarized reflections from crystals oriented in the [100] direction. Here we report a circular polarisation study of the macroscopic reflections of the wing scales of Callophrys rubi and Teinopalpus imperialis that reveals no circular dichroism, that is, we find no significant difference in the reflectance values for left- and right-circularly polarized light. The reasons for the absence of circularly polarized reflections is likely to be a compound effect of various factors, including crystallite orientation, presence of both left- and right-handed single Gyroid enantiomers, and structural disorder. Each of these factors weakens, but does not fully extinguish, the circular polarisation signal. We further find a substantial amount of blue-absorbing pigment in those wing scales of C. rubi that are structured according to the single Gyroid. Numerical simulations demonstrate that absorption, while evidently reducing overall reflectance, does generally not reduce the circular dichroism strength. The experimental findings of this paper, however, clearly demonstrate that circular dichroism is absent from the reflections of the butterfly wing scale. Henceforth, the chiro-optical response of the idealised structure does not fulfil a biological photonic function

Colour formation on the wings of the butterfly Hypolimnas salmacis by scale stacking

The butterfly genus Hypolimnas features iridescent blue colouration in some areas of its dorsal wings. Here, we analyse the mechanisms responsible for such colouration on the dorsal wings of Hypolimnas salmacis and experimentally demonstrate that the lower thin lamina in the white cover scales causes the blue iridescence. This outcome contradicts other studies reporting that the radiant blue in Hypolimnas butterflies is caused by complex ridge-lamellar architectures in the upper lamina of the cover scales. Our comprehensive optical study supported by numerical calculation however shows that scale stacking primarily induces the observed colour appearance of Hypolimnas salmacis

Nanoscale cuticle density variations correlate with pigmentation and color in butterfly wing scales

How pigment distribution correlates with cuticle density within a microscopic butterfly wing scale, and how both impact final reflected color remains unknown. We used ptychographic X-ray computed tomography to quantitatively determine, at nanoscale resolutions, the three-dimensional mass density of scales with pigmentation differences. By comparing cuticle densities with pigmentation and color within a scale, we determine that the lower lamina structure in all scales has the highest density and lowest pigmentation. Low pigment levels also correlate with sheet-like chitin structures as opposed to rod-like structures, and distinct density layers within the lower lamina help explain reflected color. We propose that pigments, in addition to absorbing specific wavelengths, can affect cuticle polymerization, density, and refractive index, thereby impacting reflected wavelengths that produce structural colors