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Structural coloration

Structural coloration in animals, and a few plants, is the production of colour by microscopically structured surfaces fine enough to interfere with visible light instead of pigments, although some structural coloration occurs in combination with pigments. For example, peacock tail feathers are pigmented brown, but their microscopic structure makes them also reflect blue, turquoise, and green light, and they are often iridescent.

History
's 1665 Micrographia contains the first observations of structural colours. In his 1665 book Micrographia, Robert Hooke described the "fantastical" colours of the peacock's feathers: In his 1704 book Opticks, Isaac Newton described the mechanism of the colours other than the brown pigment of peacock tail feathers. Newton noted that Thomas Young (1773–1829) extended Newton's particle theory of light by showing that light could also behave as a wave. He showed in 1803 that light could diffract from sharp edges or slits, creating interference patterns. In his 1892 book Animal Coloration, Frank Evers Beddard (1858–1925) acknowledged the existence of structural colours: noted that Chrysospalax golden moles' thick fur was structurally coloured. But Beddard then largely dismissed structural coloration, firstly as subservient to pigments: "in every case the [structural] colour needs for its display a background of dark pigment;" and then by asserting its rarity: "By far the commonest source of colour in invertebrate animals is the presence in the skin of definite pigments", though he does later admit that the Cape golden mole has "structural peculiarities" in its hair that "give rise to brilliant colours". == Principles ==
Principles
Structure not pigment , the waves reflected from the upper and lower surfaces travel different distances depending on the angle, so they interfere. Structural coloration is caused by interference effects rather than by pigments. Colours are produced when a material is scored with fine parallel lines, or formed of one or more parallel thin layers, or otherwise composed of microstructures on the scale of the colour's wavelength. Structural coloration is responsible for the blues and greens of the feathers of many birds (the bee-eater, kingfisher and roller, for example), as well as many butterfly wings, beetle wing-cases (elytra) and (while rare among flowers) the gloss of buttercup petals. These are often iridescent, as in peacock feathers and nacreous shells such as of pearl oysters (Pteriidae) and Nautilus. This is because the reflected colour depends on the viewing angle, which in turn governs the apparent spacing of the structures responsible. Structural colours can be combined with pigment colours: peacock feathers are pigmented brown with melanin, while buttercup petals have both carotenoid pigments for yellowness and thin films for reflectiveness. Principle of iridescence of a fractured surface of nacre showing multiple thin layers Iridescence, as explained by Thomas Young in 1803, is created when extremely thin films reflect part of the light falling on them from their top surfaces. The rest of the light goes through the films, and a further part of it is reflected from their bottom surfaces. The two sets of reflected waves travel back upwards in the same direction. But since the bottom-reflected waves travelled a little farther – controlled by the thickness and refractive index of the film, and the angle at which the light fell – the two sets of waves are out of phase. When the waves are one or more whole wavelengths apart – in other words, at certain specific angles, they add (interfere constructively), giving a strong reflection. At other angles and phase differences, they can subtract, giving weak reflections. The thin film therefore selectively reflects just one wavelength – a pure colour – at any given angle, but other wavelengths – different colours – at different angles. So, as a thin-film structure such as a butterfly's wing or bird's feather moves, it seems to change colour. == Mechanisms ==
Mechanisms
Fixed structures A number of fixed structures can create structural colours, by mechanisms including diffraction gratings, selective mirrors, photonic crystals, crystal fibres and deformed matrices. When the bird moves the colour switches sharply between these two colours, rather than drifting iridescently. During courtship, the male bird systematically makes small movements to attract females, so the structures must have evolved through sexual selection. Photonic crystals can be formed in different ways. In Parides sesostris, the emerald-patched cattleheart butterfly, photonic crystals are formed of arrays of nano-sized holes in the chitin of the wing scales. The holes have a diameter of about 150 nanometres and are about the same distance apart. The holes are arranged regularly in small patches; neighbouring patches contain arrays with differing orientations. The result is that these emerald-patched cattleheart scales reflect green light evenly at different angles instead of being iridescent. In Lamprocyphus augustus, a weevil from Brazil, the chitin exoskeleton is covered in iridescent green oval scales. These contain diamond-based crystal lattices oriented in all directions to give a brilliant green coloration that hardly varies with angle. The scales are effectively divided into pixels about a micrometre wide. Each such pixel is a single crystal and reflects light in a direction different from its neighbours. Selective mirrors to create interference effects are formed of micron-sized bowl-shaped pits lined with multiple layers of chitin in the wing scales of Papilio palinurus, the emerald swallowtail butterfly. These act as highly selective mirrors for two wavelengths of light. Yellow light is reflected directly from the centres of the pits; blue light is reflected twice by the sides of the pits. The combination appears green, but can be seen as an array of yellow spots surrounded by blue circles under a microscope. The chitin walls of the hollow bristles form a hexagonal honeycomb-shaped photonic crystal; the hexagonal holes are 0.51 μm apart. The structure behaves optically as if it consisted of a stack of 88 diffraction gratings, making Aphrodita one of the most iridescent of marine organisms. created by random nanochannels Deformed matrices, consisting of randomly oriented nanochannels in a spongelike keratin matrix, create the diffuse non-iridescent blue colour of Ara ararauna, the blue-and-yellow macaw. Since the reflections are not all arranged in the same direction, the colours, while still magnificent, do not vary much with angle, so they are not iridescent. '' berries Spiral coils, formed of helicoidally stacked cellulose microfibrils, create Bragg reflection in the "marble berries" of the African herb Pollia condensata, resulting in the most intense blue coloration known in nature. The berry's surface has four layers of cells with thick walls, containing spirals of transparent cellulose spaced so as to allow constructive interference with blue light. Below these cells is a layer two or three cells thick containing dark brown tannins. Pollia produces a stronger colour than the wings of Morpho butterflies, and is one of the first instances of structural coloration known from any plant. Each cell has its own thickness of stacked fibres, making it reflect a different colour from its neighbours, and producing a pixellated or pointillist effect with different blues speckled with brilliant green, purple, and red dots. The fibres in any one cell are either left-handed or right-handed, so each cell circularly polarizes the light it reflects in one direction or the other. Pollia is the first organism known to show such random polarization of light, which, nevertheless does not have a visual function, as the seed-eating birds who visit this plant species are not able to perceive polarised light. Spiral microstructures are also found in scarab beetles where they produce iridescent colours. petals exploit both yellow pigment and structural coloration. Thin film with diffuse reflector, based on the top two layers of a buttercup's petals. The brilliant yellow gloss derives from a combination, rare among plants, of yellow pigment and structural coloration. The very smooth upper epidermis acts as a reflective and iridescent thin film; for example, in Ranunculus acris, the layer is 2.7 micrometres thick. The unusual starch cells form a diffuse but strong reflector, enhancing the flower's brilliance. The curved petals form a paraboloidal dish which directs the sun's heat to the reproductive parts at the centre of the flower, keeping it some degrees Celsius above the ambient temperature. Interference from multiple total internal reflections can occur in microscale structures, such as sessile water droplets and biphasic oil-in-water droplets as well as polymer microstructured surfaces. In this structural coloration mechanism, light rays that travel by different paths of total internal reflection along an interface interfere to generate iridescent colour. Variable structures '' Some animals including cephalopods such as squid are able to vary their colours rapidly for both camouflage and signalling. The mechanisms include reversible proteins which can be switched between two configurations. The configuration of reflectin proteins in chromatophore cells in the skin of the Doryteuthis pealeii squid is controlled by electric charge. When charge is absent, the proteins stack together tightly, forming a thin, more reflective layer; when charge is present, the molecules stack more loosely, forming a thicker layer. Since chromatophores contain multiple reflectin layers, the switch changes the layer spacing and hence the colour of light that is reflected. == Examples ==
Examples
File:European bee eater.jpg|European bee-eaters owe their brilliant colours partly to diffraction grating microstructures in their feathers File:Butterfly Morpho rhetenor helena (M) KL.jpg|In Morpho butterflies such as Morpho helena the brilliant colours are produced by intricate firtree-shaped microstructures too small for optical microscopes. File:Parotia lawesii by Bowdler Sharpe.jpg|The male Parotia lawesii bird of paradise signals to the female with his breast feathers that switch from blue to yellow. File:Green Swallotail (Papilio palinurus) - Relic38.jpg|Brilliant green of emerald swallowtail, Papilio palinurus, is created by arrays of microscopic bowls that reflect yellow directly and blue from the sides. File:Parides sesostris MHNT dos.jpg|Emerald-patched cattleheart butterfly, Parides sesostris, creates its brilliant green using photonic crystals. File:Curculionidae - Lamprocyphus augustus.JPG|Iridescent scales of Lamprocyphus augustus weevil contain diamond-based crystal lattices oriented in all directions to give almost uniform green. File:Scales covering Entimus imperialis' elytra.jpg|Iridescent scales on Entimus imperialis weevil File:Entimus imperialis' photonic crystal.jpg|Electron micrograph of the three-dimensional photonic crystals within the scales on Entimus imperialis weevil File:Aphrodita aculeata (Sea mouse).jpg|Hollow nanofibre bristles of Aphrodita aculeata (a species of sea mouse) reflect light in yellows, reds and greens to warn off predators. File:Loligo pealeii.jpg|Longfin inshore squid, Doryteuthis pealeii, has been studied for its ability to change colour. File:Thinfilmbubble.jpg|Thin-film interference in a soap bubble. Colour varies with film thickness. File:Pepsis.jpg|Wasps of the Pepsis and Hemipepsis genera often produce a bluish tint from the sculpturing of their otherwise black chitin. File:Briarius weevil, from two slightly different angles.jpg|Two photographs of the same Eupholus weevil show the unique expression of structural colour. == In technology ==
In technology
's colour photographs, "Le Cervin", 1899, made using a monochrome photographic process (a single emulsion). The colours are structural, created by interference with light reflected from the back of the glass plate. Gabriel Lippmann won the Nobel Prize in Physics in 1908 for his work on a structural coloration method of colour photography, the Lippmann plate. This used a photosensitive emulsion fine enough for the interference caused by light waves reflecting off the back of the glass plate to be recorded in the thickness of the emulsion layer, in a monochrome (black and white) photographic process. Shining white light through the plate effectively reconstructs the colours of the photographed scene. In 2010, the dressmaker Donna Sgro made a dress from Teijin Fibers' Morphotex, an undyed fabric woven from structurally coloured fibres, mimicking the microstructure of Morpho butterfly wing scales. The fibres are composed of 61 flat alternating layers, between 70 and 100 nanometres thick, of two plastics with different refractive indices, nylon and polyester, in a transparent nylon sheath with an oval cross-section. The materials are arranged so that the colour does not vary with angle. The fibres have been produced in red, green, blue, and violet. Several countries and regions, including the U.S., European Union, and Brazil, use banknotes that include optically variable ink, which is structurally coloured, as a security feature. These pearlescent inks appear as different colours depending on the angle the banknote is viewed from. Because the ink is hard to obtain, and because a photocopier or scanner (which works from only one angle) cannot reproduce or even perceive the color-shifting effect, the ink serves to make counterfeiting more difficult. Structural coloration could be further exploited industrially and commercially, and research that could lead to such applications is under way. A direct parallel would be to create active or adaptive military camouflage fabrics that vary their colours and patterns to match their environments, just as chameleons and cephalopods do. The ability to vary reflectivity to different wavelengths of light could also lead to efficient optical switches that could function like transistors, enabling engineers to make fast optical computers and routers. Similarly, the eyes of some moths have antireflective surfaces, again using arrays of pillars smaller than the wavelength of light. "Moth-eye" nanostructures could be used to create low-reflectance glass for windows, solar cells, display devices, and military stealth technologies. Antireflective biomimetic surfaces using the "moth-eye" principle can be manufactured by first creating a mask by lithography with gold nanoparticles, and then performing reactive-ion etching. == See also ==
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