Wing colors based on arrangement of the multilayer structure of wing scales in lycaenid butterflies (Insecta: Lepidoptera)

Male wing colors and wing scale morphology were examined for three species of lycaenid butterflies: Chrysozephyrus ataxus, Favonius cognatus and F. jezoensis. Measurement of spectral reflectance on the wing surface with a spectrophotometer revealed species‐specific reflection spectra, with one or two peaks in the ultraviolet and/or green ranges. Observations of wing scales using an optical microscope revealed that light was reflected from the inter‐ridge regions, where transmission electron microscopy revealed a multilayer structure. Based on the multilayer dimensions obtained, three models were devised and compared to explain the measured reflectance spectrum. The results showed that the best fit is a model in which thicknesses of thin films of the multilayer system are not constant and air spaces between cuticle layers are more or less packed with cuticle spacers. This suggests that the specific wing colors of the species examined are produced by the species‐specific arrangement of the multilayer structure of wing scales.     

Sensitivity Analysis of a Bioinspired Refractive Index Based Gas Sensor

It was found out that the change of refractive index of ambient gas can lead to obvious change of the color of Morpho butterfly's wing. Such phenomenon has been employed as a sensing principle for detecting gas. In the present study, Rigorous Coupled-Wave Analysis (RCWA) was described briefly, and the partial derivative of optical reflection efficiency with respect to the refractive index of ambient gas, i.e., sensitivity of the sensor, was derived based on RCWA. A bioinspired grating model was constructed by mimicking the nanostructure on the ground scale of Morpho didius butterfly's wing. The analytical sensitivity was verified and the effect of the grating shape on the reflection spectra and its sensitivity were discussed. The results show that by tuning shape parameters of the grating, we can obtain desired reflection spectra and sensitivity, which can be applied to the design of the bioinspired refractive index based gas sensor.     

Pterin pigment granules are responsible for both broadband light scattering and wavelength selective absorption in the wing scales of pierid butterflies

A small but growing literature indicates that many animal colours are produced by combinations of structural and pigmentary mechanisms. We investigated one such complex colour phenotype: the highly chromatic wing colours of pierid butterflies including oranges, yellows and patterns which appear white to the human eye, but strongly absorb the ultraviolet (UV) wavelengths visible to butterflies. Pierids produce these bright colours using wing scales that contain collections of minute granules. However, to date, no work has directly characterized the molecular composition or optical properties of these granules. We present results that indicate these granules contain pterin pigments. We also find that pterin granules increase light reflection from single wing scales, such that wing scales containing denser granule arrays reflect more light than those with less dense granule collections. As male wing scales contain more pterin granules than those of females, the sexual dichromatism found in many pierid species can be explained by differences in wing scale pterin deposition. Additionally, the colour pattern elements produced by these pterins are known to be important during mating interactions in a number of pierid species. Therefore, we discuss the potential relevance of our results within the framework of sexual selection and colour signal evolution.     

Parameters of the wing coloration in the butterfly genus Colias Fabr. (Lepidoptera, Pieridae) of the steppe zone of eastern and Central Europe

On the basis of reflection measurements and colorimetric characteristics, the parameters of wing coloration for six European species of the genus Colias are established. The specificity of individual species and differentiation between the species according to the quantitative indices for wing coloration are revealed.     

Estimation of the leaf chlorophyll content using multiangular spectral reflectance factor

Chlorophyll is one of the primary pigments of plant leaves, and changes in its content can be used to characterize the physiological status of plants. Spectral indices have been devised and validated for estimating leaf chlorophyll content (LCC). However, most of the existing spectral indices do not consider the influence of angular reflection on the accuracy of the LCC estimation. In this study, the spectral reflectance factors of leaves from three plant species were measured from several observations in the principal plane. The relationship between the existing spectral indices and the LCC from different directions suggests that the directional reflection of a leaf surface impacts the accuracy of its LCC estimation. Subsequently, the ratio of reflectance differences, that is, the modified Datt index, was tested to reduce the directional reflection effect when predicting LCC. Our results indicated that the modified Datt index not only estimated LCC with high accuracy for all observation directions and plant species but also consistently predicted the LCC of each species in individual observation directions. Our method opens the possibility for optical detection of LCC using multiangular spectral reflection, which is convenient for plant science studies focused on the variation in LCC.     

Fourier analysis of wing beat signals: assessing the effects of genetic alterations of flight muscle structure in Diptera.

A method for determining and analyzing the wing beat frequency in Diptera is presented. This method uses an optical tachometer to measure Diptera wing movement during flight. The resulting signal from the optical measurement is analyzed using a Fast Fourier Transform (FFT) technique, and the dominant frequency peak in the Fourier spectrum is selected as the wing beat frequency. Also described is a method for determining quantitatively the degree of variability of the wing beat frequency about the dominant frequency. This method is based on determination of a quantity called the Hindex, which is derived using data from the FFT analysis. Calculation of the H index allows computer-based selection of the most suitable segment of recorded data for determination of the representative wing beat frequency. Experimental data suggest that the H index can also prove useful in examining wing beat frequency variability in Diptera whose flight muscle structure has been genetically altered. Examples from Drosophila indirect flight muscle studies as well as examples of artificial data are presented to illustrate the method. This method fulfills a need for a standardized method for determining wing beat frequencies and examining wing beat frequency variability in insects whose flight muscles have been altered by protein engineering methods.     

Remarkable iridescence in the hindwings of the damselfly Neurobasis chinensis chinensis (Linnaeus) (Zygoptera: Calopterygidae)

The bright green dorsal iridescence of the hindwings of Neurobasis chinensis chinensis males, very rare in Odonata, is known to play a significant role in their courtship behaviour. The mechanism responsible for such high contrast and spectrally pure colour has been investigated and found to be optical interference, producing structural colour from distinct laminations in the wing membrane cuticle. The ventral sides of these iridescent wings are dark brown in colour. In a single continuous membrane of wing cuticle, this is an effect that requires a specialized structure. It is accomplished through the presence of high optical absorption (kappa = 0.13) within two thick layers near the ventral surface of the wing, which leads to superior dorsal colour characteristics. By simultaneously fitting five sets of optical reflectivity and transmissivity spectra to theory, we were able to extract very accurate values of the complex refractive index for all three layer types present in the wing. The real parts of these are n = 1.47, 1.68 and 1.74. Although there is often similarly significant dorsal and ventral colour contrast in other structurally coloured natural systems, very few system designs comprise only a single continuous membrane.     

A New Model for the Evolution of the Pterosaur Wing--with a twist

A new hypothesis for the evolution of the pterosaur wing is presented. It is based on the observation that many aspects of pterosaur non-wing anatomy were present in their non-volant, prolacertiform sister taxa. In pterosaurs alone, metacarpal IV was twisted 90° medially, so the wing digit flexed in the plane of the metacarpus, rather than towards the palm as in other tetrapods. While grappling a tree using manual digits I-III, a pre-pterosaur with this character would have been free to flex and extend digit IV in the plane tangential to the circumference of the tree. The addition of a small dermal extension that opened like a Japanese fan would essentially have completed the development of the proto-wing. Its subsequent increase in size was brought on by selective competition. Successful powered flight would have been possible only after a critical wing size had been achieved.     

Comparative morphology and evolutionary aspects of the reflective under wing scale-pattern in Fritillary butterflies (Nymphalidae: Argynnini)

The wing scale ultrastructure of the reflective under wing pattern found in many Argynnini butterflies are examined for all recognised genera and subgenera and compared to that of some basal Heliconiini. A true reflective pattern probably evolved once within the Argynnini. But the phylogenetic information in these structures is limited due to a high degree of homoplasy in the scale ultrastructure related to the reflective patterns. The degree of specialisation is also homoplastic. The general morphological modification responsible for the reflective patterns seems to be a “closing” of the large windows, which generally occupy most of the inter-ridge space on the abwing surface in the scales of higher Lepidoptera. The fact that the Argynnis niobe morphs with a silvery pattern have scales with ‘closed windows’ whereas the Argynnis niobe morphs without a silvery pattern have typical non-reflective scales with very large windows supports this conclusion.The degrees of modification of the scales, including whether both cover and ground scales or only cover scales are modified, are to some extent correlated to the degree of reflectiveness in the wing pattern. Boloria eunomia has, as the only species, more modified ground scales than cover scales.     

Inhibiting aphid wing development: is immune challenge important?

Aphids show inhibition of wing development when parasitized by hymenopteran parasitoids, especially when parasitism takes place early in the host's development. The present study tests the hypothesis that the aphid itself redirects the resources needed for the development of wings to support the immune response, rather than the parasitoid directly promoting wing inhibition. When the aphid's immune system is challenged with bacterial antigens and a fungal pathogen, neither immune challenge alters wing development, indicating that perhaps there is no redirection by the aphid of resources required for wing development to cover possible costs of mounting an immune response.     

Anisotropism of the Non-Smooth Surface of Butterfly Wing

Twenty-nine species of butterflies were collected for observation and determination of the wing surfaces using a ScanningElectron Microscope(SEM).Butterfly wing surface displays structural anisotropism in micro-,submicro- and nano-scales.Thescales on butterfly wing surface arrange like overlapping roof tiles.There are submicrometric vertical gibbosities,horizontallinks,and nano-protuberances on the scales.First-incline-then-drip method and first-drip-then-incline method were used tomeasure the Sliding Angle(SA)of droplet on butterfly wing surface by an optical Contact Angle(CA)measuring system.Relatively smaller sliding angles indicate that the butterfly wing surface has fine self-cleaning property.Significantly differentSAs in various directions indicate the anisotropic self-cleaning property of butterfly wing surface.The SAs on the butterfly wingsurface without scales are remarkably larger than those with scales,which proves the crucial role of scales in determining theself-cleaning property.Butterfly wing surface is a template for design and fabrication ofbiomimetic materials and self-cleaningsubstrates.This work may offer insights into how to design directional self-cleaning coatings and anisotropic wetting surface.     

Direct measurements of the kinematics and dynamics of bat flight

Experimental measurements and analysis of the flight of bats are presented, including kinematic analysis of high-speed stereo videography of straight and turning flight, and measurements of the wake velocity field behind the bat. The kinematic data reveal that, at relatively slow flight speeds, wing motion is quite complex, including a sharp retraction of the wing during the upstroke and a broad sweep of the partially extended wing during the downstroke. The data also indicate that the flight speed and elevation are not constant, but oscillate in synchrony with both the horizontal and vertical movements of the wing. PIV measurements in the transverse (Trefftz) plane of the wake indicate a complex 'wake vortex' structure dominated by a strong wing tip vortex shed from the wing tip during the downstroke and either the wing tip or a more proximal joint during the upstroke. Data synthesis of several discrete realizations suggests a 'cartoon' of the wake structure during the entire wing beat cycle. Considerable work remains to be done to confirm and amplify these results.     

Sexual Dichromatism of the Damselfly Calopteryx japonica Caused by a Melanin-Chitin Multilayer in the Male Wing Veins

Mature male Calopteryx japonica damselflies have dark-blue wings, due to darkly coloured wing membranes and blue reflecting veins. The membranes contain a high melanin concentration and the veins have a multilayer of melanin and chitin. Female and immature C. japonica damselflies have brown wings. We have determined the refractive index of melanin by comparing the differently pigmented wing membranes and applying Jamin-Lebedeff interference microscopy. Together with the previously measured refractive index of chitin the blue, structural colour of the male wing veins could be quantitatively explained by an optical multilayer model. The obtained melanin refractive index data will be useful in optical studies on melanized tissues, especially where melanin is concentrated in layers, thus causing iridescence.     

Die Spiegeloptik des Flußkrebsauges

Summary In the superposition eye of the crayfish (Astacus leptodactylus), images are formed by radial plane mirrors arranged in an orthogonal pattern. The optical structure of the crayfish eye can be described as a family of virtual reflecting cone envelopes concentric around each direction in space. There exist two reflection mechanisms: total internal reflection at the sides of the crystalline cones and reflection by multilayer mirrors attached to the distal parts of the cones. Image-forming rays have to be reflected twice in the general case, and once in the case of perpendicular position of the plane of incidence and the mirror plane. For rays incident at small angles to the ommatidial axis, this condition is almost satisfied due to a particular axial variation of the refractive index of the crystalline cone, and for rays incident at large angles, due to the spectral reflecting properties of the multilayer reflector.

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Dissertation der Universität Stuttgart

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Dem Betreuer der Arbeit, Prof. Dr. P. Kunze, danke ich für wertvolle Diskussionen, Frl. M. Gonnert für unermüdliche technische Hilfe und Dipl. Ing. W. Jaschinski für das Nachprüfen von Gleichungen.     

Microscopy with Plastic Substitutes for Cover Glasses

To determine whether plastic substitutes for cover glasses on microscope slides affect the performance of the microscope, their optical constants were determined. The plastic covers and glass cover glasses were mounted also on silvered slides to form Star Test Plates which were studied by competent observers. The thicker plastic cover glasses, now on the market, are satisfactory when mounted to give a plane surface. Optically inhomogeneous materials, of irregular thickness, those that curl or do not have plane surfaces, adversely affect the performance of the microscope and should not be used. Since the substitutes are softer than glass they must be protected from abrasion. It is recommended that thicknesses of 0.18 mm., and none outside of a range of 0.12 to 0.20 mm. be used. For critical observation with unimmersed objectives of high aperture, best results are obtained with the correction collar set at the position corresponding to the actual thickness of the cover slip, just as would be done with glass cover glasses.     

Improving Solar Cells’ Light Trapping by the Low Loss Interface Photonic Crystals

Improving the silicon layer’s optical absorption is a key research point for crystalline silicon based thin film solar cells. Light trapping is a method widely adopted to achieve this research purpose. In this paper, we propose low loss interface photonic crystals layer (IPC), which is sandwiched between the crystalline silicon layer and the cover layer. The low loss interface photonic crystals layer could boost the light trapping efficiency significantly. The mechanism is that the smaller refraction index difference between silicon layer and the low loss interface photonic crystals layer could reduce the light’s interface reflection. Taking advantage of the coupling calculation by optical and electrical simulations, solar cell’s absorption efficiency and electrical performance parameters are obtained. Compared with optimized reference group, the maximum output power of the proposed solar cell could be improved by 6.44%. The result indicates that the proposed low loss interface photonic crystals layer is applicable for light’s trapping in crystalline silicon thin film solar cells.

     

Planar cell polarity and tissue design: Shaping the Drosophila wing membrane

Planar cell polarity (PCP) describes the orientation of a cell within the plane of an epithelial cell layer. During tissue development, epithelial cells normally align their PCP so that they face in the same direction. This alignment allows cells to move in a common direction, or to generate structures with a common orientation. A classic system for studying the coordination of epithelial PCP is the developing Drosophila wing. The alignment of epithelial PCP during pupal wing development allows the production of an array of cell hairs that point towards the wing tip. Multiple studies have established that the Frizzled (Fz) PCP signaling pathway coordinates wing PCP. Recently, we have found that the same pathway also controls the formation of ridges on the Drosophila wing membrane. However, in contrast to hair polarity, ridge orientation differs between the anterior and posterior wing. How can the Fz PCP pathway generate a different relationship between hair and ridge orientation in different parts of the wing? In this Extra View article, we discuss membrane ridge development drawing upon our recent PLoS Genetics paper and other, published and unpublished, data. We also speculate upon how our findings impact the ongoing debate concerning the interaction of the Fz PCP and Fat/Dachsous pathways in the control of PCP.     

Programmed cell death in the wing of Orgyia leucostigma (Lepidoptera: Lymantriidae)

Programmed cell death is an integral and ubiquitous phenomenon of development that is responsible for the reduction of wing size in female moths of Orgyia leucostigma (Lymantriidae). Throughout larval and pupal life, cells of the wing epithelium proliferate and interact to form normal imaginal discs and pupal wings in both sexes. But at the onset of adult development, most cells in female O. leucostigma wings degenerate over a brief, 2-day period. Lysosomes and autophagic vacuoles appear in cells of the wing epithelium shortly after it retracts from the pupal cuticle. Hemocytes actively participate in removing the resulting cellular debris. By contrast, epithelial cells in wings of developing adult males of O. leucostigma do not undergo massive cell death. Wing epithelium of female pupae transferred to male pupal hosts behaves autonomously in this foreign environment. By pupation, cells of the female wing apparently are committed to self-destruct even in a male pupal environment. Normal interactions among epithelial cells within the plane of a wing monolayer as well as between the upper and lower monolayers of the wing are disrupted in female O. leucostigma by massive cell degeneration. Despite this disruption, the remaining cells of the wing contribute to the formation of a diminutive, but reasonably proportioned, adult wing with scales and veins.     

Comparative characteristics of the wing apparatus and flight of Syrphid flies (Diptera: Syrphidae)

Comparative analysis of the wing apparatus and flight in nine species of flower flies (Syrphidae) has been performed. Data on the flight velocity, aerodynamic force, wing-beat frequency, stroke amplitude and stroke plane angle, wing area, body mass and volume, as well as correlations between these parameters at the intraspecific and family levels, have been obtained. Based on the obtained data, the subfamilies Syrphinae and Eristalinae have been compared.     

Sex and age-specific differences in wing pointedness and wing length in blackcaps Sylvia atricapilla migrating through the southern Baltic coast

The blackcap Sylvia atricapilla shows a complex migratory pattern and is a suitable species for the studies of morphological migratory syndrome, including adaptations of wing shape to different migratory performance. Obligate migrants of this species that breed in northern, central, and Eastern Europe differ by migration distance and some cover shorter distance to the wintering grounds in the southern part of Europe/North Africa or the British Isles, although others migrate to sub-Saharan Africa. Based on ˃40 years of ringing data on blackcaps captured during autumn migration in the Southern Baltic region, we studied age- and sex-related correlations in wing pointedness and wing length of obligate blackcap migrants to understand the differences in migratory behavior of this species. Even though the recoveries of blackcaps were scarce, we reported some evidence that individuals which differ in migration distance differed also in wing length. We found that wing pointedness significantly increased with an increasing wing length of migrating birds, and adults had longer and more pointed wings than juvenile birds. This indicates stronger antipredator adaptation in juvenile blackcaps than selection on flight efficiency, which is particularly important during migration. Moreover, we documented more pronounced differences in wing length between adult and juvenile males and females. Such differences in wing length may enhance a faster speed of adult male blackcaps along the spring migration route and may be adaptive when taking into account climatic effects, which favor earlier arrival from migration to the breeding grounds.