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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,
sometimes 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.

Structural coloration was first observed by English

scientists Robert Hooke and Isaac Newton, and its
principle – wave interference – explained by Thomas
The brilliant iridescent colors of the
Young a century later. Young described iridescence as the
peacock's tail feathers are created by
result of interference between reflections from two or
structural coloration, as first noted by
more surfaces of thin films, combined with refraction as
Isaac Newton and Robert Hooke.
light enters and leaves such films. The geometry then
determines that at certain angles, the light reflected from
both surfaces interferes constructively, while at other angles, the light interferes destructively.
Different colours therefore appear at different angles.

In animals such as on the feathers of birds and the scales of butterflies, interference is created by a
range of photonic mechanisms, including diffraction gratings, selective mirrors, photonic crystals,
crystal fibres, matrices of nanochannels and proteins that can vary their configuration. Some cuts
of meat also show structural coloration due to the exposure of the periodic arrangement of the
muscular fibres. Many of these photonic mechanisms correspond to elaborate structures visible by
electron microscopy. In the few plants that exploit structural coloration, brilliant colours are
produced by structures within cells. The most brilliant blue coloration known in any living tissue is
found in the marble berries of Pollia condensata, where a spiral structure of cellulose fibrils
produces Bragg's law scattering of light. The bright gloss of buttercups is produced by thin-film
reflection by the epidermis supplemented by yellow pigmentation, and strong diffuse scattering by
a layer of starch cells immediately beneath.

Structural coloration has potential for industrial, commercial and military application, with
biomimetic surfaces that could provide brilliant colours, adaptive camouflage, efficient optical
switches and low-reflectance glass.

Contents
History
Principles
Structure not pigment
Principle of iridescence
Mechanisms
Fixed structures
Variable structures
Examples
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In technology
See also
Bibliography
Pioneering books
Research
General books
Notes
References
External links

History
In his 1665 book Micrographia, Robert Hooke described the
"fantastical" colours of the peacock's feathers:[1]

The parts of the Feathers of this glorious Bird appear,

through the Microscope, no less gaudy then do the whole
Feathers; for, as to the naked eye 'tis evident that the stem
or quill of each Feather in the tail sends out multitudes of
Lateral branches, … so each of those threads in the
Microscope appears a large long body, consisting of a
multitude of bright reflecting parts.

… their upper sides seem to me to consist of a multitude of

thin plated bodies, which are exceeding thin, and lie very
close together, and thereby, like mother of Pearl shells, do
not onely reflect a very brisk light, but tinge that light in a Robert Hooke's 1665
most curious manner; and by means of various positions, Micrographia contains the
in respect of the light, they reflect back now one colour, first observations of
and then another, and those most vividly. Now, that these structural colours.
colours are onely fantastical ones, that is, such as arise
immediately from the refractions of the light, I found by
this, that water wetting these colour'd parts, destroy'd
their colours, which seem'd to proceed from the alteration
of the reflection and refraction.[1]

In his 1704 book Opticks, Isaac Newton described the mechanism of the colours other than the
brown pigment of peacock tail feathers.[2] Newton noted that[3]

The finely colour'd Feathers of some Birds, and particularly those of Peacocks Tails, do,
in the very same part of the Feather, appear of several Colours in several Positions of
the Eye, after the very same manner that thin Plates were found to do in the 7th and
19th Observations, and therefore their Colours arise from the thinness of the
transparent parts of the Feathers; that is, from the slenderness of the very fine Hairs, or
Capillamenta, which grow out of the sides of the grosser lateral Branches or Fibres of
those Feathers.[3]

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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.[4][5]

In his 1892 book Animal Coloration, Frank Evers Beddard (1858–1925) acknowledged the
existence of structural colours:

The colours of animals are due either solely to the

presence of definite pigments in the skin, or …
beneath the skin; or they are partly caused by
optical effects due to the scattering, diffraction or
unequal refraction of the light rays. Colours of the
latter kind are often spoken of as structural colours;
they are caused by the structure of the coloured In 1892, Frank Evers Beddard noted
surfaces. The metallic lustre of the feathers of many that Chrysospalax golden moles'
birds, such as the humming birds, is due to the thick fur was structurally coloured.
presence of excessively fine striae upon the surface
of the feathers.[6]: 1 

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;"[6]: 2  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",[6]: 2  though he does later admit that the Cape golden
mole has "structural peculiarities" in its hair that "give rise to brilliant colours".[6]: 32 

Principles

Structure not pigment

Structural coloration is caused by interference effects rather

than by pigments.[7][8] 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.[9]

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.[10][11] These are often iridescent, as in When light falls on a thin film, the
peacock feathers and nacreous shells such as of pearl oysters waves reflected from the upper and
(Pteriidae) and Nautilus. This is because the reflected colour lower surfaces travel different
depends on the viewing angle, which in turn governs the distances depending on the angle,
apparent spacing of the structures responsible. [12] Structural so they interfere.
colours can be combined with pigment colours: peacock
feathers are pigmented brown with melanin,[1][10][13][14] while
buttercup petals have both carotenoid pigments for yellowness and thin films for reflectiveness.[11]

Principle of iridescence

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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
Electron micrograph of a fractured of it is reflected from their A 3-slide series of pictures taken
surface of nacre showing multiple bottom surfaces. The two with and without a pair of
thin layers sets of reflected waves MasterImage 3D circularly polarized
travel back upwards in the movie glasses of some dead
same direction. But since European rose chafers (Cetonia
aurata) whose shiny green colour
the bottom-reflected waves travelled a little farther –
comes from left-polarized light. Note
controlled by the thickness and refractive index of the film, and
that, without glasses, both the
the angle at which the light fell – the two sets of waves are out
beetles and their images have shiny
of phase. When the waves are one or more whole wavelengths
colour. The right-polarizer removes
apart – in other words, at certain specific angles, they add
the colour of the beetles but leaves
(interfere constructively), giving a strong reflection. At other
the color of the images. The left-
angles and phase differences, they can subtract, giving weak polarizer does the opposite,
reflections. The thin film therefore selectively reflects just one showing reversal of handedness of
wavelength – a pure colour – at any given angle, but other the reflected light.
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.[2]

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.[8] Structures
can be far more elaborate than a single thin film: films can be stacked up to give strong
iridescence, to combine two colours, or to balance out the inevitable change of colour with angle to
give a more diffuse, less iridescent effect.[10] Each mechanism offers a specific solution to the
problem of creating a bright colour or combination of colours visible from different directions.

A diffraction grating constructed of layers of chitin and air gives rise to the iridescent colours of
various butterfly wing scales as well as to the tail feathers of birds such as the peacock. Hooke and
Newton were correct in their claim that the peacock's colours are created by interference, but the
structures responsible, being close to the wavelength of light in scale (see micrographs), were
smaller than the striated structures they could see with their light microscopes. Another way to
produce a diffraction grating is with tree-shaped arrays of chitin, as in the wing scales of some of
the brilliantly coloured tropical Morpho butterflies (see drawing). Yet another variant exists in
Parotia lawesii, Lawes's parotia, a bird of paradise. The barbules of the feathers of its brightly
coloured breast patch are V-shaped, creating thin-film microstructures that strongly reflect two
different colours, bright blue-green and orange-yellow. 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.[10][15]

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Photonic crystals can be formed in

different ways.[16] In Parides sesostris, the
emerald-patched cattleheart butterfly,[17]
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
Drawing of
orientations. The result is that these
'firtree' micro-
emerald-patched cattleheart scales reflect
structures in
green light evenly at different angles
Morpho butterfly
wing scale
instead of being iridescent.[10][18] 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 Butterfly wing at different
effectively divided into pixels about a micrometre wide. Each magnifications reveals
such pixel is a single crystal and reflects light in a direction microstructured chitin acting as a
different from its neighbours.[19][20] diffraction grating

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.[10]
Structural coloration through
selective mirrors in the emerald
Crystal fibres, formed of hexagonal arrays of hollow
swallowtail
nanofibres, create the bright iridescent colours of the bristles
of Aphrodita, the sea mouse, a non-wormlike genus of marine
annelids.[10] The colours are aposematic, warning predators
not to attack.[21] 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.[22]

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.[10][23]

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.[24] 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,
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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
Magnificent non-iridescent colours
polarizes the light it reflects
of blue-and-yellow macaw created The most intense blue known in
in one direction or the
by random nanochannels nature: Pollia condensata berries
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.[25] Spiral
microstructures are also found in scarab beetles where they produce iridescent colours.

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,
Buttercup petals exploit both yellow pigment and structural
enhancing the flower's brilliance. The
coloration.
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.[11]

Surface gratings, consisting of ordered surface features due to exposure of ordered muscle cells
on cuts of meat. The structural coloration on meat cuts appears only after the ordered pattern of
muscle fibrils is exposed and light is diffracted by the proteins in the fibrils. The coloration or
wavelength of the diffracted light depends on the angle of observation and can be enhanced by
covering the meat with translucent foils. Roughening the surface or removing water content by
drying causes the structure to collapse, thus, the structural coloration to disappear.[26]

Interference from multiple total internal reflections can occur in microscale structures,
such as sessile water droplets and biphasic oil-in-water droplets[27] as well as polymer
microstructured surfaces.[28] 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

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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
Variable ring patterns on mantles of Hapalochlaena lunulata
colour of light that is reflected.[10]

Blue-ringed octopuses spend much of

their time hiding in crevices whilst displaying effective camouflage patterns with their dermal
chromatophore cells. If they are provoked, they quickly change colour, becoming bright yellow
with each of the 50-60 rings flashing bright iridescent blue within a third of a second. In the
greater blue-ringed octopus (Hapalochlaena lunulata), the rings contain multi-layer iridophores.
These are arranged to reflect blue–green light in a wide viewing direction. The fast flashes of the
blue rings are achieved using muscles under neural control. Under normal circumstances, each
ring is hidden by contraction of muscles above the iridophores. When these relax and muscles
outside the ring contract, the bright blue rings are exposed.[29]

Examples

European bee-eaters In Morpho butterflies The male Parotia lawesii

owe their brilliant colours such as Morpho helena bird of paradise signals
partly to diffraction the brilliant colours are to the female with his
grating microstructures produced by intricate breast feathers that
in their feathers firtree-shaped switch from blue to
microstructures too yellow.
small for optical
microscopes.

Brilliant green of Emerald-patched Iridescent scales of

emerald swallowtail, cattleheart butterfly, Lamprocyphus augustus
Papilio palinurus, is Parides sesostris, weevil contain diamond-
created by arrays of creates its brilliant green based crystal lattices
microscopic bowls that using photonic crystals. oriented in all directions
reflect yellow directly to give almost uniform
and blue from the sides. green.

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Hollow nanofibre bristles

Iridescent scales on Electron micrograph of of Aphrodita aculeata (a
Entimus imperialis the three-dimensional species of sea mouse)
weevil photonic crystals within reflect light in yellows,
the scales on Entimus reds and greens to warn
imperialis weevil off predators.

Longfin inshore squid, Thin-film interference in

Doryteuthis pealeii, has a soap bubble. Colour
been studied for its varies with film
ability to change colour. thickness.

In technology
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.[30][31]

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 One of Gabriel Lippmann's colour
butterfly wing scales.[32][33][34] The fibres are composed of 61 photographs, "Le Cervin", 1899,
flat alternating layers, between 70 and 100 nanometres thick, made using a monochrome
of two plastics with different refractive indices, nylon and photographic process (a single
polyester, in a transparent nylon sheath with an oval cross- emulsion). The colours are
structural, created by interference
section. The materials are arranged so that the colour does not
with light reflected from the back of
vary with angle.[35] The fibres have been produced in red,
the glass plate.
green, blue, and violet.[36]

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

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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.[10]

The surface of the compound eye of the housefly is densely packed with microscopic projections
that have the effect of reducing reflection and hence increasing transmission of incident light.[37]
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.[38] 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.[39]

See also
Animal coloration
Camouflage
Patterns in nature

Bibliography

Pioneering books
Beddard, Frank Evers (1892). Animal Coloration, An Account of the Principal Facts and
Theories Relating to the Colours and Markings of Animals. Swan Sonnenschein, London.

--- 2nd Edition, 1895 (https://www.amazon.co.uk/Coloration-Account-Principal-Theories-Rela

ting/dp/0543914062/ref=sr_1_11?s=books&ie=UTF8&qid=1335557196&sr=1-11).

Hooke, Robert (1665). Micrographia, John Martyn and James Allestry, London.
Newton, Isaac (1704). Opticks, William Innys, London.

Research
Fox, D.L. (1992). Animal Biochromes and Animal Structural Colours. University of California
Press.
Johnsen, S. (2011). The Optics of Life: A Biologist's Guide to Light in Nature. Princeton
University Press.
Kolle, M. (2011). Photonic Structures Inspired by Nature . Springer.

General books
Brebbia, C.A. (2011). Colour in Art, Design and Nature. WIT Press.
Lee, D.W. (2008). Nature's Palette: The Science of Plant Color. University of Chicago Press.
Kinoshita, S. (2008). "Structural Color in the Realm of Nature". World Scientific Publishing
Mouchet, S. R., Deparis, O. (2021). "Natural Photonics and Bioinspiration". Artech House

Notes

References
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1. Hooke, Robert. Micrographia. Chapter 36 ('Observ. XXXVI. Of Peacoks, Ducks, and Other
Feathers of Changeable Colours.')
2. "Iridescence in Lepidoptera" (https://web.archive.org/web/20140407082122/http://emps.exeter.
ac.uk/physics-astronomy/research/emag/themes/natural-photonics/iridescenceinlepidoptera/).
Natural Photonics (originally in Physics Review Magazine). University of Exeter. September
1998. Archived from the original (http://emps.exeter.ac.uk/physics-astronomy/research/emag/t
hemes/natural-photonics/iridescenceinlepidoptera/) on April 7, 2014. Retrieved April 27, 2012.
3. Newton, Isaac (1730) [1704]. Opticks (http://www.gutenberg.org/files/33504/33504-h/33504-h.
htm) (4th ed.). William Innys at the West-End of St. Paul's, London. pp. Prop. V., page 251.
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Light" (https://zenodo.org/record/1432310). Philosophical Transactions of the Royal Society of
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pp. 96–101.
6. Beddard, Frank Evers (1892). Animal Coloration: an account of the principal facts and theories
relating to the colours and markings of animals (https://books.google.com/books?id=5K-nT8GI
n5AC). Swan Sonnenschein. ISBN 978-0-543-91406-4.
7. Structural colour under the microscope! Feathers, beetles and butterflie!! (https://www.youtube.
com/watch?v=8DGA9eT16ec)
8. Mouchet, Sébastien R; Deparis, Olivier (2021), Natural Photonics and Bioinspiration (https://u
s.artechhouse.com/Natural-Photonics-and-Bioinspiration-P2221.aspx) (1st ed.), Artech House,
ISBN 978-163-081-797-8
9. Parker, A.R., Martini, N. (June–September 2006). "Structural colour in animals—simple to
complex optics". Optics & Laser Technology. 38 (4–6): 315–322.
Bibcode:2006OptLT..38..315P (https://ui.adsabs.harvard.edu/abs/2006OptLT..38..315P).
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10. Ball, Philip (May 2012). "Nature's Color Tricks". Scientific American. 306 (5): 74–79.
Bibcode:2012SciAm.306e..74B (https://ui.adsabs.harvard.edu/abs/2012SciAm.306e..74B).
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74). PMID 22550931 (https://pubmed.ncbi.nlm.nih.gov/22550931).
11. van der Kooi, C.J.; Elzenga, J.T.M.; Dijksterhuis, J.; Stavenga, D.G. (2017). "Functional optics
of glossy buttercup flowers" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5332578). Journal
of the Royal Society Interface. 14 (127): 20160933. doi:10.1098/rsif.2016.0933 (https://doi.org/
10.1098%2Frsif.2016.0933). PMC 5332578 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC53
32578). PMID 28228540 (https://pubmed.ncbi.nlm.nih.gov/28228540).
12. Wallin, Margareta (2002). "Nature's Palette: How animals, including humans, produce colours"
(http://www.bioscience-explained.org/ENvol1_2/pdf/paletteEN.pdf) (PDF). Bioscience
Explained. 1 (2): 1–12. Retrieved November 17, 2011.
13. Smyth, S.; et al. (2007). "What Makes the Peacock Feather Colorful?" (http://www.nnin.org/site
s/default/files/files/2007nninREUsmyth.pdf) (PDF). NNIN REU Journal.
14. Smyth, S. (2009). "What Makes the Peacock Feather Bright and Colorful" (https://web.archive.
org/web/20160304063316/https://scholarworks.alaska.edu/handle/11122/2835). University of
Alaska, Fairbanks (Honors Thesis). Archived from the original (https://scholarworks.alaska.ed
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15. Stavenga, Doekele G.; Leertouwer, H. L.; Marshall, N. J.; Osorio, D. (2010). "Dramatic colour
changes in a bird of paradise caused by uniquely structured breast feather barbules" (http://cb
n.eldoc.ub.rug.nl/FILES/root/2011/ProcRSocBStavenga/2011ProcRSocBStavenga.pdf) (PDF).
Proceedings of the Royal Society B. 278 (1715): 2098–2104. doi:10.1098/rspb.2010.2293 (http
s://doi.org/10.1098%2Frspb.2010.2293). PMC 3107630 (https://www.ncbi.nlm.nih.gov/pmc/arti
cles/PMC3107630). PMID 21159676 (https://pubmed.ncbi.nlm.nih.gov/21159676).
16. Welch, V.L., Vigneron, J.-P. (July 2007). "Beyond butterflies—the diversity of biological
photonic crystals" (http://perso.fundp.ac.be/~jpvigner/press/BioPhotonics/Beyond.pdf) (PDF).
Opt Quant Electron. 39 (4–6): 295–303. doi:10.1007/s11082-007-9094-4 (https://doi.org/10.10
07%2Fs11082-007-9094-4). S2CID 121911730 (https://api.semanticscholar.org/CorpusID:121
911730).
17. Yablonovitch, Eli (December 2001). "Photonic Crystals: Semiconductors of Light" (http://optoel
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External links
National Geographic News: Peacock Plumage Secrets Uncovered (http://news.nationalgeogra
phic.com/news/2003/10/1016_031017_peacockcolors.html)
Doucet, S. M.; Shawkey, M. D.; Hill, G. E.; Montgomerie, R. (2006). "Iridescent plumage in
satin bowerbirds: Structure, mechanisms and nanostructural predictors of individual variation
in colour" (https://doi.org/10.1242%2Fjeb.01988). Journal of Experimental Biology. 209 (2):
380–390. doi:10.1242/jeb.01988 (https://doi.org/10.1242%2Fjeb.01988). PMID 16391360 (http
s://pubmed.ncbi.nlm.nih.gov/16391360). S2CID 14595674 (https://api.semanticscholar.org/Cor
pusID:14595674).
Causes of Color: Peacock feathers (http://www.webexhibits.org/causesofcolor/15C.html)
Butterflies and Gyroids – Numberphile (https://www.youtube.com/watch?v=VFRrzzb1dvU)

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