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First published online October 7, 2004
Journal of Experimental Biology 207, 3999-4009 (2004)
Published by The Company of Biologists 2004
doi: 10.1242/jeb.01240
Blue integumentary structural colours in dragonflies (Odonata) are not produced by incoherent Tyndall scattering
1 Department of Ecology and Evolutionary Biology, Yale University, PO Box
208105, New Haven, CT 06520, USA
2 Department of Ecology and Evolutionary Biology, University of Kansas,
Lawrence, KS 66045, USA
3 Department of Mathematics, University of Kansas, Lawrence, KS 66045,
USA
* Author for correspondence (e-mail: richard.prum{at}yale.edu)
Accepted 12 August 2004
For nearly 80 years, the non-iridescent, blue, integumentary structural
colours of dragonflies and damselflies (Odonata) have been attributed to
incoherent Tyndall or Rayleigh scattering. We investigated the production of
the integumentary structural colours of a damselfly the familiar
bluet, Enallagma civile (Coenagrionidae) and a dragonfly
the common green darner, Anax junius (Aeshnidae)
using fibre optic spectrophotometry and transmission electron microscopy
(TEM). The reflectance spectra of both species showed discrete reflectance
peaks of
30% reflectance at 475 and 460 nm, respectively. These
structural colours are produced by light scattering from closely packed arrays
of spheres in the endoplasmic reticulum of box-shaped epidermal pigment cells
underlying the cuticle. The observed reflectance spectra do not conform to the
inverse fourth power relationship predicted for Tyndall/Rayleigh scattering.
Two-dimensional (2-D) Fourier analysis of the TEM images of the
colour-producing arrays reveals ring-shaped distributions of Fourier power at
intermediate spatial frequencies, documenting a quasiordered nanostructure.
The nanostructured Fourier power spectra falsify the assumption of spatial
independence of scatterers that is required for incoherent scattering. Radial
averages of the Fourier power spectrum indicate that the spheres are
substantially nanostructured at the appropriate spatial scale to produce
visible colours by coherent scattering. However, the spatial periodicity of
the arrays is apparently too large to produce the observed colour by coherent
scattering. The nanospheres could have expanded substantially (
50%)
during preparation for TEM. Alternatively, coherent light scattering could be
occurring both from the surfaces and from structures at the centre of
the spheres. These arrays of colour-producing spheres within pigment cells
have convergently evolved at least 1114 times independently within the
Odonata. Structural colouration from arrays in living cells has also fostered
the convergent evolution of temperature-dependent colour change in numerous
odonate lineages.
Key words: structural colour, dragonfly, damselfly, coherent scattering, Tyndall scattering, Rayleigh scattering, pigment cells, Enallagma civile, Anax junius
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