Nanostructural self-assembly of iridescent feather barbules through depletion attraction of melanosomes during keratinization

Avian plumage colours are model traits in understanding the evolution of sexually selected ornamental traits. Paradoxically, iridescent structural colours, probably the most dazzling of these traits, remain the most poorly understood. Though some data suggest that expression of bright iridescent plumage colours produced by highly ordered arrays of melanosomes and keratin is condition-dependent, almost nothing is known of their ontogeny and thus of any developmental mechanisms that may be susceptible to perturbation. Here, we use light and electron microscopy to compare the ontogeny of iridescent male and non-iridescent female feathers in blue-black grassquits. Feather barbules of males contain a single layer of melanosomes bounded by a thin layer of keratin-producing blue iridescent colour, while those of females contain disorganized melanosomes and no outer layer. We found that nanostructural organization of male barbules occurs late in development, following death of the barbule cell, and is thus unlikely to be under direct cellular control, contrary to previous suggestions. Rather, organization appears to be caused by entropically driven self-assembly through depletion attraction forces that pin melanosomes to the edge of barbule cells and to one another. These forces are probably stronger in developing barbules of males than of females because their melanosomes are (i) larger, (ii) more densely packed, and (iii) more homogeneously distributed owing to the more consistent shape of barbules during keratinization. These data provide the first proposed developmental pathway for iridescent plumage colours, and suggest that any condition dependence of iridescent barbules is likely driven by factors other than direct metabolic cost.     

Analysing photonic structures in plants

The outer layers of a range of plant tissues, including flower petals, leaves and fruits, exhibit an intriguing variation of microscopic structures. Some of these structures include ordered periodic multilayers and diffraction gratings that give rise to interesting optical appearances. The colour arising from such structures is generally brighter than pigment-based colour. Here, we describe the main types of photonic structures found in plants and discuss the experimental approaches that can be used to analyse them. These experimental approaches allow identification of the physical mechanisms producing structural colours with a high degree of confidence.     

Coloration using higher order optical interference in the wing pattern of the Madagascan sunset moth

Colour patterns of animals' bodies are usually produced by the spatial distribution of pigments with different colours. However, some animals use the spatial variation of colour-producing microstructures. We have studied one distinctive example of such structurally produced colour patterns, the wing of the Madagascan sunset moth, to clarify the physical rules that underlie the colour variation. It is known that the iridescent wing scale of the sunset moth has the alternate air-cuticle multilayer structure that causes optical interference. The microscopic and optical investigations of various parts of the wing have confirmed that the thickness of the cuticle layers within the scale largely varies to produce the colour pattern. However, it varies in very different ways between the dorsal and ventral sides of the hind wing; the thickness gradually varies on the dorsal side from scale to scale, while the abrupt changes are found on the ventral side to form distinctive borders between differently coloured areas. It is also revealed that an unusual coloration mechanism is involved in the green part of the ventral hind wing: the colour is caused by higher order optical interference of the highly non-ideal multilayer structure. The physical mechanism of the colour pattern formation is briefly discussed with the several mathematical models proposed so far.     

Comprehensive analysis of retroreflection in Papilio crino Fabricius, 1792 wings

Multilayer thin‐film structures in the wings of a butterfly; Papilio crino produce a colourful iridescence from reflected light. In this investigation, scanning electron microscope images show both the concave cover scales and pigmented air‐chamber ground scales. The microstructures with the concavities retroreflect incident light, thus causing the double reflection. This gives rise to both the colour mixing and polarisation conversion clearly depicted in the optical images. The result of the numerical and theoretical analysis via the CIELAB, and optical reflection and transmission of light through the multilayer stacks with the use of transfer method show that the emerging colouration on the Papilio crino is structural and is due to the combination of colours caused by multiple bounces within the concavities. The butterfly wing structure can be used as the template for designing the photonic device.Inspec keywords: bio‐optics, scanning electron microscopy, photodiodes, optical sensors, optical images, light reflection, reflectivity, colour, optical links, multilayers, optical multilayers, light polarisationOther keywords: pigmented air‐chamber ground scales, concavities, incident light, double reflection, colour mixing, polarisation conversion, optical images, numerical analysis, theoretical analysis, optical reflection, multilayer stacks, emerging colouration, butterfly wing structure, papilio crino fabricius, thin‐film structures, colourful iridescence, reflected light, electron microscope images, concave cover scales     

Dynamic Control of Light Direction Enabled by Stimuli‐Responsive Liquid Crystal Gratings

The ability to control light direction with tailored precision via facile means is long‐desired in science and industry. With the advances in optics, a periodic structure called diffraction grating gains prominence and renders a more flexible control over light propagation when compared to prisms. Today, diffraction gratings are common components in wavelength division multiplexing devices, monochromators, lasers, spectrometers, media storage, beam steering, and many other applications. Next‐generation optical devices, however, demand nonmechanical, full and remote control, besides generating higher than 1D diffraction patterns with as few optical elements as possible. Liquid crystals (LCs) are great candidates for light control since they can form various patterns under different stimuli, including periodic structures capable of behaving as diffraction gratings. The characteristics of such gratings depend on several physical properties of the LCs such as film thickness, periodicity, and molecular orientation, all resulting from the internal constraints of the sample, and all of these are easily controllable. In this review, the authors summarize the research and development on stimuli‐controllable diffraction gratings and beam steering using LCs as the active optical materials. Dynamic gratings fabricated by applying external field forces or surface treatments and made of chiral and nonchiral LCs with and without polymer networks are described. LC gratings capable of switching under external stimuli such as light, electric and magnetic fields, heat, and chemical composition are discussed. The focus is on the materials, designs, applications, and future prospects of diffraction gratings using LC materials as active layers.     

Alignment of crystal orientations of the multi-domain photonic crystals in Parides sesostris wing scales

It is known that the wing scales of the emerald-patched cattleheart butterfly, Parides sesostris, contain gyroid-type photonic crystals, which produce a green structural colour. However, the photonic crystal is not a single crystal that spreads over the entire scale, but it is separated into many small domains with different crystal orientations. As a photonic crystal generally has band gaps at different frequencies depending on the direction of light propagation, it seems mysterious that the scale is observed to be uniformly green under an optical microscope despite the multi-domain structure. In this study, we have carefully investigated the structure of the wing scale and discovered that the crystal orientations of different domains are not perfectly random, but there is a preferred crystal orientation that is aligned along the surface normal of the scale. This finding suggests that there is an additional factor during the developmental process of the microstructure that regulates the crystal orientation.     

Additive colour mixing by surface relief gratings utilizing the power spectrum of a fluorescent lamp

Additive colour mixing produced by diffractive grating structures is studied. The gratings produce almost monochromatic primary colours to a selected viewing angle, providing thus a very large colour gamut. The colours inside the gamut are obtained by additive colour mixing in the reflected diffraction order. The amount of a single primary in the additive colour mixing within one colour pixel is determined by the area fraction of the corresponding grating. We demonstrated the concept by fabricating samples which appear white at 30° viewing angle when illuminated by 4000?K fluorescent lamp.     

Measuring and modelling the reflectance spectra of male Swinhoe's pheasant feather barbules

A range of iridescent colour appearances are presented by male Swinhoe''s pheasants'' (Lophura swinhoii) mantle feathers. Two distinct regions of the open pennaceous portion of its feathers display particularly conspicuous angle-dependent reflection. A bright blue band appears in one region at normal incidence that spatially shifts to another at higher illumination angles. The two-dimensional photonic crystal-like nanostructures inside the barbules of these two regions are similar. However, this study found that the spatial variation in their colour appearance results from a continuously changing orientation of barbules with respect to the alignment of their associated barb. A multi-layered rigorous coupled-wave analysis approach was used to model the reflections from the identified intra-barbule structures. Well-matched simulated and measured reflectance spectra, at both normal and oblique incidence, support our elucidation of the origin of the bird''s distinctive feather colour appearance.     

Out-of-plane diffraction of a two-dimensional photonic crystal with finite dielectric modulation

We present a generalized picture of out-of-plane diffraction in a two-dimensional photonic crystal using the concept of photonic bands and employing a three-dimensional, equal-frequency-surface analysis. We show that the discrete spots of diffraction pattern in a weakly modulated photonic crystal, including those of conventional diffraction gratings, become continuous when the dielectric modulation becomes finite. Furthermore, in a finite-modulated photonic crystal, the diffraction can take place even in the region prohibited by Bragg's law: there are available states for the incident light, which are evanescent in the case of a diffraction grating (weakly modulated photonic crystal).     

Effects of a butterfly scale microstructure on the iridescent color observed at different angles

Multilayer thin-film structures in butterfly wing scales produce a colorful iridescence from reflected sunlight. Because of optical phenomena, changes in the angle of incidence of light and the viewing angle of an observer result in shifts in the color of butterfly wings. Colors ranging from green to purple, which are due to nonplanar specular reflection, can be observed on Papilio blumei iridescent scales. This refers to a phenomenon in which the curved surface patterns in the thin-film structure cause the specular component of the reflected light to be directed at various angles while affecting the spectral reflectivity at the same time by changing the optical path length through the structure. We determined the spectral reflectivities of P. blumei iridescent scales numerically by using models of a butterfly scale microstructure and experimentally by using a microscale-reflectance spectrometer. The numerical models accurately predict the shifts in spectral reflectivity observed experimentally.     

Scattered-light properties of diffraction gratings

One of the many calibrations performed for a scientific-quality spectrometer is the characterization of its scattered-light properties. The scattered light can arise from any optical surface, and light leaks or scattering from baffles can also contribute to the instrumental stray-light level. For a diffraction-grating spectrometer the primary contribution to instrumental scattered light has been found to be the scattered light from the grating. The results from measuring the scattered-light properties of 10 diffraction gratings are discussed along with the application of these results in analyzing the total scattered light measured for three spectrometers. It has been found from these measurements that there are two components of the grating scattered light: a Lorentzian-type component and a constant background component. The Lorentzian component is predicted from the diffraction theory for a grating, and the constant background component is consistent with Rayleigh scattering from the microscopic surface im erfections. It was also discovered that multiple replicas of gratings from the same master grating exhibit significantly more scattered light than the preceding replica by factors of 1.1-2.     

Integration of Optical Surface Structures with Chiral Nanocellulose for Enhanced Chiroptical Properties

The integration of chiral organization with photonic structures found in many living creatures enables unique chiral photonic structures with a combination of selective light reflection, light propagation, and circular dichroism. Inspired by these natural integrated nanostructures, hierarchical chiroptical systems that combine imprinted surface optical structures with the natural chiral organization of cellulose nanocrystals are fabricated. Different periodic photonic surface structures with rich diffraction phenomena, including various optical gratings and microlenses, are replicated into nanocellulose film surfaces over large areas. The resulting films with embedded optical elements exhibit vivid, controllable structural coloration combined with highly asymmetric broadband circular dichroism and a microfocusing capability not typically found in traditional photonic bioderived materials without compromising their mechanical strength. The strategy of imprinting surface optical structures onto chiral biomaterials facilitates a range of prospective photonic applications, including stereoscopic displays, polarization encoding, chiral polarizers, and colorimetric chiral biosensing.     

Understanding the Self-Assembly of Cellulose Nanocrystals—Toward Chiral Photonic Materials

Over millions of years, animals and plants have evolved complex molecules and structures that endow them with vibrant colors. Among the sources of natural coloration, structural color is prominent in insects, bird feathers, snake skin, plants, and other organisms, where the color arises from the interaction of light with nanoscale features rather than absorption from a pigment. Cellulose nanocrystals (CNCs) are a biorenewable resource that spontaneously organize into chiral nematic liquid crystals having a hierarchical structure that resembles the Bouligand structure of arthropod shells. The periodic, chiral nematic organization of CNC films leads them to diffract light, making them appear iridescent. Over the past two decades, there have been many advances to develop the photonic properties of CNCs for applications ranging from cosmetics to sensors. Here, the origin of color in CNCs, the control of photonic properties of CNC films, the development of new composite materials of CNCs that can yield flexible photonic structures, and the future challenges in this field are discussed. In particular, recent efforts to make flexible photonic materials using CNCs are highlighted.     

Light-emitting nanocasts formed from bio-templates: FESEM and cathodoluminescent imaging studies of butterfly scale replicas

Nanocasts comprising of red-light-emitting cubic Y(2)O(3):Eu phosphors were made from butterfly wing scale bio-templates. We report herein the first cathodoluminescent images made from such nanocasts and show that valuable insights into the nature of the internal structure of the casts can be gained by the use of this technique. The casts faithfully reproduced the fine sub-micrometre size detail of the scales, as was made evident by both FESEM and cathodoluminescent images that were collected from the same sample areas using a hyphenated FESEM-CL instrument. There was excellent agreement between the FESEM and cathodoluminescent images, the image quality of the latter indicating that the Eu(3+) activator ions were evenly dispersed in the Y(2)O(3):Eu phosphor on a sub-micrometre scale. The casts were made by infilling the natural moulds with a Y(2)O(3):Eu precursor solution that was subsequently dried and fired to convert it into the phosphor material. This method provides a simple, low cost route for fabricating nanostructures having feature dimensions as small as 20?nm in size, and it has the potential to be applied to other metal oxide systems for producing nano-and micro-components for electronic, magnetic or photonic integrated systems.     

Two-dimensional gratings for low polarization dependent wavelength demultiplexing

Two-dimensional (crossed) diffraction gratings with sinusoidal or truncated pyramidal (trapezoidal) profiles are proposed to have diffraction efficiency almost independent of the incident polarization inside the optical communication spectral window 1.5-1.6 microm. The gratings are characterized by different periods in the two orthogonal directions, chosen to support only one dispersive diffraction order in addition to the zeroth (specular) one.     

Cryptic iridescence in a fossil weevil generated by single diamond photonic crystals

Nature''s most spectacular colours originate in integumentary tissue architectures that scatter light via nanoscale modulations of the refractive index. The most intricate biophotonic nanostructures are three-dimensional crystals with opal, single diamond or single gyroid lattices. Despite intense interest in their optical and structural properties, the evolution of such nanostructures is poorly understood, due in part to a lack of data from the fossil record. Here, we report preservation of single diamond (Fd-3m) three-dimensional photonic crystals in scales of a 735 000 year old specimen of the brown Nearctic weevil Hypera diversipunctata from Gold Run, Canada, and in extant conspecifics. The preserved red to green structural colours exhibit near-field brilliancy yet are inconspicuous from afar; they most likely had cryptic functions in substrate matching. The discovery of pristine fossil examples indicates that the fossil record is likely to yield further data on the evolution of three-dimensional photonic nanostructures and their biological functions.     

Almost perfect blazing by photonic crystal rod gratings

A periodic array of dielectric rods or holes, known as two-dimensional photonic crystal, is shown to have blazing properties similar to those of classical diffraction gratings. Several different optogeometric configurations are shown numerically to exhibit an almost perfect blazing in the -1st reflected order with a plateaulike spectral dependence in nonpolarized light.     

Periodic defects in 2D-PBG materials: full-wave analysis and design

In this paper, an accurate and efficient characterization of two-dimensional photonic bandgap structures with periodic defects is performed, which exploits a full-wave diffraction theory developed for one-dimensional gratings. The high convergence rate of the proposed technique is demonstrated. Results are presented for both TE and TM polarizations, showing the efficiencies as a function of wavelength, incidence angle, geometrical and physical parameters. A comparison with other theoretical results reported in the literature is shown with a good agreement. The transmission properties of photonic crystals with periodic defects are studied, investigating the effects of the variation of geometrical and physical parameters; design efficiency maps and formulas are given; moreover, the application of the analyzed structures as filters is discussed.     

Biomaterial‐Based “Structured Opals” with Programmable Combination of Diffractive Optical Elements and Photonic Bandgap Effects

Naturally occurring iridescent systems produce brilliant color displays through multiscale, hierarchical assembly of structures that combine reflective, diffractive, diffusive, or absorbing domains. The fabrication of biopolymer‐based, hierarchical 3D photonic crystals through the use of a topographical templating strategy that allows combined optical effects derived from the interplay of predesigned 2D and 3D geometries is reported here. This biomaterials‐based approach generates 2D diffractive optics composed of 3D nanophotonic lattices that allow simultaneous control over the reflection (through the 3D photonic bandgap) and the transmission (through 2D diffractive structuring) of light with the additional utility of being constituted by a biocompatible, implantable, edible commodity textile material. The use of biopolymers allows additional degrees of freedom in photonic bandgap design through directed protein conformation modulation. Demonstrator structures are presented to illustrate the lattice multifunctionality, including tunable diffractive properties, increased angle of view of photonic crystals, color‐mixing, and sensing applications.     

Bringing some photonic structures for solar cells to the fore

A review on the use of photonic structures enabling a better absorption of solar radiation within solar cells is proposed. Specific geometric configurations, such as folded solar cells or fiber-based architectures, are shown to be promising solutions to reach better light absorption. Electromagnetic optimization of thin-film solar cells and the use of angular thin-film filters, proposed by several research groups, also provide solutions to better concentrate solar radiation within the active layers of solar cells. Finally, results on "photonized" solar cells comprising gratings or more advanced photonic components, such as photonic crystals or plasmonic structures, and their effects on light-matter interaction in solar cells are highlighted.