2017-01-01
The evolution of wing pattern in Lepidoptera is a popular area of inquiry but few studies have examined microlepidoptera, with fewer still focusing on intraspecific variation. The tineid genus Moerarchis Durrant, 1914 includes two species with high intraspecific variation of wing pattern. A subset of the specimens examined here provide, to my knowledge, the first examples of wing patterns that follow both the ‘alternating wing-margin’ and ‘uniform wing-margin’ models in different regions along the costa. These models can also be evaluated along the dorsum of Moerarchis, where a similar transition between the two models can be seen. Fusion of veins is shown not to effect wing pattern, in agreement with previous inferences that the plesiomorphic location of wing veins constrains the development of colour pattern. The significant correlation between wing length and number of wing pattern elements in Moerarchis australasiella shows that wing size can act as a major determinant of wing pattern complexity. Lastly, some M. australasiella specimens have wing patterns that conform entirely to the ‘uniform wing-margin’ model and contain more than six bands, providing new empirical insight into the century-old question of how wing venation constrains wing patterns with seven or more bands. PMID:28405390
Lucas, Lauren K; Nice, Chris C; Gompert, Zachariah
2018-03-13
Patterns of phenotypic variation within and among species can be shaped and constrained by trait genetic architecture. This is particularly true for complex traits, such as butterfly wing patterns, that consist of multiple elements. Understanding the genetics of complex trait variation across species boundaries is difficult, as it necessitates mapping in structured populations and can involve many loci with small or variable phenotypic effects. Here, we investigate the genetic architecture of complex wing pattern variation in Lycaeides butterflies as a case study of mapping multivariate traits in wild populations that include multiple nominal species or groups. We identify conserved modules of integrated wing pattern elements within populations and species. We show that trait covariances within modules have a genetic basis and thus represent genetic constraints that can channel evolution. Consistent with this, we find evidence that evolutionary changes in wing patterns among populations and species occur in the directions of genetic covariances within these groups. Thus, we show that genetic constraints affect patterns of biological diversity (wing pattern) in Lycaeides, and we provide an analytical template for similar work in other systems. © 2018 John Wiley & Sons Ltd.
Conservatism and novelty in the genetic architecture of adaptation in Heliconius butterflies.
Huber, B; Whibley, A; Poul, Y L; Navarro, N; Martin, A; Baxter, S; Shah, A; Gilles, B; Wirth, T; McMillan, W O; Joron, M
2015-05-01
Understanding the genetic architecture of adaptive traits has been at the centre of modern evolutionary biology since Fisher; however, evaluating how the genetic architecture of ecologically important traits influences their diversification has been hampered by the scarcity of empirical data. Now, high-throughput genomics facilitates the detailed exploration of variation in the genome-to-phenotype map among closely related taxa. Here, we investigate the evolution of wing pattern diversity in Heliconius, a clade of neotropical butterflies that have undergone an adaptive radiation for wing-pattern mimicry and are influenced by distinct selection regimes. Using crosses between natural wing-pattern variants, we used genome-wide restriction site-associated DNA (RAD) genotyping, traditional linkage mapping and multivariate image analysis to study the evolution of the architecture of adaptive variation in two closely related species: Heliconius hecale and H. ismenius. We implemented a new morphometric procedure for the analysis of whole-wing pattern variation, which allows visualising spatial heatmaps of genotype-to-phenotype association for each quantitative trait locus separately. We used the H. melpomene reference genome to fine-map variation for each major wing-patterning region uncovered, evaluated the role of candidate genes and compared genetic architectures across the genus. Our results show that, although the loci responding to mimicry selection are highly conserved between species, their effect size and phenotypic action vary throughout the clade. Multilocus architecture is ancestral and maintained across species under directional selection, whereas the single-locus (supergene) inheritance controlling polymorphism in H. numata appears to have evolved only once. Nevertheless, the conservatism in the wing-patterning toolkit found throughout the genus does not appear to constrain phenotypic evolution towards local adaptive optima.
Schunk, Cosima; Swartz, Sharon M; Breuer, Kenneth S
2017-02-06
Aspect ratio (AR) is one parameter used to predict the flight performance of a bat species based on wing shape. Bats with high AR wings are thought to have superior lift-to-drag ratios and are therefore predicted to be able to fly faster or to sustain longer flights. By contrast, bats with lower AR wings are usually thought to exhibit higher manoeuvrability. However, the half-span ARs of most bat wings fall into a narrow range of about 2.5-4.5. Furthermore, these predictions do not take into account the wide variation in flapping motion observed in bats. To examine the influence of different stroke patterns, we measured lift and drag of highly compliant membrane wings with different bat-relevant ARs. A two degrees of freedom shoulder joint allowed for independent control of flapping amplitude and wing sweep. We tested five models with the same variations of stroke patterns, flapping frequencies and wind speed velocities. Our results suggest that within the relatively small AR range of bat wings, AR has no clear effect on force generation. Instead, the generation of lift by our simple model mostly depends on wingbeat frequency, flapping amplitude and freestream velocity; drag is mostly affected by the flapping amplitude.
The influence of aspect ratio and stroke pattern on force generation of a bat-inspired membrane wing
Swartz, Sharon M.; Breuer, Kenneth S.
2017-01-01
Aspect ratio (AR) is one parameter used to predict the flight performance of a bat species based on wing shape. Bats with high AR wings are thought to have superior lift-to-drag ratios and are therefore predicted to be able to fly faster or to sustain longer flights. By contrast, bats with lower AR wings are usually thought to exhibit higher manoeuvrability. However, the half-span ARs of most bat wings fall into a narrow range of about 2.5–4.5. Furthermore, these predictions do not take into account the wide variation in flapping motion observed in bats. To examine the influence of different stroke patterns, we measured lift and drag of highly compliant membrane wings with different bat-relevant ARs. A two degrees of freedom shoulder joint allowed for independent control of flapping amplitude and wing sweep. We tested five models with the same variations of stroke patterns, flapping frequencies and wind speed velocities. Our results suggest that within the relatively small AR range of bat wings, AR has no clear effect on force generation. Instead, the generation of lift by our simple model mostly depends on wingbeat frequency, flapping amplitude and freestream velocity; drag is mostly affected by the flapping amplitude. PMID:28163875
Alexis, Matamoro-Vidal; Isaac, Salazar-Ciudad; David, Houle
2015-01-01
One of the aims of evolutionary developmental biology is to discover the developmental origins of morphological variation. The discipline has mainly focused on qualitative morphological differences (e.g., presence or absence of a structure) between species. Studies addressing subtle, quantitative variation are less common. The Drosophila wing is a model for the study of development and evolution, making it suitable to investigate the developmental mechanisms underlying the subtle quantitative morphological variation observed in nature. Previous reviews have focused on the processes involved in wing differentiation, patterning and growth. Here, we investigate what is known about how the wing achieves its final shape, and what variation in development is capable of generating the variation in wing shape observed in nature. Three major developmental stages need to be considered: larval development, pupariation, and pupal development. The major cellular processes involved in the determination of tissue size and shape are cell proliferation, cell death, oriented cell division and oriented cell intercalation. We review how variation in temporal and spatial distribution of growth and transcription factors affects these cellular mechanisms, which in turn affects wing shape. We then discuss which aspects of the wing morphological variation are predictable on the basis of these mechanisms. PMID:25619644
The Functional Basis of Wing Patterning in Heliconius Butterflies: The Molecules Behind Mimicry
Kronforst, Marcus R.; Papa, Riccardo
2015-01-01
Wing-pattern mimicry in butterflies has provided an important example of adaptation since Charles Darwin and Alfred Russell Wallace proposed evolution by natural selection >150 years ago. The neotropical butterfly genus Heliconius played a central role in the development of mimicry theory and has since been studied extensively in the context of ecology and population biology, behavior, and mimicry genetics. Heliconius species are notable for their diverse color patterns, and previous crossing experiments revealed that much of this variation is controlled by a small number of large-effect, Mendelian switch loci. Recent comparative analyses have shown that the same switch loci control wing-pattern diversity throughout the genus, and a number of these have now been positionally cloned. Using a combination of comparative genetic mapping, association tests, and gene expression analyses, variation in red wing patterning throughout Heliconius has been traced back to the action of the transcription factor optix. Similarly, the signaling ligand WntA has been shown to control variation in melanin patterning across Heliconius and other butterflies. Our understanding of the molecular basis of Heliconius mimicry is now providing important insights into a variety of additional evolutionary phenomena, including the origin of supergenes, the interplay between constraint and evolvability, the genetic basis of convergence, the potential for introgression to facilitate adaptation, the mechanisms of hybrid speciation in animals, and the process of ecological speciation. PMID:25953905
Genomic architecture of adaptive color pattern divergence and convergence in Heliconius butterflies
Supple, Megan A.; Hines, Heather M.; Dasmahapatra, Kanchon K.; Lewis, James J.; Nielsen, Dahlia M.; Lavoie, Christine; Ray, David A.; Salazar, Camilo; McMillan, W. Owen; Counterman, Brian A.
2013-01-01
Identifying the genetic changes driving adaptive variation in natural populations is key to understanding the origins of biodiversity. The mosaic of mimetic wing patterns in Heliconius butterflies makes an excellent system for exploring adaptive variation using next-generation sequencing. In this study, we use a combination of techniques to annotate the genomic interval modulating red color pattern variation, identify a narrow region responsible for adaptive divergence and convergence in Heliconius wing color patterns, and explore the evolutionary history of these adaptive alleles. We use whole genome resequencing from four hybrid zones between divergent color pattern races of Heliconius erato and two hybrid zones of the co-mimic Heliconius melpomene to examine genetic variation across 2.2 Mb of a partial reference sequence. In the intergenic region near optix, the gene previously shown to be responsible for the complex red pattern variation in Heliconius, population genetic analyses identify a shared 65-kb region of divergence that includes several sites perfectly associated with phenotype within each species. This region likely contains multiple cis-regulatory elements that control discrete expression domains of optix. The parallel signatures of genetic differentiation in H. erato and H. melpomene support a shared genetic architecture between the two distantly related co-mimics; however, phylogenetic analysis suggests mimetic patterns in each species evolved independently. Using a combination of next-generation sequencing analyses, we have refined our understanding of the genetic architecture of wing pattern variation in Heliconius and gained important insights into the evolution of novel adaptive phenotypes in natural populations. PMID:23674305
Wing shape variation associated with mimicry in butterflies.
Jones, Robert T; Le Poul, Yann; Whibley, Annabel C; Mérot, Claire; ffrench-Constant, Richard H; Joron, Mathieu
2013-08-01
Mimetic resemblance in unpalatable butterflies has been studied by evolutionary biologists for over a century, but has largely focused on the convergence in wing color patterns. In Heliconius numata, discrete color-pattern morphs closely resemble comimics in the distantly related genus Melinaea. We examine the possibility that the shape of the butterfly wing also shows adaptive convergence. First, simple measures of forewing dimensions were taken of individuals in a cross between H. numata morphs, and showed quantitative differences between two of the segregating morphs, f. elegans and f. silvana. Second, landmark-based geometric morphometric and elliptical Fourier outline analyses were used to more fully characterize these shape differences. Extension of these techniques to specimens from natural populations suggested that, although many of the coexisting morphs could not be discriminated by shape, the differences we identified between f. elegans and f. silvana hold in the wild. Interestingly, despite extensive overlap, the shape variation between these two morphs is paralleled in their respective Melinaea comimics. Our study therefore suggests that wing-shape variation is associated with mimetic resemblance, and raises the intriguing possibility that the supergene responsible for controlling the major switch in color pattern between morphs also contributes to wing shape differences in H. numata. © 2013 The Author(s). Evolution © 2013 The Society for the Study of Evolution.
Reed, Robert D; McMillan, W Owen; Nagy, Lisa M
2008-01-07
Geographical variation in the mimetic wing patterns of the butterfly Heliconius erato is a textbook example of adaptive polymorphism; however, little is known about how this variation is controlled developmentally. Using microarrays and qPCR, we identified and compared expression of candidate genes potentially involved with a red/yellow forewing band polymorphism in H. erato. We found that transcripts encoding the pigment synthesis enzymes cinnabar and vermilion showed pattern- and polymorphism-related expression patterns, respectively. cinnabar expression was associated with the forewing band regardless of pigment colour, providing the first gene expression pattern known to be correlated with a major Heliconius colour pattern. In contrast, vermilion expression changed spatially over time in red-banded butterflies, but was not expressed at detectable levels in yellow-banded butterflies, suggesting that regulation of this gene may be involved with the red/yellow polymorphism. Furthermore, we found that the yellow pigment, 3-hydroxykynurenine, is incorporated into wing scales from the haemolymph rather than being synthesized in situ. We propose that some aspects of Heliconius colour patterns are determined by spatio-temporal overlap of pigment gene transcription prepatterns and speculate that evolutionary changes in vermilion regulation may in part underlie an adaptive colour pattern polymorphism.
2015-01-01
Geographical patterns in body size have been described across a wide range of species, leading to the development of a series of fundamental biological rules. However, shape variables are less well-described despite having substantial consequences for organism performance. Wing aspect ratio (AR) has been proposed as a key shape parameter that determines function in flying animals, with high AR corresponding to longer, thinner wings that promote high manoeuvrability, low speed flight, and low AR corresponding to shorter, broader wings that promote high efficiency long distance flight. From this principle it might be predicted that populations living in cooler areas would exhibit low AR wings to compensate for reduced muscle efficiency at lower temperatures. I test this hypothesis using the riverine damselfly, Calopteryx maculata, sampled from 34 sites across its range margin in North America. Nine hundred and seven male specimens were captured from across the 34 sites (mean = 26.7 ± 2.9 SE per site), dissected and measured to quantify the area and length of all four wings. Geometric morphometrics were employed to investigate geographical variation in wing shape. The majority of variation in wing shape involved changes in wing aspect ratio, confirmed independently by geometric morphometrics and wing measurements. There was a strong negative relationship between wing aspect ratio and the maximum temperature of the warmest month which varies from west-east in North America, creating a positive relationship with longitude. This pattern suggests that higher aspect ratio may be associated with areas in which greater flight efficiency is required: regions of lower temperatures during the flight season. I discuss my findings in light of research of the functional ecology of wing shape across vertebrate and invertebrate taxa. PMID:26336648
Rosetti, Natalia; Remis, Maria I
2018-06-06
Wing dimorphism occurs widely in insects and involves discontinuous variation in a wide variety of traits involved in fight and reproduction. In the current study, we analyzed the spatial pattern of wing dimorphism and intraspecific morphometric variation in nine natural populations of the grasshopper Dichroplus vittatus (Bruner; Orthoptera: Acrididae) in Argentina. Considerable body size differences among populations, between sexes and wing morphs were detected. As a general trend, females were larger than males and macropterous individuals showed increased thorax length over brachypterous which can be explained by the morphological requirements for the development of flight muscles in the thoracic cavity favoring dispersal. Moreover, when comparing wing morphs, a higher phenotypic variability was detected in macropterous females. The frequency of macropterous individuals showed negative correlation with longitude and positive with precipitations, indicating that the macropterous morph is more frequent in the humid eastern part of the studied area. Our results provide valuable about spatial variation of fully winged morph and revealed geographic areas in which the species would experience greater dispersal capacity.
Wnt signaling underlies evolution and development of the butterfly wing pattern symmetry systems.
Martin, Arnaud; Reed, Robert D
2014-11-15
Most butterfly wing patterns are proposed to be derived from a set of conserved pattern elements known as symmetry systems. Symmetry systems are so-named because they are often associated with parallel color stripes mirrored around linear organizing centers that run between the anterior and posterior wing margins. Even though the symmetry systems are the most prominent and diverse wing pattern elements, their study has been confounded by a lack of knowledge regarding the molecular basis of their development, as well as the difficulty of drawing pattern homologies across species with highly derived wing patterns. Here we present the first molecular characterization of symmetry system development by showing that WntA expression is consistently associated with the major basal, discal, central, and external symmetry system patterns of nymphalid butterflies. Pharmacological manipulations of signaling gradients using heparin and dextran sulfate showed that pattern organizing centers correspond precisely with WntA, wingless, Wnt6, and Wnt10 expression patterns, thus suggesting a role for Wnt signaling in color pattern induction. Importantly, this model is supported by recent genetic and population genomic work identifying WntA as the causative locus underlying wing pattern variation within several butterfly species. By comparing the expression of WntA between nymphalid butterflies representing a range of prototypical symmetry systems, slightly deviated symmetry systems, and highly derived wing patterns, we were able to infer symmetry system homologies in several challenging cases. Our work illustrates how highly divergent morphologies can be derived from modifications to a common ground plan across both micro- and macro-evolutionary time scales. Copyright © 2014 Elsevier Inc. All rights reserved.
Optical diffraction by the microstructure of the wing of a moth
NASA Astrophysics Data System (ADS)
Brink, D. J.; Smit, J. E.; Lee, M. E.; Möller, A.
1995-09-01
On the wing of the moth Trichoplusia orichalcea a prominent, apparently highly reflective, golden spot can be seen. Scales from this area of the wing exhibit a regular microstructure resembling a submicrometer herringbone pattern. We show that a diffraction process from this structure is responsible for the observed optical properties, such as directionality, brightness variations, polarization, and color.
Federal Register 2010, 2011, 2012, 2013, 2014
2011-08-10
... 1993, p. 134). Wings of males are blue above (dorsally), with a narrow black outer border and white... Krizek 1984, p. 112). Seasonal wing pattern variation may be caused by changes in humidity, temperature... the wings (Minno and Emmel 1994, p. 647). The Miami blue can be distinguished from the ceraunus blue...
Federal Register 2010, 2011, 2012, 2013, 2014
2012-04-06
.... 134). Wings of males are blue above (dorsally), with a narrow black outer border and white fringes... Krizek 1984, p. 112). Seasonal wing pattern variation may be caused by changes in humidity, temperature... on the undersides of the wings (Minno and Emmel 1994, p. 647). The Miami blue can be distinguished...
Torquato, Libéria Souza; Mattos, Daniel; Matta, Bruna Palma; Bitner-Mathé, Blanche Christine
2014-12-01
Organ shape evolves through cross-generational changes in developmental patterns at cellular and/or tissue levels that ultimately alter tissue dimensions and final adult proportions. Here, we investigated the cellular basis of an artificially selected divergence in the outline shape of Drosophila melanogaster wings, by comparing flies with elongated or rounded wing shapes but with remarkably similar wing sizes. We also tested whether cellular plasticity in response to developmental temperature was altered by such selection. Results show that variation in cellular traits is associated with wing shape differences, and that cell number may play an important role in wing shape response to selection. Regarding the effects of developmental temperature, a size-related plastic response was observed, in that flies reared at 16 °C developed larger wings with larger and more numerous cells across all intervein regions relative to flies reared at 25 °C. Nevertheless, no conclusive indication of altered phenotypic plasticity was found between selection strains for any wing or cellular trait. We also described how cell area is distributed across different intervein regions. It follows that cell area tends to decrease along the anterior wing compartment and increase along the posterior one. Remarkably, such pattern was observed not only in the selected strains but also in the natural baseline population, suggesting that it might be canalized during development and was not altered by the intense program of artificial selection for divergent wing shapes.
Neighboring genes shaping a single adaptive mimetic trait.
Pardo-Diaz, Carolina; Jiggins, Chris D
2014-01-01
The colorful wing patterns of Heliconius butterflies represent an excellent system in which to study the genetic and developmental control of adaptation and convergence. Using qRT-PCR and in situ hybridization on developing wings of the co-mimic species Heliconius melpomene and Heliconius erato, we have profiled the expression of three candidate genes located in the genomic locus controlling red color pattern variation. We found convergent domains of gene expression in H. melpomene and H. erato associated with red wing elements in the two genes optix and kinesin. During early pupal development of both species, the expression of optix perfectly associated with all red pattern elements whereas that of kinesin was specifically correlated with the presence of the red forewing band. These results provide evidence for the use of these two tightly linked patterning genes, acting together to create convergent wing phenotypes in Heliconius and constituting a hotspot of adaptation. © 2013 Wiley Periodicals, Inc.
Cavicchi, Sandro; Guerra, Daniela; Giorgi, Gianfranco; Pezzoli, Cristina
1985-01-01
The effects of environmental temperature on wing size and shape of Drosophila melanogaster were analyzed in populations derived from an Oregon laboratory strain kept at three temperatures (18°, 25°, 28°) for 4 yr. Temperature-directed selection was identified for both wing size and shape. The length of the four longitudinal veins, used as a test for wing size variations in the different populations, appears to be affected by both genetic and maternal influences. Vein expression appears to be dependent upon developmental pattern of the wing: veins belonging to the same compartment are coordinated in their expression and relative position, whereas veins belonging to different compartments are not. Both wing and cell areas show genetic divergence, particularly in the posterior compartment. Cell number seems to compensate for cell size variations. Such compensation is carried out both at the level of single organisms and at the level of population as a whole. The two compartments behave as individual units of selection. PMID:17246257
Chouteau, Mathieu; Whibley, Annabel; Joron, Mathieu; Llaurens, Violaine
2016-01-01
Identifying the genetic basis of adaptive variation is challenging in non-model organisms and quantitative real time PCR. is a useful tool for validating predictions regarding the expression of candidate genes. However, comparing expression levels in different conditions requires rigorous experimental design and statistical analyses. Here, we focused on the neotropical passion-vine butterflies Heliconius, non-model species studied in evolutionary biology for their adaptive variation in wing color patterns involved in mimicry and in the signaling of their toxicity to predators. We aimed at selecting stable reference genes to be used for normalization of gene expression data in RT-qPCR analyses from developing wing discs according to the minimal guidelines described in Minimum Information for publication of Quantitative Real-Time PCR Experiments (MIQE). To design internal RT-qPCR controls, we studied the stability of expression of nine candidate reference genes (actin, annexin, eF1α, FK506BP, PolyABP, PolyUBQ, RpL3, RPS3A, and tubulin) at two developmental stages (prepupal and pupal) using three widely used programs (GeNorm, NormFinder and BestKeeper). Results showed that, despite differences in statistical methods, genes RpL3, eF1α, polyABP, and annexin were stably expressed in wing discs in late larval and pupal stages of Heliconius numata. This combination of genes may be used as a reference for a reliable study of differential expression in wings for instance for genes involved in important phenotypic variation, such as wing color pattern variation. Through this example, we provide general useful technical recommendations as well as relevant statistical strategies for evolutionary biologists aiming to identify candidate-genes involved adaptive variation in non-model organisms. PMID:27271971
Márquez, E J; Saldamando-Benjumea, C I
2013-09-01
Habitat change in Rhodnius spp may represent an environmental challenge for the development of the species, particularly when feeding frequency and population density vary in nature. To estimate the effect of these variables in stability on development, the degree of directional asymmetry (DA) and fluctuating asymmetry (FA) in the wing size and shape of R. prolixus and R. robustus-like were measured under laboratory controlled conditions. DA and FA in wing size and shape were significant in both species, but their variation patterns showed both inter-specific and sexual dimorphic differences in FA of wing size and shape induced by nutrition stress. These results suggest different abilities of the genotypes and sexes of two sylvatic and domestic genotypes of Rhodnius to buffer these stress conditions. However, both species showed non-significant differences in the levels of FA between treatments that simulated sylvan vs domestic conditions, indicating that the developmental noise did not explain the variation in wing size and shape found in previous studies. Thus, this result confirm that the variation in wing size and shape in response to treatments constitute a plastic response of these genotypes to population density and feeding frequency.
Okada, Hirokazu; Ebhardt, H Alexander; Vonesch, Sibylle Chantal; Aebersold, Ruedi; Hafen, Ernst
2016-09-01
The manner by which genetic diversity within a population generates individual phenotypes is a fundamental question of biology. To advance the understanding of the genotype-phenotype relationships towards the level of biochemical processes, we perform a proteome-wide association study (PWAS) of a complex quantitative phenotype. We quantify the variation of wing imaginal disc proteomes in Drosophila genetic reference panel (DGRP) lines using SWATH mass spectrometry. In spite of the very large genetic variation (1/36 bp) between the lines, proteome variability is surprisingly small, indicating strong molecular resilience of protein expression patterns. Proteins associated with adult wing size form tight co-variation clusters that are enriched in fundamental biochemical processes. Wing size correlates with some basic metabolic functions, positively with glucose metabolism but negatively with mitochondrial respiration and not with ribosome biogenesis. Our study highlights the power of PWAS to filter functional variants from the large genetic variability in natural populations.
Extensive transcriptional response associated with seasonal plasticity of butterfly wing patterns.
Daniels, Emily V; Murad, Rabi; Mortazavi, Ali; Reed, Robert D
2014-12-01
In the eastern United States, the buckeye butterfly, Junonia coenia, shows seasonal wing colour plasticity where adults emerging in the spring are tan, while those emerging in the autumn are dark red. This variation can be artificially induced in laboratory colonies, thus making J. coenia a useful model system to examine the mechanistic basis of plasticity. To better understand the developmental basis of seasonal plasticity, we used RNA-seq to quantify transcription profiles associated with development of alternative seasonal wing morphs. Depending on the developmental stage, between 547 and 1420 transfrags were significantly differentially expressed between morphs. These extensive differences in gene expression stand in contrast to the much smaller numbers of differentially expressed transcripts identified in previous studies of genetic wing pattern variation in other species and suggest that environmentally induced phenotypic shifts arise from very broad systemic processes. Analyses of candidate endocrine and pigmentation transcripts revealed notable genes upregulated in the red morph, including several ecdysone-associated genes, and cinnabar, an ommochrome pigmentation gene implicated in colour pattern variation in other butterflies. We also found multiple melanin-related transcripts strongly upregulated in the red morph, including tan and yellow-family genes, leading us to speculate that dark red pigmentation in autumn J. coenia may involve nonommochrome pigments. While we identified several endocrine and pigmentation genes as obvious candidates for seasonal colour morph differentiation, we speculate that the majority of observed expression differences were due to thermal stress response. The buckeye transcriptome provides a basis for further developmental studies of phenotypic plasticity. © 2014 John Wiley & Sons Ltd.
Morphological outcomes of gynandromorphism in Lycaeides butterflies (Lepidoptera: Lycaenidae).
Jahner, Joshua P; Lucas, Lauren K; Wilson, Joseph S; Forister, Matthew L
2015-01-01
The genitalia of male insects have been widely used in taxonomic identification and systematics and are potentially involved in maintaining reproductive isolation between species. Although sexual selection has been invoked to explain patterns of morphological variation in genitalia among populations and species, developmental plasticity in genitalia likely contributes to observed variation but has been rarely examined, particularly in wild populations. Bilateral gynandromorphs are individuals that are genetically male on one side of the midline and genetically female on the other, while mosaic gynandromorphs have only a portion of their body developing as the opposite sex. Gynandromorphs might offer unique insights into developmental plasticity because individuals experience abnormal cellular interactions at the genitalic midline. In this study, we compare the genitalia and wing patterns of gynandromorphic Anna and Melissa blue butterflies, Lycaeides anna (Edwards) (formerly L. idas anna) and L. melissa (Edwards) (Lepidoptera: Lycaenidae), to the morphology of normal individuals from the same populations. Gynandromorph wing markings all fell within the range of variation of normal butterflies; however, a number of genitalic measurements were outliers when compared with normal individuals. From these results, we conclude that the gynandromorphs' genitalia, but not wing patterns, can be abnormal when compared with normal individuals and that the gynandromorphic genitalia do not deviate developmentally in a consistent pattern across individuals. Finally, genetic mechanisms are considered for the development of gynandromorphism in Lycaeides butterflies. © The Author 2015. Published by Oxford University Press on behalf of the Entomological Society of America.
Wing morphology variations in a natural population of Phlebotomus tobbi Adler and Theodor 1930.
Oguz, Gizem; Kasap, Ozge Erisoz; Alten, Bulent
2017-12-01
Cutaneous leishmaniasis (CL) is highly endemic in the Cukurova region, located on the crossroads of main refugee routes from the Middle East to Europe on the eastern Mediterranean part of Turkey. Our purpose was to investigate the phenotypic variation of Phlebotomus tobbi, the known vector of CL in the region, during one active season. Sand flies and microclimatic data were collected monthly from May to October, 2011, from five locations in six villages in the study area. A geometric morphometric approach was used to investigate wing morphology. Shape analyses revealed that males collected in May and June comprised one group, while specimens collected in August, September, and October formed a second group. Specimens from July were found to be distributed within these two groups. A similar distribution pattern was observed for females, but specimens from October were represented as the third district group. Significant size variation was detected for both sexes between months. Wing size and temperature were negatively correlated for females, but there was no temperature effect for males. Wing size of both sexes was increased in correlation to increasing relative humidity. Males were found to have smaller wings with increasing population density. © 2017 The Society for Vector Ecology.
Color pattern analysis of nymphalid butterfly wings: revision of the nymphalid groundplan.
Otaki, Joji M
2012-09-01
To better understand the developmental mechanisms of color pattern variation in butterfly wings, it is important to construct an accurate representation of pattern elements, known as the "nymphalid groundplan". However, some aspects of the current groundplan remain elusive. Here, I examined wing-wide elemental patterns of various nymphalid butterflies and confirmed that wing-wide color patterns are composed of the border, central, and basal symmetry systems. The central and basal symmetry systems can express circular patterns resembling eyespots, indicating that these systems have developmental mechanisms similar to those of the border symmetry system. The wing root band commonly occurs as a distinct symmetry system independent from the basal symmetry system. In addition, the marginal and submarginal bands are likely generated as a single system, referred to as the "marginal band system". Background spaces between two symmetry systems are sometimes light in coloration and can produce white bands, contributing significantly to color pattern diversity. When an element is enlarged with a pale central area, a visually similar (yet developmentally distinct) white band is produced. Based on the symmetric relationships of elements, I propose that both the central and border symmetry systems are comprised of "core elements" (the discal spot and the border ocelli, respectively) and a pair of "paracore elements" (the distal and proximal bands and the parafocal elements, respectively). Both core and paracore elements can be doubled, or outlined. Developmentally, this system configuration is consistent with the induction model, but not with the concentration gradient model for positional information.
Sánchez Herrera, Melissa; Kuhn, William R; Lorenzo-Carballa, Maria Olalla; Harding, Kathleen M; Ankrom, Nikole; Sherratt, Thomas N; Hoffmann, Joachim; Van Gossum, Hans; Ware, Jessica L; Cordero-Rivera, Adolfo; Beatty, Christopher D
2015-01-01
The study of color polymorphisms (CP) has provided profound insights into the maintenance of genetic variation in natural populations. We here offer the first evidence for an elaborate wing polymorphism in the Neotropical damselfly genus Polythore, which consists of 21 described species, distributed along the eastern slopes of the Andes in South America. These damselflies display highly complex wing colors and patterning, incorporating black, white, yellow, and orange in multiple wing bands. Wing colors, along with some components of the male genitalia, have been the primary characters used in species description; few other morphological traits vary within the group, and so there are few useful diagnostic characters. Previous research has indicated the possibility of a cryptic species existing in P. procera in Colombia, despite there being no significant differences in wing color and pattern between the populations of the two putative species. Here we analyze the complexity and diversity of wing color patterns of individuals from five described Polythore species in the Central Amazon Basin of Peru using a novel suite of morphological analyses to quantify wing color and pattern: geometric morphometrics, chromaticity analysis, and Gabor wavelet transformation. We then test whether these color patterns are good predictors of species by recovering the phylogenetic relationships among the 5 species using the barcode gene (COI). Our results suggest that, while highly distinct and discrete wing patterns exist in Polythore, these "wingforms" do not represent monophyletic clades in the recovered topology. The wingforms identified as P. victoria and P. ornata are both involved in a polymorphism with P. neopicta; also, cryptic speciation may have taking place among individuals with the P. victoria wingform. Only P. aurora and P. spateri represent monophyletic species with a single wingform in our molecular phylogeny. We discuss the implications of this polymorphism, and the potential evolutionary mechanisms that could maintain it.
Harding, Kathleen M.; Ankrom, Nikole; Sherratt, Thomas N.; Hoffmann, Joachim; Van Gossum, Hans; Ware, Jessica L.; Cordero-Rivera, Adolfo
2015-01-01
The study of color polymorphisms (CP) has provided profound insights into the maintenance of genetic variation in natural populations. We here offer the first evidence for an elaborate wing polymorphism in the Neotropical damselfly genus Polythore, which consists of 21 described species, distributed along the eastern slopes of the Andes in South America. These damselflies display highly complex wing colors and patterning, incorporating black, white, yellow, and orange in multiple wing bands. Wing colors, along with some components of the male genitalia, have been the primary characters used in species description; few other morphological traits vary within the group, and so there are few useful diagnostic characters. Previous research has indicated the possibility of a cryptic species existing in P. procera in Colombia, despite there being no significant differences in wing color and pattern between the populations of the two putative species. Here we analyze the complexity and diversity of wing color patterns of individuals from five described Polythore species in the Central Amazon Basin of Peru using a novel suite of morphological analyses to quantify wing color and pattern: geometric morphometrics, chromaticity analysis, and Gabor wavelet transformation. We then test whether these color patterns are good predictors of species by recovering the phylogenetic relationships among the 5 species using the barcode gene (COI). Our results suggest that, while highly distinct and discrete wing patterns exist in Polythore, these “wingforms” do not represent monophyletic clades in the recovered topology. The wingforms identified as P. victoria and P. ornata are both involved in a polymorphism with P. neopicta; also, cryptic speciation may have taking place among individuals with the P. victoria wingform. Only P. aurora and P. spateri represent monophyletic species with a single wingform in our molecular phylogeny. We discuss the implications of this polymorphism, and the potential evolutionary mechanisms that could maintain it. PMID:25923455
de Camargo, Nícholas Ferreira; Corrêa, Danilo do Carmo Vieira; de Camargo, Amabílio J. Aires; Diniz, Ivone Rezende
2015-01-01
Sexual dimorphism is a pronounced pattern of intraspecific variation in Lepidoptera. However, moths of the family Sphingidae (Lepidoptera: Bombycoidea) are considered exceptions to this rule. We used geometric morphometric techniques to detect shape and size sexual dimorphism in the fore and hindwings of seven hawkmoth species. The shape variables produced were then subjected to a discriminant analysis. The allometric effects were measured with a simple regression between the canonical variables and the centroid size. We also used the normalized residuals to assess the nonallometric component of shape variation with a t-test. The deformations in wing shape between sexes per species were assessed with a regression between the nonreduced shape variables and the residuals. We found sexual dimorphism in both wings in all analyzed species, and that the allometric effects were responsible for much of the wing shape variation between the sexes. However, when we removed the size effects, we observed shape sexual dimorphism. It is very common for females to be larger than males in Lepidoptera, so it is expected that the shape of structures such as wings suffers deformations in order to preserve their function. However, sources of variation other than allometry could be a reflection of different reproductive flight behavior (long flights in search for sexual mates in males, and flight in search for host plants in females). PMID:26206895
NASA Technical Reports Server (NTRS)
Razak, K.
1980-01-01
The question of the effect of distribution and magnitude of spanwise circulation and shed vorticity from an airplane wing on the distribution pattern of agricultural products distributed from an airplane was studied. The first step in an analysis of this question is the determination of the actual distribution of lift along an airplane wing, from which the pattern of shed vorticity can be determined. A procedure is developed to calculate the span loading for flapped and unflapped wings of arbitrary aspect ratio and taper ratio. The procedure was programmed on a small programmable calculator, the Hewlett Packard HP-97, and also was programmed in BASIC language. They could be used to explore the variations in span loading that can be secured by variable flap deflections or the effect of flying at varying air speeds at different airplane gross weights. Either an absolute evaluation of span loading can be secured or comparative span loading can be evaluated to determine their effect on swath width and swath distribution pattern. The programs are intended to assist the user in evaluating the effect of a given spanload distribution.
Chazot, Nicolas; Panara, Stephen; Zilbermann, Nicolas; Blandin, Patrick; Le Poul, Yann; Cornette, Raphaël; Elias, Marianne; Debat, Vincent
2016-01-01
Butterfly wings harbor highly diverse phenotypes and are involved in many functions. Wing size and shape result from interactions between adaptive processes, phylogenetic history, and developmental constraints, which are complex to disentangle. Here, we focus on the genus Morpho (Nymphalidae: Satyrinae, 30 species), which presents a high diversity of sizes, shapes, and color patterns. First, we generate a comprehensive molecular phylogeny of these 30 species. Next, using 911 collection specimens, we quantify the variation of wing size and shape across species, to assess the importance of shared ancestry, microhabitat use, and sexual selection in the evolution of the wings. While accounting for phylogenetic and allometric effects, we detect a significant difference in wing shape but not size among microhabitats. Fore and hindwings covary at the individual and species levels, and the covariation differs among microhabitats. However, the microhabitat structure in covariation disappears when phylogenetic relationships are taken into account. Our results demonstrate that microhabitat has driven wing shape evolution, although it has not strongly affected forewing and hindwing integration. We also found that sexual dimorphism of forewing shape and color pattern are coupled, suggesting a common selective force. © 2015 The Author(s). Evolution © 2015 The Society for the Study of Evolution.
de Camargo, Willian Rogers Ferreira; de Camargo, Nícholas Ferreira; Corrêa, Danilo do Carmo Vieira; de Camargo, Amabílio J Aires; Diniz, Ivone Rezende
2015-01-01
Sexual dimorphism is a pronounced pattern of intraspecific variation in Lepidoptera. However, moths of the family Sphingidae (Lepidoptera: Bombycoidea) are considered exceptions to this rule. We used geometric morphometric techniques to detect shape and size sexual dimorphism in the fore and hindwings of seven hawkmoth species. The shape variables produced were then subjected to a discriminant analysis. The allometric effects were measured with a simple regression between the canonical variables and the centroid size. We also used the normalized residuals to assess the nonallometric component of shape variation with a t-test. The deformations in wing shape between sexes per species were assessed with a regression between the nonreduced shape variables and the residuals. We found sexual dimorphism in both wings in all analyzed species, and that the allometric effects were responsible for much of the wing shape variation between the sexes. However, when we removed the size effects, we observed shape sexual dimorphism. It is very common for females to be larger than males in Lepidoptera, so it is expected that the shape of structures such as wings suffers deformations in order to preserve their function. However, sources of variation other than allometry could be a reflection of different reproductive flight behavior (long flights in search for sexual mates in males, and flight in search for host plants in females). © The Author 2015. Published by Oxford University Press on behalf of the Entomological Society of America.
Evolutionary Novelty in a Butterfly Wing Pattern through Enhancer Shuffling
Pardo-Diaz, Carolina; Hanly, Joseph J.; Martin, Simon H.; Mallet, James; Dasmahapatra, Kanchon K.; Salazar, Camilo; Joron, Mathieu; Nadeau, Nicola; McMillan, W. Owen; Jiggins, Chris D.
2016-01-01
An important goal in evolutionary biology is to understand the genetic changes underlying novel morphological structures. We investigated the origins of a complex wing pattern found among Amazonian Heliconius butterflies. Genome sequence data from 142 individuals across 17 species identified narrow regions associated with two distinct red colour pattern elements, dennis and ray. We hypothesise that these modules in non-coding sequence represent distinct cis-regulatory loci that control expression of the transcription factor optix, which in turn controls red pattern variation across Heliconius. Phylogenetic analysis of the two elements demonstrated that they have distinct evolutionary histories and that novel adaptive morphological variation was created by shuffling these cis-regulatory modules through recombination between divergent lineages. In addition, recombination of modules into different combinations within species further contributes to diversity. Analysis of the timing of diversification in these two regions supports the hypothesis of introgression moving regulatory modules between species, rather than shared ancestral variation. The dennis phenotype introgressed into Heliconius melpomene at about the same time that ray originated in this group, while ray introgressed back into H. elevatus much more recently. We show that shuffling of existing enhancer elements both within and between species provides a mechanism for rapid diversification and generation of novel morphological combinations during adaptive radiation. PMID:26771987
Evolutionary Novelty in a Butterfly Wing Pattern through Enhancer Shuffling.
Wallbank, Richard W R; Baxter, Simon W; Pardo-Diaz, Carolina; Hanly, Joseph J; Martin, Simon H; Mallet, James; Dasmahapatra, Kanchon K; Salazar, Camilo; Joron, Mathieu; Nadeau, Nicola; McMillan, W Owen; Jiggins, Chris D
2016-01-01
An important goal in evolutionary biology is to understand the genetic changes underlying novel morphological structures. We investigated the origins of a complex wing pattern found among Amazonian Heliconius butterflies. Genome sequence data from 142 individuals across 17 species identified narrow regions associated with two distinct red colour pattern elements, dennis and ray. We hypothesise that these modules in non-coding sequence represent distinct cis-regulatory loci that control expression of the transcription factor optix, which in turn controls red pattern variation across Heliconius. Phylogenetic analysis of the two elements demonstrated that they have distinct evolutionary histories and that novel adaptive morphological variation was created by shuffling these cis-regulatory modules through recombination between divergent lineages. In addition, recombination of modules into different combinations within species further contributes to diversity. Analysis of the timing of diversification in these two regions supports the hypothesis of introgression moving regulatory modules between species, rather than shared ancestral variation. The dennis phenotype introgressed into Heliconius melpomene at about the same time that ray originated in this group, while ray introgressed back into H. elevatus much more recently. We show that shuffling of existing enhancer elements both within and between species provides a mechanism for rapid diversification and generation of novel morphological combinations during adaptive radiation.
Rossa, Robert; Goczał, Jakub; Tofilski, Adam
2016-01-01
We have described the morphological variation of five Western Palaearctic species of Monochamus Dejean, 1821. The variation was assessed using wing measurements. Special emphasis was placed on the differences between Monochamus sartor (F., 1787) and Monochamus urussovii (Fischer-Waldheim, 1805). There was an interesting pattern of variation between the two species. Individuals of M. sartor from the Carpathians differed markedly from individuals of M. urussovii from Siberia, but individuals from north-eastern Poland were intermediate between those two populations. The intermediate individuals were more similar to the Siberian M. urussovii than to the Carpathian M. sartor despite the relatively large geographic distance between north-eastern Poland and Siberia. The occurrence of the intermediate individuals in north-eastern Poland may be the effect of hybridization between M. urussovii and M. sartor, which might have occurred after secondary contact between the two species in the Holocene. PMID:26798141
Dynamics of F-actin prefigure the structure of butterfly wing scales.
Dinwiddie, April; Null, Ryan; Pizzano, Maria; Chuong, Lisa; Leigh Krup, Alexis; Ee Tan, Hwei; Patel, Nipam H
2014-08-15
The wings of butterflies and moths consist of dorsal and ventral epidermal surfaces that give rise to overlapping layers of scales and hairs (Lepidoptera, "scale wing"). Wing scales (average length ~200 µm) are homologous to insect bristles (macrochaetes), and their colors create the patterns that characterize lepidopteran wings. The topology and surface sculpture of wing scales vary widely, and this architectural complexity arises from variations in the developmental program of the individual scale cells of the wing epithelium. One of the more striking features of lepidopteran wing scales are the longitudinal ridges that run the length of the mature (dead) cell, gathering the cuticularized scale cell surface into pleats on the sides of each scale. While also present around the periphery of other insect bristles and hairs, longitudinal ridges in lepidopteran wing scales gain new significance for their creation of iridescent color through microribs and lamellae. Here we show the dynamics of the highly organized F-actin filaments during scale cell development, and present experimental manipulations of actin polymerization that reveal the essential role of this cytoskeletal component in wing scale elongation and the positioning of longitudinal ribs. Copyright © 2014 Elsevier Inc. All rights reserved.
Polymorphism at a mimicry supergene maintained by opposing frequency-dependent selection pressures.
Chouteau, Mathieu; Llaurens, Violaine; Piron-Prunier, Florence; Joron, Mathieu
2017-08-01
Explaining the maintenance of adaptive diversity within populations is a long-standing goal in evolutionary biology, with important implications for conservation, medicine, and agriculture. Adaptation often leads to the fixation of beneficial alleles, and therefore it erodes local diversity so that understanding the coexistence of multiple adaptive phenotypes requires deciphering the ecological mechanisms that determine their respective benefits. Here, we show how antagonistic frequency-dependent selection (FDS), generated by natural and sexual selection acting on the same trait, maintains mimicry polymorphism in the toxic butterfly Heliconius numata Positive FDS imposed by predators on mimetic signals favors the fixation of the most abundant and best-protected wing-pattern morph, thereby limiting polymorphism. However, by using mate-choice experiments, we reveal disassortative mate preferences of the different wing-pattern morphs. The resulting negative FDS on wing-pattern alleles is consistent with the excess of heterozygote genotypes at the supergene locus controlling wing-pattern variation in natural populations of H. numata The combined effect of positive and negative FDS on visual signals is sufficient to maintain a diversity of morphs displaying accurate mimicry with other local prey, although some of the forms only provide moderate protection against predators. Our findings help understand how alternative adaptive phenotypes can be maintained within populations and emphasize the need to investigate interactions between selective pressures in other cases of puzzling adaptive polymorphism.
Polymorphism at a mimicry supergene maintained by opposing frequency-dependent selection pressures
Chouteau, Mathieu; Llaurens, Violaine; Piron-Prunier, Florence; Joron, Mathieu
2017-01-01
Explaining the maintenance of adaptive diversity within populations is a long-standing goal in evolutionary biology, with important implications for conservation, medicine, and agriculture. Adaptation often leads to the fixation of beneficial alleles, and therefore it erodes local diversity so that understanding the coexistence of multiple adaptive phenotypes requires deciphering the ecological mechanisms that determine their respective benefits. Here, we show how antagonistic frequency-dependent selection (FDS), generated by natural and sexual selection acting on the same trait, maintains mimicry polymorphism in the toxic butterfly Heliconius numata. Positive FDS imposed by predators on mimetic signals favors the fixation of the most abundant and best-protected wing-pattern morph, thereby limiting polymorphism. However, by using mate-choice experiments, we reveal disassortative mate preferences of the different wing-pattern morphs. The resulting negative FDS on wing-pattern alleles is consistent with the excess of heterozygote genotypes at the supergene locus controlling wing-pattern variation in natural populations of H. numata. The combined effect of positive and negative FDS on visual signals is sufficient to maintain a diversity of morphs displaying accurate mimicry with other local prey, although some of the forms only provide moderate protection against predators. Our findings help understand how alternative adaptive phenotypes can be maintained within populations and emphasize the need to investigate interactions between selective pressures in other cases of puzzling adaptive polymorphism. PMID:28673971
Adhikari, Kiran; Otaki, Joji M
2016-02-01
It is often desirable but difficult to retrieve information on the mature phenotype of an immature tissue sample that has been subjected to gene expression analysis. This problem cannot be ignored when individual variation within a species is large. To circumvent this problem in the butterfly wing system, we developed a new surgical method for removing a single forewing from a pupa using Junonia orithya; the operated pupa was left to develop to an adult without eclosion. The removed right forewing was subjected to gene expression analysis, whereas the non-removed left forewing was examined for color patterns. As a test case, we focused on Distal-less (Dll), which likely plays an active role in inducing elemental patterns, including eyespots. The Dll expression level in forewings was paired with eyespot size data from the same individual. One third of the operated pupae survived and developed wing color patterns. Dll expression levels were significantly higher in males than in females, although male eyespots were smaller in size than female eyespots. Eyespot size data showed weak but significant correlations with the Dll expression level in females. These results demonstrate that a single-wing removal method was successfully applied to the butterfly wing system and suggest the weak and non-exclusive contribution of Dll to eyespot size determination in this butterfly. Our novel methodology for establishing correspondence between gene expression and phenotype can be applied to other candidate genes for color pattern development in butterflies. Conceptually similar methods may also be applicable in other developmental systems.
NASA Astrophysics Data System (ADS)
Pecháček, Pavel; Stella, David; Keil, Petr; Kleisner, Karel
2014-12-01
The males of the Brimstone butterfly ( Gonepteryx rhamni) have ultraviolet pattern on the dorsal surfaces of their wings. Using geometric morphometrics, we have analysed correlations between environmental variables (climate, productivity) and shape variability of the ultraviolet pattern and the forewing in 110 male specimens of G. rhamni collected in the Palaearctic zone. To start with, we subjected the environmental variables to principal component analysis (PCA). The first PCA axis (precipitation, temperature, latitude) significantly correlated with shape variation of the ultraviolet patterns across the Palaearctic. Additionally, we have performed two-block partial least squares (PLS) analysis to assess co-variation between intraspecific shape variation and the variation of 11 environmental variables. The first PLS axis explained 93 % of variability and represented the effect of precipitation, temperature and latitude. Along this axis, we observed a systematic increase in the relative area of ultraviolet colouration with increasing temperature and precipitation and decreasing latitude. We conclude that the shape variation of ultraviolet patterns on the forewings of male Brimstones is correlated with large-scale environmental factors.
A major gene controls mimicry and crypsis in butterflies and moths
Nadeau, Nicola J.; Pardo-Diaz, Carolina; Whibley, Annabel; Supple, Megan; Saenko, Suzanne V.; Wallbank, Richard W. R.; Wu, Grace C.; Maroja, Luana; Ferguson, Laura; Hanly, Joseph J.; Hines, Heather; Salazar, Camilo; Merrill, Richard; Dowling, Andrea; ffrench-Constant, Richard; Llaurens, Violaine; Joron, Mathieu; McMillan, W. Owen; Jiggins, Chris D.
2016-01-01
The wing patterns of butterflies and moths (Lepidoptera) are diverse and striking examples of evolutionary diversification by natural selection1,2. Lepidopteran wing colour patterns are a key innovation, consisting of arrays of coloured scales. We still lack a general understanding of how these patterns are controlled and if there is any commonality across the 160,000 moth and 17,000 butterfly species. Here, we identify a gene, cortex, through fine-scale mapping using population genomics and gene expression analyses, which regulates pattern switches in multiple species across the mimetic radiation in Heliconius butterflies. cortex belongs to a fast evolving subfamily of the otherwise highly conserved fizzy family of cell cycle regulators3, suggesting that it most likely regulates pigmentation patterning through regulation of scale cell development. In parallel with findings in the peppered moth (Biston betularia)4, our results suggest that this mechanism is common within Lepidoptera and that cortex has become a major target for natural selection acting on colour and pattern variation in this group of insects. PMID:27251285
The gene cortex controls mimicry and crypsis in butterflies and moths.
Nadeau, Nicola J; Pardo-Diaz, Carolina; Whibley, Annabel; Supple, Megan A; Saenko, Suzanne V; Wallbank, Richard W R; Wu, Grace C; Maroja, Luana; Ferguson, Laura; Hanly, Joseph J; Hines, Heather; Salazar, Camilo; Merrill, Richard M; Dowling, Andrea J; ffrench-Constant, Richard H; Llaurens, Violaine; Joron, Mathieu; McMillan, W Owen; Jiggins, Chris D
2016-06-02
The wing patterns of butterflies and moths (Lepidoptera) are diverse and striking examples of evolutionary diversification by natural selection. Lepidopteran wing colour patterns are a key innovation, consisting of arrays of coloured scales. We still lack a general understanding of how these patterns are controlled and whether this control shows any commonality across the 160,000 moth and 17,000 butterfly species. Here, we use fine-scale mapping with population genomics and gene expression analyses to identify a gene, cortex, that regulates pattern switches in multiple species across the mimetic radiation in Heliconius butterflies. cortex belongs to a fast-evolving subfamily of the otherwise highly conserved fizzy family of cell-cycle regulators, suggesting that it probably regulates pigmentation patterning by regulating scale cell development. In parallel with findings in the peppered moth (Biston betularia), our results suggest that this mechanism is common within Lepidoptera and that cortex has become a major target for natural selection acting on colour and pattern variation in this group of insects.
Environmental predictors of dispersal traits across a species' geographic range.
LaRue, Elizabeth A; Holland, Jeffrey D; Emery, Nancy C
2018-05-30
Variation in habitat quality and quantity drive selection on dispersal traits in heterogeneous environments, but the extent to which environmental conditions predict geographic variation in dispersal is rarely evaluated. We assessed dispersal trait variation across the range of Cakile edentula var. lacustris, an annual herb that occupies beaches of the Great Lakes. Cakile edentula has dimorphic fruits that each contain one dispersive and one non-dispersive seed. Previous work showed that plant height, branching density, and dispersive fruit wing-loading can determine the distance that seeds disperse locally by wind, while pericarp thickness influences the distance they disperse by water. We tested if these traits vary predictably with latitude across the species' geographic range, and if variation in dispersal characteristics can be predicted by the quality and quantity of habitat available at a site. We observed that the dispersive fruits from northern and southern populations had thinner pericarps than those from the interior of the species' range, reflecting reduced long-distance dispersal by water at both range limits. Plants at the northern range limit were shorter with less dense branching and lower wing-loading than populations elsewhere in the range, suggesting that these populations have enhanced local wind dispersal. In contrast, southern populations exhibited traits with inconsistent effects on wind dispersal: plants tended to be short, which facilitates wind dispersal in C. edentula, but also had relatively higher branching density and distal segment wing-loading that reduce wind dispersal. Geographic variation in maternal plant height and branching density was partially explained by variation in habitat quality, which declined at the species' range limits. In addition, population differences in branching density, fruit wing-loading, and pericarp thickness were predicted by the abundance and distribution of beach habitat. Finally, a common garden analysis recovered latitudinal patterns for the dispersal traits associated with fruits, but not those associated with maternal architecture. Thus, the geographic patterns of dispersal trait variation that we observed likely reflect responses to past selection by the distribution, abundance, and quality of habitat, strong plasticity in dispersal traits, and the effects dispersal itself has in shaping local adaptation by driving gene flow among populations. This article is protected by copyright. All rights reserved. This article is protected by copyright. All rights reserved.
NASA Astrophysics Data System (ADS)
Seino, M.; Kakazu, Y.
The vein and cell patterns for the fore and hind wing of Lepidoptera, Hemiptera, Orthoptera and Odonata are analyzed and discussed. For vein patterns of them, the fractal properties are shown and the inequality between four orders is obtained. The nature of wings observed by mass distributions for fractal dimensions of the vein pattern is presented.
Stable structural color patterns displayed on transparent insect wings.
Shevtsova, Ekaterina; Hansson, Christer; Janzen, Daniel H; Kjærandsen, Jostein
2011-01-11
Color patterns play central roles in the behavior of insects, and are important traits for taxonomic studies. Here we report striking and stable structural color patterns--wing interference patterns (WIPs)--in the transparent wings of small Hymenoptera and Diptera, patterns that have been largely overlooked by biologists. These extremely thin wings reflect vivid color patterns caused by thin film interference. The visibility of these patterns is affected by the way the insects display their wings against various backgrounds with different light properties. The specific color sequence displayed lacks pure red and matches the color vision of most insects, strongly suggesting that the biological significance of WIPs lies in visual signaling. Taxon-specific color patterns are formed by uneven membrane thickness, pigmentation, venation, and hair placement. The optically refracted pattern is also stabilized by microstructures of the wing such as membrane corrugations and spherical cell structures that reinforce the pattern and make it essentially noniridescent over a large range of light incidences. WIPs can be applied to map the micromorphology of wings through direct observation and are useful in several fields of biology. We demonstrate their usefulness as identification patterns to solve cases of cryptic species complexes in tiny parasitic wasps, and indicate their potentials for research on the genetic control of wing development through direct links between the transregulatory wing landscape and interference patterns we observe in Drosophila model species. Some species display sexually dimorphic WIPs, suggesting sexual selection as one of the driving forces for their evolution.
Stable structural color patterns displayed on transparent insect wings
Shevtsova, Ekaterina; Hansson, Christer; Janzen, Daniel H.; Kjærandsen, Jostein
2011-01-01
Color patterns play central roles in the behavior of insects, and are important traits for taxonomic studies. Here we report striking and stable structural color patterns—wing interference patterns (WIPs)—in the transparent wings of small Hymenoptera and Diptera, patterns that have been largely overlooked by biologists. These extremely thin wings reflect vivid color patterns caused by thin film interference. The visibility of these patterns is affected by the way the insects display their wings against various backgrounds with different light properties. The specific color sequence displayed lacks pure red and matches the color vision of most insects, strongly suggesting that the biological significance of WIPs lies in visual signaling. Taxon-specific color patterns are formed by uneven membrane thickness, pigmentation, venation, and hair placement. The optically refracted pattern is also stabilized by microstructures of the wing such as membrane corrugations and spherical cell structures that reinforce the pattern and make it essentially noniridescent over a large range of light incidences. WIPs can be applied to map the micromorphology of wings through direct observation and are useful in several fields of biology. We demonstrate their usefulness as identification patterns to solve cases of cryptic species complexes in tiny parasitic wasps, and indicate their potentials for research on the genetic control of wing development through direct links between the transregulatory wing landscape and interference patterns we observe in Drosophila model species. Some species display sexually dimorphic WIPs, suggesting sexual selection as one of the driving forces for their evolution. PMID:21199954
Schmieder, Daniela A.; Benítez, Hugo A.; Borissov, Ivailo M.; Fruciano, Carmelo
2015-01-01
External morphology is commonly used to identify bats as well as to investigate flight and foraging behavior, typically relying on simple length and area measures or ratios. However, geometric morphometrics is increasingly used in the biological sciences to analyse variation in shape and discriminate among species and populations. Here we compare the ability of traditional versus geometric morphometric methods in discriminating between closely related bat species – in this case European horseshoe bats (Rhinolophidae, Chiroptera) – based on morphology of the wing, body and tail. In addition to comparing morphometric methods, we used geometric morphometrics to detect interspecies differences as shape changes. Geometric morphometrics yielded improved species discrimination relative to traditional methods. The predicted shape for the variation along the between group principal components revealed that the largest differences between species lay in the extent to which the wing reaches in the direction of the head. This strong trend in interspecific shape variation is associated with size, which we interpret as an evolutionary allometry pattern. PMID:25965335
Comparative genomics of the mimicry switch in Papilio dardanus.
Timmermans, Martijn J T N; Baxter, Simon W; Clark, Rebecca; Heckel, David G; Vogel, Heiko; Collins, Steve; Papanicolaou, Alexie; Fukova, Iva; Joron, Mathieu; Thompson, Martin J; Jiggins, Chris D; ffrench-Constant, Richard H; Vogler, Alfried P
2014-07-22
The African Mocker Swallowtail, Papilio dardanus, is a textbook example in evolutionary genetics. Classical breeding experiments have shown that wing pattern variation in this polymorphic Batesian mimic is determined by the polyallelic H locus that controls a set of distinct mimetic phenotypes. Using bacterial artificial chromosome (BAC) sequencing, recombination analyses and comparative genomics, we show that H co-segregates with an interval of less than 500 kb that is collinear with two other Lepidoptera genomes and contains 24 genes, including the transcription factor genes engrailed (en) and invected (inv). H is located in a region of conserved gene order, which argues against any role for genomic translocations in the evolution of a hypothesized multi-gene mimicry locus. Natural populations of P. dardanus show significant associations of specific morphs with single nucleotide polymorphisms (SNPs), centred on en. In addition, SNP variation in the H region reveals evidence of non-neutral molecular evolution in the en gene alone. We find evidence for a duplication potentially driving physical constraints on recombination in the lamborni morph. Absence of perfect linkage disequilibrium between different genes in the other morphs suggests that H is limited to nucleotide positions in the regulatory and coding regions of en. Our results therefore support the hypothesis that a single gene underlies wing pattern variation in P. dardanus.
The impact of circulation control on rotary aircraft controls systems
NASA Technical Reports Server (NTRS)
Kingloff, R. F.; Cooper, D. E.
1987-01-01
Application of circulation to rotary wing systems is a new development. Efforts to determine the near and far field flow patterns and to analytically predict those flow patterns have been underway for some years. Rotary wing applications present a new set of challenges in circulation control technology. Rotary wing sections must accommodate substantial Mach number, free stream dynamic pressure and section angle of attack variation at each flight condition within the design envelope. They must also be capable of short term circulation blowing modulation to produce control moments and vibration alleviation in addition to a lift augmentation function. Control system design must provide this primary control moment, vibration alleviation and lift augmentation function. To accomplish this, one must simultaneously control the compressed air source and its distribution. The control law algorithm must therefore address the compressor as the air source, the plenum as the air pressure storage and the pneumatic flow gates or valves that distribute and meter the stored pressure to the rotating blades. Also, mechanical collective blade pitch, rotor shaft angle of attack and engine power control must be maintained.
Development and design of flexible Fowler flaps for an adaptive wing
NASA Astrophysics Data System (ADS)
Monner, Hans P.; Hanselka, Holger; Breitbach, Elmar J.
1998-06-01
Civil transport airplanes fly with fixed geometry wings optimized only for one design point described by altitude, Mach number and airplane weight. These parameters vary continuously during flight, to which means the wing geometry seldom is optimal. According to aerodynamic investigations a chordwide variation of the wing camber leads to improvements in operational flexibility, buffet boundaries and performance resulting in reduction of fuel consumption. A spanwise differential camber variation allows to gain control over spanwise lift distributions reducing wing root bending moments. This paper describes the design of flexible Fowler flaps for an adaptive wing to be used in civil transport aircraft that allows both a chordwise as well as spanwise differential camber variation during flight. Since both lower and upper skins are flexed by active ribs, the camber variation is achieved with a smooth contour and without any additional gaps.
Connahs, Heidi; Rhen, Turk; Simmons, Rebecca B
2016-03-31
Butterfly wing color patterns are an important model system for understanding the evolution and development of morphological diversity and animal pigmentation. Wing color patterns develop from a complex network composed of highly conserved patterning genes and pigmentation pathways. Patterning genes are involved in regulating pigment synthesis however the temporal expression dynamics of these interacting networks is poorly understood. Here, we employ next generation sequencing to examine expression patterns of the gene network underlying wing development in the nymphalid butterfly, Vanessa cardui. We identified 9, 376 differentially expressed transcripts during wing color pattern development, including genes involved in patterning, pigmentation and gene regulation. Differential expression of these genes was highest at the pre-ommochrome stage compared to early pupal and late melanin stages. Overall, an increasing number of genes were down-regulated during the progression of wing development. We observed dynamic expression patterns of a large number of pigment genes from the ommochrome, melanin and also pteridine pathways, including contrasting patterns of expression for paralogs of the yellow gene family. Surprisingly, many patterning genes previously associated with butterfly pattern elements were not significantly up-regulated at any time during pupation, although many other transcription factors were differentially expressed. Several genes involved in Notch signaling were significantly up-regulated during the pre-ommochrome stage including slow border cells, bunched and pebbles; the function of these genes in the development of butterfly wings is currently unknown. Many genes involved in ecdysone signaling were also significantly up-regulated during early pupal and late melanin stages and exhibited opposing patterns of expression relative to the ecdysone receptor. Finally, a comparison across four butterfly transcriptomes revealed 28 transcripts common to all four species that have no known homologs in other metazoans. This study provides a comprehensive list of differentially expressed transcripts during wing development, revealing potential candidate genes that may be involved in regulating butterfly wing patterns. Some differentially expressed genes have no known homologs possibly representing genes unique to butterflies. Results from this study also indicate that development of nymphalid wing patterns may arise not only from melanin and ommochrome pigments but also the pteridine pigment pathway.
Jiggins, Chris D; Wallbank, Richard W R; Hanly, Joseph J
2017-02-05
A major challenge is to understand how conserved gene regulatory networks control the wonderful diversity of form that we see among animals and plants. Butterfly wing patterns are an excellent example of this diversity. Butterfly wings form as imaginal discs in the caterpillar and are constructed by a gene regulatory network, much of which is conserved across the holometabolous insects. Recent work in Heliconius butterflies takes advantage of genomic approaches and offers insights into how the diversification of wing patterns is overlaid onto this conserved network. WntA is a patterning morphogen that alters spatial information in the wing. Optix is a transcription factor that acts later in development to paint specific wing regions red. Both of these loci fit the paradigm of conserved protein-coding loci with diverse regulatory elements and developmental roles that have taken on novel derived functions in patterning wings. These discoveries offer insights into the 'Nymphalid Ground Plan', which offers a unifying hypothesis for pattern formation across nymphalid butterflies. These loci also represent 'hotspots' for morphological change that have been targeted repeatedly during evolution. Both convergent and divergent evolution of a great diversity of patterns is controlled by complex alleles at just a few genes. We suggest that evolutionary change has become focused on one or a few genetic loci for two reasons. First, pre-existing complex cis-regulatory loci that already interact with potentially relevant transcription factors are more likely to acquire novel functions in wing patterning. Second, the shape of wing regulatory networks may constrain evolutionary change to one or a few loci. Overall, genomic approaches that have identified wing patterning loci in these butterflies offer broad insight into how gene regulatory networks evolve to produce diversity.This article is part of the themed issue 'Evo-devo in the genomics era, and the origins of morphological diversity'. © 2016 The Author(s).
Wallbank, Richard W. R.; Hanly, Joseph J.
2017-01-01
A major challenge is to understand how conserved gene regulatory networks control the wonderful diversity of form that we see among animals and plants. Butterfly wing patterns are an excellent example of this diversity. Butterfly wings form as imaginal discs in the caterpillar and are constructed by a gene regulatory network, much of which is conserved across the holometabolous insects. Recent work in Heliconius butterflies takes advantage of genomic approaches and offers insights into how the diversification of wing patterns is overlaid onto this conserved network. WntA is a patterning morphogen that alters spatial information in the wing. Optix is a transcription factor that acts later in development to paint specific wing regions red. Both of these loci fit the paradigm of conserved protein-coding loci with diverse regulatory elements and developmental roles that have taken on novel derived functions in patterning wings. These discoveries offer insights into the ‘Nymphalid Ground Plan’, which offers a unifying hypothesis for pattern formation across nymphalid butterflies. These loci also represent ‘hotspots’ for morphological change that have been targeted repeatedly during evolution. Both convergent and divergent evolution of a great diversity of patterns is controlled by complex alleles at just a few genes. We suggest that evolutionary change has become focused on one or a few genetic loci for two reasons. First, pre-existing complex cis-regulatory loci that already interact with potentially relevant transcription factors are more likely to acquire novel functions in wing patterning. Second, the shape of wing regulatory networks may constrain evolutionary change to one or a few loci. Overall, genomic approaches that have identified wing patterning loci in these butterflies offer broad insight into how gene regulatory networks evolve to produce diversity. This article is part of the themed issue ‘Evo-devo in the genomics era, and the origins of morphological diversity’. PMID:27994126
Habitat variation and wing coloration affect wing shape evolution in dragonflies.
Outomuro, D; Dijkstra, K-D B; Johansson, F
2013-09-01
Habitats are spatially and temporally variable, and organisms must be able to track these changes. One potential mechanism for this is dispersal by flight. Therefore, we would expect flying animals to show adaptations in wing shape related to habitat variation. In this work, we explored variation in wing shape in relation to preferred water body (flowing water or standing water with tolerance for temporary conditions) and landscape (forested to open) using 32 species of dragonflies of the genus Trithemis (80% of the known species). We included a potential source of variation linked to sexual selection: the extent of wing coloration on hindwings. We used geometric morphometric methods for studying wing shape. We also explored the phenotypic correlation of wing shape between the sexes. We found that wing shape showed a phylogenetic structure and therefore also ran phylogenetic independent contrasts. After correcting for the phylogenetic effects, we found (i) no significant effect of water body on wing shape; (ii) male forewings and female hindwings differed with regard to landscape, being progressively broader from forested to open habitats; (iii) hindwings showed a wider base in wings with more coloration, especially in males; and (iv) evidence for phenotypic correlation of wing shape between the sexes across species. Hence, our results suggest that natural and sexual selection are acting partially independently on fore- and hindwings and with differences between the sexes, despite evidence for phenotypic correlation of wing shape between males and females. © 2013 The Authors. Journal of Evolutionary Biology © 2013 European Society For Evolutionary Biology.
The timing of wing molt in tundra swans: energetic and non-energetic constraints
Earnst, S.L.
1992-01-01
Date of wing molt initiation, based on the regression of tenth primary length on capture date, was calculated for breeding and nonbreeding Tundra Swans (Cygnus columbianus columbianus) on the Colville River Delta, Alaska. Breeding females initiated wing molt significantly later than breeding males and nonbreeding males and females; the molt of breeding females was correlated with the date on which their eggs hatched. Breeding males did not differ significantly from nonbreeding males and females in the date of molt initiation. Timing of molt in breeding males and females was consistent with the views that females delay molt while replenishing energy spent on reproduction, but was also consistent with the breeding pair's need for primaries to defend territories and to defend and brood young. Other results, including an increase in an index of female body condition throughout most of the molt period, and a positive correlation between clutch size and female hatch-to-molt interval, were not predicted by the hypothesis that past energy expenditures constrain the timing of molt. Patterns of wing molt within and among other Northern Hemisphere geese and swans are also difficult to explain on the basis of energetics alone. For example, breeding females initiate molt before breeding males in many species. Also, there is extreme asynchrony between mates in two swan species; one of those species also exhibits variation in which sex initiates wing molt first. Both patterns suggest that asynchrony, per se, is important, probably to facilitate brood protection or territory defense. In Tundra Swans and other northern breeding geese and swans, the non-energetic demands of territory defense, brood defense, and brooding are probably important constraints on the timing of wing molt.
Genetic Basis of Melanin Pigmentation in Butterfly Wings
Zhang, Linlin; Martin, Arnaud; Perry, Michael W.; van der Burg, Karin R. L.; Matsuoka, Yuji; Monteiro, Antónia; Reed, Robert D.
2017-01-01
Despite the variety, prominence, and adaptive significance of butterfly wing patterns, surprisingly little is known about the genetic basis of wing color diversity. Even though there is intense interest in wing pattern evolution and development, the technical challenge of genetically manipulating butterflies has slowed efforts to functionally characterize color pattern development genes. To identify candidate wing pigmentation genes, we used RNA sequencing to characterize transcription across multiple stages of butterfly wing development, and between different color pattern elements, in the painted lady butterfly Vanessa cardui. This allowed us to pinpoint genes specifically associated with red and black pigment patterns. To test the functions of a subset of genes associated with presumptive melanin pigmentation, we used clustered regularly interspaced short palindromic repeats (CRISPR)/Cas9 genome editing in four different butterfly genera. pale, Ddc, and yellow knockouts displayed reduction of melanin pigmentation, consistent with previous findings in other insects. Interestingly, however, yellow-d, ebony, and black knockouts revealed that these genes have localized effects on tuning the color of red, brown, and ochre pattern elements. These results point to previously undescribed mechanisms for modulating the color of specific wing pattern elements in butterflies, and provide an expanded portrait of the insect melanin pathway. PMID:28193726
Allen, Cerisse E; Beldade, Patrícia; Zwaan, Bas J; Brakefield, Paul M
2008-03-26
There is spectacular morphological diversity in nature but lineages typically display a limited range of phenotypes. Because developmental processes generate the phenotypic variation that fuels natural selection, they are a likely source of evolutionary biases, facilitating some changes and limiting others. Although shifts in developmental regulation are associated with morphological differences between taxa, it is unclear how underlying mechanisms affect the rate and direction of evolutionary change within populations under selection. Here we focus on two ecologically relevant features of butterfly wing color patterns, eyespot size and color composition, which are similarly and strongly correlated across the serially repeated eyespots. Though these two characters show similar patterns of standing variation and covariation within a population, they differ in key features of their underlying development. We targeted pairs of eyespots with artificial selection for coordinated (concerted selection) versus independent (antagonistic selection) change in their color composition and size and compared evolutionary responses of the two color pattern characters. The two characters respond to selection in strikingly different ways despite initially similar patterns of variation in all directions present in the starting population. Size (determined by local properties of a diffusing inductive signal) evolves flexibly in all selected directions. However, color composition (determined by a tissue-level response to the signal concentration gradient) evolves only in the direction of coordinated change. There was no independent evolutionary change in the color composition of two eyespots in response to antagonistic selection. Moreover, these differences in the directions of short-term evolutionary change in eyespot size and color composition within a single species are consistent with the observed wing pattern diversity in the genus. Both characters respond rapidly to selection for coordinated change, but there are striking differences in their response to selection for antagonistic, independent change across eyespots. While many additional factors may contribute to both short- and long-term evolutionary response, we argue that the compartmentalization of developmental processes can influence the diversification of serial repeats such as butterfly eyespots, even under strong selection.
Nijhout, H Frederik; Cinderella, Margaret; Grunert, Laura W
2014-03-01
The wings of butterflies and moths develop from imaginal disks whose structure is always congruent with the final adult wing. It is therefore possible to map every point on the imaginal disk to a location on the adult wing throughout ontogeny. We studied the growth patterns of the wings of two distantly related species with very different adult wing shapes, Junonia coenia and Manduca sexta. The shape of the wing disks change throughout their growth phase in a species-specific pattern. We measured mitotic densities and mitotic orientation in successive stages of wing development approximately one cell division apart. Cell proliferation was spatially patterned, and the density of mitoses was highly correlated with local growth. Unlike other systems in which the direction of mitoses has been viewed as the primary determinant of directional growth, we found that in these two species the direction of growth was only weakly correlated with the orientation of mitoses. Directional growth appears to be imposed by a constantly changing spatial pattern of cell division coupled with a weak bias in the orientation of cell division. Because growth and cell division in imaginal disk require ecdysone and insulin signaling, the changing spatial pattern of cell division may due to a changing pattern of expression of receptors or downstream elements in the signaling pathways for one or both of these hormones. Evolution of wing shape comes about by changes in the progression of spatial patterns of cell division. © 2014 Wiley Periodicals, Inc.
A Model for Selection of Eyespots on Butterfly Wings.
Sekimura, Toshio; Venkataraman, Chandrasekhar; Madzvamuse, Anotida
2015-01-01
The development of eyespots on the wing surface of butterflies of the family Nympalidae is one of the most studied examples of biological pattern formation.However, little is known about the mechanism that determines the number and precise locations of eyespots on the wing. Eyespots develop around signaling centers, called foci, that are located equidistant from wing veins along the midline of a wing cell (an area bounded by veins). A fundamental question that remains unsolved is, why a certain wing cell develops an eyespot, while other wing cells do not. We illustrate that the key to understanding focus point selection may be in the venation system of the wing disc. Our main hypothesis is that changes in morphogen concentration along the proximal boundary veins of wing cells govern focus point selection. Based on previous studies, we focus on a spatially two-dimensional reaction-diffusion system model posed in the interior of each wing cell that describes the formation of focus points. Using finite element based numerical simulations, we demonstrate that variation in the proximal boundary condition is sufficient to robustly select whether an eyespot focus point forms in otherwise identical wing cells. We also illustrate that this behavior is robust to small perturbations in the parameters and geometry and moderate levels of noise. Hence, we suggest that an anterior-posterior pattern of morphogen concentration along the proximal vein may be the main determinant of the distribution of focus points on the wing surface. In order to complete our model, we propose a two stage reaction-diffusion system model, in which an one-dimensional surface reaction-diffusion system, posed on the proximal vein, generates the morphogen concentrations that act as non-homogeneous Dirichlet (i.e., fixed) boundary conditions for the two-dimensional reaction-diffusion model posed in the wing cells. The two-stage model appears capable of generating focus point distributions observed in nature. We therefore conclude that changes in the proximal boundary conditions are sufficient to explain the empirically observed distribution of eyespot focus points on the entire wing surface. The model predicts, subject to experimental verification, that the source strength of the activator at the proximal boundary should be lower in wing cells in which focus points form than in those that lack focus points. The model suggests that the number and locations of eyespot foci on the wing disc could be largely controlled by two kinds of gradients along two different directions, that is, the first one is the gradient in spatially varying parameters such as the reaction rate along the anterior-posterior direction on the proximal boundary of the wing cells, and the second one is the gradient in source values of the activator along the veins in the proximal-distal direction of the wing cell.
Mutation predicts 40 million years of fly wing evolution.
Houle, David; Bolstad, Geir H; van der Linde, Kim; Hansen, Thomas F
2017-08-24
Mutation enables evolution, but the idea that adaptation is also shaped by mutational variation is controversial. Simple evolutionary hypotheses predict such a relationship if the supply of mutations constrains evolution, but it is not clear that constraints exist, and, even if they do, they may be overcome by long-term natural selection. Quantification of the relationship between mutation and phenotypic divergence among species will help to resolve these issues. Here we use precise data on over 50,000 Drosophilid fly wings to demonstrate unexpectedly strong positive relationships between variation produced by mutation, standing genetic variation, and the rate of evolution over the last 40 million years. Our results are inconsistent with simple constraint hypotheses because the rate of evolution is very low relative to what both mutational and standing variation could allow. In principle, the constraint hypothesis could be rescued if the vast majority of mutations are so deleterious that they cannot contribute to evolution, but this also requires the implausible assumption that deleterious mutations have the same pattern of effects as potentially advantageous ones. Our evidence for a strong relationship between mutation and divergence in a slowly evolving structure challenges the existing models of mutation in evolution.
Genetic Basis of Melanin Pigmentation in Butterfly Wings.
Zhang, Linlin; Martin, Arnaud; Perry, Michael W; van der Burg, Karin R L; Matsuoka, Yuji; Monteiro, Antónia; Reed, Robert D
2017-04-01
Despite the variety, prominence, and adaptive significance of butterfly wing patterns, surprisingly little is known about the genetic basis of wing color diversity. Even though there is intense interest in wing pattern evolution and development, the technical challenge of genetically manipulating butterflies has slowed efforts to functionally characterize color pattern development genes. To identify candidate wing pigmentation genes, we used RNA sequencing to characterize transcription across multiple stages of butterfly wing development, and between different color pattern elements, in the painted lady butterfly Vanessa cardui This allowed us to pinpoint genes specifically associated with red and black pigment patterns. To test the functions of a subset of genes associated with presumptive melanin pigmentation, we used clustered regularly interspaced short palindromic repeats (CRISPR)/Cas9 genome editing in four different butterfly genera. pale , Ddc , and yellow knockouts displayed reduction of melanin pigmentation, consistent with previous findings in other insects. Interestingly, however, yellow-d , ebony , and black knockouts revealed that these genes have localized effects on tuning the color of red, brown, and ochre pattern elements. These results point to previously undescribed mechanisms for modulating the color of specific wing pattern elements in butterflies, and provide an expanded portrait of the insect melanin pathway. Copyright © 2017 by the Genetics Society of America.
NASA Technical Reports Server (NTRS)
Parsons-Wingerter, Patricia A.; Hosamani, Ravikumar; Bhattacharya, Sharmila
2015-01-01
Imaginal wing discs of Drosophila melanogaster (fruit fly) defined during embryogenesis ultimately result in mature wings of stereotyped (specific) venation patterning. Major regulators of wing disc development are the epidermal growth factor receptor (EGF), Notch, Hedgehog (Hh), Wingless (Wg), and Dpp signaling pathways. Highly stereotyped vascular patterning is also characteristic of tissues in other organisms flown in space such as the mouse retina and leaves of Arabidopsis thaliana. Genetic and other adaptations of vascular patterning to space environmental factors have not yet been systematically quantified, despite widespread recognition of their critical importance for terrestrial and microgravity applications. Here we report changes in gene expression with space flight related to Drosophila wing morphogenesis and vein patterning. In addition, genetically modified phenotypes of increasingly abnormal ectopic wing venation in the Drosophila wing1 were analyzed by NASA's VESsel GENeration Analysis (VESGEN) software2. Our goal is to further develop insightful vascular mappings associated with bioinformatic dimensions of genetic or other molecular phenotypes for correlation with genetic and other molecular profiling relevant to NASA's GeneLab and other Space Biology exploration initiatives.
Ancient homology underlies adaptive mimetic diversity across butterflies
Gallant, Jason R.; Imhoff, Vance E.; Martin, Arnaud; Savage, Wesley K.; Chamberlain, Nicola L.; Pote, Ben L.; Peterson, Chelsea; Smith, Gabriella E.; Evans, Benjamin; Reed, Robert D.; Kronforst, Marcus R.; Mullen, Sean P.
2014-01-01
Convergent evolution provides a rare, natural experiment with which to test the predictability of adaptation at the molecular level. Little is known about the molecular basis of convergence over macro-evolutionary timescales. Here we use a combination of positional cloning, population genomic resequencing, association mapping and developmental data to demonstrate that positionally orthologous nucleotide variants in the upstream region of the same gene, WntA, are responsible for parallel mimetic variation in two butterfly lineages that diverged >65 million years ago. Furthermore, characterization of spatial patterns of WntA expression during development suggests that alternative regulatory mechanisms underlie wing pattern variation in each system. Taken together, our results reveal a strikingly predictable molecular basis for phenotypic convergence over deep evolutionary time. PMID:25198507
The digits of the wing of birds are 1, 2, and 3. A review.
Vargas, Alexander O; Fallon, John F
2005-05-15
Fossil evidence documenting the evolutionary transition from theropod dinosaurs to birds indicates unambiguously that the digits of the wing of birds are digits 1, 2, and 3. However, some embryological evidence suggests that these digits are 2, 3, and 4. This apparent lack of correspondence has been described as the greatest challenge to the widely accepted theropod-bird link (Zhou 2004. Naturwissenschaften 91:455-471). Here we review the pertinent literature regarding the debate on the origin of birds and wing digital identity and the evidence in favor of a 1, 2, 3 identity of the wing digits. Recent molecular evidence shows that the expression of Hoxd12 and Hoxd13 in the developing wing supports the theropod-bird link. In the chicken foot and in the mouse hand and foot, digit 1 is the only digit to combine the expression of Hoxd13 with the absence of expression of Hoxd12. The same is observed in the anterior digit of the wing, suggesting it is a digit 1, as expected for a theropod. Nevertheless, Galis et al. (2005. J Exp Zool (Mol Dev Evol) in press), argue that Hoxd12 and Hoxd13 expression patterns in mutant limbs do not allow distinguishing the most anterior digit in the bird wing from digit 2. They also argue that constraints to the evolution of limb development support the 2, 3, 4 identity of the wing digits. However, the case put forward by Galis et al. is biased and flawed with regard to interpretation of mutant limbs, developmental mechanisms, stages observed, and the description of the evolutionary variation of limb development. Importantly, Galis et al. do not present evidence from wild-type limbs that counters the conclusions of Vargas and Fallon (2005. J Exp Zool (Mol Dev Evol) 304B(1):85-89), and fail to provide molecular evidence to specifically support the hypothesis that the wing digits are 2, 3, and 4. The expression of Hoxd12 and Hoxd13 in the developing wing is consistent with the hypothesis that birds are living dinosaurs; this view can lead to a greater understanding of the actual limits to the evolutionary variation of limb development. Copyright 2005 Wiley-Liss, Inc
Bowsher, Julia H; Wray, Gregory A; Abouheif, Ehab
2007-12-15
Over the last decade, it has become clear that organismal form is largely determined by developmental and evolutionary changes in the growth and pattern formation of tissues. Yet, there is little known about how these two integrated processes respond to environmental cues or how they evolve relative to one another. Here, we present the discovery of vestigial wing imaginal discs in worker larvae of the red imported fire ant, Solenopsis invicta. These vestigial wing discs are present in all worker larvae, which is uncommon for a species with a large worker size distribution. Furthermore, the growth trajectory of these vestigial discs is distinct from all of the ant species examined to date because they grow at a rate slower than the leg discs. We predicted that the growth trajectory of the vestigial wing discs would be mirrored by evolutionary changes in their patterning. We tested this prediction by examining the expression of three patterning genes, extradenticle, ultrabithorax, and engrailed, known to underlie the wing polyphenism in ants. Surprisingly, the expression patterns of these three genes in the vestigial wing discs was the same as those found in ant species with different worker size distributions and wing disc growth than fire ants. We conclude that growth and patterning are evolutionarily dissociated in the vestigial wing discs of S. invicta because patterning in these discs is conserved, whereas their growth trajectories are not. The evolutionary dissociation of growth and patterning may be an important feature of gene networks that underlie polyphenic traits. 2007 Wiley-Liss, Inc
Integration of wings and their eyespots in the speckled wood butterfly Pararge aegeria.
Breuker, Casper J; Gibbs, Melanie; Van Dyck, Hans; Brakefield, Paul M; Klingenberg, Christian Peter; Van Dongen, Stefan
2007-07-15
We investigated both the phenotypic and developmental integration of eyespots on the fore- and hindwings of speckled wood butterflies Pararge aegeria. Eyespots develop within a framework of wing veins, which may not only separate eyespots developmentally, but may at the same time also integrate them by virtue of being both signalling sources and barriers during eyespot development. We therefore specifically investigated the interaction between wing venation patterns and eyespot integration. Phenotypic covariation among eyespots was very high, but only eyespots in neighbouring wing cells and in homologous wing cells on different wing surfaces were developmentally integrated. This can be explained by the fact that the wing cells of these eyespots share one or more wing veins. The wing venation patterns of fore- and hindwings were highly integrated, both phenotypically and developmentally. This did not affect overall developmental integration of the eyespots. The adaptive significance of integration patterns is discussed and more specifically we stress the need to conduct studies on phenotypic plasticity of integration.
Wing patterning gene redefines the mimetic history of Heliconius butterflies.
Hines, Heather M; Counterman, Brian A; Papa, Riccardo; Albuquerque de Moura, Priscila; Cardoso, Marcio Z; Linares, Mauricio; Mallet, James; Reed, Robert D; Jiggins, Chris D; Kronforst, Marcus R; McMillan, W Owen
2011-12-06
The mimetic butterflies Heliconius erato and Heliconius melpomene have undergone parallel radiations to form a near-identical patchwork of over 20 different wing-pattern races across the Neotropics. Previous molecular phylogenetic work on these radiations has suggested that similar but geographically disjunct color patterns arose multiple times independently in each species. The neutral markers used in these studies, however, can move freely across color pattern boundaries, and therefore might not represent the history of the adaptive traits as accurately as markers linked to color pattern genes. To assess the evolutionary histories across different loci, we compared relationships among races within H. erato and within H. melpomene using a series of unlinked genes, genes linked to color pattern loci, and optix, a gene recently shown to control red color-pattern variation. We found that although unlinked genes partition populations by geographic region, optix had a different history, structuring lineages by red color patterns and supporting a single origin of red-rayed patterns within each species. Genes closely linked (80-250 kb) to optix exhibited only weak associations with color pattern. This study empirically demonstrates the necessity of examining phenotype-determining genomic regions to understand the history of adaptive change in rapidly radiating lineages. With these refined relationships, we resolve a long-standing debate about the origins of the races within each species, supporting the hypothesis that the red-rayed Amazonian pattern evolved recently and expanded, causing disjunctions of more ancestral patterns.
2013-01-01
Background One of the most intriguing questions in evolutionary developmental biology is how an insect acquires a mimicry pattern within its body parts. A striking example of pattern mimicry is found in the pattern diversity of moth and butterfly wings, which is thought to evolve from preexisting elements illustrated by the nymphalid ground plan (NGP). Previous studies demonstrated that individuality of the NGP facilitates the decoupling of associated common elements, leading to divergence. In contrast, recent studies on the concept of modularity have argued the importance of a combination of coupling and decoupling of the constituent elements. Here, we examine the modularity of a mimicry wing pattern in a moth and explore an evolvable characteristic of the NGP. Results This study examined the wings of the noctuid moth Oraesia excavata, which closely resemble leaves with a leaf venation pattern. Based on a comparative morphological procedure, we found that this leaf pattern was formed by the NGP common elements. Using geometric morphometrics combined with network analysis, we found that each of the modules in the leaf pattern integrates the constituent components of the leaf venation pattern (i.e., the main and lateral veins). Moreover, the detected modules were established by coupling different common elements and decoupling even a single element into different modules. The modules of the O. excavata wing pattern were associated with leaf mimicry, not with the individuality of the NGP common elements. For comparison, we also investigated the modularity of a nonmimetic pattern in the noctuid moth Thyas juno. Quantitative analysis demonstrated that the modules of the T. juno wing pattern regularly corresponded to the individuality of the NGP common elements, unlike those in the O. excavata wing pattern. Conclusions This study provides the first evidence for modularity in a leaf mimicry pattern. The results suggest that the evolution of this pattern involves coupling and decoupling processes to originate these modules, free from the individuality of the NGP system. We propose that this evolution has been facilitated by a versatile characteristic of the NGP, allowing the association of freely modifiable subordinate common elements to make modules. PMID:23890367
Convergent evolution in the genetic basis of Müllerian mimicry in heliconius butterflies.
Baxter, Simon W; Papa, Riccardo; Chamberlain, Nicola; Humphray, Sean J; Joron, Mathieu; Morrison, Clay; ffrench-Constant, Richard H; McMillan, W Owen; Jiggins, Chris D
2008-11-01
The neotropical butterflies Heliconius melpomene and H. erato are Müllerian mimics that display the same warningly colored wing patterns in local populations, yet pattern diversity between geographic regions. Linkage mapping has previously shown convergent red wing phenotypes in these species are controlled by loci on homologous chromosomes. Here, AFLP bulk segregant analysis using H. melpomene crosses identified genetic markers tightly linked to two red wing-patterning loci. These markers were used to screen a H. melpomene BAC library and a tile path was assembled spanning one locus completely and part of the second. Concurrently, a similar strategy was used to identify a BAC clone tightly linked to the locus controlling the mimetic red wing phenotypes in H. erato. A methionine rich storage protein (MRSP) gene was identified within this BAC clone, and comparative genetic mapping shows red wing color loci are in homologous regions of the genome of H. erato and H. melpomene. Subtle differences in these convergent phenotypes imply they evolved independently using somewhat different developmental routes, but are nonetheless regulated by the same switch locus. Genetic mapping of MRSP in a third related species, the "tiger" patterned H. numata, has no association with wing patterning and shows no evidence for genomic translocation of wing-patterning loci.
2011-01-01
Background Heliconius butterflies are an excellent system for studies of adaptive convergent and divergent phenotypic traits. Wing colour patterns are used as signals to both predators and potential mates and are inherited in a Mendelian manner. The underlying genetic mechanisms of pattern formation have been studied for many years and shed light on broad issues, such as the repeatability of evolution. In Heliconius melpomene, the yellow hindwing bar is controlled by the HmYb locus. MicroRNAs (miRNAs) are important post-transcriptional regulators of gene expression that have key roles in many biological processes, including development. miRNAs could act as regulators of genes involved in wing development, patterning and pigmentation. For this reason we characterised miRNAs in developing butterfly wings and examined differences in their expression between colour pattern races. Results We sequenced small RNA libraries from two colour pattern races and detected 142 Heliconius miRNAs with homology to others found in miRBase. Several highly abundant miRNAs were differentially represented in the libraries between colour pattern races. These candidates were tested further using Northern blots, showing that differences in expression were primarily due to developmental stage rather than colour pattern. Assembly of sequenced reads to the HmYb region identified hme-miR-193 and hme-miR-2788; located 2380 bp apart in an intergenic region. These two miRNAs are expressed in wings and show an upregulation between 24 and 72 hours post-pupation, indicating a potential role in butterfly wing development. A search for miRNAs in all available H. melpomene BAC sequences (~ 2.5 Mb) did not reveal any other miRNAs and no novel miRNAs were predicted. Conclusions Here we describe the first butterfly miRNAs and characterise their expression in developing wings. Some show differences in expression across developing pupal stages and may have important functions in butterfly wing development. Two miRNAs were located in the HmYb region and were expressed in developing pupal wings. Future work will examine the expression of these miRNAs in different colour pattern races and identify miRNA targets among wing patterning genes. PMID:21266089
Surridge, Alison K; Lopez-Gomollon, Sara; Moxon, Simon; Maroja, Luana S; Rathjen, Tina; Nadeau, Nicola J; Dalmay, Tamas; Jiggins, Chris D
2011-01-26
Heliconius butterflies are an excellent system for studies of adaptive convergent and divergent phenotypic traits. Wing colour patterns are used as signals to both predators and potential mates and are inherited in a Mendelian manner. The underlying genetic mechanisms of pattern formation have been studied for many years and shed light on broad issues, such as the repeatability of evolution. In Heliconius melpomene, the yellow hindwing bar is controlled by the HmYb locus. MicroRNAs (miRNAs) are important post-transcriptional regulators of gene expression that have key roles in many biological processes, including development. miRNAs could act as regulators of genes involved in wing development, patterning and pigmentation. For this reason we characterised miRNAs in developing butterfly wings and examined differences in their expression between colour pattern races. We sequenced small RNA libraries from two colour pattern races and detected 142 Heliconius miRNAs with homology to others found in miRBase. Several highly abundant miRNAs were differentially represented in the libraries between colour pattern races. These candidates were tested further using Northern blots, showing that differences in expression were primarily due to developmental stage rather than colour pattern. Assembly of sequenced reads to the HmYb region identified hme-miR-193 and hme-miR-2788; located 2380 bp apart in an intergenic region. These two miRNAs are expressed in wings and show an upregulation between 24 and 72 hours post-pupation, indicating a potential role in butterfly wing development. A search for miRNAs in all available H. melpomene BAC sequences (~2.5 Mb) did not reveal any other miRNAs and no novel miRNAs were predicted. Here we describe the first butterfly miRNAs and characterise their expression in developing wings. Some show differences in expression across developing pupal stages and may have important functions in butterfly wing development. Two miRNAs were located in the HmYb region and were expressed in developing pupal wings. Future work will examine the expression of these miRNAs in different colour pattern races and identify miRNA targets among wing patterning genes.
Ferguson, Laura C; Maroja, Luana; Jiggins, Chris D
2011-12-01
The evolution of pigmentation in vertebrates and flies has involved repeated divergence at a small number of genes related to melanin synthesis. Here, we study insect melanin synthesis genes in Heliconius butterflies, a group characterised by its diversity of wing patterns consisting of black (melanin), and yellow and red (ommochrome) pigmented scales. Consistent with their respective biochemical roles in Drosophila melanogaster, ebony is upregulated in non-melanic wing regions destined to be pigmented red whilst tan is upregulated in melanic regions. Wing regions destined to be pigmented yellow, however, are downregulated for both genes. This pattern is conserved across multiple divergent and convergent phenotypes within the Heliconii, suggesting a conserved mechanism for the development of black, red and yellow pattern elements across the genus. Linkage mapping of five melanin biosynthesis genes showed that, in contrast to other organisms, these genes do not control pattern polymorphism. Thus, the pigmentation genes themselves are not the locus of evolutionary change but lie downstream of a wing pattern regulatory factor. The results suggest a modular system in which particular combinations of genes are switched on whenever red, yellow or black pattern elements are favoured by natural selection for diverse and mimetic wing patterns. © Springer-Verlag 2011
Distal-less induces elemental color patterns in Junonia butterfly wings.
Dhungel, Bidur; Ohno, Yoshikazu; Matayoshi, Rie; Iwasaki, Mayo; Taira, Wataru; Adhikari, Kiran; Gurung, Raj; Otaki, Joji M
2016-01-01
The border ocellus, or eyespot, is a conspicuous color pattern element in butterfly wings. For two decades, it has been hypothesized that transcription factors such as Distal-less (Dll) are responsible for eyespot pattern development in butterfly wings, based on their expression in the prospective eyespots. In particular, it has been suggested that Dll is a determinant for eyespot size. However, functional evidence for this hypothesis has remained incomplete, due to technical difficulties. Here, we show that ectopically expressed Dll induces ectopic elemental color patterns in the adult wings of the blue pansy butterfly, Junonia orithya (Lepidoptera, Nymphalidae). Using baculovirus-mediated gene transfer, we misexpressed Dll protein fused with green fluorescent protein (GFP) in pupal wings, resulting in ectopic color patterns, but not the formation of intact eyespots. Induced changes included clusters of black and orange scales (a basic feature of eyespot patterns), black and gray scales, and inhibition of cover scale development. In contrast, ectopic expression of GFP alone did not induce any color pattern changes using the same baculovirus-mediated gene transfer system. These results suggest that Dll plays an instructive role in the development of color pattern elements in butterfly wings, although Dll alone may not be sufficient to induce a complete eyespot. This study thus experimentally supports the hypothesis of Dll function in eyespot development.
NASA Technical Reports Server (NTRS)
Seidman, Oscar; Neihouse, A I
1940-01-01
The reported tests are a continuation of an NACA investigation being made in the free-spinning wind tunnel to determine the effects of independent variations in load distribution, wing and tail arrangement, and control disposition on the spin characteristics of airplanes. The standard series of tests was repeated to determine the effect of airplane relative density. Tests were made at values of the relative-density parameter of 6.8, 8.4 (basic), and 12.0; and the results were analyzed. The tested variations in the relative-density parameter may be considered either as variations in the wing loading of an airplane spun at a given altitude, with the radii of gyration kept constant, or as a variation of the altitude at which the spin takes place for a given airplane. The lower values of the relative-density parameter correspond to the lower wing loadings or to the lower altitudes of the spin.
Wing geometry of Culex coronator (Diptera: Culicidae) from South and Southeast Brazil
2014-01-01
Background The Coronator Group encompasses Culex coronator Dyar & Knab, Culex camposi Dyar, Culex covagarciai Forattini, Culex ousqua Dyar, Culex usquatissimus Dyar, Culex usquatus Dyar and Culex yojoae Strickman. Culex coronator has the largest geographic distribution, occurring in North, Central and South America. Moreover, it is a potential vector-borne mosquito species because females have been found naturally infected with several arboviruses, i.e., Saint Louis Encephalitis Virus, Venezuelan Equine Encephalitis Virus and West Nile Virus. Considering the epidemiological importance of Cx. coronator, we investigated the wing shape diversity of Cx. coronator from South and Southeast Brazil, a method to preliminarily estimate population diversity. Methods Field-collected immature stages of seven populations from a large geographical area in Brazil were maintained in the laboratory to obtain both females and males linked with pupal and/or larval exuviae. For each individual female, 18 landmarks of left wings were marked and digitalized. After Procrustes superimposition, discriminant analysis of shape was employed to quantify wing shape variation among populations. The isometric estimator centroid size was calculated to assess the overall wing size and allometry. Results Wing shape was polymorphic among populations of Cx. coronator. However, dissimilarities among populations were higher than those observed within each population, suggesting populational differentiation in Cx. coronator. Morphological distances between populations were not correlated to geographical distances, indicating that other factors may act on wing shape and thus, determining microevolutionary patterns in Cx. coronator. Despite the population differentiation, intrapopulational wing shape variability was equivalent among all seven populations. Conclusion The wing variability found in Cx. coronator populations brings to light a new biological problem to be investigated: the population genetics of Cx. coronator. Because of differences in the male genitalia, we also transferred Cx. yojoae to the Apicinus Subgroup. PMID:24721508
apterous A specifies dorsal wing patterns and sexual traits in butterflies
2018-01-01
Butterflies have evolved different colour patterns on their dorsal and ventral wing surfaces to serve different signalling functions, yet the developmental mechanisms controlling surface-specific patterning are still unknown. Here, we mutate both copies of the transcription factor apterous in Bicyclus anynana butterflies using CRISPR/Cas9 and show that apterous A, expressed dorsally, functions both as a repressor and modifier of ventral wing colour patterns, as well as a promoter of dorsal sexual ornaments in males. We propose that the surface-specific diversification of wing patterns in butterflies proceeded via the co-option of apterous A or its downstream effectors into various gene regulatory networks involved in the differentiation of discrete wing traits. Further, interactions between apterous and sex-specific factors such as doublesex may have contributed to the origin of sexually dimorphic surface-specific patterns. Finally, we discuss the evolution of eyespot number diversity in the family Nymphalidae within the context of developmental constraints due to apterous regulation. PMID:29467265
Chromosomal rearrangements maintain a polymorphic supergene controlling butterfly mimicry
Joron, Mathieu; Frezal, Lise; Jones, Robert T.; Chamberlain, Nicola L.; Lee, Siu F.; Haag, Christoph R.; Whibley, Annabel; Becuwe, Michel; Baxter, Simon W.; Ferguson, Laura; Wilkinson, Paul A.; Salazar, Camilo; Davidson, Claire; Clark, Richard; Quail, Michael A.; Beasley, Helen; Glithero, Rebecca; Lloyd, Christine; Sims, Sarah; Jones, Matthew C.; Rogers, Jane; Jiggins, Chris D.; ffrench-Constant, Richard H.
2013-01-01
Supergenes are tight clusters of loci that facilitate the co-segregation of adaptive variation, providing integrated control of complex adaptive phenotypes1. Polymorphic supergenes, in which specific combinations of traits are maintained within a single population, were first described for ‘pin’ and ‘thrum’ floral types in Primula1 and Fagopyrum2, but classic examples are also found in insect mimicry3–5 and snail morphology6. Understanding the evolutionary mechanisms that generate these co-adapted gene sets, as well as the mode of limiting the production of unfit recombinant forms, remains a substantial challenge7–10. Here we show that individual wing-pattern morphs in the polymorphic mimetic butterfly Heliconius numata are associated with different genomic rearrangements at the supergene locus P. These rearrangements tighten the genetic linkage between at least two colour-pattern loci that are known to recombine in closely related species9–11, with complete suppression of recombination being observed in experimental crosses across a 400-kilobase interval containing at least 18 genes. In natural populations, notable patterns of linkage disequilibrium (LD) are observed across the entire P region. The resulting divergent haplotype clades and inversion breakpoints are found in complete association with wing-pattern morphs. Our results indicate that allelic combinations at known wing-patterning loci have become locked together in a polymorphic rearrangement at the Plocus, forming a supergene that acts as a simple switch between complex adaptive phenotypes found in sympatry. These findings highlight how genomic rearrangements can have a central role in the coexistence of adaptive phenotypes involving several genes acting in concert, by locally limiting recombination and gene flow. PMID:21841803
De Keyser, Rien; Breuker, Casper J.; Hails, Rosemary S.; Dennis, Roger L. H.; Shreeve, Tim G.
2015-01-01
We examined the roles of wing melanisation, weight, and basking posture in thermoregulation in Polyommatus Icarus, a phenotypically variable and protandrous member of the diverse Polyommatinae (Lycaenidae). Under controlled experimental conditions, approximating to marginal environmental conditions for activity in the field (= infrequent flight, long duration basking periods), warming rates are maximised with fully open wings and maximum body temperatures are dependent on weight. Variation in wing melanisation within and between sexes has no effect on warming rates; males and females which differ in melanisation had similar warming rates. Posture also affected cooling rates, consistent with cooling being dependent on convective heat loss. We hypothesise that for this small sized butterfly, melanisation has little or no effect on thermoregulation. This may be a factor contributing to the diversity of wing colours in the Polyommatinae. Because of the importance of size for thermoregulation in this small butterfly, requirements for attaining a suitable size to confer thermal stability in adults may also be a factor influencing larval feeding rates, development time and patterns of voltinism. Our findings indicate that commonly accepted views of the importance of melanisation, posture and size to thermoregulation, developed using medium and large sized butterflies, are not necessarily applicable to small sized butterflies. PMID:25923738
De Keyser, Rien; Breuker, Casper J; Hails, Rosemary S; Dennis, Roger L H; Shreeve, Tim G
2015-01-01
We examined the roles of wing melanisation, weight, and basking posture in thermoregulation in Polyommatus Icarus, a phenotypically variable and protandrous member of the diverse Polyommatinae (Lycaenidae). Under controlled experimental conditions, approximating to marginal environmental conditions for activity in the field (= infrequent flight, long duration basking periods), warming rates are maximised with fully open wings and maximum body temperatures are dependent on weight. Variation in wing melanisation within and between sexes has no effect on warming rates; males and females which differ in melanisation had similar warming rates. Posture also affected cooling rates, consistent with cooling being dependent on convective heat loss. We hypothesise that for this small sized butterfly, melanisation has little or no effect on thermoregulation. This may be a factor contributing to the diversity of wing colours in the Polyommatinae. Because of the importance of size for thermoregulation in this small butterfly, requirements for attaining a suitable size to confer thermal stability in adults may also be a factor influencing larval feeding rates, development time and patterns of voltinism. Our findings indicate that commonly accepted views of the importance of melanisation, posture and size to thermoregulation, developed using medium and large sized butterflies, are not necessarily applicable to small sized butterflies.
Traits and evolution of wing venation pattern in paraneopteran insects.
Nel, André; Prokop, Jakub; Nel, Patricia; Grandcolas, Philippe; Huang, Di-Ying; Roques, Patrick; Guilbert, Eric; Dostál, Ondřej; Szwedo, Jacek
2012-05-01
Two different patterns of wing venation are currently supposed to be present in each of the three orders of Paraneoptera. This is unlikely compared with the situation in other insects where only one pattern exists per order. We propose for all Paraneoptera a new and unique interpretation of wing venation pattern, assuming that the convex cubitus anterior gets fused with the common stem of median and radial veins at or very near to wing base, after separation from concave cubitus posterior, and re-emerges more distally from R + M stem. Thereafter, the vein between concave cubitus posterior and CuA is a specialized crossvein called "cua-cup," proximally concave and distally convex. We show that despite some variations, that is, cua-cup can vary from absent to hypertrophic; CuA can re-emerge together with M or not, or even completely disappear, this new interpretation explains all situations among all fossil and recent paraneopteran lineages. We propose that the characters "CuA fused in a common stem with R and M"and "presence of specialized crossvein cua-cup" are venation apomorphies that support the monophyly of the Paraneoptera. In the light of these characters, we reinterpret several Palaeozoic and early Mesozoic fossils that were ascribed to Paraneoptera, and confirm the attribution of several to this superorder as well as possible attribution of Zygopsocidae (Zygopsocus permianus Tillyard, 1935) as oldest Psocodea. We discuss the situation in extinct Hypoperlida and Miomoptera, suggesting that both orders could well be polyphyletic, with taxa related to Archaeorthoptera, Paraneoptera, or even Holometabola. The Carboniferous Protoprosbolidae is resurrected and retransferred into the Paraneoptera. The genus Lithoscytina is restored. The miomopteran Eodelopterum priscum Schmidt, 1962 is newly revised and considered as a fern pinnule. In addition, the new paraneopteran Bruayaphis oudardi gen. nov. et sp. nov. is described fromthe Upper Carboniferous of France (see Supporting Information). Copyright © 2011 Wiley Periodicals, Inc.
Asymmetric hindwing foldings in rove beetles.
Saito, Kazuya; Yamamoto, Shuhei; Maruyama, Munetoshi; Okabe, Yoji
2014-11-18
Foldable wings of insects are the ultimate deployable structures and have attracted the interest of aerospace engineering scientists as well as entomologists. Rove beetles are known to fold their wings in the most sophisticated ways that have right-left asymmetric patterns. However, the specific folding process and the reason for this asymmetry remain unclear. This study reveals how these asymmetric patterns emerge as a result of the folding process of rove beetles. A high-speed camera was used to reveal the details of the wing-folding movement. The results show that these characteristic asymmetrical patterns emerge as a result of simultaneous folding of overlapped wings. The revealed folding mechanisms can achieve not only highly compact wing storage but also immediate deployment. In addition, the right and left crease patterns are interchangeable, and thus each wing internalizes two crease patterns and can be folded in two different ways. This two-way folding gives freedom of choice for the folding direction to a rove beetle. The use of asymmetric patterns and the capability of two-way folding are unique features not found in artificial structures. These features have great potential to extend the design possibilities for all deployable structures, from space structures to articles of daily use.
Investigation of Surface Enhanced Coherent Raman Scattering on Nano-patterned Insect Wings
NASA Astrophysics Data System (ADS)
Ujj, Laszlo; Lawhead, Carlos
2015-03-01
Many insect wings (cicadas, butterflies, mosquitos) poses nano-patterned surface structure. Characterization of surface morphology and chemical composition of insect wings is important to understand the extreme mechanical properties and the biophysical functionalities of the wings. We have measured the image of the membrane of a cicada's wing with the help of Scanning Electron Microscopy (SEM). The results confirm the existing periodic structure of the wing measured previously. In order to identify the chemical composition of the wing, we have deposited silver nanoparticles on it and applied Coherent anti-Stokes Raman Spectroscopy to measure the vibrational spectra of the molecules comprising the wing for the first time. The measured spectra are consistent with the original assumption that the wing membrane is composed of protein, wax, and chitin. The results of these studies can be used to measure other nano-patterned surfaces and to make artificial materials in the future. Authors grateful for financial support from the Department of Physics of the College of Sciences Engineering and Health of UWF and the Pall Corporation for SEM imaging.
Yoshioka, Shinya; Kinoshita, Shuichi
2006-01-22
A few species of Morpho butterflies have a distinctive white stripe pattern on their structurally coloured blue wings. Since the colour pattern of a butterfly wing is formed as a mosaic of differently coloured scales, several questions naturally arise: are the microstructures the same between the blue and white scales? How is the distinctive whiteness produced, structurally or by means of pigmentation? To answer these questions, we have performed structural and optical investigations of the stripe pattern of a butterfly, Morpho cypris. It is found that besides the dorsal and ventral scale layers, the wing substrate also has the corresponding stripe pattern. Quantitative optical measurements and analysis using a simple model for the wing structure reveal the origin of the higher reflectance which makes the white stripe brighter.
NASA Technical Reports Server (NTRS)
Jones, R.; Creel, T. R., Jr.; Lawing, P.; Quan, M.; Dye, W.; Cummings, J.; Gorowitz, H.; Craig, C.; Rich, G.
1973-01-01
Phase change paint tests of a Rockwell International .00593-scale space shuttle orbiter were conducted in the Langley Research Center's Variable Density Wind Tunnel. The test objectives were to determine the effects of various wing/underbody configurations on the aerodynamic heating rates and boundary layer transition during simulated entry conditions. Several models were constructed. Each varied from the other in either wing cuff radius, airfoil thickness, or wing-fuselage underbody blending. Two ventral fins were glued to the fuselage underside of one model to test the interference heating effects. Simulated Mach 8 entry data were obtained for each configuration at angles of attack ranging from 25 to 40 deg, and a Reynolds number variation of one million to eight million. Elevon, bodyflap, and rudder flare deflections were tested. Oil flow visualization and Schlieren photographs were obtained to aid in reducing the phase change paint data as well as to observe the flow patterns peculiar to each configuration.
Prunier, Jérôme G.; Dewulf, Alexandre; Kuhlmann, Michael; Michez, Denis
2017-01-01
Morphological traits can be highly variable over time in a particular geographical area. Different selective pressures shape those traits, which is crucial in evolutionary biology. Among these traits, insect wing morphometry has already been widely used to describe phenotypic variability at the inter-specific level. On the contrary, fewer studies have focused on intra-specific wing morphometric variability. Yet, such investigations are relevant to study potential convergences of variation that could highlight micro-evolutionary processes. The recent sampling and sequencing of three solitary bees of the genus Melitta across their entire species range provides an excellent opportunity to jointly analyse genetic and morphometric variability. In the present study, we first aim to analyse the spatial distribution of the wing shape and centroid size (used as a proxy for body size) variability. Secondly, we aim to test different potential predictors of this variability at both the intra- and inter-population levels, which includes genetic variability, but also geographic locations and distances, elevation, annual mean temperature and precipitation. The comparison of spatial distribution of intra-population morphometric diversity does not reveal any convergent pattern between species, thus undermining the assumption of a potential local and selective adaptation at the population level. Regarding intra-specific wing shape differentiation, our results reveal that some tested predictors, such as geographic and genetic distances, are associated with a significant correlation for some species. However, none of these predictors are systematically identified for the three species as an important factor that could explain the intra-specific morphometric variability. As a conclusion, for the three solitary bee species and at the scale of this study, our results clearly tend to discard the assumption of the existence of a common pattern of intra-specific signal/structure within the intra-specific wing shape and body size variability. PMID:28273178
NASA Technical Reports Server (NTRS)
Hwang, Kyu C.; Tiwari, Surrendra N.; Miley, Stanley J.
1995-01-01
In recent years, active research has been conducted to study the technological feasibility of supersonic laminar flow control on the wing of the High Speed Civil Transport (HSCT). For this study, the F-16XL has been chosen due to its highly swept crank wing planform that closely resembles the HSCT configurations. During flights, it is discovered that the shock wave generated from the aircraft inlet introduces disturbances on the wing where the data acquisition is conducted. The flow field about a supersonic inlet is characterized by a complex three dimensional pattern of shock waves generated by the geometrical configuration of a deflector and a cowl lip. Hence, in this study, experimental method is employed to investigate the effects of the variation of deflector configuration on the flow field, and consequently, the possibility of diverting the incoming shock-disturbances away from the test section. In the present experiments, a model composed of a simple circular tube with a triangular deflector is designed to study the deflector length and the deflector base width variation in the flow field. Experimental results indicate that the lowest external pressure ratio is observed at the junction where the deflector lip and the inlet cowl lip merge. Also, it is noted that the external pressure ratio, the internal pressure ratio, the coefficient of spillage drag, and the shock standoff distance decrease as the deflector length increases. In addition, the Redefined Total Pressure Recovery Ratio (RTPRR) increases with an increase in the deflector length. Results from the study of the effect of the deflector's base width variation on the flow field indicate that the lowest external pressure ratio is observed at the junction between the inlet cowl lip and the deflector lip. As the base width of the deflector increases, the external pressure ratio at 0 rotation increases, whereas the external pressure ratio at 180 rotation decreases. In addition, the internal pressure ratio and the coefficient of spillage drag decrease as the base width of the deflector increases. However, RTPRR and shock standoff distance increase as the base width increases. In conclusion, as deflector dimensions vary, distinctive patterns in the pressure variation around the inlet deflector are observed. With an increase in the deflector length and base width, the magnitude of shock-disturbances are weakened due to a decrease in the external pressure ratio. Also, as the deflector length and base width increase, a smaller bow shock angle is formed. Therefore, the inlet shock wave formation would be significantly altered, and consequently, shock disturbances on the wing test section can be avoided through appropriately designing the deflector.
The phenetic structure of Aedes albopictus.
Morales Vargas, Ronald Enrique; Phumala-Morales, Noppawan; Tsunoda, Takashi; Apiwathnasorn, Chamnan; Dujardin, Jean-Pierre
2013-01-01
The large and recent migrations of the main vector of the Chikungunya virus, Aedes albopictus, raise questions about the epidemiological impact of possible microevolutionary changes in new territories. Quantitative traits are suitable to detect such changes as induced by environmental adaptations, local competition and founder effects. Using landmark-based geometric morphometrics, we compared the size and shape of 22 populations (1572 females) of Ae. albopictus. The collection sites covered five countries around the world, with special emphasis on Asia, which is assumed to be the origin of the vector. Some collections came from places where an important epidemic outbreak of Chikungunya has recently occurred. Quantitative comparisons were based on 18 anatomical landmarks on the wing. To contrast geographic and possible interspecific shape variation, close species were introduced in the sample, namely five Aedes riversi and nine Ae. flavopictus from Japan. The three species had similar size, but they were clearly separated by shape. Within Ae. albopictus, there was general agreement on size variation with the available climatic data. Shape variation was less related to climatic data than to either geography or a known itinerary of past territorial expansion. Thus, two main clusters were distinguished by wing shape variation: the first one composed of the Southeast Asian sample, the second one grouping Japan, Florida, Hawaii and the Reunion Island samples. The Southeast Asian countries, assumed to be at the origin of the geographic expansion of the mosquito, had similar wings and constituted a distinct group where localities clustered into northern and southern localities. Contrasting with this homogeneous group, very distant localities such as United States (US) and Japan shared a common shape pattern. The US Ae. albopictus samples (Hawaii and Florida) were indeed very similar to the Japanese samples, with Florida behaving exactly like a northern Japanese locality, close to Tokyo and Yokohama. Shape proximity among these distant areas could be explained by a common and relatively recent ancestor, as generally suggested in the literature. The three Reunion Island samples conformed a group external to the Southeast Asian countries, loosely connected to the remaining localities. Thus, areas where recent epidemics took place, the Reunion Island and the southernmost localities of Thailand, did not cluster together, but represented separate groups. In sum, the size of the wing was apparently under the influence of climatic factors, while its shape could contribute information on species and geographic differences. Apparent departures from expectations based on geography were explained by known past and present migratory routes. These patterns of shape variation were compatible with genetic drift, suggesting microevolutionary changes probably induced by the expansion of the Ae. albopictus mosquito. Copyright © 2012 Elsevier B.V. All rights reserved.
[Transverse folding and the evolution of hind wings in beetles (Insecta, Coleoptera)].
Fedorenko, D N
2013-01-01
Strong intensification of the protective function of the fore wing in Coleoptera has made their flight apparatus a posteromotoric one and invited an apparatus responsible for folding the hindwings beneath the elytra to develop. Folding apparatus could hardly develop without higher deformability of veins or their parts, which diminished strength properties of the wing support. The effect was stressed by folds that intersected veins. Organization of the folds into a system confined this negative influence to a few wing regions and some veinal sections. This having happened, wing support and folding pattern evolved interrelated, the former into being more flexible, with no or minimum loss of rigidity, and the latter towards being less harmful for the supporting elements, especially axial ones. Monofunctionality, together with very simple structure and little specialization of constituent parts, made the folding pattern very labile during evolution. The folding pattern evolved more rapidly than wing venation, thus defining transformations of the latter. Evolutionary conservatism of wing venation stemmed from that many veins were strongly specialized in performing two conflicting functions. An adaptive compromise was necessary for the conflict to be solved, which determined the wing to orthogenetic development. The main evolutionary trends for wing venation and folding pattern were those towards simplification and a higher complexity, respectively. The beetle wing has passed through two main evolutionary stages. Among them, the first resulted in the development of the "Archostemata" wing type, the second started from the "cantharoid" structural plan. The main evolutionary factors were the infancies of wing posteromotorism at the first stage while the wing strongly influenced by size evolution, with the main trend towards miniaturization, at the second. The archostematan and "cantharoid" morphofunctional wing types differ fundamentally. In the wing of the former kind, folding and flight apparatus, because of considerably overlapping supporting systems, constitute a lasting coadaptive ensemble, with only minor deviations from the ground-plan occurring through evolution. The uprise of the "cantharoid" wing type was an upgrade of morpho-functional organization. The region of maximum transverse deformations having been extruded from the remigium basal part, chief supporting axes of the wing increased their rigid properties. The supporting systems of the two wing apparatus became more autonomous, having been separated. This expanded the adaptive zone for the wing strongly, which a great variety of derived wing types have emerged from.
Variation in wing pattern and palatability in a female-limited polymorphic mimicry system
Long, Elizabeth C; Hahn, Thomas P; Shapiro, Arthur M
2014-01-01
Checkerspot butterflies in the genera Euphydryas and Chlosyne exhibit phenotypic polymorphisms along a well-defined latitudinal and elevational gradient in California. The patterns of phenotypic variation in Euphydryas chalcedona, Chlosyne palla, and Chlosyne hoffmanni suggest a mimetic relationship; in addition, the specific patterns of variation in C. palla suggest a female-limited polymorphic mimicry system (FPM). However, the existence of polymorphic models runs counter to predictions of mimicry theory. Palatability trials were undertaken to assess whether or not the different color morphs of each species were distasteful or toxic to a generalized avian predator, the European starling (Sturnus vulgaris). Results indicate that the black morph of E. chalcedona is distasteful, but not toxic, to predators, while the red morph is palatable. C . hoffmanni and both color morphs of C. palla are palatable to predators. Predators that learn to reject black E. chalcedona also reject black C. palla, suggesting that the latter is a FPM of the former. C. hoffmanni does not appear to be involved in this mimetic relationship. PMID:25512850
Correlated evolution of host and parasite body size: tests of Harrison's rule using birds and lice.
Johnson, Kevin P; Bush, Sarah E; Clayton, Dale H
2005-08-01
Large-bodied species of hosts often harbor large-bodied parasites, a pattern known as Harrison's rule. Harrison's rule has been documented for a variety of animal parasites and herbivorous insects, yet the adaptive basis of the body-size correlation is poorly understood. We used phylogenetically independent methods to test for Harrison's rule across a large assemblage of bird lice (Insecta: Phthiraptera). The analysis revealed a significant relationship between louse and host size, despite considerable variation among taxa. We explored factors underlying this variation by testing Harrison's rule within two groups of feather-specialist lice that share hosts (pigeons and doves). The two groups, wing lice (Columbicola spp.) and body lice (Physconelloidinae spp.), have similar life histories, despite spending much of their time on different feather tracts. Wing lice showed strong support for Harrison's rule, whereas body lice showed no significant correlation with host size. Wing louse size was correlated with wing feather size, which was in turn correlated with overall host size. In contrast, body louse size showed no correlation with body feather size, which also was not correlated with overall host size. The reason why body lice did not fit Harrison's rule may be related to the fact that different species of body lice use different microhabitats within body feathers. More detailed measurements of body feathers may be needed to explore the precise relationship of body louse size to relevant components of feather size. Whatever the reason, Harrison's rule does not hold in body lice, possibly because selection on body size is mediated by community-level interactions between body lice.
Character displacement in the fighting colours of Hetaerina damselflies.
Anderson, Christopher N; Grether, Gregory F
2010-12-07
Aggression between species is a seldom-considered but potentially widespread mechanism of character displacement in secondary sexual characters. Based on previous research showing that similarity in wing coloration directly influences interspecific territorial aggression in Hetaerina damselflies, we predicted that wing coloration would show a pattern of character displacement (divergence in sympatry). A geographical survey of four Hetaerina damselfly species in Mexico and Texas showed evidence for character displacement in both species pairs that regularly occurs sympatrically. Hetaerina titia, a species that typically has large black wing spots and small red wing spots, shifted to having even larger black spots and smaller red wing spots at sites where a congener with large red wing spots is numerically dominant (Hetaerina americana or Hetaerina occisa). Hetaerina americana showed the reverse pattern, shifting towards larger red wing spots where H. titia is numerically dominant. This pattern is consistent with the process of agonistic character displacement, but the ontogenetic basis of the shift remains to be demonstrated.
Le, Tuyen Quang; Truong, Tien Van; Park, Soo Hyung; Quang Truong, Tri; Ko, Jin Hwan; Park, Hoon Cheol; Byun, Doyoung
2013-01-01
In this work, the aerodynamic performance of beetle wing in free-forward flight was explored by a three-dimensional computational fluid dynamics (CFDs) simulation with measured wing kinematics. It is shown from the CFD results that twist and camber variation, which represent the wing flexibility, are most important when determining the aerodynamic performance. Twisting wing significantly increased the mean lift and camber variation enhanced the mean thrust while the required power was lower than the case when neither was considered. Thus, in a comparison of the power economy among rigid, twisting and flexible models, the flexible model showed the best performance. When the positive effect of wing interaction was added to that of wing flexibility, we found that the elytron created enough lift to support its weight, and the total lift (48.4 mN) generated from the simulation exceeded the gravity force of the beetle (47.5 mN) during forward flight. PMID:23740486
Patterns of variation in clutch sizes in a guild of temperate-nesting dabbling ducks
Krapu, G.L.; Reynolds, R.E.; Sargeant, G.A.; Renner, R.W.
2004-01-01
Clutch sizes of temperate-nesting dabbling ducks vary widely within and between years. Biologists have long been interested in why such patterns exist but have had difficulty separating intrinsic effects from environmental influences. In an attempt to gain greater insight into the roles of intrinsic and environmental influences on clutch sizes of dabbling ducks, we compared clutch-size patterns of mallard, northern pintail, gadwall, northern shoveler, and blue-winged teal in North Dakota and South Dakota over a 3-year period. The study was conducted during 1993-1995 when wetland habitat conditions were exceptionally consistent within time intervals but varied widely from the onset to the end of the study. We found that clutch sizes of gadwall and blue-winged teal were significantly smaller at the onset of nesting in 1993, when water conditions remained relatively poor following a 5-year drought (1988-1992), and pintail clutch sizes were larger late in the breeding season in 1993 after wetland conditions became exceptionally productive following heavy rains in late May and June. These findings were associated with gadwall and teal relying primarily on nutrients acquired on the breeding grounds to produce early clutches, and pintails using an exceptionally high proportion of their body lipids to produce early clutches, making this species highly dependent on abundant local sources of lipids for clutches produced later. Mallard, pintail, and shoveler rely largely on endogenous lipids carried to the breeding grounds to produce early clutches, and at least in the years of study, clutch sizes exhibited minimal variation at the onset of nesting. Our findings, when examined in context with existing information, suggest that interspecific variation in clutch sizes results from innate differences in several traits -- including body size, diet, timing of lipid acquisition, and nesting-- that affect the amount of lipid available for egg production. Annual differences in clutch sizes of all five species were not significant when effects of annual variation in nest initiation dates was accounted for, reflecting the key role of environmental influences on intraspecific variation among years.
McFarlane, Laura; Altringham, John D.; Askew, Graham N.
2016-01-01
ABSTRACT Diurnal and seasonal increases in body mass and seasonal reductions in wing area may compromise a bird's ability to escape, as less of the power available from the flight muscles can be used to accelerate and elevate the animal's centre of mass. Here, we investigated the effects of intra-specific variation in wing morphology on escape take-off performance in blue tits (Cyanistes caeruleus). Flights were recorded using synchronised high-speed video cameras and take-off performance was quantified as the sum of the rates of change of the kinetic and potential energies of the centre of mass. Individuals with a lower wing loading, WL (WL=body weight/wing area) had higher escape take-off performance, consistent with the increase in lift production expected from relatively larger wings. Unexpectedly, it was found that the total power available from the flight muscles (estimated using an aerodynamic analysis) was inversely related to WL. This could simply be because birds with a higher WL have relatively smaller flight muscles. Alternatively or additionally, variation in the aerodynamic load on the wing resulting from differences in wing morphology will affect the mechanical performance of the flight muscles via effects on the muscle's length trajectory. Consistent with this hypothesis is the observation that wing beat frequency and relative downstroke duration increase with decreasing WL; both are factors that are expected to increase muscle power output. Understanding how wing morphology influences take-off performance gives insight into the potential risks associated with feather loss and seasonal and diurnal fluctuations in body mass. PMID:26994175
Independently Controlled Wing Stroke Patterns in the Fruit Fly Drosophila melanogaster
Chakraborty, Soma; Bartussek, Jan; Fry, Steven N.; Zapotocky, Martin
2015-01-01
Flies achieve supreme flight maneuverability through a small set of miniscule steering muscles attached to the wing base. The fast flight maneuvers arise from precisely timed activation of the steering muscles and the resulting subtle modulation of the wing stroke. In addition, slower modulation of wing kinematics arises from changes in the activity of indirect flight muscles in the thorax. We investigated if these modulations can be described as a superposition of a limited number of elementary deformations of the wing stroke that are under independent physiological control. Using a high-speed computer vision system, we recorded the wing motion of tethered flying fruit flies for up to 12 000 consecutive wing strokes at a sampling rate of 6250 Hz. We then decomposed the joint motion pattern of both wings into components that had the minimal mutual information (a measure of statistical dependence). In 100 flight segments measured from 10 individual flies, we identified 7 distinct types of frequently occurring least-dependent components, each defining a kinematic pattern (a specific deformation of the wing stroke and the sequence of its activation from cycle to cycle). Two of these stroke deformations can be associated with the control of yaw torque and total flight force, respectively. A third deformation involves a change in the downstroke-to-upstroke duration ratio, which is expected to alter the pitch torque. A fourth kinematic pattern consists in the alteration of stroke amplitude with a period of 2 wingbeat cycles, extending for dozens of cycles. Our analysis indicates that these four elementary kinematic patterns can be activated mutually independently, and occur both in isolation and in linear superposition. The results strengthen the available evidence for independent control of yaw torque, pitch torque, and total flight force. Our computational method facilitates systematic identification of novel patterns in large kinematic datasets. PMID:25710715
Independently controlled wing stroke patterns in the fruit fly Drosophila melanogaster.
Chakraborty, Soma; Bartussek, Jan; Fry, Steven N; Zapotocky, Martin
2015-01-01
Flies achieve supreme flight maneuverability through a small set of miniscule steering muscles attached to the wing base. The fast flight maneuvers arise from precisely timed activation of the steering muscles and the resulting subtle modulation of the wing stroke. In addition, slower modulation of wing kinematics arises from changes in the activity of indirect flight muscles in the thorax. We investigated if these modulations can be described as a superposition of a limited number of elementary deformations of the wing stroke that are under independent physiological control. Using a high-speed computer vision system, we recorded the wing motion of tethered flying fruit flies for up to 12,000 consecutive wing strokes at a sampling rate of 6250 Hz. We then decomposed the joint motion pattern of both wings into components that had the minimal mutual information (a measure of statistical dependence). In 100 flight segments measured from 10 individual flies, we identified 7 distinct types of frequently occurring least-dependent components, each defining a kinematic pattern (a specific deformation of the wing stroke and the sequence of its activation from cycle to cycle). Two of these stroke deformations can be associated with the control of yaw torque and total flight force, respectively. A third deformation involves a change in the downstroke-to-upstroke duration ratio, which is expected to alter the pitch torque. A fourth kinematic pattern consists in the alteration of stroke amplitude with a period of 2 wingbeat cycles, extending for dozens of cycles. Our analysis indicates that these four elementary kinematic patterns can be activated mutually independently, and occur both in isolation and in linear superposition. The results strengthen the available evidence for independent control of yaw torque, pitch torque, and total flight force. Our computational method facilitates systematic identification of novel patterns in large kinematic datasets.
Yang, Ke; He, Peng; Dong, Shuang-Lin
2014-01-01
Abstract Chemosensory proteins (CSPs) play various roles in insect physiology including olfaction and development. The brown planthopper, Nilaparvata lugens Stål , is one of the most notorious rice pests worldwide. The wing-from variation and annually long distance migration imply that olfaction would play a key role in N. lugens behavior. In this study, full-length cDNAs of nine CSPs were cloned by the rapid amplification of cDNA ends procedure, and their expression profiles were determined by the quantitative real-time Polymerase Chain Reaction (qPCR), with regard to developmental stage, wing-form, gender, and tissues of short-wing adult. These NlugCSP genes showed distinct expression patterns, indicating different roles they play. In particular, NlugCSP5 was long wing form biased and highly expressed in female wings among tissues; NlugCSP1 was mainly expressed in male adults and abdomen; NlugCSP7 was widely expressed in chemosensory tissues but little in the nonchemosensory abdomen. The function of NlugCSP7 in olfaction was further explored by the competitive fluorescence binding assay using the recombinant protein. However, the recombinant NlugCSP7 showed no obvious binding with all tested volatile compounds, suggesting that it may participate in physiological processes other than olfaction. Our results provide bases and some important clues for the function of NlugCSPs . PMID:25527582
Wang, Yongji; Wang, Jianjian; Lai, Liming; Jiang, Lianhe; Zhuang, Ping; Zhang, Lehua; Zheng, Yuanrun; Baskin, Jerry M; Baskin, Carol C
2014-01-01
Seed mass and morphology are plant life history traits that influence seed dispersal ability, seeding establishment success, and population distribution pattern. Southeastern Tibet is a diversity center for Rhododendron species, which are distributed from a few hundred meters to 5500 m above sea level. We examined intra- and interspecific variation in seed mass and morphology in relation to altitude, habitat, plant height, and phylogeny. Seed mass decreased significantly with the increasing altitude and increased significantly with increasing plant height among populations of the same species. Seed mass differed significantly among species and subsections, but not among sections and subgenera. Seed length, width, surface area, and wing length were significantly negative correlated with altitude and significantly positive correlated with plant height. Further, these traits differed significantly among habitats and varied among species and subsection, but not among sections and subgenera. Species at low elevation had larger seeds with larger wings, and seeds became smaller and the wings of seeds tended to be smaller with the increasing altitude. Morphology of the seed varied from flat round to long cylindrical with increasing altitude. We suggest that seed mass and morphology have evolved as a result of both long-term adaptation and constraints of the taxonomic group over their long evolutionary history. PMID:24963385
Ribak, Gal
2017-01-01
Intraspecific variation in adult body mass can be particularly high in some insect species, mandating adjustment of the wing's structural properties to support the weight of the larger body mass in air. Insect wings elastically deform during flapping, dynamically changing the twist and camber of the relatively thin and flat aerofoil. We examined how wing deformations during free flight scale with body mass within a species of rose chafers (Coleoptera: Protaetia cuprea) in which individuals varied more than threefold in body mass (0.38–1.29 g). Beetles taking off voluntarily were filmed using three high-speed cameras and the instantaneous deformation of their wings during the flapping cycle was analysed. Flapping frequency decreased in larger beetles but, otherwise, flapping kinematics remained similar in both small and large beetles. Deflection of the wing chord-wise varied along the span, with average deflections at the proximal trailing edge higher by 0.2 and 0.197 wing lengths compared to the distal trailing edge in the downstroke and the upstroke, respectively. These deflections scaled with wing chord to the power of 1.0, implying a constant twist and camber despite the variations in wing and body size. This suggests that the allometric growth in wing size includes adjustment of the flexural stiffness of the wing structure to preserve wing twist and camber during flapping. PMID:29134103
Meresman, Yonatan; Ribak, Gal
2017-10-01
Intraspecific variation in adult body mass can be particularly high in some insect species, mandating adjustment of the wing's structural properties to support the weight of the larger body mass in air. Insect wings elastically deform during flapping, dynamically changing the twist and camber of the relatively thin and flat aerofoil. We examined how wing deformations during free flight scale with body mass within a species of rose chafers (Coleoptera: Protaetia cuprea ) in which individuals varied more than threefold in body mass (0.38-1.29 g). Beetles taking off voluntarily were filmed using three high-speed cameras and the instantaneous deformation of their wings during the flapping cycle was analysed. Flapping frequency decreased in larger beetles but, otherwise, flapping kinematics remained similar in both small and large beetles. Deflection of the wing chord-wise varied along the span, with average deflections at the proximal trailing edge higher by 0.2 and 0.197 wing lengths compared to the distal trailing edge in the downstroke and the upstroke, respectively. These deflections scaled with wing chord to the power of 1.0, implying a constant twist and camber despite the variations in wing and body size. This suggests that the allometric growth in wing size includes adjustment of the flexural stiffness of the wing structure to preserve wing twist and camber during flapping.
Garzón, Maximiliano J; Schweigmann, Nicolás
2018-05-16
Gene flow restrictions between populations of Aedes albifasciatus, the vector of Western equine encephalitis and Dirophilaria immitis, have been described in the central region of Argentina. Genetic and eco-physiological variations usually result in local forms reflecting the climatic regions. Mosquito wings and their different parts have ecological functions in flight and communication. Therefore, wing shape could be considered an aspect of sexual dimorphism, and its eco-physiological responses can be expressed as morphological changes induced by the environment. To compare the geographical and sexual variations with respect to wing shape and size in two Ae. albifasciatus populations from contrasting climates of Argentina (temperate: Buenos Aires, and the arid steppe of Patagonia: Sarmiento), the wings of adults reared in thermal trays at different constant temperatures (10-29 °C) were analyzed. The wing size of Ae. albifasciatus showed inverse linear relationships with the rearing thermal condition and higher slope for Buenos Aires. In the cool range (10-17 °C), geographical size variations responded to the converse Bergmann's rule, where Buenos Aires individuals were larger than those from Sarmiento. Sexual shape dimorphism occurred in both populations while geographical variation in shape was observed in both sexes. Buenos Aires individuals showed greater response sensitivity with respect to the size-temperature relation than those from Sarmiento. The converse Bergmann's rule in size variation could be due to a higher development rate in Sarmiento to produce more cohorts in the limited favorable season. The shape could be more relevant with respect to the size in the study of population structures due to the size being more liable to vary due to changes in the environment. The geographical variations with respect to morphology could be favored by the isolation between populations and adaptations to the environmental conditions. Our results demonstrate that the shape and size of wing provide useful phenotypic information for studies related to sexual and environmental adaptations.
NASA Technical Reports Server (NTRS)
Byrdsong, T. A.; Brooks, C. W., Jr.
1983-01-01
Wind-tunnel measurements were made of the wing-surface static-pressure distributions on a 0.237 scale model of a remotely piloted research vehicle equipped with a thick, high-aspect-ratio supercritical wing. Data are presented for two model configurations (with and without a ventral pod) at Mach numbers from 0.70 to 0.92 at angles of attack from -4 deg to 8 deg. Large variations of wing-surface local pressure distributions were developed; however, the characteristic supercritical-wing pressure distribution occurred near the design condition of 0.80 Mach number and 2 deg angle of attack. The significant variations of the local pressure distributions indicated pronounced shock-wave movements that were highly sensitive to angle of attack and Mach number. The effect of the vertical pod varied with test conditions; however at the higher Mach numbers, the effects on wing flow characteristics were significant at semispan stations as far outboard as 0.815. There were large variations of the wing loading in the range of test conditions, both model configurations exhibited a well-defined peak value of normal-force coefficient at the cruise angle of attack (2 deg) and Mach number (0.80).
USDA-ARS?s Scientific Manuscript database
The nymphalid groundplan (NGP) is an idealized system used to classify and interpret wing pattern elements of butterflies. Nearly a century ago, the principles of the NGP were applied to the wing patterns of higher moths (Macroheterocera). Recent advances in phylogeny and in the comparative morpholo...
Koch, P Bernhardt; Nijhout, H Frederik
1990-05-01
A set of stage specific proteins of approximally 86 to 90 kDal are synthesized by isolated wings ofPrecis coenia on day 5 of the pupal stage. They are named "B proteins". They are synthesized in presumptive black wing areas in higher amounts than in presumptive white wing areas and are the major proteins synthesized on day 5. Wings from 5 days old pupae, which were incubated with 35 S-methionine for 2 or 4 hours, incorporate radiolabel into presumptive black pattern elements. This is probably due to the localized synthesis of the above mentioned proteins. Injection of 35 S-methionine into whole pupae on day 5 resulted in the labelling of the mature black and grey scales but not white scales. This radiolabel incorporation pattern corresponds exactly to the incorporation of the melanin precursor 14 C-tyrosine into the scales. The results indicate that the "B proteins" are specifically related to the formation of black and grey portions of thePrecis wing pattern. Injection of 35 S-methionine into whole pupae on day 6 resulted in the labelling of the mature red scales probably due to labelling of "R proteins", which may be involved in the formation of red pattern elements.
Bischoff, Kara; Ballew, Anna C.; Simon, Michael A.; O'Reilly, Alana M.
2009-01-01
Background The coordinated action of genes that control patterning, cell fate determination, cell size, and cell adhesion is required for proper wing formation in Drosophila. Defects in any of these basic processes can lead to wing aberrations, including blisters. The xenicid mutation was originally identified in a screen designed to uncover regulators of adhesion between wing surfaces [1]. Principal Findings Here, we demonstrate that expression of the βPS integrin or the patterning protein Engrailed are not affected in developing wing imaginal discs in xenicid mutants. Instead, expression of the homeotic protein Ultrabithorax (Ubx) is strongly increased in xenicid mutant cells. Conclusion Our results suggest that upregulation of Ubx transforms cells from a wing blade fate to a haltere fate, and that the presence of haltere cells within the wing blade is the primary defect leading to the adult wing phenotypes observed. PMID:19956620
Multiple capacitors for natural genetic variation in Drosophila melanogaster.
Takahashi, Kazuo H
2013-03-01
Cryptic genetic variation (CGV) or a standing genetic variation that is not ordinarily expressed as a phenotype is released when the robustness of organisms is impaired under environmental or genetic perturbations. Evolutionary capacitors modulate the amount of genetic variation exposed to natural selection and hidden cryptically; they have a fundamental effect on the evolvability of traits on evolutionary timescales. In this study, I have demonstrated the effects of multiple genomic regions of Drosophila melanogaster on CGV in wing shape. I examined the effects of 61 genomic deficiencies on quantitative and qualitative natural genetic variation in the wing shape of D. melanogaster. I have identified 10 genomic deficiencies that do not encompass a known candidate evolutionary capacitor, Hsp90, exposing natural CGV differently depending on the location of the deficiencies in the genome. Furthermore, five genomic deficiencies uncovered qualitative CGV in wing morphology. These findings suggest that CGV in wing shape of wild-type D. melanogaster is regulated by multiple capacitors with divergent functions. Future analysis of genes encompassed by these genomic regions would help elucidate novel capacitor genes and better understand the general features of capacitors regarding natural genetic variation. © 2012 Blackwell Publishing Ltd.
Comparative population genetics of a mimicry locus among hybridizing Heliconius butterfly species.
Chamberlain, N L; Hill, R I; Baxter, S W; Jiggins, C D; Kronforst, M R
2011-09-01
The comimetic Heliconius butterfly species pair, H. erato and H. melpomene, appear to use a conserved Mendelian switch locus to generate their matching red wing patterns. Here we investigate whether H. cydno and H. pachinus, species closely related to H. melpomene, use this same switch locus to generate their highly divergent red and brown color pattern elements. Using an F2 intercross between H. cydno and H. pachinus, we first map the genomic positions of two novel red/brown wing pattern elements; the G locus, which controls the presence of red vs brown at the base of the ventral wings, and the Br locus, which controls the presence vs absence of a brown oval pattern on the ventral hind wing. The results reveal that the G locus is tightly linked to markers in the genomic interval that controls red wing pattern elements of H. erato and H. melpomene. Br is on the same linkage group but approximately 26 cM away. Next, we analyze fine-scale patterns of genetic differentiation and linkage disequilibrium throughout the G locus candidate interval in H. cydno, H. pachinus and H. melpomene, and find evidence for elevated differentiation between H. cydno and H. pachinus, but no localized signature of association. Overall, these results indicate that the G locus maps to the same interval as the locus controlling red patterning in H. melpomene and H. erato. This, in turn, suggests that the genes controlling red pattern elements may be homologous across Heliconius, supporting the hypothesis that Heliconius butterflies use a limited suite of conserved genetic switch loci to generate both convergent and divergent wing patterns.
An illustrated checklist of the genus Elymnias Hübner, 1818 (Nymphalidae, Satyrinae)
Wei, Chia-Hsuan; Lohman, David J.; Peggie, Djunijanti; Yen, Shen-Horn
2017-01-01
Abstract We review the genus Elymnias Hübner, 1818, a morphologically diverse satyrine butterfly clade involved in multifarious Batesian mimicry relationships throughout Asia and Africa. A variety of different model species are mimicked, and many Elymnias species are sexually dimorphic mimics, with males and females resembling different model species. We revise species and subspecies delimitations in light of an integrative taxonomic investigation using external morphology, male and female genital morphology, and a multi-locus molecular phylogeny. There is little interspecific genitalic variation among species in this group, and previous taxonomists therefore relied almost entirely on wing patterns. Our molecular phylogenetic analysis reveals several examples of polymorphism or wing pattern divergence within a single species currently classified as two or more different species. We also found examples of wing pattern convergence among disparate lineages that mimic the same widespread model species. Frequently, two or more phenotypically similar species were classified as a single species. This comprehensive checklist reviews all names associated with Elymnias to align its taxonomy with the evolutionary history of the group. All available information on nomenclature, type localities, repositories of type specimens, and geographical distributions is summarized, and images of adult specimens and genitalia are provided along with distribution maps of all species and selected subspecies. We identify 2 species incertae sedis, establish 15 monophyletic species groups (including 1 species unplaced in any species group), and make 49 taxonomic changes, including 35 new synonyms, 7 new combinations (2 of which have new status), 1 resurrected combination, 1 resurrected subspecies, and 7 status changes. PMID:28769686
Numerical simulation of incidence and sweep effects on delta wing vortex breakdown
NASA Technical Reports Server (NTRS)
Ekaterinaris, J. A.; Schiff, Lewis B.
1994-01-01
The structure of the vortical flowfield over delta wings at high angles of attack was investigated. Three-dimensional Navier-Stokes numerical simulations were carried out to predict the complex leeward-side flowfield characteristics, including leading-edge separation, secondary separation, and vortex breakdown. Flows over a 75- and a 63-deg sweep delta wing with sharp leading edges were investigated and compared with available experimental data. The effect of variation of circumferential grid resolution grid resolution in the vicinity of the wing leading edge on the accuracy of the solutions was addressed. Furthermore, the effect of turbulence modeling on the solutions was investigated. The effects of variation of angle of attack on the computed vortical flow structure for the 75-deg sweep delta wing were examined. At moderate angles of attack no vortex breakdown was observed. When a critical angle of attack was reached, bubble-type vortex breakdown was found. With further increase in angle of attack, a change from bubble-type breakdown to spiral-type vortex breakdown was predicted by the numerical solution. The effects of variation of sweep angle and freestream Mach number were addressed with the solutions on a 63-deg sweep delta wing.
Schäfer, Martin A; Berger, David; Rohner, Patrick T; Kjaersgaard, Anders; Bauerfeind, Stephanie S; Guillaume, Frédéric; Fox, Charles W; Blanckenhorn, Wolf U
2018-06-17
Geographic clines offer insights about putative targets and agents of natural selection as well as tempo and mode of adaptation. However, demographic processes can lead to clines that are indistinguishable from adaptive divergence. Using the widespread yellow dung fly Scathophaga stercoraria (Diptera: Scathophagidae), we examine quantitative genetic differentiation (Q ST ) of wing shape across North America, Europe and Japan, and compare this differentiation with that of ten microsatellites (F ST ). Morphometric analyses of 28 populations reared at three temperatures revealed significant thermal plasticity, sexual dimorphism and geographic differentiation in wing shape. In North America morphological differentiation followed the decline in microsatellite variability along the presumed route of recent colonization from the southeast to the northwest. Across Europe, where S. stercoraria presumably existed for much longer time and where no molecular pattern of isolation by distance was evident, clinal variation was less pronounced despite significant morphological differentiation (Q ST >F ST ). Shape vector comparisons further indicate that thermal plasticity (hot-to-cold) does not mirror patterns of latitudinal divergence (south-to-north), as might have been expected under a scenario with temperature as the major agent of selection. Our findings illustrate the importance of detailed phylogeographic information when interpreting geographic clines of dispersal traits in an adaptive evolutionary framework. This article is protected by copyright. All rights reserved. This article is protected by copyright. All rights reserved.
Some observations of separated flow on finite wings
NASA Technical Reports Server (NTRS)
Winkelmann, A. E.; Ngo, H. T.; De Seife, R. C.
1982-01-01
Wind tunnel test results for aspects of flow over airfoils exhibiting single and multiple trailing edge stall 'mushroom' cells are reported. Rectangular wings with aspect ratios of 4.0 and 9.0 were tested at Reynolds numbers of 480,000 and 257,000, respectively. Surface flow patterns were visualized by means of a fluorescent oil flow technique, separated flow was observed with a tuft wand and a water probe, spanwise flow was studied with hot-wire anemometry, smoke flow and an Ar laser illuminated the centerplane flow, and photographs were made of the oil flow patterns. Swirl patterns on partially and fully stalled wings suggested vortex flow attachments in those regions, and a saddle point on the fully stalled AR=4.0 wing indicated a secondary vortex flow at the forward region of the separation bubble. The separation wake decayed downstream, while the tip vortex interacted with the separation bubble on the fully stalled wing. Three mushroom cells were observed on the AR=9.0 wing.
The evolution of avian wing shape and previously unrecognized trends in covert feathering
Wang, Xia; Clarke, Julia A.
2015-01-01
Avian wing shape has been related to flight performance, migration, foraging behaviour and display. Historically, linear measurements of the feathered aerofoil and skeletal proportions have been used to describe this shape. While the distribution of covert feathers, layered over the anterior wing, has long been assumed to contribute to aerofoil properties, to our knowledge no previous studies of trends in avian wing shape assessed their variation. Here, these trends are explored using a geometric–morphometric approach with landmarks describing the wing outline as well as the extent of dorsal and ventral covert feathers for 105 avian species. We find that most of the observed variation is explained by phylogeny and ecology but shows only a weak relationship with previously described flight style categories, wing loading and an investigated set of aerodynamic variables. Most of the recovered variation is in greater primary covert feather extent, followed by secondary feather length and the shape of the wing tip. Although often considered a plastic character strongly linked to flight style, the estimated ancestral wing morphology is found to be generally conservative among basal parts of most major avian lineages. The radiation of birds is characterized by successive diversification into largely distinct areas of morphospace. However, aquatic taxa show convergence in feathering despite differences in flight style, and songbirds move into a region of morphospace also occupied by basal taxa but at markedly different body sizes. These results have implications for the proposed inference of flight style in extinct taxa. PMID:26446812
Genomic Signatures of Speciation in Sympatric and Allopatric Hawaiian Picture-Winged Drosophila
Kang, Lin; Settlage, Robert; McMahon, Wyatt; Michalak, Katarzyna; Tae, Hongseok; Garner, Harold R.; Stacy, Elizabeth A.; Price, Donald K.; Michalak, Pawel
2016-01-01
The Hawaiian archipelago provides a natural arena for understanding adaptive radiation and speciation. The Hawaiian Drosophila are one of the most diverse endemic groups in Hawaiì with up to 1,000 species. We sequenced and analyzed entire genomes of recently diverged species of Hawaiian picture-winged Drosophila, Drosophila silvestris and Drosophila heteroneura from Hawaiì Island, in comparison with Drosophila planitibia, their sister species from Maui, a neighboring island where a common ancestor of all three had likely occurred. Genome-wide single nucleotide polymorphism patterns suggest the more recent origin of D. silvestris and D. heteroneura, as well as a pervasive influence of positive selection on divergence of the three species, with the signatures of positive selection more prominent in sympatry than allopatry. Positively selected genes were significantly enriched for functional terms related to sensory detection and mating, suggesting that sexual selection played an important role in speciation of these species. In particular, sequence variation in Olfactory receptor and Gustatory receptor genes seems to play a major role in adaptive radiation in Hawaiian pictured-winged Drosophila. PMID:27189993
Dhungel, Bidur; Otaki, Joji M
2009-11-01
Butterfly wing color patterns can be changed by the application of a temperature shock or pharmacological agents such as tungstate, producing a distinctive type of elemental modification called the TS (temperature shock) type. Heterochronic uncoupling between the signaling and reception steps during the color-pattern determination process has been proposed as a mechanism for TS-type changes. As an extension of this hypothesis, both the parafocal element (PFE) and the eyespot in the same wing compartment are considered to be determined by morphogenic signal(s) emitted from the same eyespot focus. However, these models need to be examined with additional experimental data. Furthermore, there is controversy as to whether the action of tungstate on wing color patterns is direct or Indirect. Using a species of nymphalid butterfly (Junonia orithya), we have devised a simple method for the local application of pharmacological agents directly on developing wings of pupae. Local tungstate application resulted in reduced eyespots and circular dislocated PFEs in the eyespot-less compartments only on the treated wing, demonstrating that tungstate directly induces color-pattern changes on wings. We further examined the eyespot-PFE relationship in normal and cold-shocked Individuals, showing that an eyespot can be superimposed on a PFE and vice versa, probably depending on the timing of their fate determination. Taken together, we propose a two-morphogen model for the normal color-pattern determination, in which the morphogenic signals for the eyespot and PFE are different from each other despite their Identical origin. This two-morphogen model is compatible with the heterochronic uncoupling model for TS-type changes.
Otaki, Joji M; Ogasawara, Tsuyoshi; Yamamoto, Haruhiko
2005-06-01
Systemic injections of sodium tungstate, a protein-tyrosine phosphatase (PTPase) inhibitor, to pupae immediately after pupation have been shown to efficiently produce characteristic color-pattern modifications on the wings of many species of butterflies. Here we demonstrated that the tungstate-induced modification pattern was entirely different from other chemically-induced ones in a species of nymphalid butterfly Junonia (Precis) orithya. In this species, the systemic injections of tungstate produced characteristic expansion of black area and shrinkage of white area together with the move of parafocal elements toward the wing base. Overall, pattern boundaries became obscure. In contrast, an entirely different modification pattern, overall darkening of wings, was observed by the injections of stress-inducing chemicals, thapsigargin, ionomycin, or geldanamycin, to pupae under the rearing conditions for the adult summer form. On the ventral wings, this darkening was due to an increase of the proportion of peppered dark scales, which was reminiscent of the natural fall form of this species. Under the same rearing conditions, the injections of ecdysteroid, which is a well-known hormone being responsible for the seasonal polyphenism of nymphalid butterflies, yielded overall expansion of orange area especially around eyespots. Taken together, we conclude that the tungstate-induced modifications are clearly distinguishable from those of stress response and ecdysteroid effect. This conclusion then suggests that the putative PTPase signaling pathway that is sensitive to tungstate uniquely contributes to the wing-wide color-pattern development in butterflies.
Fanara, Juan Jose; Werenkraut, Victoria
2017-08-01
Changes in the environmental conditions experienced by naturally occurring populations are frequently accompanied by changes in adaptive traits allowing the organism to cope with environmental unpredictability. Phenotypic plasticity is a major aspect of adaptation and it has been involved in population dynamics of interacting species. In this study, phenotypic plasticity (i.e., environmental sensitivity) of morphological adaptive traits were analyzed in the cactophilic species Drosophila buzzatii and Drosophila koepferae (Diptera: Drosophilidae) considering the effect of crowding conditions (low and high density), type of competition (intraspecific and interspecific competition) and cacti hosts (Opuntia and Columnar cacti). All traits (wing length, wing width, thorax length, wing loading and wing aspect) showed significant variation for each environmental factor considered in both Drosophila species. The phenotypic plasticity pattern observed for each trait was different within and between these cactophilic Drosophila species depending on the environmental factor analyzed suggesting that body size-related traits respond almost independently to environmental heterogeneity. The effects of ecological factors analyzed in this study are discussed in order to elucidate the causal factors investigated (type of competition, crowding conditions and alternative host) affecting the election of the breeding site and/or the range of distribution of these cactophilic species. © 2016 Institute of Zoology, Chinese Academy of Sciences.
Determinants of breeding distributions of ducks
Johnson, D.H.; Grier, J.W.
1988-01-01
The settling of breeding habitat by migratory waterfowl is a topic of both theoretical and practical interest. We use the results of surveys conducted annually during 1955-81 in major breeding areas to examine the factors that affect the distributions of 10 common North American duck species. Three patterns of settling are described: homing, opportunistic, and flexible. Homing is generally more pronounced among species that use more stable (more predictable) wetlands, such as the redhead (Aythya americana), canvasback (A. valisineria), lesser scaup (A. affinis), mallard (Anas platyrhynchos), gadwall (Anas strepera), and northern shoveler (Anas clypeata). Opportunistic settling is more prevalent among species that use less stable (less predictable) wetlands, such as northern pintail (Anas acuta) and blue-winged teal (Anas discors). Flexible settling is exhibited to various degrees by most species.The 10 species are shown to fall along a natural ordination reflecting different life history characteristics. Average values of indices of r- and K-selection indicated that pintail, mallard, blue-winged teal, and shoveler have the most features associated with unstable or unpredictable environments. Gadwall, American wigeon (Anas americana), and green-winged teal (Anas crecca) were intermediate, and attributes of the diving ducks were associated with the use of stable or predictable environments.Some species--notably mallard, gadwall, blue-winged teal, redhead, and canvasback--tend to fill available breeding habitat first in the central portions of their range, and secondly in peripheral areas. Other species--American wigeon, green-winged teal, northern shoveler, northern pintail, and lesser scaup--fill their habitat in the order it is encountered during spring migration.Age and sex classes within species vary in their settling pattern. Some of this variation can be predicted from the mating systems of ducks in which breeding females, especially successful ones, have a greater investment in habitat resources and are more likely to return to the same area in subsequent years.
Nanostructured Antireflective and Thermoisolative Cicada Wings.
Morikawa, Junko; Ryu, Meguya; Seniutinas, Gediminas; Balčytis, Armandas; Maximova, Ksenia; Wang, Xuewen; Zamengo, Massimiliano; Ivanova, Elena P; Juodkazis, Saulius
2016-05-10
Inter-related mechanical, thermal, and optical macroscopic properties of biomaterials are defined at the nanoscale by their constituent structures and patterns, which underpin complex functions of an entire bio-object. Here, the temperature diffusivity of a cicada (Cyclochila australasiae) wing with nanotextured surfaces was measured using two complementary techniques: a direct contact method and IR imaging. The 4-6-μm-thick wing section was shown to have a thermal diffusivity of α⊥ = (0.71 ± 0.15) × 10(-7) m(2)/s, as measured by the contact temperature wave method along the thickness of the wing; it corresponds to the inherent thermal property of the cuticle. The in-plane thermal diffusivity value of the wing was determined by IR imaging and was considerably larger at α∥ = (3.6 ± 0.2) × 10(-7) m(2)/s as a result of heat transport via air. Optical properties of wings covered with nanospikes were numerically simulated using an accurate 3D model of the wing pattern and showed that light is concentrated between spikes where intensity is enhanced by up to 3- to 4-fold. The closely packed pattern of nanospikes reduces the reflectivity of the wing throughout the visible light spectrum and over a wide range of incident angles, hence acting as an antireflection coating.
Ancient Wings: animating the evolution of butterfly wing patterns.
Arbesman, Samuel; Enthoven, Leo; Monteiro, Antónia
2003-10-01
Character optimization methods can be used to reconstruct ancestral states at the internal nodes of phylogenetic trees. However, seldom are these ancestral states visualized collectively. Ancient Wings is a computer program that provides a novel method of visualizing the evolution of several morphological traits simultaneously. It allows users to visualize how the ventral hindwing pattern of 54 butterflies in the genus Bicyclus may have changed over time. By clicking on each of the nodes within the evolutionary tree, the user can see an animation of how wing size, eyespot size, and eyespot position relative the wing margin, have putatively evolved as a collective whole. Ancient Wings may be used as a pedagogical device as well as a research tool for hypothesis-generation in the fields of evolutionary, ecological, and developmental biology.
Morphological Variations of Leading-Edge Serrations in Owls (Strigiformes).
Weger, Matthias; Wagner, Hermann
2016-01-01
Owls have developed serrations, comb-like structures, along the leading edge of their wings. Serrations were investigated from a morphological and a mechanical point of view, but were not yet quantitatively compared for different species. Such a comparative investigation of serrations from species of different sizes and activity patterns may provide new information about the function of the serrations. Serrations on complete wings and on tenth primary remiges of seven owl species were investigated. Small, middle-sized, and large owl species were investigated as well as species being more active during the day and owls being more active during the night. Serrations occurred at the outer parts of the wings, predominantly at tenth primary remiges, but also on further wing feathers in most species. Serration tips were oriented away from the feather rachis so that they faced into the air stream during flight. The serrations of nocturnal owl species were higher developed as demonstrated by a larger inclination angle (the angle between the base of the barb and the rachis), a larger tip displacement angle (the angle between the tip of the serration and the base of the serration) and a longer length. Putting the measured data into a clustering algorithm yielded dendrograms that suggested a strong influence of activity pattern, but only a weak influence of size on the development of the serrations. Serrations are supposed to be involved in noise reduction during flight and also depend on the aerodynamic properties that in turn depend on body size. Since especially nocturnal owls have to rely on hearing during prey capture, the more pronounced serrations of nocturnal species lend further support to the notion that serrations have an important function in noise reduction. The differences in shape of the serrations investigated indicate that a silent flight requires well-developed serrations.
Acoustic Surveys of a Scaled-Model CESTOL Transport Aircraft in Static and Forward Speed Conditions
NASA Technical Reports Server (NTRS)
Burnside, Nathan; Horne, Clifton
2012-01-01
An 11% scale-model of a Cruise-Efficient Short Take-off and Landing (CESTOL) scalemodel test was recently completed. The test was conducted in the AEDC National Full-Scale Aerodynamic Complex (NFAC) 40- by 80-Foot Wind Tunnel at NASA Ames Research Center. The model included two over-wing pod-mounted turbine propulsion simulators (TPS). The hybrid blended wing-body used a circulation control wing (CCW) with leadingand trailing-edge blowing. The bulk of the test matrix included three forward velocities (40 kts, 60 kts, and 100kts), angle-of-attack variation between -5 and 25 , and CCW mass flow variation. Seven strut-mounted microphones outboard of the left wing provided source directivity. A phased microphone array was mounted outboard of the right wing for source location. The goal of this paper is to provide a preliminary look at the acoustic data acquired during the Advanced Model for Extreme Lift and Improved Aeroacoustics (AMELIA) test for 0 angle-of-attack and 0 sideslip conditions. Data presented provides a good overview of the test conditions and the signal-to-noise quality of the data. TPS height variation showed a difference of 2 dB to 3 dB due to wing shielding. Variation of slot mass flow showed increases of 12 dB to 26 dB above the airframe noise and the TPS increased the overall levels an additional 5 dB to 10 dB.
Convergent evolution of sexual shape dimorphism in Diptera.
Bonduriansky, Russell
2006-05-01
Several patterns of sexual shape dimorphism, such as male body elongation, eye stalks, or extensions of the exoskeleton, have evolved repeatedly in the true flies (Diptera). Although these dimorphisms may have evolved in response to sexual selection on male body shape, conserved genetic factors may have contributed to this convergent evolution, resulting in stronger phenotypic convergence than might be expected from functional requirements alone. I compared phenotypic variation in body shape in two distantly related species exhibiting sexually dimorphic body elongation: Prochyliza xanthostoma (Piophilidae) and Telostylinus angusticollis (Neriidae). Although sexual selection appears to act differently on male body shape in these species, they exhibited strikingly similar patterns of sexual dimorphism. Likewise, patterns of within-sex shape variation were similar in the two species, particularly in males: relative elongation of the male head capsule, antenna, and legs was associated with reduced head capsule width and wing length, but was nearly independent of variation in thorax length. However, the two species presented contrasting patterns of static allometry: male sexual traits exhibited elevated allometric slopes in T. angusticollis, but not in P. xanthostoma. These results suggest that a shared pattern of covariation among traits may have channeled the evolution of sexually dimorphic body elongation in these species. Nonetheless, static allometries may have been shaped by species-specific selection pressures or genetic architectures. Copyright 2006 Wiley-Liss, Inc.
Spontaneous long-range calcium waves in developing butterfly wings.
Ohno, Yoshikazu; Otaki, Joji M
2015-03-25
Butterfly wing color patterns emerge as the result of a regular arrangement of scales produced by epithelial scale cells at the pupal stage. These color patterns and scale arrangements are coordinated throughout the wing. However, the mechanism by which the development of scale cells is controlled across the entire wing remains elusive. In the present study, we used pupal wings of the blue pansy butterfly, Junonia orithya, which has distinct eyespots, to examine the possible involvement of Ca(2+) waves in wing development. Here, we demonstrate that the developing pupal wing tissue of the blue pansy butterfly displayed spontaneous low-frequency Ca(2+) waves in vivo that propagated slowly over long distances. Some waves appeared to be released from the immediate peripheries of the prospective eyespot and discal spot, though it was often difficult to identify the specific origins of these waves. Physical damage, which is known to induce ectopic eyespots, led to the radiation of Ca(2+) waves from the immediate periphery of the damaged site. Thapsigargin, which is a specific inhibitor of Ca(2+)-ATPases in the endoplasmic reticulum, induced an acute increase in cytoplasmic Ca(2+) levels and halted the spontaneous Ca(2+) waves. Additionally, thapsigargin-treated wings showed incomplete scale development as well as other scale and color pattern abnormalities. We identified a novel form of Ca(2+) waves, spontaneous low-frequency slow waves, which travel over exceptionally long distances. Our results suggest that spontaneous Ca(2+) waves play a critical role in the coordinated development of scale arrangements and possibly in color pattern formation in butterflies.
Hatadani, Luciane Mendes; Klaczko, Louis Bernard
2008-07-01
The second chromosome of Drosophila mediopunctata is highly polymorphic for inversions. Previous work reported a significant interaction between these inversions and collecting date on wing size, suggesting the presence of genotype-environment interaction. We performed experiments in the laboratory to test for the joint effects of temperature and chromosome inversions on size and shape of the wing in D. mediopunctata. Size was measured as the centroid size, and shape was analyzed using the generalized least squares Procrustes superimposition followed by discriminant analysis and canonical variates analysis of partial warps and uniform components scores. Our findings show that wing size and shape are influenced by temperature, sex, and karyotype. We also found evidence suggestive of an interaction between the effects of karyotype and temperature on wing shape, indicating the existence of genotype-environment interaction for this trait in D. mediopunctata. In addition, the association between wing size and chromosome inversions is in agreement with previous results indicating that these inversions might be accumulating alleles adapted to different temperatures. However, no significant interaction between temperature and karyotype for size was found--in spite of the significant presence of temperature-genotype (cross) interaction. We suggest that other ecological factors--such as larval crowding--or seasonal variation of genetic content within inversions may explain the previous results.
The evolution of avian wing shape and previously unrecognized trends in covert feathering.
Wang, Xia; Clarke, Julia A
2015-10-07
Avian wing shape has been related to flight performance, migration, foraging behaviour and display. Historically, linear measurements of the feathered aerofoil and skeletal proportions have been used to describe this shape. While the distribution of covert feathers, layered over the anterior wing, has long been assumed to contribute to aerofoil properties, to our knowledge no previous studies of trends in avian wing shape assessed their variation. Here, these trends are explored using a geometric-morphometric approach with landmarks describing the wing outline as well as the extent of dorsal and ventral covert feathers for 105 avian species. We find that most of the observed variation is explained by phylogeny and ecology but shows only a weak relationship with previously described flight style categories, wing loading and an investigated set of aerodynamic variables. Most of the recovered variation is in greater primary covert feather extent, followed by secondary feather length and the shape of the wing tip. Although often considered a plastic character strongly linked to flight style, the estimated ancestral wing morphology is found to be generally conservative among basal parts of most major avian lineages. The radiation of birds is characterized by successive diversification into largely distinct areas of morphospace. However, aquatic taxa show convergence in feathering despite differences in flight style, and songbirds move into a region of morphospace also occupied by basal taxa but at markedly different body sizes. These results have implications for the proposed inference of flight style in extinct taxa. © 2015 The Author(s).
Dhungel, Bidur; Ohno, Yoshikazu; Matayoshi, Rie; Otaki, Joji M
2013-03-25
Candidate genes for color pattern formation in butterfly wings have been known based on gene expression patterns since the 1990s, but their functions remain elusive due to a lack of a functional assay. Several methods of transferring and expressing a foreign gene in butterfly wings have been reported, but they have suffered from low success rates or low expression levels. Here, we developed a simple, practical method to efficiently deliver and express a foreign gene using baculovirus-mediated gene transfer in butterfly wings in vivo. A recombinant baculovirus containing a gene for green fluorescent protein (GFP) was injected into pupae of the blue pansy butterfly Junonia orithya (Nymphalidae). GFP fluorescence was detected in the pupal wings and other body parts of the injected individuals three to five days post-injection at various degrees of fluorescence. We obtained a high GFP expression rate at relatively high virus titers, but it was associated with pupal death before color pattern formation in wings. To reduce the high mortality rate caused by the baculovirus treatment, we administered an anti-gp64 antibody, which was raised against baculovirus coat protein gp64, to infected pupae after the baculovirus injection. This treatment greatly reduced the mortality rate of the infected pupae. GFP fluorescence was observed in pupal and adult wings and other body parts of the antibody-treated individuals at various degrees of fluorescence. Importantly, we obtained completely developed wings with a normal color pattern, in which fluorescent signals originated directly from scales or the basal membrane after the removal of scales. GFP fluorescence in wing tissues spatially coincided with anti-GFP antibody staining, confirming that the fluorescent signals originated from the expressed GFP molecules. Our baculovirus-mediated gene transfer system with an anti-gp64 antibody is reasonably efficient, and it can be an invaluable tool to transfer, express, and functionally examine foreign genes in butterfly wings and also in other non-model insect systems.
A computational study on the influence of insect wing geometry on bee flight mechanics
Feaster, Jeffrey; Bayandor, Javid
2017-01-01
ABSTRACT Two-dimensional computational fluid dynamics (CFD) is applied to better understand the effects of wing cross-sectional morphology on flow field and force production. This study investigates the influence of wing cross-section on insect scale flapping flight performance, for the first time, using a morphologically representative model of a bee (Bombus pensylvanicus) wing. The bee wing cross-section was determined using a micro-computed tomography scanner. The results of the bee wing are compared with flat and elliptical cross-sections, representative of those used in modern literature, to determine the impact of profile variation on aerodynamic performance. The flow field surrounding each cross-section and the resulting forces are resolved using CFD for a flight speed range of 1 to 5 m/s. A significant variation in vortex formation is found when comparing the ellipse and flat plate with the true bee wing. During the upstroke, the bee and approximate wing cross-sections have a much shorter wake structure than the flat plate or ellipse. During the downstroke, the flat plate and elliptical cross-sections generate a single leading edge vortex, while the approximate and bee wings generate numerous, smaller structures that are shed throughout the stroke. Comparing the instantaneous aerodynamic forces on the wing, the ellipse and flat plate sections deviate progressively with velocity from the true bee wing. Based on the present findings, a simplified cross-section of an insect wing can misrepresent the flow field and force production. We present the first aerodynamic study using a true insect wing cross-section and show that the wing corrugation increases the leading edge vortex formation frequency for a given set of kinematics. PMID:29061734
A computational study on the influence of insect wing geometry on bee flight mechanics.
Feaster, Jeffrey; Battaglia, Francine; Bayandor, Javid
2017-12-15
Two-dimensional computational fluid dynamics (CFD) is applied to better understand the effects of wing cross-sectional morphology on flow field and force production. This study investigates the influence of wing cross-section on insect scale flapping flight performance, for the first time, using a morphologically representative model of a bee ( Bombus pensylvanicus ) wing. The bee wing cross-section was determined using a micro-computed tomography scanner. The results of the bee wing are compared with flat and elliptical cross-sections, representative of those used in modern literature, to determine the impact of profile variation on aerodynamic performance. The flow field surrounding each cross-section and the resulting forces are resolved using CFD for a flight speed range of 1 to 5 m/s. A significant variation in vortex formation is found when comparing the ellipse and flat plate with the true bee wing. During the upstroke, the bee and approximate wing cross-sections have a much shorter wake structure than the flat plate or ellipse. During the downstroke, the flat plate and elliptical cross-sections generate a single leading edge vortex, while the approximate and bee wings generate numerous, smaller structures that are shed throughout the stroke. Comparing the instantaneous aerodynamic forces on the wing, the ellipse and flat plate sections deviate progressively with velocity from the true bee wing. Based on the present findings, a simplified cross-section of an insect wing can misrepresent the flow field and force production. We present the first aerodynamic study using a true insect wing cross-section and show that the wing corrugation increases the leading edge vortex formation frequency for a given set of kinematics. © 2017. Published by The Company of Biologists Ltd.
Yang, Shuyi; Grall, Aurélie; Chapman, Mark A
2018-05-01
For many crops, research into the origin and partitioning of genetic variation is limited and this can slow or prevent crop improvement programs. Many of these underutilized crops have traits that could be of benefit in a changing climate due to stress tolerance or nutritional properties. Winged bean (Psophocarpus tetragonolobus (L.) DC.) is one such crop. All parts of the plant can be eaten, from the roots to the seeds, and is high in protein as well as other micronutrients. The goal of our study was to identify the wild progenitor and analyze the partitioning of genetic variation in the crop. We used molecular phylogenetic analyses (cpDNA and nuclear ITS sequencing) to resolve relationships between all species in the genus, and population genetics (utilizing microsatellites) to identify genetic clusters of winged bean accessions and compare this to geography. We find that winged bean is genetically distinct from all other members of the genus. We also provide support for four groups of species in the genus, largely, but not completely, corresponding to the results of previous morphological analyses. Within winged bean, population genetic analysis using 10 polymorphic microsatellite markers suggests four genetic groups; however, there is little correspondence between the genetic variation and the geography of the accessions. The true wild progenitor of winged bean remains unknown (or is extinct). There has likely been large-scale cross-breeding, trade, and transport of winged bean and/or multiple origins of the crop. © 2018 Botanical Society of America.
Baxter, S W; Hoffman, J I; Tregenza, T; Wedell, N; Hosken, D J
2017-01-01
Understanding selection in the wild remains a major aim of evolutionary ecology and work by Ford and colleagues on the meadow brown butterfly Maniola jurtina did much to ignite this agenda. A great deal of their work was conducted during the 1950s on the Isles of Scilly. They documented island-specific wing-spot patterns that remained consistent over about a decade, but patterns on some islands changed after environmental perturbation. It was suggested that these wing-spot patterns reflected island-specific selection and that there was little migration between islands. However, genetic studies to test the underlying assumption of restricted migration are lacking and it is also unknown whether the originally described wing-spot patterns have persisted over time. We therefore collected female butterflies from five of Ford's original study locations, including three large islands (St Mary's, St Martin's and Tresco) and two small islands (Tean and St Helen's). Wing-spot patterns had not changed appreciably over time on three of the islands (two large and one small), but were significantly different on the other two. Furthermore, analysis of 176 amplified fragment length polymorphisms revealed significant genome-wide differentiation among the five islands. Our findings are consistent with Ford's conclusions that despite the close proximity of these islands, there is restricted gene flow among them. PMID:27804964
What shapes the continuum of reproductive isolation? Lessons from Heliconius butterflies.
Mérot, C; Salazar, C; Merrill, R M; Jiggins, C D; Joron, M
2017-06-14
The process by which species evolve can be illuminated by investigating barriers that limit gene flow between taxa. Recent radiations, such as Heliconius butterflies, offer the opportunity to compare isolation between pairs of taxa at different stages of ecological, geographical, and phylogenetic divergence. Here, we report a comparative analysis of existing and novel data in order to quantify the strength and direction of isolating barriers within a well-studied clade of Heliconius Our results highlight that increased divergence is associated with the accumulation of stronger and more numerous barriers to gene flow. Wing pattern is both under natural selection for Müllerian mimicry and involved in mate choice, and therefore underlies several isolating barriers. However, pairs which share a similar wing pattern also display strong reproductive isolation mediated by traits other than wing pattern. This suggests that, while wing pattern is a key factor for early stages of divergence, it may become facultative at later stages of divergence. Additional factors including habitat partitioning, hybrid sterility, and chemically mediated mate choice are associated with complete speciation. Therefore, although most previous work has emphasized the role of wing pattern, our comparative results highlight that speciation is a multi-dimensional process, whose completion is stabilized by many factors. © 2017 The Author(s).
Interplay of cell dynamics and epithelial tension during morphogenesis of the Drosophila pupal wing
Etournay, Raphaël; Popović, Marko; Merkel, Matthias; Nandi, Amitabha; Blasse, Corinna; Aigouy, Benoît; Brandl, Holger; Myers, Gene; Salbreux, Guillaume; Jülicher, Frank; Eaton, Suzanne
2015-01-01
How tissue shape emerges from the collective mechanical properties and behavior of individual cells is not understood. We combine experiment and theory to study this problem in the developing wing epithelium of Drosophila. At pupal stages, the wing-hinge contraction contributes to anisotropic tissue flows that reshape the wing blade. Here, we quantitatively account for this wing-blade shape change on the basis of cell divisions, cell rearrangements and cell shape changes. We show that cells both generate and respond to epithelial stresses during this process, and that the nature of this interplay specifies the pattern of junctional network remodeling that changes wing shape. We show that patterned constraints exerted on the tissue by the extracellular matrix are key to force the tissue into the right shape. We present a continuum mechanical model that quantitatively describes the relationship between epithelial stresses and cell dynamics, and how their interplay reshapes the wing. DOI: http://dx.doi.org/10.7554/eLife.07090.001 PMID:26102528
Color pattern evolution in Vanessa butterflies (Nymphalidae: Nymphalini): non-eyespot characters.
Abbasi, Roohollah; Marcus, Jeffrey M
2015-01-01
A phylogenetic approach was used to study color pattern evolution in Vanessa butterflies. Twenty-four color pattern elements from the Nymphalid ground plan were identified on the dorsal and ventral surfaces of the fore- and hind wings. Eyespot characters were excluded and will be examined elsewhere. The evolution of each character was traced over a Bayesian phylogeny of Vanessa reconstructed from 7750 DNA base pairs from 10 genes. Generally, the correspondence between character states on the same surface of the two wings is stronger on the ventral side compared to the dorsal side. The evolution of character states on both sides of a wing correspond with each other in most extant species, but the correspondence between dorsal and ventral character states is much stronger in the forewing than in the hindwing. The dorsal hindwing of many species of Vanessa is covered with an extended Basal Symmetry System and the Discalis I pattern element is highly variable between species, making this wing surface dissimilar to the other wing surfaces. The Basal Symmetry System and Discalis I may contribute to behavioral thermoregulation in Vanessa. Overall, interspecific directional character state evolution of non-eyespot color patterns is relatively rare in Vanessa, with a majority of color pattern elements showing non-variable, non-directional, or ambiguous character state evolution. The ease with which the development of color patterns can be modified, including character state reversals, has likely made important contributions to the production of color pattern diversity in Vanessa and other butterfly groups. © 2014 Wiley Periodicals, Inc.
Size and shape in Melipona quadrifasciata anthidioides Lepeletier, 1836 (Hymenoptera; Meliponini).
Nunes, L A; Passos, G B; Carvalho, C A L; Araújo, E D
2013-11-01
This study aimed to identify differences in wing shape among populations of Melipona quadrifasciata anthidioides obtained in 23 locations in the semi-arid region of Bahia state (Brazil). Analysis of the Procrustes distances among mean wing shapes indicated that population structure did not determine shape variation. Instead, populations were structured geographically according to wing size. The Partial Mantel Test between morphometric (shape and size) distance matrices and altitude, taking geographic distances into account, was used for a more detailed understanding of size and shape determinants. A partial Mantel test between morphometris (shape and size) variation and altitude, taking geographic distances into account, revealed that size (but not shape) is largely influenced by altitude (r = 0.54 p < 0.01). These results indicate greater evolutionary constraints for the shape variation, which must be directly associated with aerodynamic issues in this structure. The size, however, indicates that the bees tend to have larger wings in populations located at higher altitudes.
Encountering bird alarms in full-stare IRSTs
NASA Astrophysics Data System (ADS)
de Jong, Arie N.; Winkel, Hans; Kemp, Rob A. W.
2000-12-01
Birds are a potential source of frequent false alarms in Infrared Search and Track (IRST) systems. One reason is that the signals, generated by birds at short ranges (1-2 km) in IR sensors may be of the same magnitude as the signals generated by real targets (missiles) at long ranges (10-20 km). Another reason is that new generations of IRSTs have more sensitivity which brings more birds within the detection range. Furthermore military operations tend to be held more and more in coastal zones, where the frequency of occurrence of birds is greater than in the open ocean. Finally, the variety in type of birds and their flight characteristics and signature is larger. In the paper attention is spent on the IR signatures of birds in various backgrounds, including rapid variations in signature due to wing motions. Basically, these fluctuations and the flight pattern of a bird provide opportunities to encounter bird alarms in next generation IRSTs, using multiple Focal Plan Array cameras with high frame rates. One has to take into account in this process the difference between signal variations due to wing motions and scintillation for long range targets above the horizon.
Patterning of a compound eye on an extinct dipteran wing.
Dinwiddie, April; Rachootin, Stan
2011-04-23
We have discovered unexpected similarities between a novel and characteristic wing organ in an extinct biting midge from Baltic amber, Eohelea petrunkevitchi, and the surface of a dipteran's compound eye. Scanning electron microscope images now reveal vestigial mechanoreceptors between the facets of the organ. We interpret Eohelea's wing organ as the blending of these two developmental systems: the formation and patterning of the cuticle in the eye and of the wing. Typically, only females in the genus carry this distinctive, highly organized structure. Two species were studied (E. petrunkevitchi and E. sinuosa), and the structure differs in form between them. We examine Eohelea's wing structures for modes of fabrication, material properties and biological functions, and the effective ecological environment in which these midges lived. We argue that the current view of the wing organ's function in stridulation has been misconstrued since it was described half a century ago.
NASA Technical Reports Server (NTRS)
Thompson, D.; Mogili, P.; Chalasani, S.; Addy, H.; Choo, Y.
2004-01-01
Steady-state solutions of the Reynolds-averaged Navier-Stokes (RANS) equations were computed using the Colbalt flow solver for a constant-section, rectangular wing based on an extruded two-dimensional glaze ice shape. The one equation Spalart-Allmaras turbulence model was used. The results were compared with data obtained from a recent wind tunnel test. Computed results indicate that the steady RANS solutions do not accurately capture the recirculating region downstream of the ice accretion, even after a mesh refinement. The resulting predicted reattachment is farther downstream than indicated by the experimental data. Additionally, the solutions computed on a relatively coarse baseline mesh had detailed flow characteristics that were different from those computed on the refined mesh or the experimental data. Steady RANS solutions were also computed to investigate the effects of spanwise variation in the ice shape. The spanwise variation was obtained via a bleeding function that merged the ice shape with the clean wing using a sinusoidal spanwise variation. For these configurations, the results predicted for the extruded shape provided conservative estimates for the performance degradation of the wing. Additionally, the spanwise variation in the ice shape and the resulting differences in the flow fields did not significantly change the location of the primary reattachment.
New neotropical species of Trupanea (Diptera: Tephritidae) with unusual wing patterns
USDA-ARS?s Scientific Manuscript database
Four species of Trupanea (Diptera: Tephritidae) with unusual wing patterns are described from the Neotropical Region: T. dimorphica (Argentina), T. fasciata (Argentina), T. polita (Argentina and Bolivia), and T. trivittata (Argentina). Celidosphenella Hendel, 1914 and Melanotrypana Hering, 1944 are ...
Giordani, B F; Andrade, A J; Galati, E A B; Gurgel-Gonçalves, R
2017-12-01
The Lutzomyia subgenus (Diptera: Psychodidae) includes sibling species with morphologically indistinguishable females. The aims of this study were to analyse variations in the size and shape of wings of species within the Lutzomyia subgenus and to assess whether these analyses might be useful in their identification. Wings (n = 733) of 18 species deposited in Brazilian collections were analysed by geometric morphometrics, using other genera and subgenera as outgroups. Shape variation was summarized in multivariate analyses and differences in wing size among species were tested by analysis of variance. The results showed significant variation in the sizes and shapes of wings of different Lutzomyia species. Two clusters within the Lutzomyia subgenus were distinguished in analyses of both males and females. In Cluster 1 (Lutzomyia ischnacantha, Lutzomyia cavernicola, Lutzomyia almerioi, Lutzomyia forattinii, Lutzomyia renei and Lutzomyia battistinii), scores for correct reclassification were high (females, kappa = 0.91; males, kappa = 0.90), whereas in Cluster 2 (Lutzomyia alencari, Lutzomyia ischyracantha, Lutzomyia cruzi, Lutzomyia longipalpis, Lutzomyia gaminarai and Lutzomyia lichyi), scores for correct reclassification were low (females, kappa = 0.42; males, kappa = 0.48). Wing geometry was useful in the identification of some species of the Lutzomyia subgenus, but did not allow the identification of sibling species such as L. longipalpis and L. cruzi. © 2017 The Royal Entomological Society.
Peabody, Nathan C.; Pohl, Jascha B.; Diao, Fengqiu; Vreede, Andrew P.; Sandstrom, David J.; Wang, Howard; Zelensky, Paul K.; White, Benjamin H.
2009-01-01
After emergence, adult flies and other insects select a suitable perch and expand their wings. Wing expansion is governed by the hormone bursicon and can be delayed under adverse environmental conditions. How environmental factors delay bursicon release and alter perch selection and expansion behaviors has not been investigated in detail. Here we provide evidence that in Drosophila the motor programs underlying perch selection and wing expansion have different environmental dependencies. Using physical manipulations, we demonstrate that the decision to perch is based primarily on environmental valuations and is incrementally delayed under conditions of increasing perturbation and confinement. In contrast, the all-or-none motor patterns underlying wing expansion are relatively invariant in length regardless of environmental conditions. Using a novel technique for targeted activation of neurons, we show that the highly stereotyped wing expansion motor patterns can be initiated by stimulation of NCCAP, a small network of central neurons that regulates the release of bursicon. Activation of this network using the cold-sensitive rat TRPM8 channel is sufficient to trigger all essential behavioral and somatic processes required for wing expansion. The delay of wing expansion under adverse circumstances thus couples an environmentally-sensitive decision network to a command-like network that initiates a fixed action pattern. Because NCCAP mediates environmentally-insensitive ecdysis-related behaviors in Drosophila development prior to adult emergence, the study of wing expansion promises insights not only into how networks mediate behavioral choices, but also into how decision networks develop. PMID:19295141
Yamashita, Shinpei; Takigahira, Tomohiro; Takahashi, Kazuo H
2018-06-01
Accumulating evidence suggests that genotype of host insects influences the development of koinobiont endoparasitoids. Although there are many potential genetic variations that lead to the internal body environmental variations of host insects, association between the host genotype and the parasitoid development has not been examined in a genome-wide manner. In the present study, we used highly inbred whole genome sequenced strains of Drosophila melanogaster to associate single nucleotide polymorphisms (SNPs) of host flies with morphological traits of Asobara japonica, a larval-pupal parasitoid wasp that infected those hosts. We quantified the outline shape of the forewings of A. japonica with two major principal components (PC1 and PC2) calculated from Fourier coefficients obtained from elliptic Fourier analysis. We also quantified wing size and estimated wasp survival. We then examined the association between the PC scores, wing size and 1,798,561 SNPs and the association between the estimated wasp survival and 1,790,544 SNPs. As a result, we obtained 22, 24 and 14 SNPs for PC1, PC2 and wing size and four SNPs for the estimated survival with P values smaller than 10 -5 . Based on the location of the SNPs, 12, 17, 11 and five protein coding genes were identified as potential candidates for PC1, PC2, wing size and the estimated survival, respectively. Based on the function of the candidate genes, it is suggested that the host genetic variation associated with the cell growth and morphogenesis may influence the wasp's morphogenetic variation.
NASA Astrophysics Data System (ADS)
Guillermo-Ferreira, Rhainer; Gorb, Stanislav N.; Appel, Esther; Kovalev, Alexander; Bispo, Pitágoras C.
2015-04-01
Wing pigmentation is a trait that predicts the outcome of male contests in some damselflies. Thus, it is reasonable to suppose that males would have the ability to assess wing pigmentation and adjust investment in a fight according to the costs that the rival may potentially impose. Males of the damselfly Mnesarete pudica exhibit red-coloured wings and complex courtship behaviour and engage in striking male-male fights. In this study, we investigated male assessment behaviour during aerial contests. Theory suggests that the relationship between male resource-holding potential (RHP) and contest duration describes the kind of assessment adopted by males: self-assessment, opponent-only assessment or mutual assessment. A recent theory also suggests that weak and strong males exhibit variations in the assessment strategies adopted. We estimated male RHP through male body size and wing colouration (i.e. pigmentation, wing reflectance spectra and transmission spectra) and studied the relationship between male RHP and contest duration from video-documented behavioural observations of naturally occurring individual contests in the field. The results showed that males with more opaque wings and larger red spots were more likely to win contests. The relationships between RHP and contest durations partly supported the self-assessment and the mutual assessment models. We then experimentally augmented the pigmented area of the wings, in order to evaluate whether strong and weak males assess rivals' RHP through wing pigmentation. Our experimental manipulation, however, clearly demonstrated that strong males assess rivals' wing pigmentation. We finally suggest that there is a variation in the assessment strategy adopted by males.
Guillermo-Ferreira, Rhainer; Gorb, Stanislav N; Appel, Esther; Kovalev, Alexander; Bispo, Pitágoras C
2015-04-01
Wing pigmentation is a trait that predicts the outcome of male contests in some damselflies. Thus, it is reasonable to suppose that males would have the ability to assess wing pigmentation and adjust investment in a fight according to the costs that the rival may potentially impose. Males of the damselfly Mnesarete pudica exhibit red-coloured wings and complex courtship behaviour and engage in striking male-male fights. In this study, we investigated male assessment behaviour during aerial contests. Theory suggests that the relationship between male resource-holding potential (RHP) and contest duration describes the kind of assessment adopted by males: self-assessment, opponent-only assessment or mutual assessment. A recent theory also suggests that weak and strong males exhibit variations in the assessment strategies adopted. We estimated male RHP through male body size and wing colouration (i.e. pigmentation, wing reflectance spectra and transmission spectra) and studied the relationship between male RHP and contest duration from video-documented behavioural observations of naturally occurring individual contests in the field. The results showed that males with more opaque wings and larger red spots were more likely to win contests. The relationships between RHP and contest durations partly supported the self-assessment and the mutual assessment models. We then experimentally augmented the pigmented area of the wings, in order to evaluate whether strong and weak males assess rivals' RHP through wing pigmentation. Our experimental manipulation, however, clearly demonstrated that strong males assess rivals' wing pigmentation. We finally suggest that there is a variation in the assessment strategy adopted by males.
Keeping the band together: evidence for false boundary disruptive coloration in a butterfly.
Seymoure, B M; Aiello, A
2015-09-01
There is a recent surge of evidence supporting disruptive coloration, in which patterns break up the animal's outline through false edges or boundaries, increasing survival in animals by reducing predator detection and/or preventing recognition. Although research has demonstrated that false edges are successful for reducing predation of prey, research into the role of internal false boundaries (i.e. stripes and bands) in reducing predation remains warranted. Many animals have stripes and bands that may function disruptively. Here, we test the possible disruptive function of wing band patterning in a butterfly, Anartia fatima, using artificial paper and plasticine models in Panama. We manipulated the band so that one model type had the band shifted to the wing margin (nondisruptive treatment) and another model had a discontinuous band located on the wing margin (discontinuous edge treatment). We kept the natural wing pattern to represent the false boundary treatment. Across all treatment groups, we standardized the area of colour and used avian visual models to confirm a match between manipulated and natural wing colours. False boundary models had higher survival than either the discontinuous edge model or the nondisruptive model. There was no survival difference between the discontinuous edge model and the nondisruptive model. Our results demonstrate the importance of wing bands in reducing predation on butterflies and show that markings set in from the wing margin can reduce predation more effectively than marginal bands and discontinuous marginal patterns. This study demonstrates an adaptive benefit of having stripes and bands. © 2015 European Society For Evolutionary Biology.
Zhang, Linlin
2017-01-01
The optix gene has been implicated in butterfly wing pattern adaptation by genetic association, mapping, and expression studies. The actual developmental function of this gene has remained unclear, however. Here we used CRISPR/Cas9 genome editing to show that optix plays a fundamental role in nymphalid butterfly wing pattern development, where it is required for determination of all chromatic coloration. optix knockouts in four species show complete replacement of color pigments with melanins, with corresponding changes in pigment-related gene expression, resulting in black and gray butterflies. We also show that optix simultaneously acts as a switch gene for blue structural iridescence in some butterflies, demonstrating simple regulatory coordination of structural and pigmentary coloration. Remarkably, these optix knockouts phenocopy the recurring “black and blue” wing pattern archetype that has arisen on many independent occasions in butterflies. Here we demonstrate a simple genetic basis for structural coloration, and show that optix plays a deeply conserved role in butterfly wing pattern development. PMID:28923944
Zhang, Linlin; Mazo-Vargas, Anyi; Reed, Robert D
2017-10-03
The optix gene has been implicated in butterfly wing pattern adaptation by genetic association, mapping, and expression studies. The actual developmental function of this gene has remained unclear, however. Here we used CRISPR/Cas9 genome editing to show that optix plays a fundamental role in nymphalid butterfly wing pattern development, where it is required for determination of all chromatic coloration. optix knockouts in four species show complete replacement of color pigments with melanins, with corresponding changes in pigment-related gene expression, resulting in black and gray butterflies. We also show that optix simultaneously acts as a switch gene for blue structural iridescence in some butterflies, demonstrating simple regulatory coordination of structural and pigmentary coloration. Remarkably, these optix knockouts phenocopy the recurring "black and blue" wing pattern archetype that has arisen on many independent occasions in butterflies. Here we demonstrate a simple genetic basis for structural coloration, and show that optix plays a deeply conserved role in butterfly wing pattern development.
Wing shape allometry and aerodynamics in calopterygid damselflies: a comparative approach.
Outomuro, David; Adams, Dean C; Johansson, Frank
2013-06-07
Wing size and shape have important aerodynamic implications on flight performance. We explored how wing size was related to wing shape in territorial males of 37 taxa of the damselfly family Calopterygidae. Wing coloration was also included in the analyses because it is sexually and naturally selected and has been shown to be related to wing shape. We studied wing shape using both the non-dimensional radius of the second moment of wing area (RSM) and geometric morphometrics. Lower values of the RSM result in less energetically demanding flight and wider ranges of flight speed. We also re-analyzed previously published data on other damselflies and dragonflies. The RSM showed a hump-shaped relationship with wing size. However, after correcting for phylogeny using independent contrast, this pattern changed to a negative linear relationship. The basal genus of the study family, Hetaerina, was mainly driving that change. The obtained patterns were specific for the study family and differed from other damselflies and dragonflies. The relationship between the RSM and wing shape measured by geometric morphometrics was linear, but relatively small changes along the RSM axis can result in large changes in wing shape. Our results also showed that wing coloration may have some effect on RSM. We found that RSM showed a complex relationship with size in calopterygid damselflies, probably as a result of other selection pressures besides wing size per se. Wing coloration and specific behavior (e.g. courtship) are potential candidates for explaining the complexity. Univariate measures of wing shape such as RSM are more intuitive but lack the high resolution of other multivariate techniques such as geometric morphometrics. We suggest that the relationship between wing shape and size are taxa-specific and differ among closely-related insect groups.
NASA Technical Reports Server (NTRS)
Bobbitt, Percy J.
1959-01-01
Equations for the downwash and sidewash due to supersonic yawed and unswept horseshoe vortices have been utilized in formulating tables and charts to permit a rapid estimation of the flow velocities behind wings performing various steady motions. Tabulations are presented of the downwash and sidewash in the wing vertical plane of symmetry due to a unit-strength yawed horseshoe vortex located at 20 equally spaced spanwise positions along lifting lines of various sweeps. (The bound portion of the yawed vortex is coincident with the lifting line.) Charts are presented for the purpose of estimating the spanwise variations of the flow-field velocities and give longitudinal variations of the downwash and sidewash at a nuMber of vertical and spanwise locations due to a unit-strength unswept horseshoe vortex. Use of the tables and charts to calculate wing downwash or sidewash requires a knowledge of the wing spanwise distribution of circulation. Sample computations for the rolling sidewash and angle-of-attack downwash behind a typical swept wing are presented to demonstrate the use of the tables and charts.
Liu, Gui-Feng; Zang, Run-Guo; Liu, Hua; Bai, Zhi-Qiang; Guo, Zhong-Jun; Ding, Yi
2012-06-01
Taking the Picea schrenkiana var. tianschanica forests at three sites with different longitudes (Zhaosu, Tianchi, and Qitai) in Tianshan Mountains as the objects, the cones were collected along an altitudinal gradient to analyze the variation of their seed morphological traits (seed scale length and width, seed scale length/width ratio, seed wing length and width, seed wing length/ width ratio, seed length and width, and seed length/width ratio). All the seed traits except seed width tended to decrease with increasing altitude. The seed traits except seed wing width, seed width, and seed length/width ratio all had significant negative correlations with altitude. Seed scale length and width and seed scale length/width ratio had significant positive correlations with longitude. Seed scale length, seed scale length/width ratio, and seed wing length/width ratio had significant negative correlations with slope degree. No significant correlations were observed between the seed traits except seed wing width and the slope aspect. Altitude was the main factor affecting the seed scale length, seed scale length/width ratio, and seed wing length/width ratio.
Genomic Signatures of Speciation in Sympatric and Allopatric Hawaiian Picture-Winged Drosophila.
Kang, Lin; Settlage, Robert; McMahon, Wyatt; Michalak, Katarzyna; Tae, Hongseok; Garner, Harold R; Stacy, Elizabeth A; Price, Donald K; Michalak, Pawel
2016-05-30
The Hawaiian archipelago provides a natural arena for understanding adaptive radiation and speciation. The Hawaiian Drosophila are one of the most diverse endemic groups in Hawaiì with up to 1,000 species. We sequenced and analyzed entire genomes of recently diverged species of Hawaiian picture-winged Drosophila, Drosophila silvestris and Drosophila heteroneura from Hawaiì Island, in comparison with Drosophila planitibia, their sister species from Maui, a neighboring island where a common ancestor of all three had likely occurred. Genome-wide single nucleotide polymorphism patterns suggest the more recent origin of D. silvestris and D. heteroneura, as well as a pervasive influence of positive selection on divergence of the three species, with the signatures of positive selection more prominent in sympatry than allopatry. Positively selected genes were significantly enriched for functional terms related to sensory detection and mating, suggesting that sexual selection played an important role in speciation of these species. In particular, sequence variation in Olfactory receptor and Gustatory receptor genes seems to play a major role in adaptive radiation in Hawaiian pictured-winged Drosophila. © The Author 2016. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution.
Donovan, Edward R; Keeney, Brooke K; Kung, Eric; Makan, Sirish; Wild, J Martin; Altshuler, Douglas L
2013-01-01
Flying animals exhibit profound transformations in anatomy, physiology, and neural architecture. Although much is known about adaptations in the avian skeleton and musculature, less is known about neuroanatomy and motor unit integration for bird flight. Hummingbirds are among the most maneuverable and specialized of vertebrate fliers, and two unusual neuromuscular features have been previously reported: (1) the pectoralis major has a unique distribution pattern of motor end plates (MEPs) compared with all other birds and (2) electromyograms (EMGs) from the hummingbird's pectoral muscles, the pectoralis major and the supracoracoideus, show activation bursts composed of one or a few spikes that appear to have a very consistent pattern. Here, we place these findings in a broader context by comparing the MEPs, EMGs, and organization of the spinal motor neuron pools of flight muscles of Anna's hummingbird Calypte anna, zebra finches Taeniopygia guttata, and, for MEPs, several other species. The previously shown MEP pattern of the hummingbird pectoralis major is not shared with its closest taxonomic relative, the swift, and appears to be unique to hummingbirds. MEP arrangements in previously undocumented wing muscles show patterns that differ somewhat from other avian muscles. In the parallel-fibered strap muscles of the shoulder, MEP patterns appear to relate to muscle length, with the smallest muscles having fibers that span the entire muscle. MEP patterns in pennate distal wing muscles were the same regardless of size, with tightly clustered bands in the middle portion of the muscle, not evenly distributed bands over the muscle's entire length. Muscle activations were examined during slow forward flight in both species, during hovering in hummingbirds, and during slow ascents in zebra finches. The EMG bursts of a wing muscle, the pronator superficialis, were highly variable in peak number, size, and distribution across wingbeats for both species. In the pectoralis major, although the individual EMG bursts were much shorter in duration in hummingbirds relative to zebra finches, the variables describing the normalized amplitude and area of the activation bursts were otherwise indistinguishable between taxa during these flight modes. However, the degree of variation in the time intervals between EMG peaks was much lower in hummingbirds, which is a plausible explanation for the "patterned" EMG signals reported previously.
Schöneich, Stefan; Hedwig, Berthold
2012-01-01
The singing behavior of male crickets allows analyzing a central pattern generator (CPG) that was shaped by sexual selection for reliable production of species-specific communication signals. After localizing the essential ganglia for singing in Gryllus bimaculatus, we now studied the calling song CPG at the cellular level. Fictive singing was initiated by pharmacological brain stimulation. The motor pattern underlying syllables and chirps was recorded as alternating spike bursts of wing-opener and wing-closer motoneurons in a truncated wing nerve; it precisely reflected the natural calling song. During fictive singing, we intracellularly recorded and stained interneurons in thoracic and abdominal ganglia and tested their impact on the song pattern by intracellular current injections. We identified three interneurons of the metathoracic and first unfused abdominal ganglion that rhythmically de- and hyperpolarized in phase with the syllable pattern and spiked strictly before the wing-opener motoneurons. Depolarizing current injection in two of these opener interneurons caused additional rhythmic singing activity, which reliably reset the ongoing chirp rhythm. The closely intermeshing arborizations of the singing interneurons revealed the dorsal midline neuropiles of the metathoracic and three most anterior abdominal neuromeres as the anatomical location of singing pattern generation. In the same neuropiles, we also recorded several closer interneurons that rhythmically hyper- and depolarized in the syllable rhythm and spiked strictly before the wing-closer motoneurons. Some of them received pronounced inhibition at the beginning of each chirp. Hyperpolarizing current injection in the dendrite revealed postinhibitory rebound depolarization as one functional mechanism of central pattern generation in singing crickets. PMID:23170234
2013-01-01
Background Candidate genes for color pattern formation in butterfly wings have been known based on gene expression patterns since the 1990s, but their functions remain elusive due to a lack of a functional assay. Several methods of transferring and expressing a foreign gene in butterfly wings have been reported, but they have suffered from low success rates or low expression levels. Here, we developed a simple, practical method to efficiently deliver and express a foreign gene using baculovirus-mediated gene transfer in butterfly wings in vivo. Results A recombinant baculovirus containing a gene for green fluorescent protein (GFP) was injected into pupae of the blue pansy butterfly Junonia orithya (Nymphalidae). GFP fluorescence was detected in the pupal wings and other body parts of the injected individuals three to five days post-injection at various degrees of fluorescence. We obtained a high GFP expression rate at relatively high virus titers, but it was associated with pupal death before color pattern formation in wings. To reduce the high mortality rate caused by the baculovirus treatment, we administered an anti-gp64 antibody, which was raised against baculovirus coat protein gp64, to infected pupae after the baculovirus injection. This treatment greatly reduced the mortality rate of the infected pupae. GFP fluorescence was observed in pupal and adult wings and other body parts of the antibody-treated individuals at various degrees of fluorescence. Importantly, we obtained completely developed wings with a normal color pattern, in which fluorescent signals originated directly from scales or the basal membrane after the removal of scales. GFP fluorescence in wing tissues spatially coincided with anti-GFP antibody staining, confirming that the fluorescent signals originated from the expressed GFP molecules. Conclusions Our baculovirus-mediated gene transfer system with an anti-gp64 antibody is reasonably efficient, and it can be an invaluable tool to transfer, express, and functionally examine foreign genes in butterfly wings and also in other non-model insect systems. PMID:23522444
Interaction of a trailing vortex with an oscillating wing
NASA Astrophysics Data System (ADS)
McKenna, C.; Fishman, G.; Rockwell, D.
2018-01-01
A technique of particle image velocimetry is employed to characterize the flow structure of a trailing vortex incident upon the tip region of an oscillating wing (plate). The amplitude and velocity of the wing are nearly two orders of magnitude smaller than the wing chord and free stream velocity, respectively. Depending upon the outboard displacement of the incident vortex relative to the wing tip, distinctive patterns of upwash, downwash, and shed vorticity are observed. These patterns are a strong function of the phase of the wing motion during its oscillation cycle. At a given phase, the wing oscillation induces upwash that is reinforced by the upwash of the incident vortex, giving a maximum net upwash. Conversely, when these two origins of upwash counteract, rather than reinforce, one another during the oscillation cycle, the net upwash attains minimum value. Analogous interpretations hold for regions of maximum and minimum net downwash located outboard of the regions of upwash. The magnitude and scale of the vorticity shed from the tip of the wing are directly correlated with the net upwash, which takes different forms related to the outboard displacement of the incident vortex. As the location of the incident vortex is displaced towards the wing tip, both the maximum upwash and the maximum vorticity of the tip vortex initially increase and then decrease. For the limiting case where the incident vortex impinges directly upon the tip of the wing, there is no tip vortex or induced region of upwash. Furthermore, at small values of vortex displacement from the wing tip, the position of the incident vortex varies significantly from its nominal position during the oscillation cycle. All of the foregoing features are interpreted in conjunction with the flow topology in the form of streamlines and critical points, superposed on patterns of vorticity. It is shown that despite the small amplitude of the wing motion, the flow topology is fundamentally different at maximum positive and negative values of the velocity of the wing tip, that is, they are not symmetric.
6. DETAIL OF MASONRY ON SOUTHWEST WING WALL. MASONRY ON ...
6. DETAIL OF MASONRY ON SOUTHWEST WING WALL. MASONRY ON WING WALLS IS LAID IN A RANDOM RUBBLE PATTERN. - Core Creek County Bridge, Spanning Core Creek, approximately 1 mile South of State Route 332 (Newtown Bypass), Newtown, Bucks County, PA
Biological plasticity in penguin heat-retention structures.
Thomas, Daniel B; Fordyce, R Ewan
2012-02-01
Insulation and vascular heat-retention mechanisms allow penguins to forage for a prolonged time in water that is much cooler than core body temperature. Wing-based heat retention involves a plexus of humeral arteries and veins, which redirect heat to the body core rather than to the wing periphery. The humeral arterial plexus is described here for Eudyptes and Megadyptes, the only extant penguin genera for which wing vascular anatomy had not previously been reported. The erect-crested (Eudyptes sclateri) and yellow-eyed (Megadyptes antipodes) penguins both have a plexus of three humeral arteries on the ventral surface of the humerus. The wing vascular system shows little variation between erect-crested and yellow-eyed penguins, and is generally conserved across the six extant genera of penguins, with the exception of the humeral arterial plexus. The number of humeral arteries within the plexus demonstrates substantial variation and correlates well with wing surface area. Little penguins (Eudyptula minor) have two humeral arteries and a wing surface area of ∼ 75 cm(2) , whereas emperor penguins (Aptenodytes forsteri) have up to 15 humeral arteries and a wing surface area of ∼ 203 cm(2) . Further, the number of humeral arteries has a stronger correlation with wing surface area than with sea water temperature. We propose that thermoregulation has placed the humeral arterial plexus under a strong selection pressure, driving penguins with larger wing surface areas to compensate for heat loss by developing additional humeral arteries. Copyright © 2011 Wiley Periodicals, Inc.
Derivation of charts for determining the horizontal tail load variation with any elevator motion
NASA Technical Reports Server (NTRS)
Pearson, Henry A
1943-01-01
The equations relating the wing and tail loads are derived for a unit elevator displacement. These equations are then converted into a nondimensional form and charts are given by which the wing- and tail-load-increment variation may be determined under dynamic conditions for any type of elevator motion and for various degrees of airplane stability. In order to illustrate the use of the charts, several examples are included in which the wing and tail loads are evaluated for a number of types of elevator motion. Methods are given for determining the necessary derivatives from results of wind-tunnel tests when such tests are available.
NASA Technical Reports Server (NTRS)
Mugler, John P., Jr.
1959-01-01
Pressure distributions obtained in the Langley 8-foot transonic pressure tunnel on a thin, highly tapered, twisted, 45 deg sweptback wing in combination with a body are presented. The wing has a linear span-wise twist variation from 0 deg at 10 percent of the semispan to 6 deg at the tip. The tip is at a lower angle of attack than the root. Tests were made at stagnation pressures of 1.0 and 0.5 atmosphere, at Mach numbers from 0.800 to 1.200, and at angles of attack from -4 to 12 deg.
Effect of external jet-flow deflector geometry on OTW aero-acoustic characteristics
NASA Technical Reports Server (NTRS)
Vonglahn, U.; Groesbeck, D.
1976-01-01
The effect of geometry variations in the design of external deflectors for use with over-the-wing (OTW) configurations was studied at model scale and subsonic jet velocities. Included in the variations were deflector size and angle as well as wing size and flap setting. A conical nozzle (5.2-cm diameter) mounted at 0.1 chord above and downstream of the wing leading edges was used. The data indicate that external deflectors provide satisfactory takeoff and approach aerodynamic performance and acoustic characteristics for OTW configurations. These characteristics together with expected good cruise aerodynamics, since external deflectors are storable, may provide optimum OTW design configurations.
Sweet, Andrew D.; Chesser, R. Terry; Johnson, Kevin P.
2017-01-01
Host–parasite coevolutionary histories can differ among multiple groups of parasites associated with the same group of hosts. For example, parasitic wing and body lice (Insecta: Phthiraptera) of New World pigeons and doves (Aves: Columbidae) differ in their cophylogenetic patterns, with body lice exhibiting higher phylogenetic congruence with their hosts than wing lice. In this study, we focus on the wing and body lice of Australian phabine pigeons and doves to determine whether the patterns in New World pigeons and doves are consistent with those of pigeons and doves from other regions. Using molecular sequence data for most phabine species and their lice, we estimated phylogenetic trees for all three groups (pigeons and doves, wing lice and body lice), and compared the phabine (host) tree with both parasite trees using multiple cophylogenetic methods. We found a pattern opposite to that found for New World pigeons and doves, with Australian wing lice showing congruence with their hosts, and body lice exhibiting a lack of congruence. There are no documented records of hippoboscid flies associated with Australian phabines, thus these lice may lack the opportunity to disperse among host species by attaching to hippoboscid flies (phoresis), which could explain these patterns. However, additional sampling for flies is needed to confirm this hypothesis. Large differences in body size among phabine pigeons and doves may also help to explain the congruence of the wing lice with their hosts. It may be more difficult for wing lice than body lice to switch among hosts that vary more dramatically in size. The results from this study highlight how host–parasite coevolutionary histories can vary by region, and how local factors can shape the relationship.
Sweet, Andrew D; Chesser, R Terry; Johnson, Kevin P
2017-05-01
Host-parasite coevolutionary histories can differ among multiple groups of parasites associated with the same group of hosts. For example, parasitic wing and body lice (Insecta: Phthiraptera) of New World pigeons and doves (Aves: Columbidae) differ in their cophylogenetic patterns, with body lice exhibiting higher phylogenetic congruence with their hosts than wing lice. In this study, we focus on the wing and body lice of Australian phabine pigeons and doves to determine whether the patterns in New World pigeons and doves are consistent with those of pigeons and doves from other regions. Using molecular sequence data for most phabine species and their lice, we estimated phylogenetic trees for all three groups (pigeons and doves, wing lice and body lice), and compared the phabine (host) tree with both parasite trees using multiple cophylogenetic methods. We found a pattern opposite to that found for New World pigeons and doves, with Australian wing lice showing congruence with their hosts, and body lice exhibiting a lack of congruence. There are no documented records of hippoboscid flies associated with Australian phabines, thus these lice may lack the opportunity to disperse among host species by attaching to hippoboscid flies (phoresis), which could explain these patterns. However, additional sampling for flies is needed to confirm this hypothesis. Large differences in body size among phabine pigeons and doves may also help to explain the congruence of the wing lice with their hosts. It may be more difficult for wing lice than body lice to switch among hosts that vary more dramatically in size. The results from this study highlight how host-parasite coevolutionary histories can vary by region, and how local factors can shape the relationship. Copyright © 2017 Australian Society for Parasitology. All rights reserved.
2014-01-01
Background The colorful wing patterns of butterflies, a prime example of biodiversity, can change dramatically within closely related species. Wing pattern diversity is specifically present among papilionid butterflies. Whether a correlation between color and the evolution of these butterflies exists so far remained unsolved. Results We here investigate the Cattlehearts, Parides, a small Neotropical genus of papilionid butterflies with 36 members, the wings of which are marked by distinctly colored patches. By applying various physical techniques, we investigate the coloration toolkit of the wing scales. The wing scales contain two different, wavelength-selective absorbing pigments, causing pigmentary colorations. Scale ridges with multilayered lamellae, lumen multilayers or gyroid photonic crystals in the scale lumen create structural colors that are variously combined with these pigmentary colors. Conclusions The pigmentary and structural traits strongly correlate with the taxonomical distribution of Parides species. The experimental findings add crucial insight into the evolution of butterfly wing scales and show the importance of morphological parameter mapping for butterfly phylogenetics. PMID:25064167
Wu, Jun; Yu, Zhijing; Wang, Tao; Zhuge, Jingchang; Ji, Yue; Xue, Bin
2017-06-01
Airplane wing deformation is an important element of aerodynamic characteristics, structure design, and fatigue analysis for aircraft manufacturing, as well as a main test content of certification regarding flutter for airplanes. This paper presents a novel real-time detection method for wing deformation and flight flutter detection by using three-dimensional speckle image correlation technology. Speckle patterns whose positions are determined through the vibration characteristic of the aircraft are coated on the wing; then the speckle patterns are imaged by CCD cameras which are mounted inside the aircraft cabin. In order to reduce the computation, a matching technique based on Geodetic Systems Incorporated coded points combined with the classical epipolar constraint is proposed, and a displacement vector map for the aircraft wing can be obtained through comparing the coordinates of speckle points before and after deformation. Finally, verification experiments containing static and dynamic tests by using an aircraft wing model demonstrate the accuracy and effectiveness of the proposed method.
Three-Dimensional Piecewise-Continuous Class-Shape Transformation of Wings
NASA Technical Reports Server (NTRS)
Olson, Erik D.
2015-01-01
Class-Shape Transformation (CST) is a popular method for creating analytical representations of the surface coordinates of various components of aerospace vehicles. A wide variety of two- and three-dimensional shapes can be represented analytically using only a modest number of parameters, and the surface representation is smooth and continuous to as fine a degree as desired. This paper expands upon the original two-dimensional representation of airfoils to develop a generalized three-dimensional CST parametrization scheme that is suitable for a wider range of aircraft wings than previous formulations, including wings with significant non-planar shapes such as blended winglets and box wings. The method uses individual functions for the spanwise variation of airfoil shape, chord, thickness, twist, and reference axis coordinates to build up the complete wing shape. An alternative formulation parameterizes the slopes of the reference axis coordinates in order to relate the spanwise variation to the tangents of the sweep and dihedral angles. Also discussed are methods for fitting existing wing surface coordinates, including the use of piecewise equations to handle discontinuities, and mathematical formulations of geometric continuity constraints. A subsonic transport wing model is used as an example problem to illustrate the application of the methodology and to quantify the effects of piecewise representation and curvature constraints.
Otaki, Joji M
2008-07-01
A mechanistic understanding of the butterfly wing color-pattern determination can be facilitated by experimental pattern changes. Here I review physiologically induced color-pattern changes in nymphalid butterflies and their mechanistic and evolutionary implications. A type of color-pattern change can be elicited by elemental changes in size and position throughout the wing, as suggested by the nymphalid groundplan. These changes of pattern elements are bi-directional and bi-sided dislocation toward or away from eyespot foci and in both proximal and distal sides of the foci. The peripheral elements are dislocated even in the eyespot-less compartments. Anterior spots are more severely modified, suggesting the existence of an anterior-posterior gradient. In one species, eyespots are transformed into white spots with remnant-like orange scales, and such patterns emerge even at the eyespot-less "imaginary" foci. A series of these color-pattern modifications probably reveal "snap-shots" of a dynamic morphogenic signal due to heterochronic uncoupling between the signaling and reception steps. The conventional gradient model can be revised to account for these observed color-pattern changes.
Takahashi, Kazuo H
2017-02-01
Drosophila wings have been a model system to study the effect of HSP90 on quantitative trait variation. The effect of HSP90 inhibition on environmental buffering of wing morphology varies among studies while the genetic buffering effect of it was examined in only one study and was not detected. Variable results so far might show that the genetic background influences the environmental and genetic buffering effect of HSP90. In the previous studies, the number of the genetic backgrounds used is limited. To examine the effect of HSP90 inhibition with a larger number of genetic backgrounds than the previous studies, 20 wild-type strains of Drosophila melanogaster were used in this study. Here I investigated the effect of HSP90 inhibition on the environmental buffering of wing shape and size by assessing within-individual and among-individual variations, and as a result, I found little or very weak effects on environmental and genetic buffering. The current results suggest that the role of HSP90 as a global regulator of environmental and genetic buffering is limited at least in quantitative traits.
Evolution of antero‐posterior patterning of the limb: Insights from the chick
2017-01-01
Summary The developing limbs of chicken embryos have served as pioneering models for understanding pattern formation for over a century. The ease with which chick wing and leg buds can be experimentally manipulated, while the embryo is still in the egg, has resulted in the discovery of important developmental organisers, and subsequently, the signals that they produce. Sonic hedgehog (Shh) is produced by mesenchyme cells of the polarizing region at the posterior margin of the limb bud and specifies positional values across the antero‐posterior axis (the axis running from the thumb to the little finger). Detailed experimental embryology has revealed the fundamental parameters required to specify antero‐posterior positional values in response to Shh signaling in chick wing and leg buds. In this review, the evolution of the avian wing and leg will be discussed in the broad context of tetrapod paleontology, and more specifically, ancestral theropod dinosaur paleontology. How the parameters that dictate antero‐posterior patterning could have been modulated to produce the avian wing and leg digit patterns will be considered. Finally, broader speculations will be made regarding what the antero‐posterior patterning of chick limbs can tell us about the evolution of other digit patterns, including those that were found in the limbs of the earliest tetrapods. PMID:28734068
Theoretical antisymmetric span loading for wings of arbitrary plan form at subsonic speeds
NASA Technical Reports Server (NTRS)
Deyoung, John
1951-01-01
A simplified lifting-surface theory that includes effects of compressibility and spanwise variation of section lift-curve slope is used to provide charts with which antisymmetric loading due to arbitrary antisymmetric angle of attack can be found for wings having symmetric plan forms with a constant spanwise sweep angle of the quarter-chord line. Consideration is given to the flexible wing in roll. Aerodynamic characteristics due to rolling, deflected ailerons, and sideslip of wings with dihedral are considered. Solutions are presented for straight-tapered wings for a range of swept plan forms.
Chordwise and compressibility corrections to slender-wing theory
NASA Technical Reports Server (NTRS)
Lomax, Harvard; Sluder, Loma
1952-01-01
Corrections to slender-wing theory are obtained by assuming a spanwise distribution of loading and determining the chordwise variation which satisfies the appropriate integral equation. Such integral equations are set up in terms of the given vertical induced velocity on the center line or, depending on the type of wing plan form, its average value across the span at a given chord station. The chordwise distribution is then obtained by solving these integral equations. Results are shown for flat-plate rectangular, and triangular wings.
Lift Production on Flapping and Rotary Wings at Low Reynolds Numbers
2016-02-26
though parameter variations were also performed. For the rotating cases, the wing was an aspect ratio 2 rectangular flat plate , and the root cutout (i.e...rectangular flat plate . 2 U (Side View) (a) 1A: Rectilinear pitch U (Side View) (b) 1B: Rectilinear surge (Top View) (Side View) (c) 2A: Rotational...0.5c φ (b) A=2 flat plate wing Figure 2: Schematic of the AVT-202 rotating wing kinematics and geometry, from Ref. 12. 3.2 Experimental Setup Rotating
Adaptive 3D single-block grids for the computation of viscous flows around wings
DOE Office of Scientific and Technical Information (OSTI.GOV)
Hagmeijer, R.; Kok, J.C.
1996-12-31
A robust algorithm for the adaption of a 3D single-block structured grid suitable for the computation of viscous flows around a wing is presented and demonstrated by application to the ONERA M6 wing. The effects of grid adaption on the flow solution and accuracy improvements is analyzed. Reynolds number variations are studied.
The marginal band system in nymphalid butterfly wings.
Taira, Wataru; Kinjo, Seira; Otaki, Joji M
2015-01-01
Butterfly wing color patterns are highly complex and diverse, but they are believed to be derived from the nymphalid groundplan, which is composed of several color pattern systems. Among these pattern systems, the marginal band system, including marginal and submarginal bands, has rarely been studied. Here, we examined the color pattern diversity of the marginal band system among nymphalid butterflies. Marginal and submarginal bands are usually expressed as a pair of linear bands aligned with the wing margin. However, a submarginal band can be expressed as a broken band, an elongated oval, or a single dot. The marginal focus, usually a white dot at the middle of a wing compartment along the wing edge, corresponds to the pupal edge spot, one of the pupal cuticle spots that signify the locations of color pattern organizing centers. A marginal band can be expressed as a semicircle, an elongated oval, or a pair of eyespot-like structures, which suggest the organizing activity of the marginal focus. Physical damage at the pupal edge spot leads to distal dislocation of the submarginal band in Junonia almana and in Vanessa indica, suggesting that the marginal focus functions as an organizing center for the marginal band system. Taken together, we conclude that the marginal band system is developmentally equivalent to other symmetry systems. Additionally, the marginal band is likely a core element and the submarginal band a paracore element of the marginal band system, and both bands are primarily specified by the marginal focus organizing center.
Ricca, Mark A.; Miles, A. Keith; Anthony, Robert G.
2008-01-01
Persistent organochlorine compounds and mercury (Hg) have been detected in numerous coastal organisms of the Aleutian archipelago of Alaska, yet sources of these contaminants are unclear. We collected glaucous-winged gulls, northern fulmars, and tufted puffins along a natural longitudinal gradient across the western and central Aleutian Islands (Buldir, Kiska, Amchitka, Adak), and an additional 8 seabird species representing different foraging and migratory guilds from Buldir Island to evaluate: 1) point source input from former military installations, 2) westward increases in contaminant concentrations suggestive of distant source input, and 3) effects of trophic status (δ15N) and carbon source (δ13C) on contaminant accumulation. Concentrations of Σ polychlorinated biphenyls (PCBs) and most chlorinated pesticides in glaucous-winged gulls consistently exhibited a ‘U’-shaped pattern of high levels at Buldir and the east side of Adak and low levels at Kiska and Amchitka. In contrast, concentrations of Σ PCBs and chlorinated pesticides in northern fulmars and tufted puffins did not differ among islands. Hg concentrations increased westward in glaucous-winged gulls and were highest in northern fulmars from Buldir. Among species collected only at Buldir, Hg was notably elevated in pelagic cormorants, and relatively high Σ PCBs were detected in black-legged kittiwakes. Concentrations of Σ PCBs, dichlorodiphenyldichloroethylene (p,p′ DDE), and Hg were positively correlated with δ15N across all seabird species, indicating biomagnification across trophic levels. The east side of Adak Island (a former military installation) was a likely point source of Σ PCBs and p,p′ DDE, particularly in glaucous-winged gulls. In contrast, elevated levels of these contaminants and Hg, along with PCB congener and chlorinated pesticide compositional patterns detected at Buldir Island indicated exposure from distant sources influenced by a combination of atmospheric–oceanic processes and the migratory movements of seabirds.
Morphological Variations of Leading-Edge Serrations in Owls (Strigiformes)
Weger, Matthias; Wagner, Hermann
2016-01-01
Background Owls have developed serrations, comb-like structures, along the leading edge of their wings. Serrations were investigated from a morphological and a mechanical point of view, but were not yet quantitatively compared for different species. Such a comparative investigation of serrations from species of different sizes and activity patterns may provide new information about the function of the serrations. Results Serrations on complete wings and on tenth primary remiges of seven owl species were investigated. Small, middle-sized, and large owl species were investigated as well as species being more active during the day and owls being more active during the night. Serrations occurred at the outer parts of the wings, predominantly at tenth primary remiges, but also on further wing feathers in most species. Serration tips were oriented away from the feather rachis so that they faced into the air stream during flight. The serrations of nocturnal owl species were higher developed as demonstrated by a larger inclination angle (the angle between the base of the barb and the rachis), a larger tip displacement angle (the angle between the tip of the serration and the base of the serration) and a longer length. Putting the measured data into a clustering algorithm yielded dendrograms that suggested a strong influence of activity pattern, but only a weak influence of size on the development of the serrations. Conclusions Serrations are supposed to be involved in noise reduction during flight and also depend on the aerodynamic properties that in turn depend on body size. Since especially nocturnal owls have to rely on hearing during prey capture, the more pronounced serrations of nocturnal species lend further support to the notion that serrations have an important function in noise reduction. The differences in shape of the serrations investigated indicate that a silent flight requires well-developed serrations. PMID:26934104
An analysis of aircrew communication patterns and content
NASA Astrophysics Data System (ADS)
Oser, Randall L.; Prince, Carolyn; Morgan, Ben B., Jr.; Simpson, Steven S.
1991-09-01
The findings reported here represent a detailed analysis of tactical rotary-wing aircrew communication patterns and content. This research is part of an extensive effort to investigate the nature of tactical aircrew coordination and to develop effective mission-oriented aircrew coordination training. The primary objectives of this research were to answer the following questions: (1) What specific communication patterns and content are demonstrated by different helicopter crewmembers (i.e., Helicopter Aircraft Commander - HAC and Helicopter 2nd Pilot - H2P)? (2) Do tactical aircrew communication patterns and content vary as a function of the performance demands and requirements of different flight conditions (i.e., routine and non-routine)? (3) Are the communication patterns and content of more effective aircrews different from those of less effective aircrews? (4) What similarities exist between the communication patterns and content of military rotary-wing aircrews and commercial fixed-wing aircrews? and (5) Can the results of the communication analyses have an impact on aircrew coordination training?
Numerical simulation of X-wing type biplane flapping wings in 3D using the immersed boundary method.
Tay, W B; van Oudheusden, B W; Bijl, H
2014-09-01
The numerical simulation of an insect-sized 'X-wing' type biplane flapping wing configuration is performed in 3D using an immersed boundary method solver at Reynolds numbers equal to 1000 (1 k) and 5 k, based on the wing's root chord length. This X-wing type flapping configuration draws its inspiration from Delfly, a bio-inspired ornithopter MAV which has two pairs of wings flapping in anti-phase in a biplane configuration. The objective of the present investigation is to assess the aerodynamic performance when the original Delfly flapping wing micro-aerial vehicle (FMAV) is reduced to the size of an insect. Results show that the X-wing configuration gives more than twice the average thrust compared with only flapping the upper pair of wings of the X-wing. However, the X-wing's average thrust is only 40% that of the upper wing flapping at twice the stroke angle. Despite this, the increased stability which results from the smaller lift and moment variation of the X-wing configuration makes it more suited for sharp image capture and recognition. These advantages make the X-wing configuration an attractive alternative design for insect-sized FMAVS compared to the single wing configuration. In the Reynolds number comparison, the vorticity iso-surface plot at a Reynolds number of 5 k revealed smaller, finer vortical structures compared to the simulation at 1 k, due to vortices' breakup. In comparison, the force output difference is much smaller between Re = 1 k and 5 k. Increasing the body inclination angle generates a uniform leading edge vortex instead of a conical one along the wingspan, giving higher lift. Understanding the force variation as the body inclination angle increases will allow FMAV designers to optimize the thrust and lift ratio for higher efficiency under different operational requirements. Lastly, increasing the spanwise flexibility of the wings increases the thrust slightly but decreases the efficiency. The thrust result is similar to one of the spanwise studies, but the efficiency result contradicts it, indicating that other flapping parameters are involved as well. Results from this study provide a deeper understanding of the underlying aerodynamics of the X-wing type, which will help to improve the performance of insect-sized FMAVs using this unique configuration.
Determination of stores pointing error due to wing flexibility under flight load
NASA Technical Reports Server (NTRS)
Lokos, William A.; Bahm, Catherine M.; Heinle, Robert A.
1995-01-01
The in-flight elastic wing twist of a fighter-type aircraft was studied to provide for an improved on-board real-time computed prediction of pointing variations of three wing store stations. This is an important capability to correct sensor pod alignment variation or to establish initial conditions of iron bombs or smart weapons prior to release. The original algorithm was based upon coarse measurements. The electro-optical Flight Deflection Measurement System measured the deformed wing shape in flight under maneuver loads to provide a higher resolution database from which an improved twist prediction algorithm could be developed. The FDMS produced excellent repeatable data. In addition, a NASTRAN finite-element analysis was performed to provide additional elastic deformation data. The FDMS data combined with the NASTRAN analysis indicated that an improved prediction algorithm could be derived by using a different set of aircraft parameters, namely normal acceleration, stores configuration, Mach number, and gross weight.
Miocene honey bees from the Randeck Maar of southwestern Germany (Hymenoptera, Apidae)
Kotthoff, Ulrich; Wappler, Torsten; Engel, Michael S.
2011-01-01
Abstract The Miocene Randeck Maar (southwestern Germany) is one of the only sites with abundant material of fossil honey bees. The fauna has been the focus of much scrutiny by early authors who recognized multiple species or subspecies within the fauna. The history of work on the Randeck Maar is briefly reviewed and these fossils placed into context with other Tertiary and living species of the genus Apis Linnaeus (Apinae: Apini). Previously unrecorded specimens from Randeck Maar were compared with earlier series in an attempt to evaluate the observed variation. A morphometric analysis of forewing venation angles across representative Recent and Tertiary species of Apis as well as various non-Apini controls was undertaken to evaluate the distribution of variation in fossil honey bees. The resulting dendrogram shows considerable variation concerning the wing venation of Miocene Apini, but intergradation of other morphological characters reveals no clear pattern of separate species. This suggests that a single, highly variable species was present in Europe during the Miocene. The pattern also supports the notion that the multiple species and subspecies proposed by earlier authors for the Randeck Maar honey bee fauna are not valid, and all are accordingly recognized as Apis armbrusteri Zeuner. PMID:21594072
NASA Technical Reports Server (NTRS)
Goodykoontz, J. H.; Wagner, J. M.; Sargent, N. B.
1973-01-01
Noise data were taken for variations to a large scale model of an externally blown flap lift augmentation system. The variations included two different mixer nozzles (7 and 8 lobes), two different wing models (2 and 3 flaps), and different lateral distances between the wing chord line and the nozzle centerline. When the seven lobe was used with the trailing flap in the 60 deg position, increasing the wing to nozzle distance had no effect on the sound level. When the eight lobe nozzle was used there was a decrease in sound level. With the 20 deg flap setting the noise level decreased when the distance was increased using either nozzle.
Experimental and Theoretical Study of a Rectangular Wing in a Vortical Wake at Low Speed
NASA Technical Reports Server (NTRS)
Smith, Willard G.; Lazzeroni, Frank A.
1960-01-01
A systematic study has been made, experimentally and theoretically, of the effects of a vortical wake on the aerodynamic characteristics of a rectangular wing at subsonic speed. The vortex generator and wing were mounted on a reflection plane to avoid body-wing interference. Vortex position, relative to the wing, was varied both in the spanwise direction and normal to the wing. Angle of attack of the wing was varied from -40 to +60. Both chordwise and spanwise pressure distributions were obtained with the wing in uniform and vortical flow fields. Stream surveys were made to determine the flow characteristics in the vortical wake. The vortex-induced lift was calculated by several theoretical methods including strip theory, reverse-flow theory, and reverse-flow theory including a finite vortex core. In addition, the Prandtl lifting-line theory and the Weissinger theory were used to calculate the spanwise distribution of vortex-induced loads. With reverse-flow theory, predictions of the interference lift were generally good, and with Weissinger's theory the agreement between the theoretical spanwise variation of induced load and the experimental variation was good. Results of the stream survey show that the vortex generated by a lifting surface of rectangular plan form tends to trail back streamwise from the tip and does not approach the theoretical location, or centroid of circulation, given by theory. This discrepancy introduced errors in the prediction of vortex interference, especially when the vortex core passed immediately outboard of the wing tip. The wake produced by the vortex generator in these tests was not fully rolled up into a circular vortex, and so lacked symmetry in the vertical direction of the transverse plane. It was found that the direction of circulation affected the induced loads on the wing either when the wing was at angle of attack or when the vortex was some distance away from the plane of the wing.
Hypothesis testing in evolutionary developmental biology: a case study from insect wings.
Jockusch, E L; Ober, K A
2004-01-01
Developmental data have the potential to give novel insights into morphological evolution. Because developmental data are time-consuming to obtain, support for hypotheses often rests on data from only a few distantly related species. Similarities between these distantly related species are parsimoniously inferred to represent ancestral aspects of development. However, with limited taxon sampling, ancestral similarities in developmental patterning can be difficult to distinguish from similarities that result from convergent co-option of developmental networks, which appears to be common in developmental evolution. Using a case study from insect wings, we discuss how these competing explanations for similarity can be evaluated. Two kinds of developmental data have recently been used to support the hypothesis that insect wings evolved by modification of limb branches that were present in ancestral arthropods. This support rests on the assumption that aspects of wing development in Drosophila, including similarities to crustacean epipod patterning, are ancestral for winged insects. Testing this assumption requires comparisons of wing development in Drosophila and other winged insects. Here we review data that bear on this assumption, including new data on the functions of wingless and decapentaplegic during appendage allocation in the red flour beetle Tribolium castaneum.
Live Cell Imaging of Butterfly Pupal and Larval Wings In Vivo
Ohno, Yoshikazu; Otaki, Joji M.
2015-01-01
Butterfly wing color patterns are determined during the late larval and early pupal stages. Characterization of wing epithelial cells at these stages is thus critical to understand how wing structures, including color patterns, are determined. Previously, we successfully recorded real-time in vivo images of developing butterfly wings over time at the tissue level. In this study, we employed similar in vivo fluorescent imaging techniques to visualize developing wing epithelial cells in the late larval and early pupal stages 1 hour post-pupation. Both larval and pupal epithelial cells were rich in mitochondria and intracellular networks of endoplasmic reticulum, suggesting high metabolic activities, likely in preparation for cellular division, polyploidization, and differentiation. Larval epithelial cells in the wing imaginal disk were relatively large horizontally and tightly packed, whereas pupal epithelial cells were smaller and relatively loosely packed. Furthermore, larval cells were flat, whereas pupal cells were vertically elongated as deep as 130 μm. In pupal cells, many endosome-like or autophagosome-like structures were present in the cellular periphery down to approximately 10 μm in depth, and extensive epidermal feet or filopodia-like processes were observed a few micrometers deep from the cellular surface. Cells were clustered or bundled from approximately 50 μm in depth to deeper levels. From 60 μm to 80 μm in depth, horizontal connections between these clusters were observed. The prospective eyespot and marginal focus areas were resistant to fluorescent dyes, likely because of their non-flat cone-like structures with a relatively thick cuticle. These in vivo images provide important information with which to understand processes of epithelial cell differentiation and color pattern determination in butterfly wings. PMID:26107809
Live Cell Imaging of Butterfly Pupal and Larval Wings In Vivo.
Ohno, Yoshikazu; Otaki, Joji M
2015-01-01
Butterfly wing color patterns are determined during the late larval and early pupal stages. Characterization of wing epithelial cells at these stages is thus critical to understand how wing structures, including color patterns, are determined. Previously, we successfully recorded real-time in vivo images of developing butterfly wings over time at the tissue level. In this study, we employed similar in vivo fluorescent imaging techniques to visualize developing wing epithelial cells in the late larval and early pupal stages 1 hour post-pupation. Both larval and pupal epithelial cells were rich in mitochondria and intracellular networks of endoplasmic reticulum, suggesting high metabolic activities, likely in preparation for cellular division, polyploidization, and differentiation. Larval epithelial cells in the wing imaginal disk were relatively large horizontally and tightly packed, whereas pupal epithelial cells were smaller and relatively loosely packed. Furthermore, larval cells were flat, whereas pupal cells were vertically elongated as deep as 130 μm. In pupal cells, many endosome-like or autophagosome-like structures were present in the cellular periphery down to approximately 10 μm in depth, and extensive epidermal feet or filopodia-like processes were observed a few micrometers deep from the cellular surface. Cells were clustered or bundled from approximately 50 μm in depth to deeper levels. From 60 μm to 80 μm in depth, horizontal connections between these clusters were observed. The prospective eyespot and marginal focus areas were resistant to fluorescent dyes, likely because of their non-flat cone-like structures with a relatively thick cuticle. These in vivo images provide important information with which to understand processes of epithelial cell differentiation and color pattern determination in butterfly wings.
Ramey, Andrew M.; Poulson, Rebecca L.; González-Reiche, Ana S.; Wilcox, Benjamin R.; Walther, Patrick; Link, Paul; Carter, Deborah L.; Newsome, George M.; Müller, Maria L.; Berghaus, Roy D.; Perez, Daniel R.; Hall, Jeffrey S.; Stallknecht, David E.
2014-01-01
Seasonal dynamics of influenza A viruses (IAVs) are driven by host density and population immunity. Through an analysis of subtypic data for IAVs isolated from Blue-winged Teal (Anas discors), we present evidence for seasonal patterns in the relative abundance of viral subtypes in spring and summer/autumn.
Fukutomi, Yuichi; Matsumoto, Keiji; Agata, Kiyokazu; Funayama, Noriko; Koshikawa, Shigeyuki
2017-06-01
Various organisms have color patterns on their body surfaces, and these color patterns are thought to contribute to physiological regulation, communication with conspecifics, and signaling with the environment. An adult fly of Drosophila guttifera (Insecta: Diptera: Drosophilidae) has melanin pigmentation patterns on its body and wings. Though D. guttifera has been used for research into color pattern formation, how its pupal development proceeds and when the pigmentation starts have not been well studied. In this study, we defined the pupal stages of D. guttifera and measured the pigment content of wing spots from the pupal period to the period after eclosion. Using a transgenic line which carries eGFP connected with an enhancer of yellow, a gene necessary for melanin synthesis, we analyzed the timing at which the yellow enhancer starts to drive eGFP. We also analyzed the distribution of Yellow-producing cells, as indicated by the expression of eGFP during pupal and young adult periods. The results suggested that Yellow-producing cells were removed from wings within 3 h after eclosion, and wing pigmentation continued without epithelial cells. Furthermore, the results of vein cutting experiments showed that the transport of melanin precursors through veins was necessary for wing pigmentation. These results showed the importance of melanin precursors transported through veins and of extracellular factors which were secreted from epithelial cells and left in the cuticle.
Johnson, Haley; Solensky, Michelle J.; Satterfield, Dara A.; Davis, Andrew K.
2014-01-01
In animals with complex life cycles, all resources needed to form adult tissues are procured at the larval stage. For butterflies, the proper development of wings involves synthesizing tissue during metamorphosis based on the raw materials obtained by larvae. Similarly, manufacture of pigment for wing scales also requires resources acquired by larvae. We conducted an experiment to test the effects of food deprivation in the larval stage on multiple measures of adult wing morphology and coloration of monarch butterflies (Danaus plexippus), a species in which long-distance migration makes flight efficiency critical. In a captive setting, we restricted food (milkweed) from late-stage larvae for either 24 hrs or 48 hrs, then after metamorphosis we used image analysis methods to measure forewing surface area and elongation (length/width), which are both important for migration. We also measured the brightness of orange pigment and the intensity of black on the wing. There were correlations between several wing features, including an unexpected association between wing elongation and melanism, which will require further study to fully understand. The clearest effect of food restriction was a reduction in adult wing size in the high stress group (by approximately 2%). Patterns observed for other wing traits were ambiguous: monarchs in the low stress group (but not the high) had less elongated and paler orange pigmentation. There was no effect on wing melanism. Although some patterns obtained in this study were unclear, our results concerning wing size have direct bearing on the monarch migration. We show that if milkweed is limited for monarch larvae, their wings become stunted, which could ultimately result in lower migration success. PMID:24695643
Johnson, Haley; Solensky, Michelle J; Satterfield, Dara A; Davis, Andrew K
2014-01-01
In animals with complex life cycles, all resources needed to form adult tissues are procured at the larval stage. For butterflies, the proper development of wings involves synthesizing tissue during metamorphosis based on the raw materials obtained by larvae. Similarly, manufacture of pigment for wing scales also requires resources acquired by larvae. We conducted an experiment to test the effects of food deprivation in the larval stage on multiple measures of adult wing morphology and coloration of monarch butterflies (Danaus plexippus), a species in which long-distance migration makes flight efficiency critical. In a captive setting, we restricted food (milkweed) from late-stage larvae for either 24 hrs or 48 hrs, then after metamorphosis we used image analysis methods to measure forewing surface area and elongation (length/width), which are both important for migration. We also measured the brightness of orange pigment and the intensity of black on the wing. There were correlations between several wing features, including an unexpected association between wing elongation and melanism, which will require further study to fully understand. The clearest effect of food restriction was a reduction in adult wing size in the high stress group (by approximately 2%). Patterns observed for other wing traits were ambiguous: monarchs in the low stress group (but not the high) had less elongated and paler orange pigmentation. There was no effect on wing melanism. Although some patterns obtained in this study were unclear, our results concerning wing size have direct bearing on the monarch migration. We show that if milkweed is limited for monarch larvae, their wings become stunted, which could ultimately result in lower migration success.
Prediction of added noise due to the effect of unsteady flow on pusher propellers
NASA Technical Reports Server (NTRS)
Takallu, M. A.; Block, P. J. W.
1987-01-01
An analytical/computational study has been conducted to predict the effect of an upstream wing or pylon on the noise of an operating propeller. The wing trailing edge was placed at variable distances (0.1 and 0.3 chord) upstream of a scaled model propeller (SR-2). The wake was modeled using a similarity formulation. The instantaneous pressure distribution on the propeller blades during the passage through the wake was formulated in terms of a time-dependent variation of each blade section's angle of attack and in terms of the shed vortices from the blade trailing edge. It was found that the final expressions for the unsteady loads considerably altered the radiated noise pattern. Predicted noise for various observer positions, rotational speeds and propeller/pylon distances were computed and are presented in terms of the pressure time history. It has been shown that the positioning of a pylon upstream of a propeller indeed increases the noise. Some comparisons with experimental results are also given.
NASA Technical Reports Server (NTRS)
Bamber, M J
1935-01-01
General methods of theoretical analysis of airplane spinning characteristics have been available for some time. Some of these methods of analysis might be used by designers to predict the spinning characteristics of proposed airplane designs if the necessary aerodynamic data were known. The present investigation, to determine the spinning characteristics of wings, is planned to include variations in airfoil sections, plan forms, and tip shapes of monoplane wings and variations in stagger, gap, and decalage for biplane cellules. The first series of tests, made on a rectangular Clark Y monoplane wing, are reported in reference 1. That report also gives an analysis of the data for predicting the probable effects of various important parameters on the spin for normal airplanes using such a wing. The present report is the second of the series. It gives the aerodynamic characteristics of a rectangular Clark Y biplane cellule in spinning attitudes and includes a discussion of the data, using the method of analysis given in reference 1.
Sumruayphol, Suchada; Chittsamart, Boonruam; Polseela, Raxsina; Sriwichai, Patchara; Samung, Yudthana; Apiwathnasorn, Chamnarn; Dujardin, Jean-Pierre
2017-01-01
Geographic populations of the two main sandflies genera present in Thailand were studied for species and population identification. Size and shape of Phlebotomus stantoni and Sergentomyia hodgsoni from different island and mainland locations were examined by landmark-based geometric morphometrics. Intraspecific and interspecific wing comparison was carried out based on 12 anatomical landmarks. The wing centroid size of P. stantoni was generally larger than that of S. hodgsoni. Within both species, wings from the continent were significantly larger than those from island populations. Size variation could be significant between geographic locations, but could also overlap between genera. The wing venation geometry showed non-overlapping differences between two species. The within-species variation of geometric shape between different geographical locations was highly significant, but it could not interfere with the interspecies difference. The lack of species overlapping in shape, and the high discrimination between geographic populations, make geometric shape a promising character for future taxonomic and epidemiological studies. Copyright © 2016 Académie des sciences. Published by Elsevier SAS. All rights reserved.
Of hummingbirds and helicopters: hovering costs, competitive ability, and foraging strategies.
Altshuler, Douglas L
2004-01-01
Wing morphology and flight kinematics profoundly influence foraging costs and the overall behavioral ecology of hummingbirds. By analogy with helicopters, previous energetic studies have applied the momentum theory of aircraft propellers to estimate hovering costs from wing disc loading (WDL), a parameter incorporating wingspan (or length) and body mass. Variation in WDL has been used to elucidate differences either among hummingbird species in nectar-foraging strategies (e.g., territoriality, traplining) and dominance relations or among gender-age categories within species. We first demonstrate that WDL, as typically calculated, is an unreliable predictor of hovering (induced power) costs; predictive power is increased when calculations use wing length instead of wingspan and when actual wing stroke amplitudes are incorporated. We next evaluate the hypotheses that foraging strategy and competitive ability are functions of WDL, using our data in combination with those of published sources. Variation in hummingbird behavior cannot be easily classified using WDL and instead is correlated with a diversity of morphological and physiological traits. Evaluating selection pressures on hummingbird wings will require moving beyond wing and body mass measurements to include the assessment of the aerodynamic forces, power requirements, and power reserves of hovering, forward flight, and maneuvering. However, the WDL-helicopter dynamics model has been instrumental in calling attention to the importance of comparative wing morphology and related aerodynamics for understanding the behavioral ecology of hummingbirds.
NASA Technical Reports Server (NTRS)
Ayers, T. G.
1973-01-01
An investigation was conducted in the Langley 8 foot transonic pressure tunnel and the Langley Unitary Plan wind tunnel to evaluate the effectiveness of three variations of the NASA supercritical airfoil as applied to a model of a variable wing sweep fighter airplane. Wing panels incorporating conventional NACA 64A series airfoil with 0.20 and 0.40 camber were used as bases of reference for this evaluation. Static force and moment measurements were obtained for wing leading edge sweep angles of 26, 33, 39, and 72.5 degrees. Fluctuating wing root bending moment data were obtained at subsonic speeds to determine buffet characteristics. Subsonic data were also obtained for determining the effects of wing transition location and spoiler deflection. Limited lateral directional data are included for the conventional 0.20 cambered wing and the supercritical wing.
Winged Scapula: A Comprehensive Review of Surgical Treatment
Charran, Ordessia; Yilmaz, Emre; Edwards, Bryan; Muhleman, Mitchel A; Oskouian, Rod J; Tubbs, R. Shane; Loukas, Marios
2017-01-01
Winged scapula is caused by paralysis of the serratus anterior or trapezius muscles due to damage to the long thoracic or accessory nerves, resulting in loss of strength and range of motion of the shoulder. Because this nerve damage can happen in a variety of ways, initial diagnosis may be overlooked. This paper discusses the anatomical structures involved in several variations of winged scapula, the pathogenesis of winged scapula, and several historical and contemporary surgical procedures used to treat this condition. Additionally, this review builds upon the conclusions of several studies in order to suggest areas for continued research regarding the treatment of winged scapula. PMID:29456903
Spectrally tuned structural and pigmentary coloration of birdwing butterfly wing scales.
Wilts, Bodo D; Matsushita, Atsuko; Arikawa, Kentaro; Stavenga, Doekele G
2015-10-06
The colourful wing patterns of butterflies play an important role for enhancing fitness; for instance, by providing camouflage, for interspecific mate recognition, or for aposematic display. Closely related butterfly species can have dramatically different wing patterns. The phenomenon is assumed to be caused by ecological processes with changing conditions, e.g. in the environment, and also by sexual selection. Here, we investigate the birdwing butterflies, Ornithoptera, the largest butterflies of the world, together forming a small genus in the butterfly family Papilionidae. The wings of these butterflies are marked by strongly coloured patches. The colours are caused by specially structured wing scales, which act as a chirped multilayer reflector, but the scales also contain papiliochrome pigments, which act as a spectral filter. The combined structural and pigmentary effects tune the coloration of the wing scales. The tuned colours are presumably important for mate recognition and signalling. By applying electron microscopy, (micro-)spectrophotometry and scatterometry we found that the various mechanisms of scale coloration of the different birdwing species strongly correlate with the taxonomical distribution of Ornithoptera species. © 2015 The Author(s).
Spectrally tuned structural and pigmentary coloration of birdwing butterfly wing scales
Wilts, Bodo D.; Matsushita, Atsuko; Arikawa, Kentaro; Stavenga, Doekele G.
2015-01-01
The colourful wing patterns of butterflies play an important role for enhancing fitness; for instance, by providing camouflage, for interspecific mate recognition, or for aposematic display. Closely related butterfly species can have dramatically different wing patterns. The phenomenon is assumed to be caused by ecological processes with changing conditions, e.g. in the environment, and also by sexual selection. Here, we investigate the birdwing butterflies, Ornithoptera, the largest butterflies of the world, together forming a small genus in the butterfly family Papilionidae. The wings of these butterflies are marked by strongly coloured patches. The colours are caused by specially structured wing scales, which act as a chirped multilayer reflector, but the scales also contain papiliochrome pigments, which act as a spectral filter. The combined structural and pigmentary effects tune the coloration of the wing scales. The tuned colours are presumably important for mate recognition and signalling. By applying electron microscopy, (micro-)spectrophotometry and scatterometry we found that the various mechanisms of scale coloration of the different birdwing species strongly correlate with the taxonomical distribution of Ornithoptera species. PMID:26446560
Hussain, Abid; Tian, Ming-Yi; Wen, Shuo-Yang
2017-12-12
The survival and foraging of Coptotermes formosanus Shiraki in a microbe-rich environment reflect the adaptation of an extraordinary, sophisticated defense mechanism by the nest-mates. We aimed to explore the host pathogen interaction by studying caste-specific volatile chemistry and genes encoding the antioxidant defense of winged imagoes, nymphs, soldiers and workers of Formosan subterranean termites. Qualitative analyses of C. formosanus Shiraki performed by HS-SPME/GC-MS showed considerable variations in the chemical composition of volatile organic compounds (VOCs) and their proportions among all the castes. Winged imagoes produced the most important compounds such as naphthalene and n- hexanoic acid. The antifungal activity of these compounds along with nonanal, n -pentadecane, n- tetradecane, n -heptadecane and methyl octanoate against the conidial suspensions of Metarhizium anisopliae and Beauveria bassiana isolates enable us to suggest that the failure of natural fungal infection in the nest is due to the antiseptic environment of the nest, which is mainly controlled by the VOCs of nest-mates. In addition, conidial germination of M. anisopliae and B. bassiana isolates evaluated on the cuticle of each caste showed significant variations among isolates and different castes. Our results showed that the conidia of M. anisopliae 02049 exhibited the highest germination on the cuticle of all the inoculated castes. Moreover, we recorded the lowest germination of the conidia of B. bassiana 200436. Caste-specific germination variations enabled us to report for the first time that the cuticle of winged imagoes was found to be the most resistant cuticle. The analysis of the transcriptome of C. formosanus Shiraki revealed the identification of 17 genes directly involved in antioxidant defense. Expression patterns of the identified antioxidant genes by quantitative real-time PCR (qPCR) revealed the significantly highest upregulation of CAT , GST , PRXSL , Cu/Zn-SOD2 , TXN1 , TXN2 , TXNL1 , TXNL2 , TXNL4A and TPx genes among winged imagoes upon infection with the most virulent isolate, M. anisopliae 02049. Furthermore, soldiers showed the least expression of genes encoding antioxidant defense. Our findings indicated that the volatile chemistry of nest-mates and genes encoding antioxidant defense greatly contribute to the survival and foraging of Formosan subterranean termites in a microbe-rich habitat.
Flow rate and trajectory of water spray produced by an aircraft tire
NASA Technical Reports Server (NTRS)
Daugherty, Robert H.; Stubbs, Sandy M.
1986-01-01
One of the risks associated with wet runway aircraft operation is the ingestion of water spray produced by an aircraft's tires into its engines. This problem can be especially dangerous at or near rotation speed on the takeoff roll. An experimental investigation was conducted in the NASA Langley Research Center Hydrodynamics Research Facility to measure the flow rate and trajectory of water spray produced by an aircraft nose tire operating on a flooded runway. The effects of various parameters on the spray patterns including distance aft of nosewheel, speed, load, and water depth were evaluated. Variations in the spray pattern caused by the airflow about primary structure such as the fuselage and wing are discussed. A discussion of events in and near the tire footprint concerning spray generation is included.
Ely, Craig R.; Fox, A.D.; Alisauskas, R.T.; Andreev, A.; Bromley, R.G.; Degtyarev, Andrei G.; Ebbinge, B.; Gurtovaya, E.N.; Kerbes, R.; Kondratyev, Alexander V.; Kostin, I.; Krechmar, A.V.; Litvin, K.E.; Miyabayashi, Y.; Moou, J.H.; Oates, R.M.; Orthmeyer, D.L.; Sabano, Yutaka; Simpson, S.G.; Solovieva, D.V.; Spindler, Michael A.; Syroechkovsky, Y.V.; Takekawa, John Y.; Walsh, A.
2005-01-01
Capsule: Greater White-fronted Geese show significant variation in body size from sampling locations throughout their circumpolar breeding range. Aims: To determine the degree of geographical variation in body size of Greater White-fronted Geese and identify factors contributing to any apparent patterns in variation. Methods: Structural measures of >3000 geese from 16 breeding areas throughout the Holarctic breeding range of the species were compared statistically. Results: Palearctic forms varied clinally, and increased in size from the smallest forms on the Kanin and Taimyr peninsulas in western Eurasia to the largest forms breeding in the Anadyr Lowlands of eastern Chukotka. Clinal variation was less apparent in the Nearctic, as both the smallest form in the Nearctic and the largest form overall (the Tule Goose) were from different breeding areas in Alaska. The Tule Goose was 25% larger than the smallest form. Birds from Greenland (A. a. flavirostris) were the second largest, although only slightly larger than geese from several North American populations. Body size was not correlated with breeding latitude but was positively correlated with temperature on the breeding grounds, breeding habitat, and migration distance. Body mass of Greater White-fronted Geese from all populations remained relatively constant during the period of wing moult. Morphological distinctness of eastern and western Palearctic forms concurs with earlier findings of complete range disjunction. Conclusions: Patterns of morphological variation in Greater White-fronted Geese across the Holarctic can be generally attributed to adaptation to variable breeding environments, migration requirements, and phylo-geographical histories.
Garzón, Maximiliano J; Schweigmann, Nicolás
2018-06-23
Shape variability among individuals is important to understand some ecological relationships, since it provides the nexus between the genotype and the environment. Geometric morphometrics based on generalized procrustes analysis was applied on 17 landmarks of the wings of Aedes albifasciatus (Macquart 1838) (Diptera: Culicidae) females collected from three ecoregions of Argentina (Delta and islands of the Paraná River, Pampa, and Patagonian steppe). This methodology was used to discriminate the shapes of individuals belonging to different regions. The population of the Patagonian steppe, which was the most geographically distant, showed the most dissimilar shape. Different local variations in wing shape could have been selected according to the environmental characteristics and maintained by geographic isolation. The individuals of the two ecoregions closest to each other (Delta and islands of the Paraná River and Pampa) showed differences in shape that can be explained by a lower gene flow due to the effect of geographic isolation (by the Paraná River) and the limited dispersive capacity of Ae. albifasciatus. The results allow concluding that both environmental diversity and geographic barriers could contribute to local variations in wing shape.
Pirih, Primož; Wilts, Bodo D; Stavenga, Doekele G
2011-10-01
The males of many pierid butterflies have iridescent wings, which presumably function in intraspecific communication. The iridescence is due to nanostructured ridges of the cover scales. We have studied the iridescence in the males of a few members of Coliadinae, Gonepteryx aspasia, G. cleopatra, G. rhamni, and Colias croceus, and in two members of the Colotis group, Hebomoia glaucippe and Colotis regina. Imaging scatterometry demonstrated that the pigmentary colouration is diffuse whereas the structural colouration creates a directional, line-shaped far-field radiation pattern. Angle-dependent reflectance measurements demonstrated that the directional iridescence distinctly varies among closely related species. The species-dependent scale curvature determines the spatial properties of the wing iridescence. Narrow beam illumination of flat scales results in a narrow far-field iridescence pattern, but curved scales produce broadened patterns. The restricted spatial visibility of iridescence presumably plays a role in intraspecific signalling.
Iwata, Masaki; Hiyama, Atsuki; Otaki, Joji M.
2013-01-01
Developmental studies on wing colour patterns have been performed in nymphalid butterflies, but efficient genetic manipulations, including mutagenesis, have not been well established. Here, we have performed mutagenesis experiments in a lycaenid butterfly, the pale grass blue Zizeeria maha, to produce colour-pattern mutants. We fed the P-generation larvae an artificial diet containing the mutagen ethyl methane sulfonate (EMS), and the F1- and F2-generation adults showed various aberrant colour patterns: dorsoventral transformation, anterioposterior background colouration gap, weak contrast, disarrangement of spots, reduction of the size of spots, loss of spots, fusion of spots, and ectopic spots. Among them, the disarrangement, reduction, and loss of spots were likely produced by the coordinated changes of many spots of a single wing around the discal spot in a system-dependent manner, demonstrating the existence of the central symmetry system. The present study revealed multiple genetic regulations for system-dependent and wing-wide colour-pattern determination in lycaenid butterflies. PMID:23917124
Wing optimization for space shuttle orbiter vehicles
NASA Technical Reports Server (NTRS)
Surber, T. E.; Bornemann, W. E.; Miller, W. D.
1972-01-01
The results were presented of a parametric study performed to determine the optimum wing geometry for a proposed space shuttle orbiter. The results of the study establish the minimum weight wing for a series of wing-fuselage combinations subject to constraints on aerodynamic heating, wing trailing edge sweep, and wing over-hang. The study consists of a generalized design evaluation which has the flexibility of arbitrarily varying those wing parameters which influence the vehicle system design and its performance. The study is structured to allow inputs of aerodynamic, weight, aerothermal, structural and material data in a general form so that the influence of these parameters on the design optimization process can be isolated and identified. This procedure displays the sensitivity of the system design of variations in wing geometry. The parameters of interest are varied in a prescribed fashion on a selected fuselage and the effect on the total vehicle weight is determined. The primary variables investigated are: wing loading, aspect ratio, leading edge sweep, thickness ratio, and taper ratio.
Flight Speeds among Bird Species: Allometric and Phylogenetic Effects
Alerstam, Thomas; Rosén, Mikael; Bäckman, Johan; Ericson, Per G. P; Hellgren, Olof
2007-01-01
Flight speed is expected to increase with mass and wing loading among flying animals and aircraft for fundamental aerodynamic reasons. Assuming geometrical and dynamical similarity, cruising flight speed is predicted to vary as (body mass)1/6 and (wing loading)1/2 among bird species. To test these scaling rules and the general importance of mass and wing loading for bird flight speeds, we used tracking radar to measure flapping flight speeds of individuals or flocks of migrating birds visually identified to species as well as their altitude and winds at the altitudes where the birds were flying. Equivalent airspeeds (airspeeds corrected to sea level air density, U e) of 138 species, ranging 0.01–10 kg in mass, were analysed in relation to biometry and phylogeny. Scaling exponents in relation to mass and wing loading were significantly smaller than predicted (about 0.12 and 0.32, respectively, with similar results for analyses based on species and independent phylogenetic contrasts). These low scaling exponents may be the result of evolutionary restrictions on bird flight-speed range, counteracting too slow flight speeds among species with low wing loading and too fast speeds among species with high wing loading. This compression of speed range is partly attained through geometric differences, with aspect ratio showing a positive relationship with body mass and wing loading, but additional factors are required to fully explain the small scaling exponent of U e in relation to wing loading. Furthermore, mass and wing loading accounted for only a limited proportion of the variation in U e. Phylogeny was a powerful factor, in combination with wing loading, to account for the variation in U e. These results demonstrate that functional flight adaptations and constraints associated with different evolutionary lineages have an important influence on cruising flapping flight speed that goes beyond the general aerodynamic scaling effects of mass and wing loading. PMID:17645390
Flow Structure and Force Variation with Aspect Ratio for a Two-Degree-of-Freedom Flapping Wing
NASA Astrophysics Data System (ADS)
Burge, Matthew; Favale, James; Ringuette, Matthew
2014-11-01
We investigate experimentally the effect of aspect ratio (AR) on the flow structure and forces of a two-degree-of-freedom flapping wing. Flapping wings are known to produce complex and unsteady vortex loop structures, and the objective is to characterize their variation with AR and how this influences the lift force. Previous results on rotating wings demonstrated that changes in AR significantly affect the three-dimensional flow structure and lift coefficient. This is primarily due to the relatively greater influence of the tip vortex for lower AR. At Reynolds number of order O(103) we test wings of AR = 2-4, values typically found in nature, with simplified planform shapes. The lift force is measured using a submersible transducer at the base of the wing in a glycerin-water mixture. The qualitative, three-dimensional vortex loop structure for different ARs is obtained using multi-color dye flow visualization. Guided by this, quantitative three-component flow information, namely vorticity, the Q-criterion, and circulation, is acquired from stereoscopic particle image velocimetry in key planes. Of interest is how these parameters and the vortex loop topology vary with AR, and their connection to features in the unsteady force signal. This work is supported by the National Science Foundation, Award Number 1336548, supervised by Dr. Dimitrios Papavassiliou.
Kang, Chang-ku; Moon, Jong-yeol; Lee, Sang-im; Jablonski, Piotr G.
2013-01-01
Many moths have wing patterns that resemble bark of trees on which they rest. The wing patterns help moths to become camouflaged and to avoid predation because the moths are able to assume specific body orientations that produce a very good match between the pattern on the bark and the pattern on the wings. Furthermore, after landing on a bark moths are able to perceive stimuli that correlate with their crypticity and are able to re-position their bodies to new more cryptic locations and body orientations. However, the proximate mechanisms, i.e. how a moth finds an appropriate resting position and orientation, are poorly studied. Here, we used a geometrid moth Jankowskia fuscaria to examine i) whether a choice of resting orientation by moths depends on the properties of natural background, and ii) what sensory cues moths use. We studied moths’ behavior on natural (a tree log) and artificial backgrounds, each of which was designed to mimic one of the hypothetical cues that moths may perceive on a tree trunk (visual pattern, directional furrow structure, and curvature). We found that moths mainly used structural cues from the background when choosing their resting position and orientation. Our findings highlight the possibility that moths use information from one type of sensory modality (structure of furrows is probably detected through tactile channel) to achieve crypticity in another sensory modality (visual). This study extends our knowledge of how behavior, sensory systems and morphology of animals interact to produce crypsis. PMID:24205118
Kang, Chang-Ku; Moon, Jong-Yeol; Lee, Sang-Im; Jablonski, Piotr G
2013-01-01
Many moths have wing patterns that resemble bark of trees on which they rest. The wing patterns help moths to become camouflaged and to avoid predation because the moths are able to assume specific body orientations that produce a very good match between the pattern on the bark and the pattern on the wings. Furthermore, after landing on a bark moths are able to perceive stimuli that correlate with their crypticity and are able to re-position their bodies to new more cryptic locations and body orientations. However, the proximate mechanisms, i.e. how a moth finds an appropriate resting position and orientation, are poorly studied. Here, we used a geometrid moth Jankowskia fuscaria to examine i) whether a choice of resting orientation by moths depends on the properties of natural background, and ii) what sensory cues moths use. We studied moths' behavior on natural (a tree log) and artificial backgrounds, each of which was designed to mimic one of the hypothetical cues that moths may perceive on a tree trunk (visual pattern, directional furrow structure, and curvature). We found that moths mainly used structural cues from the background when choosing their resting position and orientation. Our findings highlight the possibility that moths use information from one type of sensory modality (structure of furrows is probably detected through tactile channel) to achieve crypticity in another sensory modality (visual). This study extends our knowledge of how behavior, sensory systems and morphology of animals interact to produce crypsis.
The effect of butterfly scales on flight efficiency and leading edge vortex formation
NASA Astrophysics Data System (ADS)
Lang, Amy; Wilroy, Jacob; Wahidi, Redha; Slegers, Nathan; Heilman, Micahel; Cranford, Jacob
2016-11-01
It is hypothesized that the scales on a butterfly wing lead to increased flight efficiency. Recent testing of live butterflies tracked their motion over 246 flights for 11 different specimens. Results show a 37.8 percent mean decrease in flight efficiency and a flapping amplitude reduction of 6.7 percent once the scales were removed. This change could be largely a result of how the leading edge vortex (LEV) interacts with the wing. To simplify this complex flow problem, an experiment was designed to focus on the alteration of 2-D vortex development with a variation in surface patterning. Specifically, the secondary vorticity generated by the LEV interacting at the patterned surface was studied, as well as the subsequent effect on the LEV's growth rate and peak circulation. For this experiment butterfly inspired grooves were created using additive manufacturing and were attached to a flat plate with a chordwise orientation, thus increasing plate surface area. The vortex generated by the grooved plate was then compared to a smooth case as the plate translated vertically through a tow tank at Re = 1500, 3000, and 6000. Using DPIV, the vortex formation was documented and a maximum vortex formation time of 4.22 was found based on the flat plate travel distance and chord length. Results indicate that the patterned surface slows down the growth of the vortex which corroborates the flight test results. Funding from NSF CBET Fluid Dynamcis is gratefully acknowledged.
Waterproof and translucent wings at the same time: problems and solutions in butterflies.
Goodwyn, Pablo Perez; Maezono, Yasunori; Hosoda, Naoe; Fujisaki, Kenji
2009-07-01
Although the colour of butterflies attracts the most attention, the waterproofing properties of their wings are also extremely interesting. Most butterfly wings are considered "super-hydrophobic" because the contact angle (CA) with a water drop exceeds 150 degrees. Usually, butterfly wings are covered with strongly overlapping scales; however, in the case of transparent or translucent wings, scale cover is reduced; thus, the hydrophobicity could be affected. Here, we present a comparative analysis of wing hydrophobicity and its dependence on morphology for two species with translucent wings Parantica sita (Nymphalidae) and Parnassius glacialis (Papilionidae). These species have very different life histories: P. sita lives for up to 6 months as an adult and migrates over long distance, whereas P. glacialis lives for less than 1 month and does not migrate. We measured the water CA and analysed wing morphology with scanning electron microscopy and atomic force microscopy. P. sita has super-hydrophobic wing surfaces, with CA > 160 degrees, whereas P. glacialis did not (CA = 100-135 degrees). Specialised scales were found on the translucent portions of P. sita wings. These scales were ovoid and much thinner than common scales, erect at about 30 degrees, and leaving up to 80% of the wing surface uncovered. The underlying bare wing surface had a remarkable pattern of ridges and knobs. P. glacialis also had over 80% of the wing surface uncovered, but the scales were either setae-like or spade-like. The bare surface of the wing had an irregular wavy smooth pattern. We suggest a mode of action that allows this super-hydrophobic effect with an incompletely covered wing surface. The scales bend, but do not collapse, under the pressure of a water droplet, and the elastic recovery of the structure at the borders of the droplet allows a high apparent CA. Thus, P. sita can be translucent without losing its waterproof properties. This characteristic is likely necessary for the long life and migration of this species. This is the first study of some of the effects on the hydrophobicity of translucency through scales' cover reduction in butterfly wings and on the morphology associated with improved waterproofing.
Molecular basis of wing coloration in a Batesian mimic butterfly, Papilio polytes
Nishikawa, Hideki; Iga, Masatoshi; Yamaguchi, Junichi; Saito, Kazuki; Kataoka, Hiroshi; Suzuki, Yutaka; Sugano, Sumio; Fujiwara, Haruhiko
2013-01-01
Batesian mimicry protects animals from predators through resemblance with distasteful models in shape, color pattern, or behavior. To elucidate the wing coloration mechanisms involved in the mimicry, we investigated chemical composition and gene expression of the pale yellow and red pigments of a swallowtail butterfly, Papilio polytes, whose females mimic the unpalatable butterfly Pachliopta aristolochiae. Using LC/MS, we showed that the pale yellow wing regions in non-mimetic females consist of kynurenine and N-β-alanyldopamine (NBAD). Moreover, qRT-PCR showed that kynurenine/NBAD biosynthetic genes were upregulated in these regions in non-mimetic females. However, these pigments were absent in mimetic females. RNA-sequencing showed that kynurenine/NBAD synthesis and Toll signaling genes were upregulated in the red spots specific to mimetic female wings. These results demonstrated that drastic changes in gene networks in the red and pale yellow regions can switch wing color patterns between non-mimetic and mimetic females of P. polytes. PMID:24212474
Molecular basis of wing coloration in a Batesian mimic butterfly, Papilio polytes.
Nishikawa, Hideki; Iga, Masatoshi; Yamaguchi, Junichi; Saito, Kazuki; Kataoka, Hiroshi; Suzuki, Yutaka; Sugano, Sumio; Fujiwara, Haruhiko
2013-11-11
Batesian mimicry protects animals from predators through resemblance with distasteful models in shape, color pattern, or behavior. To elucidate the wing coloration mechanisms involved in the mimicry, we investigated chemical composition and gene expression of the pale yellow and red pigments of a swallowtail butterfly, Papilio polytes, whose females mimic the unpalatable butterfly Pachliopta aristolochiae. Using LC/MS, we showed that the pale yellow wing regions in non-mimetic females consist of kynurenine and N-β-alanyldopamine (NBAD). Moreover, qRT-PCR showed that kynurenine/NBAD biosynthetic genes were upregulated in these regions in non-mimetic females. However, these pigments were absent in mimetic females. RNA-sequencing showed that kynurenine/NBAD synthesis and Toll signaling genes were upregulated in the red spots specific to mimetic female wings. These results demonstrated that drastic changes in gene networks in the red and pale yellow regions can switch wing color patterns between non-mimetic and mimetic females of P. polytes.
Pinto, N S; Juen, L; Cabette, H S R; De Marco, P
2012-06-01
Effects of riparian vegetation removal on body size and wing fluctuating asymmetry (FA) of Argia tinctipennis Selys (Odonata: Coenagrionidae) were studied in the River Suiá-Miçú basin, which is part of the Xingu basin in Brazilian Amazonia. A total of 70 specimens (n = 33 from preserved and n = 37 from degraded areas) was measured. Five wing measures of each wing (totalizing ten measured characters) were taken. Preserved and degraded points presented non-overlapped variations of a Habitat Integrity Index, supporting the environmental differentiation between these two categories. FA increases in degraded areas approximately four times for the width between the nodus and proximal portion of the pterostigma of forewings (FW), two times for the width of the wing in the region of nodus of FW, and approximately 1.7 times for the number of postnodal cells of FW. The increase is almost five times for the width between the nodus and the proximal portion of the pterostigma of hind wings (HW), three times for the number of postnodal cells of HW, and approximately 1.6 times the width between quadrangle and nodus of HW. Individuals of preserved sites were nearly 3.3% larger than for degraded sites, based on mean hind wing length. Our results supports that the development of A. tinctipennis in degraded areas is affected by riparian vegetation removal and may reflect in wing FA variations. Consequently, these FA measures may be a useful tool for bioassessment using Odonata insects as a model.
Climate variability slows evolutionary responses of Colias butterflies to recent climate change.
Kingsolver, Joel G; Buckley, Lauren B
2015-03-07
How does recent climate warming and climate variability alter fitness, phenotypic selection and evolution in natural populations? We combine biophysical, demographic and evolutionary models with recent climate data to address this question for the subalpine and alpine butterfly, Colias meadii, in the southern Rocky Mountains. We focus on predicting patterns of selection and evolution for a key thermoregulatory trait, melanin (solar absorptivity) on the posterior ventral hindwings, which affects patterns of body temperature, flight activity, adult and egg survival, and reproductive success in Colias. Both mean annual summer temperatures and thermal variability within summers have increased during the past 60 years at subalpine and alpine sites. At the subalpine site, predicted directional selection on wing absorptivity has shifted from generally positive (favouring increased wing melanin) to generally negative during the past 60 years, but there is substantial variation among years in the predicted magnitude and direction of selection and the optimal absorptivity. The predicted magnitude of directional selection at the alpine site declined during the past 60 years and varies substantially among years, but selection has generally been positive at this site. Predicted evolutionary responses to mean climate warming at the subalpine site since 1980 is small, because of the variability in selection and asymmetry of the fitness function. At both sites, the predicted effects of adaptive evolution on mean population fitness are much smaller than the fluctuations in mean fitness due to climate variability among years. Our analyses suggest that variation in climate within and among years may strongly limit evolutionary responses of ectotherms to mean climate warming in these habitats. © 2015 The Author(s) Published by the Royal Society. All rights reserved.
Van Truong, Tien; Byun, Doyoung; Kim, Min Jun; Yoon, Kwang Joon; Park, Hoon Cheol
2013-09-01
The aim of this work is to provide an insight into the aerodynamic performance of the beetle during takeoff, which has been estimated in previous investigations. We employed a scaled-up electromechanical model flapping wing to measure the aerodynamic forces and the three-dimensional flow structures on the flapping wing. The ground effect on the unsteady forces and flow structures were also characterized. The dynamically scaled wing model could replicate the general stroke pattern of the beetle's hind wing kinematics during takeoff flight. Two wing kinematic models have been studied to examine the influences of wing kinematics on unsteady aerodynamic forces. In the first model, the angle of attack is asymmetric and varies during the translational motion, which is the flapping motion of the beetle's hind wing. In the second model, the angle of attack is constant during the translational motion. The instantaneous aerodynamic forces were measured for four strokes during the beetle's takeoff by the force sensor attached at the wing base. Flow visualization provided a general picture of the evolution of the three-dimensional leading edge vortex (LEV) on the beetle hind wing model. The LEV is stable during each stroke, and increases radically from the root to the tip, forming a leading-edge spiral vortex. The force measurement results show that the vertical force generated by the hind wing is large enough to lift the beetle. For the beetle hind wing kinematics, the total vertical force production increases 18.4% and 8.6% for the first and second strokes, respectively, due to the ground effect. However, for the model with a constant angle of attack during translation, the vertical force is reduced during the first stroke. During the third and fourth strokes, the ground effect is negligible for both wing kinematic patterns. This finding suggests that the beetle's flapping mechanism induces a ground effect that can efficiently lift its body from the ground during takeoff.
Morehouse, Nathan I; Vukusic, Peter; Rutowski, Ron
2006-01-01
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. PMID:17164199
Investigation of Full-Scale Split Trailing-Edge Wing Flaps with Various Chords and Hinge Locations
NASA Technical Reports Server (NTRS)
Wallace, Rudolf
1936-01-01
This report gives the results of an investigation conducted in the NACA full-scale wind tunnel on a small parasol monoplane equipped with three different split trailing-edge wing flaps. The object of the investigation was to determine and correlate data on the characteristics of the airplane and flaps as affected by variation in flap chord, flap deflection, and flap location along the wing chord. The results give the lift, the drag, and the pitching moment characteristics of the airplane, and the flap forces and moments, the pressure distribution over the flaps and wing at one section, and the downwash characteristics of the flap and wing combinations.
Decapentaplegic and growth control in the developing Drosophila wing.
Akiyama, Takuya; Gibson, Matthew C
2015-11-19
As a central model for morphogen action during animal development, the bone morphogenetic protein 2/4 (BMP2/4)-like ligand Decapentaplegic (Dpp) is proposed to form a long-range signalling gradient that directs both growth and pattern formation during Drosophila wing disc development. While the patterning role of Dpp secreted from a stripe of cells along the anterior-posterior compartmental boundary is well established, the mechanism by which a Dpp gradient directs uniform cell proliferation remains controversial and poorly understood. Here, to determine the precise spatiotemporal requirements for Dpp during wing disc development, we use CRISPR-Cas9-mediated genome editing to generate a flippase recognition target (FRT)-dependent conditional null allele. By genetically removing Dpp from its endogenous stripe domain, we confirm the requirement of Dpp for the activation of a downstream phospho-Mothers against dpp (p-Mad) gradient and the regulation of the patterning targets spalt (sal), optomotor blind (omb; also known as bifid) and brinker (brk). Surprisingly, however, third-instar wing blade primordia devoid of compartmental dpp expression maintain relatively normal rates of cell proliferation and exhibit only mild defects in growth. These results indicate that during the latter half of larval development, the Dpp morphogen gradient emanating from the anterior-posterior compartment boundary is not directly required for wing disc growth.
The design of supercritical wings by the use of three-dimensional transonic theory
NASA Technical Reports Server (NTRS)
Mann, M. J.
1979-01-01
A procedure was developed for the design of transonic wings by the iterative use of three dimensional, inviscid, transonic analysis methods. The procedure was based on simple principles of supersonic flow and provided the designer with a set of guidelines for the systematic alteration of wing profile shapes to achieve some desired pressure distribution. The method was generally applicable to wing design at conditions involving a large region of supercriterical flow. To illustrate the method, it was applied to the design of a wing for a supercritical maneuvering fighter that operates at high lift and transonic Mach number. The wing profiles were altered to produce a large region of supercritical flow which was terminated by a weak shock wave. The spanwise variation of drag of this wing and some principles for selecting the streamwise pressure distribution are also discussed.
NASA Technical Reports Server (NTRS)
Steckel, D. K.; Dahlin, J. A.; Henne, P. A.
1980-01-01
These basic characteristics of critical wings included wing area, aspect ratio, average thickness, and sweep as well as practical constraints on the planform and thickness near the wing root to allow for the landing gear. Within these constraints, a large matrix of wing designs was studied with spanwise variations in the types of airfoils and distribution of lift as well as some small planform changes. The criteria by which the five candidate wings were chosen for testing were the cruise and buffet characteristics in the transonic regime and the compatibility of the design with low speed (high-lift) requirements. Five wing-wide-body configurations were tested in the NASA Ames 11-foot transonic wind tunnel. Nacelles and pylons, flap support fairings, tail surfaces, and an outboard aileron were also tested on selected configurations.
Kinematics of flap-bounding flight in the zebra finch over a wide range of speeds
Tobalske; Peacock; Dial
1999-07-01
It has been proposed elsewhere that flap-bounding, an intermittent flight style consisting of flapping phases interspersed with flexed-wing bounds, should offer no savings in average mechanical power relative to continuous flapping unless a bird flies 1.2 times faster than its maximum range speed (Vmr). Why do some species use intermittent bounds at speeds slower than 1.2Vmr? The 'fixed-gear hypothesis' suggests that flap-bounding is used to vary mean power output in small birds that are otherwise constrained by muscle physiology and wing anatomy to use a fixed muscle shortening velocity and pattern of wing motion at all flight speeds; the 'body-lift hypothesis' suggests that some weight support during bounds could make flap-bounding flight aerodynamically advantageous in comparison with continuous flapping over most forward flight speeds. To test these predictions, we studied high-speed film recordings (300 Hz) of wing and body motion in zebra finches (Taenopygia guttata, mean mass 13.2 g, N=4) taken as the birds flew in a variable-speed wind tunnel (0-14 m s-1). The zebra finches used flap-bounding flight at all speeds, so their flight style was unique compared with that of birds that facultatively shift from continuous flapping or flap-gliding at slow speeds to flap-bounding at fast speeds. There was a significant effect of flight speed on all measured aspects of wing motion except percentage of the wingbeat spent in downstroke. Changes in angular velocity of the wing indicated that contractile velocity in the pectoralis muscle changed with flight speed, which is not consistent with the fixed-gear hypothesis. Although variation in stroke-plane angle relative to the body, pronation angle of the wing and wing span at mid-upstroke showed that the zebra finch changed within-wingbeat geometries according to speed, a vortex-ring gait with a feathered upstroke appeared to be the only gait used during flapping. In contrast, two small species that use continuous flapping during slow flight (0-4 m s-1) either change wingbeat gait according to flight speed or exhibit more variation in stroke-plane and pronation angles relative to the body. Differences in kinematics among species appear to be related to wing design (aspect ratio, skeletal proportions) rather than to pectoralis muscle fiber composition, indicating that the fixed-gear hypothesis should perhaps be modified to exclude muscle physiology and to emphasize constraints due to wing anatomy. Body lift was produced during bounds at speeds from 4 to 14 m s-1. Maximum body lift was 0.0206 N (15.9 % of body weight) at 10 m s-1; body lift:drag ratio declined with increasing air speed. The aerodynamic function of bounds differed with increasing speed from an emphasis on lift production (4-10 m s-1) to an emphasis on drag reduction with a slight loss in lift (12 and 14 m s-1). From a mathematical model of aerodynamic costs, it appeared that flap-bounding offered the zebra finch an aerodynamic advantage relative to continuous flapping at moderate and fast flight speeds (6-14 m s-1), with body lift augmenting any savings offered solely by flap-bounding at speeds faster than 7.1 m s-1. The percentage of time spent flapping during an intermittent flight cycle decreased with increasing speed, so the mechanical cost of transport was likely to be lowest at faster flight speeds (10-14 m s-1).
NASA Technical Reports Server (NTRS)
Stanford, Bret K.; Jutte, Christine V.
2014-01-01
Several minimum-mass aeroelastic optimization problems are solved to evaluate the effectiveness of a variety of novel tailoring schemes for subsonic transport wings. Aeroelastic strength and panel buckling constraints are imposed across a variety of trimmed maneuver loads. Tailoring with metallic thickness variations, functionally graded materials, composite laminates, tow steering, and distributed trailing edge control effectors are all found to provide reductions in structural wing mass with varying degrees of success. The question as to whether this wing mass reduction will offset the increased manufacturing cost is left unresolved for each case.
Bat wing biometrics: using collagen-elastin bundles in bat wings as a unique individual identifier.
Amelon, Sybill K; Hooper, Sarah E; Womack, Kathryn M
2017-05-29
The ability to recognize individuals within an animal population is fundamental to conservation and management. Identification of individual bats has relied on artificial marking techniques that may negatively affect the survival and alter the behavior of individuals. Biometric systems use biological characteristics to identify individuals. The field of animal biometrics has expanded to include recognition of individuals based upon various morphologies and phenotypic variations including pelage patterns, tail flukes, and whisker arrangement. Biometric systems use 4 biologic measurement criteria: universality, distinctiveness, permanence, and collectability. Additionally, the system should not violate assumptions of capture-recapture methods that include no increased mortality or alterations of behavior. We evaluated whether individual bats could be uniquely identified based upon the collagen-elastin bundles that are visible with gross examination of their wings. We examined little brown bats ( Myotis lucifugus ), northern long-eared bats ( M. septentrionalis ), big brown bats ( Eptesicus fuscus ), and tricolored bats ( Perimyotis subflavus ) to determine whether the "wing prints" from the bundle network would satisfy the biologic measurement criteria. We evaluated 1,212 photographs from 230 individual bats comparing week 0 photos with those taken at weeks 3 or 6 and were able to confirm identity of individuals over time. Two blinded evaluators were able to successfully match 170 individuals in hand to photographs taken at weeks 0, 3, and 6. This study suggests that bats can be successfully re-identified using photographs taken at previous times. We suggest further evaluation of this methodology for use in a standardized system that can be shared among bat conservationists.
Bat wing biometrics: using collagen–elastin bundles in bat wings as a unique individual identifier
Hooper, Sarah E.; Womack, Kathryn M.
2017-01-01
Abstract The ability to recognize individuals within an animal population is fundamental to conservation and management. Identification of individual bats has relied on artificial marking techniques that may negatively affect the survival and alter the behavior of individuals. Biometric systems use biological characteristics to identify individuals. The field of animal biometrics has expanded to include recognition of individuals based upon various morphologies and phenotypic variations including pelage patterns, tail flukes, and whisker arrangement. Biometric systems use 4 biologic measurement criteria: universality, distinctiveness, permanence, and collectability. Additionally, the system should not violate assumptions of capture–recapture methods that include no increased mortality or alterations of behavior. We evaluated whether individual bats could be uniquely identified based upon the collagen–elastin bundles that are visible with gross examination of their wings. We examined little brown bats (Myotis lucifugus), northern long-eared bats (M. septentrionalis), big brown bats (Eptesicus fuscus), and tricolored bats (Perimyotis subflavus) to determine whether the “wing prints” from the bundle network would satisfy the biologic measurement criteria. We evaluated 1,212 photographs from 230 individual bats comparing week 0 photos with those taken at weeks 3 or 6 and were able to confirm identity of individuals over time. Two blinded evaluators were able to successfully match 170 individuals in hand to photographs taken at weeks 0, 3, and 6. This study suggests that bats can be successfully re-identified using photographs taken at previous times. We suggest further evaluation of this methodology for use in a standardized system that can be shared among bat conservationists. PMID:29674784
Flow Modulation and Force Control of Flapping Wings
2014-10-29
evolution of which reflect the wing morphology and kinematics. While the near-wake vortex system directly reflects the action of the wing on the...at 8 different stroke positions, which demonstrate the evolution of the vortex wake structure. The contour plot of Z vorticity at X-Y plane (Z...20 Figure 14. Smoke patterns showing the evolution of the flow structure in an
Application of the adjoint optimisation of shock control bump for ONERA-M6 wing
NASA Astrophysics Data System (ADS)
Nejati, A.; Mazaheri, K.
2017-11-01
This article is devoted to the numerical investigation of the shock wave/boundary layer interaction (SWBLI) as the main factor influencing the aerodynamic performance of transonic bumped airfoils and wings. The numerical analysis is conducted for the ONERA-M6 wing through a shock control bump (SCB) shape optimisation process using the adjoint optimisation method. SWBLI is analyzed for both clean and bumped airfoils and wings, and it is shown how the modified wave structure originating from upstream of the SCB reduces the wave drag, by improving the boundary layer velocity profile downstream of the shock wave. The numerical simulation of the turbulent viscous flow and a gradient-based adjoint algorithm are used to find the optimum location and shape of the SCB for the ONERA-M6 airfoil and wing. Two different geometrical models are introduced for the 3D SCB, one with linear variations, and another with periodic variations. Both configurations result in drag reduction and improvement in the aerodynamic efficiency, but the periodic model is more effective. Although the three-dimensional flow structure involves much more complexities, the overall results are shown to be similar to the two-dimensional case.
NASA Technical Reports Server (NTRS)
Suarez, Carlos J.; Smith, Brooke C.; Malcolm, Gerald N.
1993-01-01
Free-to-roll wind tunnel tests were conducted and a computer simulation exercise was performed in an effort to investigate in detail the mechanism of wing rock on a configuration that consisted of a highly-slender forebody and a 78 deg swept delta wing. In the wind tunnel test, the roll angle and wing surface pressures were measured during the wing rock motion. A limit cycle oscillation was observed for angles of attack between 22 deg and 30 deg. In general, the wind tunnel test confirmed that the main flow phenomena responsible for the wing-body-tail wing rock are the interactions between the forebody and the wing vortices. The variation of roll acceleration (determined from the second derivative of the roll angle time history) with roll angle clearly showed the energy balance necessary to sustain the limit cycle oscillation. Pressure measurements on the wing revealed the hysteresis of the wing rock process. First, second and nth order models for the aerodynamic damping were developed and examined with a one degree of freedom computer simulation. Very good agreement with the observed behavior from the wind tunnel was obtained.
Beneficial aerodynamic effect of wing scales on the climbing flight of butterflies.
Slegers, Nathan; Heilman, Michael; Cranford, Jacob; Lang, Amy; Yoder, John; Habegger, Maria Laura
2017-01-30
It is hypothesized that butterfly wing scale geometry and surface patterning may function to improve aerodynamic efficiency. In order to investigate this hypothesis, a method to measure butterfly flapping kinematics optically over long uninhibited flapping sequences was developed. Statistical results for the climbing flight flapping kinematics of 11 butterflies, based on a total of 236 individual flights, both with and without their wing scales, are presented. Results show, that for each of the 11 butterflies, the mean climbing efficiency decreased after scales were removed. Data was reduced to a single set of differences of climbing efficiency using are paired t-test. Results show a mean decrease in climbing efficiency of 32.2% occurred with a 95% confidence interval of 45.6%-18.8%. Similar analysis showed that the flapping amplitude decreased by 7% while the flapping frequency did not show a significant difference. Results provide strong evidence that butterfly wing scale geometry and surface patterning improve butterfly climbing efficiency. The authors hypothesize that the wing scale's effect in measured climbing efficiency may be due to an improved aerodynamic efficiency of the butterfly and could similarly be used on flapping wing micro air vehicles to potentially achieve similar gains in efficiency.
NASA Technical Reports Server (NTRS)
Groesbeck, D.; Vonglahn, U.
1979-01-01
The effects on acoustic characteristics of nozzle type and location on a wing for STOL engine over-the-wing configurations are assessed at full scale on the basis of model-scale data. Three types of nozzle configurations are evaluated: a circular nozzle with external deflector mounted above the wing, a slot nozzle with external deflector mounted on the wing and a slot nozzle mounted on the wing. Nozzle exhaust plane locations with respect to the wing leading edge are varied from 10 to 46 percent chord (flaps retracted) with flap angles of 20 (takeoff altitude) and 60 (approach attitude). Perceived noise levels (PNL) are calculated as a function of flyover distance at 152 m altitude. From these plots, static EPNL values, defined as flyover relative noise levels, are calculated and plotted as a function of lift and thrust ratios. From such plots the acoustic benefits attributable to variations in nozzle/deflector/wing geometry at full scale are assessed for equal aerodynamic performance.
Observation of the wing deformation and the CFD study of cicada
NASA Astrophysics Data System (ADS)
Dai, Hu; Mohd Adam Das, Shahrizan; Luo, Haoxiang
2011-11-01
We studied the wing properties and kinematics of cicada when the 13-year species emerged in amazingly large numbers in middle Tennessee during May 2011. Using a high-speed camera, we recorded the wing motion of the insect and then reconstructed the three-dimensional wing kinematics using a video digitization software. Like many other insects, the deformation of the cicada wing is asymmetric between the downstroke and upstroke half cycles, and this particular deformation pattern would benefit production of the lift and propulsive forces. Both two-dimensional and three-dimensional CFD studies are carried out based on the reconstructed wing motion. The implication of the study on the role of the aerodynamic force in the wing deformation will be discussed. This work is sponsored by the NSF.
Zhao, Lü-Quan; Zhu, Dao-Hong
2014-01-01
Abstract The effects of environmental factors and appendage injury on the wing variation in Velarifictorus ornatus (Shiraki) (Orthoptera: Gryllidae) were investigated. The percentage of micropters was more than 95% when the nymphs were reared at constant photoperiods, and changing photoperiod did not affect wing variation in V. ornatus at 25 or 30°C. In the crowding experiment, the percentage of macropters was only 11.2% when the nymphs were reared separately at 25°C. In contrast, the percentage of macropters was significantly higher when the rearing density was increased to two nymphs per container and lower when the rearing density was increased to five or 10 nymphs per container. These results indicate that low and high rearing densities induce micropters, but intermediate rearing density stimulates the formation of macropters. Meanwhile, severance of appendages, such as antennae, femora, and tibiae, in the nymph stage exerted a micropterizing effect. The period sensitive to such stresses ranged from 35 to 60 days of nymph development. PMID:25368061
Harbison, Christopher W.; Clayton, Dale H.
2011-01-01
Reciprocal selective effects between coevolving species are often influenced by interactions with the broader ecological community. Community-level interactions may also influence macroevolutionary patterns of coevolution, such as cospeciation, but this hypothesis has received little attention. We studied two groups of ecologically similar feather lice (Phthiraptera: Ischnocera) that differ in their patterns of association with a single group of hosts. The two groups, “body lice” and “wing lice,” are both parasites of pigeons and doves (Columbiformes). Body lice are more host-specific and show greater population genetic structure than wing lice. The macroevolutionary history of body lice also parallels that of their columbiform hosts more closely than does the evolutionary history of wing lice. The closer association of body lice with hosts, compared with wing lice, can be explained if body lice are less capable of switching hosts than wing lice. Wing lice sometimes disperse phoretically on parasitic flies (Diptera: Hippoboscidae), but body lice seldom engage in this behavior. We tested the hypothesis that wing lice switch host species more often than body lice, and that the difference is governed by phoresis. Our results show that, where flies are present, wing lice switch to novel host species in sufficient numbers to establish viable populations on the new host. Body lice do not switch hosts, even where flies are present. Thus, differences in the coevolutionary history of wing and body lice can be explained by differences in host-switching, mediated by a member of the broader parasite community. PMID:21606369
Divergence with gene flow across a speciation continuum of Heliconius butterflies.
Supple, Megan A; Papa, Riccardo; Hines, Heather M; McMillan, W Owen; Counterman, Brian A
2015-09-24
A key to understanding the origins of species is determining the evolutionary processes that drive the patterns of genomic divergence during speciation. New genomic technologies enable the study of high-resolution genomic patterns of divergence across natural speciation continua, where taxa pairs with different levels of reproductive isolation can be used as proxies for different stages of speciation. Empirical studies of these speciation continua can provide valuable insights into how genomes diverge during speciation. We examine variation across a handful of genomic regions in parapatric and allopatric populations of Heliconius butterflies with varying levels of reproductive isolation. Genome sequences were mapped to 2.2-Mb of the H. erato genome, including 1-Mb across the red color pattern locus and multiple regions unlinked to color pattern variation. Phylogenetic analyses reveal a speciation continuum of pairs of hybridizing races and incipient species in the Heliconius erato clade. Comparisons of hybridizing pairs of divergently colored races and incipient species reveal that genomic divergence increases with ecological and reproductive isolation, not only across the locus responsible for adaptive variation in red wing coloration, but also at genomic regions unlinked to color pattern. We observe high levels of divergence between the incipient species H. erato and H. himera, suggesting that divergence may accumulate early in the speciation process. Comparisons of genomic divergence between the incipient species and allopatric races suggest that limited gene flow cannot account for the observed high levels of divergence between the incipient species. Our results provide a reconstruction of the speciation continuum across the H. erato clade and provide insights into the processes that drive genomic divergence during speciation, establishing the H. erato clade as a powerful framework for the study of speciation.
Iwata, Masaki; Otaki, Joji M
2016-02-01
Complex butterfly wing color patterns are coordinated throughout a wing by unknown mechanisms that provide undifferentiated immature scale cells with positional information for scale color. Because there is a reasonable level of correspondence between the color pattern element and scale size at least in Junonia orithya and Junonia oenone, a single morphogenic signal may contain positional information for both color and size. However, this color-size relationship has not been demonstrated in other species of the family Nymphalidae. Here, we investigated the distribution patterns of scale size in relation to color pattern elements on the hindwings of the peacock pansy butterfly Junonia almana, together with other nymphalid butterflies, Vanessa indica and Danaus chrysippus. In these species, we observed a general decrease in scale size from the basal to the distal areas, although the size gradient was small in D. chrysippus. Scales of dark color in color pattern elements, including eyespot black rings, parafocal elements, and submarginal bands, were larger than those of their surroundings. Within an eyespot, the largest scales were found at the focal white area, although there were exceptional cases. Similarly, ectopic eyespots that were induced by physical damage on the J. almana background area had larger scales than in the surrounding area. These results are consistent with the previous finding that scale color and size coordinate to form color pattern elements. We propose a ploidy hypothesis to explain the color-size relationship in which the putative morphogenic signal induces the polyploidization (genome amplification) of immature scale cells and that the degrees of ploidy (gene dosage) determine scale color and scale size simultaneously in butterfly wings. Copyright © 2015 Elsevier Ltd. All rights reserved.
Vortex coupling in trailing vortex-wing interactions
NASA Astrophysics Data System (ADS)
Chen, C.; Wang, Z.; Gursul, I.
2018-03-01
The interaction of trailing vortices of an upstream wing with rigid and flexible downstream wings has been investigated experimentally in a wind tunnel, using particle image velocimetry, hot-wire, force, and deformation measurements. Counter-rotating upstream vortices exhibit increased meandering when they are close to the tip of the downstream wing. The upstream vortex forms a pair with the vortex shed from the downstream wing and then exhibits large displacements around the wing tip. This coupled motion of the pair has been found to cause large lift fluctuations on the downstream wing. The meandering of the vortex pair occurs at the natural meandering frequency of the isolated vortex, with a low Strouhal number, and is not affected by the frequency of the large-amplitude wing oscillations if the downstream wing is flexible. The displacement of the leading vortex is larger than that of the trailing vortex; however, it causes highly correlated variations of the core radius, core vorticity, and circulation of the trailing vortex with the coupled meandering motion. In contrast, co-rotating vortices do not exhibit any increased meandering.
NASA Technical Reports Server (NTRS)
Groesbeck, D.; Von Glahn, U.
1979-01-01
The effects on acoustic characteristics of nozzle type and location on a wing for STOL engine over-the-wing configurations are assessed at full scale on the basis of model-scale data. Three types of nozzle configurations are evaluated: a circular nozzle with external deflector mounted above the wing, a slot nozzle with external deflector mounted on the wing and a slot nozzle mounted on the wing. Nozzle exhaust plane locations with respect to the wing leading edge are varied from 10 to 46 percent chord (flaps retracted) with flap angles of 20 deg (take-off attitude) and 60 deg (approach attitude). Perceived noise levels (PNL) are calculated as a function of flyover distance at 152 m altitude. From these plots, static EPNL values, defined as flyover relative noise levels, are calculated and plotted as a function of lift and thrust ratios. From such plots the acoustic benefits attributable to variations in nozzle/deflector/wing geometry at full scale are assessed for equal aerodynamic performance.
NASA Technical Reports Server (NTRS)
Merten, Kenneth F; Beck, Edgar B
1951-01-01
A smooth-water-landing investigation was conducted with a small seaplane to obtain experimental wing-bending-moment time histories together with time histories of the various parameters necessary for the prediction of wing bending moments during hydrodynamic forcing functions. The experimental results were compared with calculated results which include inertia-load effects and the effects of air-load variation during impact. The responses of the fundamental mode were calculated with the use of the measured hydrodynamic forcing functions. From these responses, the wing bending moments due to the hydrodynamic load were calculated according to the procedure given in R.M. No. 2221. The comparison of the time histories of the experimental and calculated wing bending moments showed good agreement both in phase relationship of the oscillations and in numerical values.
Close-Range Photogrammetric Measurement of Static Deflections for an Aeroelastic Supercritical Wing
NASA Technical Reports Server (NTRS)
Byrdsong, Thomas A.; Adams, Richard R.; Sandford, Maynard C.
1990-01-01
Close range photogrammetric measurements were made for the lower wing surface of a full span aspect ratio 10.3 aeroelastic supercritical research wing. The measurements were made during wind tunnel tests for quasi-steady pressure distributions on the wing. The tests were conducted in the NASA Langley Transonic Dynamics Tunnel at Mach numbers up to 0.90 and dynamic pressures up to 300 pounds per square foot. Deflection data were obtained for 57 locations on the wing lower surface using dual non-metric cameras. Representative data are presented as graphical overview to show variations and trends of spar deflection with test variables. Comparative data are presented for photogrammetric and cathetometric results of measurements for the wing tip deflections. A tabulation of the basic measurements is presented in a supplement to this report.
An experimental investigation of the subcritical and supercritical flow about a swept semispan wing
NASA Technical Reports Server (NTRS)
Lockman, W. K.; Seegmiller, H. L.
1983-01-01
An experimental investigation of the turbulent, subcritical and supercritical flow over a swept, semispan wing in a solid wall wind tunnel is described. The program was conducted over a range of Mach numbers, Reynolds numbers, and angles of attack to provide a variety of test cases for assessment of wing computer codes and tunnel wall interference effects. Wing flows both without and with three dimensional flow separation are included. Data include mean surface pressures for both the wing and tunnel walls; surface oil flow patterns on the wing; and mean velocity, flow field surveys. The results are given in tabular form and presented graphically to illustrate some of the effects of the test parameters. Comparisons of the wing pressure data with the results from two inviscid wing codes are also shown to assess the importance of viscous flow and tunnel wall effects.
Multiple scaled disorder in the photonic structure of Morpho rhetenor butterfly
NASA Astrophysics Data System (ADS)
Boulenguez, J.; Berthier, S.; Leroy, F.
2012-03-01
The iridescence of Morpho rhetenor butterfly is known to result from a photonic structure on wing scales, where multilayer interference and grating diffraction occur simultaneously. We characterize the disorder at the photonic structure length scale and at the butterfly scale. We measure the scattering pattern of the wing. Through RCWA and 1st Born approximation models, we link the different disorders to different features in the scattering patterns.
Flies compensate for unilateral wing damage through modular adjustments of wing and body kinematics
Iwasaki, Nicole A.; Elzinga, Michael J.; Melis, Johan M.; Dickinson, Michael H.
2017-01-01
Using high-speed videography, we investigated how fruit flies compensate for unilateral wing damage, in which loss of area on one wing compromises both weight support and roll torque equilibrium. Our results show that flies control for unilateral damage by rolling their body towards the damaged wing and by adjusting the kinematics of both the intact and damaged wings. To compensate for the reduction in vertical lift force due to damage, flies elevate wingbeat frequency. Because this rise in frequency increases the flapping velocity of both wings, it has the undesired consequence of further increasing roll torque. To compensate for this effect, flies increase the stroke amplitude and advance the timing of pronation and supination of the damaged wing, while making the opposite adjustments on the intact wing. The resulting increase in force on the damaged wing and decrease in force on the intact wing function to maintain zero net roll torque. However, the bilaterally asymmetrical pattern of wing motion generates a finite lateral force, which flies balance by maintaining a constant body roll angle. Based on these results and additional experiments using a dynamically scaled robotic fly, we propose a simple bioinspired control algorithm for asymmetric wing damage. PMID:28163885
Flies compensate for unilateral wing damage through modular adjustments of wing and body kinematics.
Muijres, Florian T; Iwasaki, Nicole A; Elzinga, Michael J; Melis, Johan M; Dickinson, Michael H
2017-02-06
Using high-speed videography, we investigated how fruit flies compensate for unilateral wing damage, in which loss of area on one wing compromises both weight support and roll torque equilibrium. Our results show that flies control for unilateral damage by rolling their body towards the damaged wing and by adjusting the kinematics of both the intact and damaged wings. To compensate for the reduction in vertical lift force due to damage, flies elevate wingbeat frequency. Because this rise in frequency increases the flapping velocity of both wings, it has the undesired consequence of further increasing roll torque. To compensate for this effect, flies increase the stroke amplitude and advance the timing of pronation and supination of the damaged wing, while making the opposite adjustments on the intact wing. The resulting increase in force on the damaged wing and decrease in force on the intact wing function to maintain zero net roll torque. However, the bilaterally asymmetrical pattern of wing motion generates a finite lateral force, which flies balance by maintaining a constant body roll angle. Based on these results and additional experiments using a dynamically scaled robotic fly, we propose a simple bioinspired control algorithm for asymmetric wing damage.
The Aerodynamics of Hovering Insect Flight. III. Kinematics
NASA Astrophysics Data System (ADS)
Ellington, C. P.
1984-02-01
Insects in free flight were filmed at 5000 frames per second to determine the motion of their wings and bodies. General comments are offered on flight behaviour and manoeuvrability. Changes in the tilt of the stroke plane with respect to the horizontal provides kinematic control of manoeuvres, analogous to the type of control used for helicopters. A projection analysis technique is described that solves for the orientation of the animal with respect to a camera-based coordinate system, giving full kinematic details for the longitudinal wing and body axes from single-view films. The technique can be applied to all types of flight where the wing motions are bilaterally symmetrical: forward, backward and hovering flight, as well as properly banked turns. An analysis of the errors of the technique is presented, and shows that the reconstructed angles for wing position should be accurate to within 1-2^circ in general. Although measurement of the angles of attack was not possible, visual estimations are given. Only 11 film sequences show flight velocities and accelerations that are small enough for the flight to be considered as `hovering'. Two sequences are presented for a hover-fly using an inclined stroke plane, and nine sequences of hovering with a horizontal stroke plane by another hover-fly, two crane-flies, a drone-fly, a ladybird beetle, a honey bee, and two bumble bees. In general, oscillations in the body position from its mean motion are within measurement error, about 1-2% of the wing length. The amplitudes of oscillation for the body angle are only a few degrees, but the phase relation of this oscillation to the wingbeat cycle could be determined for a few sequences. The phase indicates that the pitching moments governing the oscillations result from the wing lift at the ends of the wingbeat, and not from the wing drag or inertial forces. The mean pitching moment of the wings, which determines the mean body angle, is controlled by shifting the centre of lift over the cycle by changing the mean positional angle of the flapping wings. Deviations of the wing tip path from the stroke plane are never large, and no consistent pattern could be found for the wing paths of different insects; indeed, variations in the path were even observed for individual insects. The wing motion is not greatly different from simple harmonic motion, but does show a general trend towards higher accelerations and decelerations at either end of the wingbeat, with constant velocities during the middle of half-strokes. Root mean square and cube root mean cube angular velocities are on average about 4 and 9% lower than simple harmonic motion. Angles of attack are nearly constant during the middle of half-strokes, typically 35^circ at a position 70% along the wing length. The wing is twisted along its length, with angles of attack at the wing base some 10-20^circ greater than at the tip. The wings rotate through about 110^circ at either end of the wingbeat during 10-20% of the cycle period. The mean velocity of the wing edges during rotation is similar to the mean flapping velocity of the wing tip and greater than the flapping velocity for more proximal wing regions, which indicates that vortex shedding during rotation is comparable with that during flapping. The wings tend to rotate as a flat plate during the first half of rotation, which ends just before, or at, the end of the half-stroke. The hover-fly using an inclined stroke plane provides a notable exception to this general pattern: pronation is delayed and overlaps the beginning of the downstroke. The wing profile flexes along a more or less localized longitudinal axis during the second half of rotation, generating the `flip' profile postulated by Weis-Fogh for the hover-flies. This profile occurs to some extent for all of the insects, and is not exceptionally pronounced for the hover-fly. By the end of rotation the wings are nearly flat again, although a slight camber can sometimes be seen. Weis-Fogh showed that beneficial aerodynamic interference can result when the left and right wings come into contact during rotation at the end of the wingbeat. His `fling' mechanism creates the circulation required for wing lift on the subsequent half-stroke, and can be seen on my films of the Large Cabbage White butterfly, a plume moth, and the Mediterranean flour moth. However, their wings `peel' apart like two pieces of paper being separated, rather than fling open rigidly about the trailing edges. A `partial fling' was found for some insects, with the wings touching only along posterior wing areas. A `near fling' with the wings separated by a fraction of the chord was also observed for many insects. There is a continuous spectrum for the separation distance between the wings, in fact, and the separation can vary for a given insect during different manoeuvres. It is suggested that these variants on Weis-Fogh's fling mechanism also generate circulation for wing lift, although less effectively than a complete fling, and that changes in the separation distance may provide a fine control over the amount of lift produced.
4D metrology of flapping-wing micro air vehicle based on fringe projection
NASA Astrophysics Data System (ADS)
Zhang, Qican; Huang, Lei; Chin, Yao-Wei; Keong, Lau-Gih; Asundi, Anand
2013-06-01
Inspired by dominant flight of the natural flyers and driven by civilian and military purposes, micro air vehicle (MAV) has been developed so far by passive wing control but still pales in aerodynamic performance. Better understanding of flapping wing flight mechanism is eager to improve MAV's flight performance. In this paper, a simple and effective 4D metrology technique to measure full-field deformation of flapping membrane wing is presented. Based on fringe projection and 3D Fourier analysis, the fast and complex dynamic deformation, including wing rotation and wing stroke, of a flapping wing during its flight can be accurately reconstructed from the deformed fringe patterns recorded by a highspeed camera. An experiment was carried on a flapping-wing MAV with 5-cm span membrane wing beating at 30 Hz, and the results show that this method is effective and will be useful to the aerodynamicist or micro aircraft designer for visualizing high-speed complex wing deformation and consequently aid the design of flapping wing mechanism to enhanced aerodynamic performance.
Kusaba, Kiseki; Otaki, Joji M
2009-02-01
Butterfly wing color-patterns are a phenotypically coordinated array of scales whose color is determined as cellular interpretation outputs for morphogenic signals. Here we investigated distribution patterns of scale shape and size in relation to position and coloration on the hindwings of a nymphalid butterfly Junonia orithya. Most scales had a smooth edge but scales at and near the natural and ectopic eyespot foci and in the postbasal area were jagged. Scale size decreased regularly from the postbasal to distal areas, and eyespots occasionally had larger scales than the background. Reasonable correlations were obtained between the eyespot size and focal scale size in females. Histological and real-time individual observations of the color-pattern developmental sequence showed that the background brown and blue colors expanded from the postbasal to distal areas independently from the color-pattern elements such as eyespots. These data suggest that morphogenic signals for coloration directly or indirectly influence the scale shape and size and that the blue "background" is organized by a long-range signal from an unidentified organizing center in J. orithya.
A preliminary investigation of bird classification by Doppler radar
NASA Technical Reports Server (NTRS)
Martinson, L. W.
1973-01-01
A preliminary study of the application of Doppler radar to the classification of birds is reported. The desirability for improvements in bird classification stems primarily from the hazards they present to jet aircraft in flight and in the vicinity of airports. A secondary need exists in the study of bird migration. The wing body and tail motion of a bird in flight reflect signals which, when analyzed properly present a signature of wing beat pattern which is unique for each bird species. Although the results of this investigation did not validate the feasibility of classifying bird species, they do indicate that a more thorough investigation is warranted. Certain gross characteristics such as wing beat rates, multiple bird patterns, and bird maneuverability, were indicated clearly in the results. Large birds with slow wing beat rates appear to be the most optimum subject for further study with the X-band Doppler radar used in this investigation.
Latitudinal distribution of soft X-ray flares and dispairty in butterfly diagram
NASA Astrophysics Data System (ADS)
Pandey, K. K.; Yellaiah, G.; Hiremath, K. M.
2015-04-01
We present statistical analysis of about 63000 soft X-ray flare (class≥C) observed by geostationary operational environmental satellite (GOES) during the period 1976-2008. Class wise occurrence of soft X-ray (SXR) flare is in declining trend since cycle 21. The distribution pattern of cycle 21 shows the transit of hemispheric dominance of flare activity from northern to southern hemisphere and remains there during cycle 22 and 23. During the three cycles, 0-100, 21-300 latitude belts in southern hemisphere (SH) and 31-400 latitude belt in northern hemisphere (NH) are mightier. The 11-200 latitude belt of both hemisphere is mightiest. Correlation coefficient between consecutive latitude appears to be increasing from equator to poleward in northern hemisphere whereas pole to equatorward in southern hemisphere. Slope of the regression line fitted with asymmetry time series of daily flare counts is negative in all three cycles for different classes of flares. The yearly asymmetry curve fitted by a sinusoidal function varies from 5.6 to 11 years period and depends upon the intensity of flare. Variation, of curve fitted with wings of butterfly diagram, from first to second order polynomial suggests that latitudinal migration of flare activity varies from cycle to cycle, northern to southern hemisphere. The variation in slope of the butterfly wing of different flare class indicates the non uniform migration of flare activity.
Briolat, Emmanuelle Sophie; Zagrobelny, Mika; Olsen, Carl Erik; Blount, Jonathan D; Stevens, Martin
2018-05-16
The distinctive black and red wing pattern of six-spot burnet moths (Zygaena filipendulae, L.) is a classic example of aposematism, advertising their potent cyanide-based defences. While such warning signals provide a qualitatively honest signal of unprofitability, the evidence for quantitative honesty, whereby variation in visual traits could provide accurate estimates of individual toxicity, is more equivocal. Combining measures of cyanogenic glucoside content and wing colour from the perspective of avian predators, we investigate the relationship between coloration and defences in Z. filipendulae, to test signal honesty both within and across populations. There were no significant relationships between mean cyanogenic glucoside concentration and metrics of wing coloration across populations in males, yet in females higher cyanogenic glucoside levels were associated with smaller and lighter red forewing markings. Trends within populations were similarly inconsistent with quantitative honesty, and persistent differences between the sexes were apparent: larger females, carrying a greater total cyanogenic glucoside load, displayed larger but less conspicuous markings than smaller males, according to several colour metrics. The overall high aversiveness of cyanogenic glucosides and fluctuations in colour and toxin levels during an individual's lifetime may contribute to these results, highlighting generally important reasons why signal honesty should not always be expected in aposematic species. This article is protected by copyright. All rights reserved. This article is protected by copyright. All rights reserved.
Low Reynolds Number Wing Transients in Rotation and Translation
NASA Astrophysics Data System (ADS)
Jones, Anya; Schlueter, Kristy
2012-11-01
The unsteady aerodynamic forces and flow fields generated by a wing undergoing transient motions in both rotation and translation were investigated. An aspect ratio 2 flat plate wing at a 45 deg angle of attack was driven over 84 deg of rotation (3 chord-lengths of travel at 3/4 span) and 3 and 10 chord-lengths of translation in quiescent water at Reynolds numbers between 2,500 and 15,000. Flow visualization on the rotating wing revealed a leading edge vortex that lifted off of the wing surface, but remained in the vicinity of the wing for the duration of the wing stroke. A second spanwise vortex with strong axial flow was also observed. As the tip vortex grew, the leading edge vortex joined the tip vortex in a loop-like structure over the aft half of the wing. Near the leading edge, spanwise flow in the second vortex became entrained in the tip vortex near the corner of the wing. Unsteady force measurements revealed that lift coefficient increased through the constant-velocity portion of the wing stroke. Forces were compared for variations in wing acceleration and Reynolds number for both rotational and translational motions. The effect of tank blockage was investigated by repeating the experiments on multiple wings, varying the distance between the wing tip and tank wall. U.S. Air Force Research Laboratory, Summer Faculty Fellowship Program.
Control of Flow Structure on Low Swept Delta Wing with Steady Leading Edge Blowing
NASA Astrophysics Data System (ADS)
Ozturk, Ilhan; Zharfa, Mohammadreza; Yavuz, Mehmet Metin
2014-11-01
Interest in unmanned combat air vehicles (UCAVs) and micro air vehicles (MAVs) has stimulated investigation of the flow structure, as well as its control, on delta wings having low and moderate values of sweep angle. In the present study, the flow structure is characterized on a delta wing of low sweep 35-degree angle, which is subjected to steady leading edge blowing. The techniques of laser illuminated smoke visualization, laser Doppler anemometry (LDA), and surface pressure measurements are employed to investigate the steady and unsteady nature of the flow structure on delta wing, in relation to the dimensionless magnitude of the blowing coefficient. Using statistics and spectral analysis, unsteadiness of the flow structure is studied in detail. Different injection locations are utilized to apply different blowing patterns in order to identify the most efficient control, which provides the upmost change in the flow structure with the minimum energy input. The study aims to find the optimum flow control strategy to delay or to prevent the stall and possibly to reduce the buffeting on the wing surface. Since the blowing set-up is computer controlled, the unsteady blowing patterns compared to the present steady blowing patterns will be studied next. This project was supported by the Scientific and Technological Research Council of Turkey (Project Number: 3501 111M732).
Hypersonic aerodynamic characteristics of a family of power-law, wing body configurations
NASA Technical Reports Server (NTRS)
Townsend, J. C.
1973-01-01
The configurations analyzed are half-axisymmetric, power-law bodies surmounted by thin, flat wings. The wing planform matches the body shock-wave shape. Analytic solutions of the hypersonic small disturbance equations form a basis for calculating the longitudinal aerodynamic characteristics. Boundary-layer displacement effects on the body and the wing upper surface are approximated. Skin friction is estimated by using compressible, laminar boundary-layer solutions. Good agreement was obtained with available experimental data for which the basic theoretical assumptions were satisfied. The method is used to estimate the effects of power-law, fineness ratio, and Mach number variations at full-scale conditions. The computer program is included.
Aeroelastic Tailoring of Transport Wings Including Transonic Flutter Constraints
NASA Technical Reports Server (NTRS)
Stanford, Bret K.; Wieseman, Carol D.; Jutte, Christine V.
2015-01-01
Several minimum-mass optimization problems are solved to evaluate the effectiveness of a variety of novel tailoring schemes for subsonic transport wings. Aeroelastic stress and panel buckling constraints are imposed across several trimmed static maneuver loads, in addition to a transonic flutter margin constraint, captured with aerodynamic influence coefficient-based tools. Tailoring with metallic thickness variations, functionally graded materials, balanced or unbalanced composite laminates, curvilinear tow steering, and distributed trailing edge control effectors are all found to provide reductions in structural wing mass with varying degrees of success. The question as to whether this wing mass reduction will offset the increased manufacturing cost is left unresolved for each case.
Baier, David B; Gatesy, Stephen M; Dial, Kenneth P
2013-01-01
Past studies have shown that birds use their wings not only for flight, but also when ascending steep inclines. Uphill flap-running or wing-assisted incline running (WAIR) is used by both flight-incapable fledglings and flight-capable adults to retreat to an elevated refuge. Despite the broadly varying direction of travel during WAIR, level, and descending flight, recent studies have found that the basic wing path remains relatively invariant with reference to gravity. If so, joints undergo disparate motions to maintain a consistent wing path during those specific flapping modes. The underlying skeletal motions, however, are masked by feathers and skin. To improve our understanding of the form-functional relationship of the skeletal apparatus and joint morphology with a corresponding locomotor behavior, we used XROMM (X-ray Reconstruction of Moving Morphology) to quantify 3-D skeletal kinematics in chukars (Alectoris chukar) during WAIR (ascending with legs and wings) and ascending flight (AF, ascending with wings only) along comparable trajectories. Evidence here from the wing joints demonstrates that the glenohumeral joint controls the vast majority of wing movements. More distal joints are primarily involved in modifying wing shape. All bones are in relatively similar orientations at the top of upstroke during both behaviors, but then diverge through downstroke. Total excursion of the wing is much smaller during WAIR and the tip of the manus follows a more vertical path. The WAIR stroke appears "truncated" relative to ascending flight, primarily stemming from ca. 50% reduction in humeral depression. Additionally, the elbow and wrist exhibit reduced ranges of angular excursions during WAIR. The glenohumeral joint moves in a pattern congruent with being constrained by the acrocoracohumeral ligament. Finally, we found pronounced lateral bending of the furcula during the wingbeat cycle during ascending flight only, though the phasic pattern in chukars is opposite of that observed in starlings (Sturnus vulgaris).
Cellular behavior controlled by bio-inspired and geometry-tunable nanohairs.
Heo, Chaejeong; Jeong, Chanho; Im, Hyeon Seong; Kim, Jong Uk; Woo, Juhyun; Lee, Ji Yeon; Park, Byeonghak; Suh, Minah; Kim, Tae-Il
2017-11-23
A cicada wing has a biocidal feature of rupturing the membrane of cells, while the cactus spine can transmit a water drop to the stem of the plant. Both of these properties have evolved from their respective unique structures. Here, we endeavor to develop geometry-controllable nanohairs that mimic the cicada's wing-like vertical hairs and the cactus spine-like stooped hairs, and to quantitatively characterize the cell migration behavior of the hairy structures. It was found that the neuroblastoma cells are highly sensitive to the variation of surfaces: flat, vertical, and stooped nanohairs (100 nm diameter and 900 nm height). The cells on the vertical hairs showed significantly decreased proliferation. It was found that the behavior of cells cultured on stooped nanohairs is strongly influenced by the direction of the stooped pattern of hairs when we quantitatively measured the migration of cells on flat, vertical, and stooped structures. However, the cells on the flat structures showed random movement and the cells on the vertical nanohairs restricted the nanohair movement. Cells on the stooped structure showed higher forward migration preference compared to that of the other structures. Furthermore, we found that these cellular behaviors on the different patterns of nanohairs were affected by intracellular actin flament change. Consistent with these results, the vertical and stooped structures can facilitate the control of cell viability and guide directional migration for biomedical applications such as organogenesis.
Mettler, Raeann; Segelbacher, Gernot; Schaefer, H Martin
2015-01-01
Avian research has begun to reveal associations between candidate genes and migratory behaviors of captive birds, yet few studies utilize genotypic, morphometric, and phenological data from wild individuals. Previous studies have identified an association between ADCYAP1 polymorphism and autumn migratory behavior (restlessness, or zugunruhe), but little is known about the relationship between ADCYAP1 and spring migratory behavior. The timing of spring migration and arrival to the breeding ground are phenological traits which could be particularly favorable for establishing territories and acquiring mates, thus important to fitness and reproductive success. Here, we investigated how individual genotypic ADCYAP1 variation and phenotypic variation (wing length and shape) of blackcaps (Sylvia atricapilla) affect spring arrival date across nine natural populations in Europe. We hypothesized that longer alleles should be associated with earlier spring arrival dates and expected the effect on arrival date to be stronger for males as they arrive earlier. However, we found that longer wings were associated with earlier spring arrival to the breeding grounds for females, but not for males. Another female-specific effect indicated an interaction between ADCYAP1 allele size and wing pointedness on the response of spring arrival: greater allele size had a positive effect on spring arrival date for females with rounder wings, while a negative effect was apparent for females with more pointed wings. Also, female heterozygotes with pointed wing tips arrived significantly earlier than both homozygotes with pointed wings and heterozygotes with round wings. Stable isotope ratios (δ2H) of a subset of blackcaps captured in Freiburg in 2011 allowed us also to assign individuals to their main overwintering areas in northwest (NW) and southwest (SW) Europe. NW males arrived significantly earlier to the Freiburg breeding site than both SW males and females in 2011. NW females had more pointed wing tips compared to SW females, but no difference in ADCYAP1 allele size was found between the different migration routes.
NASA Technical Reports Server (NTRS)
Alford, William J , Jr
1957-01-01
The flow-field characteristics beneath swept and unswept wings as determined by potential-flow theory are compared with the experimentally determined flow fields beneath swept and unswept wing-fuselage combinations. The potential-flow theory utilized considered both spanwise and chordwise distributions of vorticity as well as the wing-thickness effects. The perturbation velocities induced by a unit horseshoe vortex are included in tabular form. The theoretical predictions of the flow-field characteristics were qualitatively correct in all cases considered, although there were indications that the magnitudes of the downwash angles tended to be overpredicted as the tip of the swept wing was approached and that the sidewash angles ahead of the unswept wing were underpredicted. The calculated effects of compressibility indicated that significant increases in the chordwise variation of flow angles and dynamic-pressure ratios should be expected in going from low to high subsonic speeds.
Marcus, Jeffrey M.; Evans, Travis M.
2008-01-01
The color patterns on the wings of butterflies have been an important model system in evolutionary developmental biology. A recent computational model tested genetic regulatory hierarchies hypothesized to underlie the formation of butterfly eyespot foci (Evans and Marcus, 2006). The computational model demonstrated that one proposed hierarchy was incapable of reproducing the known patterns of gene expression associated with eyespot focus determination in wild-type butterflies, but that two slightly modified alternative hierarchies were capable of reproducing all of the known gene expressions patterns. Here we extend the computational models previously implemented in Delphi 2.0 to two mutants derived from the squinting bush brown butterfly (Bicyclus anynana). These two mutants, comet and Cyclops, have aberrantly shaped eyespot foci that are produced by different mechanisms. The comet mutation appears to produce a modified interaction between the wing margin and the eyespot focus that results in a series of comet-shaped eyespot foci. The Cyclops mutation causes the failure of wing vein formation between two adjacent wing-cells and the fusion of two adjacent eyespot foci to form a single large elongated focus in their place. The computational approach to modeling pattern formation in these mutants allows us to make predictions about patterns of gene expression, which are largely unstudied in butterfly mutants. It also suggests a critical experiment that will allow us to distinguish between two hypothesized genetic regulatory hierarchies that may underlie all butterfly eyespot foci. PMID:18586070
NASA Technical Reports Server (NTRS)
Bobbitt, Percy J
1957-01-01
The lifting-surface sidewash behind rolling triangular wings has been derived for a range of supersonic Mach numbers for which the wing leading edges remain swept behind the mark cone emanating from the wing apex. Variations of the sidewash with longitudinal distance in the vertical plane of symmetry are presented in graphical form. An approximate expression for the sidewash has been developed by means of an approach using a horseshoe-vortex approximate-lifting-line theory. By use of this approximate expression, sidewash may be computed for wings of arbitrary plan form and span loading. A comparison of the sidewash computed by lifting-surface and lifting-line expressions for the triangular wing showed good agreement except in the vicinity of the trailing edge when the leading edge approached the sonic condition. An illustrative calculation has been made of the force induced by the wing sidewash on a vertical tail located in various longitudinal positions.
Pyrowolakis, George; Bergmann, Sven; Affolter, Markus
2011-01-01
The wing of the fruit fly, Drosophila melanogaster, with its simple, two-dimensional structure, is a model organ well suited for a systems biology approach. The wing arises from an epithelial sac referred to as the wing imaginal disc, which undergoes a phase of massive growth and concomitant patterning during larval stages. The Decapentaplegic (Dpp) morphogen plays a central role in wing formation with its ability to co-coordinately regulate patterning and growth. Here, we asked whether the Dpp signaling activity scales, i.e. expands proportionally, with the growing wing imaginal disc. Using new methods for spatial and temporal quantification of Dpp activity and its scaling properties, we found that the Dpp response scales with the size of the growing tissue. Notably, scaling is not perfect at all positions in the field and the scaling of target gene domains is ensured specifically where they define vein positions. We also found that the target gene domains are not defined at constant concentration thresholds of the downstream Dpp activity gradients P-Mad and Brinker. Most interestingly, Pentagone, an important secreted feedback regulator of the pathway, plays a central role in scaling and acts as an expander of the Dpp gradient during disc growth. PMID:22039350
Flow Structure on a Flapping Wing: Quasi-Steady Limit
NASA Astrophysics Data System (ADS)
Ozen, Cem; Rockwell, Donald
2011-11-01
The flapping motion of an insect wing typically involves quasi-steady motion between extremes of unsteady motion. This investigation characterizes the flow structure for the quasi-steady limit via a rotating wing in the form of a thin rectangular plate having a low aspect ratio (AR =1). Particle Image Velocimetry (PIV) is employed, in order to gain insight into the effects of centripetal and Coriolis forces. Vorticity, velocity and streamline patterns are used to describe the overall flow structure with an emphasis on the leading-edge vortex. A stable leading-edge vortex is maintained over effective angles of attack from 30° to 75° and it is observed that at each angle of attack the flow structure remains relatively same over the Reynolds number range from 3,600 to 14,500. The dimensionless circulation of the leading edge vortex is found to be proportional to the effective angle of attack. Quasi-three-dimensional construction of the flow structure is used to identify the different regimes along the span of the wing which is then complemented by patterns on cross flow planes to demonstrate the influence of root and tip swirls on the spanwise flow. The rotating wing results are also compared with the equivalent of translating wing to further illustrate the effects of the rotation.
NASA Technical Reports Server (NTRS)
Morris, O. A.; Fuller, D. E.; Watson, C. B.
1978-01-01
Tests were conducted in the Langley Unitary Plan wind tunnel at Mach numbers of 2.30. 2.70, and 2.95 to determine the performance, static stability, and control characteristics of a model of a fixed-wing supersonic cruise aircraft with a design Mach Number of 2.70 (SCAT 15-F-9898). The configuration had a 74 deg swept warped wing with a reflexed trailing edge and four engine nacelles mounted below the reflexed portion of the wing. A number of variations in the basic configuration were investigated; they included the effect of wing leading edge radius, the effect of various model components, and the effect of model control deflections.
NASA Technical Reports Server (NTRS)
Adamson, David; Boatright, William B
1957-01-01
An investigation of the nature of the flow field behind a rectangular wing of circular arc cross section has been conducted in the Langley 9-inch supersonic tunnel. Pitot- and static-pressure surveys covering a region of flow behind the wing have been made together with detailed pitot surveys throughout the region of the wake. In addition, the flow direction has been measured by means of a weathercocking vane. Theoretical calculations have been made to obtain the variation of both downwash and sidewash with angle of attack by using the superposition method of Lagerstrom, Graham, and Grosslight. In addition, the effect of wing thickness on the sidewash with the wing at 0 degree angle of attack has been evaluated.
Unique wing scale photonics of male Rajah Brooke's birdwing butterflies.
Wilts, Bodo D; Giraldo, Marco A; Stavenga, Doekele G
2016-01-01
Ultrastructures in butterfly wing scales can take many shapes, resulting in the often striking coloration of many butterflies due to interference of light. The plethora of coloration mechanisms is dazzling, but often only single mechanisms are described for specific animals. We have here investigated the male Rajah Brooke's birdwing, Trogonoptera brookiana, a large butterfly from Malaysia, which is marked by striking, colorful wing patterns. The dorsal side is decorated with large, iridescent green patterning, while the ventral side of the wings is primarily brown-black with small white, blue and green patches on the hindwings. Dense arrays of red hairs, creating a distinct collar as well as contrasting areas ventrally around the thorax, enhance the butterfly's beauty. The remarkable coloration is realized by a diverse number of intricate and complicated nanostructures in the hairs as well as the wing scales. The red collar hairs contain a broad-band absorbing pigment as well as UV-reflecting multilayers resembling the photonic structures of Morpho butterflies; the white wing patches consist of scales with prominent thin film reflectors; the blue patches have scales with ridge multilayers and these scales also have centrally concentrated melanin. The green wing areas consist of strongly curved scales, which possess a uniquely arranged photonic structure consisting of multilayers and melanin baffles that produces highly directional reflections. Rajah Brooke's birdwing employs a variety of structural and pigmentary coloration mechanisms to achieve its stunning optical appearance. The intriguing usage of order and disorder in related photonic structures in the butterfly wing scales may inspire novel optical materials as well as investigations into the development of these nanostructures in vivo.
Differential involvement of Hedgehog signaling in butterfly wing and eyespot development.
Tong, Xiaoling; Lindemann, Anna; Monteiro, Antónia
2012-01-01
Butterfly eyespots may have evolved from the recruitment of pre-existent gene circuits or regulatory networks into novel locations on the wing. Gene expression data suggests one such circuit, the Hedgehog (Hh) signaling pathway and its target gene engrailed (en), was recruited from a role in patterning the anterior-posterior insect wing axis to a role patterning butterfly eyespots. However, while Junonia coenia expresses hh and en both in the posterior compartment of the wing and in eyespot centers, Bicyclus anynana lacks hh eyespot-specific expression. This suggests that Hh signaling may not be functioning in eyespot development in either species or that it functions in J. coenia but not in B. anynana. In order to test these hypotheses, we performed functional tests of Hh signaling in these species. We investigated the effects of Hh protein sequestration during the larval stage on en expression levels, and on wing size and eyespot size in adults. Hh sequestration led to significantly reduced en expression and to significantly smaller wings and eyespots in both species. But while eyespot size in B. anynana was reduced proportionately to wing size, in J. coenia, eyespots were reduced disproportionately, indicating an independent role of Hh signaling in eyespot development in J. coenia. We conclude that while Hh signaling retains a conserved role in promoting wing growth across nymphalid butterflies, it plays an additional role in eyespot development in some, but not all, lineages of nymphalid butterflies. We discuss our findings in the context of alternative evolutionary scenarios that led to the differential expression of hh and other Hh pathway signaling members across nymphalid species.
Artificial insect wings of diverse morphology for flapping-wing micro air vehicles.
Shang, J K; Combes, S A; Finio, B M; Wood, R J
2009-09-01
The development of flapping-wing micro air vehicles (MAVs) demands a systematic exploration of the available design space to identify ways in which the unsteady mechanisms governing flapping-wing flight can best be utilized for producing optimal thrust or maneuverability. Mimicking the wing kinematics of biological flight requires examining the potential effects of wing morphology on flight performance, as wings may be specially adapted for flapping flight. For example, insect wings passively deform during flight, leading to instantaneous and potentially unpredictable changes in aerodynamic behavior. Previous studies have postulated various explanations for insect wing complexity, but there lacks a systematic approach for experimentally examining the functional significance of components of wing morphology, and for determining whether or not natural design principles can or should be used for MAVs. In this work, a novel fabrication process to create centimeter-scale wings of great complexity is introduced; via this process, a wing can be fabricated with a large range of desired mechanical and geometric characteristics. We demonstrate the versatility of the process through the creation of planar, insect-like wings with biomimetic venation patterns that approximate the mechanical properties of their natural counterparts under static loads. This process will provide a platform for studies investigating the effects of wing morphology on flight dynamics, which may lead to the design of highly maneuverable and efficient MAVs and insight into the functional morphology of natural wings.
Ontogeny of aerodynamics in mallards: comparative performance and developmental implications.
Dial, Terry R; Heers, Ashley M; Tobalske, Bret W
2012-11-01
Wing morphology correlates with flight performance and ecology among adult birds, yet the impact of wing development on aerodynamic capacity is not well understood. Recent work using chukar partridge (Alectoris chukar), a precocial flier, indicates that peak coefficients of lift and drag (C(L) and C(D)) and lift-to-drag ratio (C(L):C(D)) increase throughout ontogeny and that these patterns correspond with changes in feather microstructure. To begin to place these results in a comparative context that includes variation in life-history strategy, we used a propeller and force-plate model to study aerodynamic force production across a developmental series of the altricial-flying mallard (Anas platyrhynchos). We observed the same trend in mallards as reported for chukar in that coefficients of vertical (C(V)) and horizontal force (C(H)) and C(V):C(H) ratio increased with age, and that measures of gross-wing morphology (aspect ratio, camber and porosity) in mallards did not account for intraspecific trends in force production. Rather, feather microstructure (feather unfurling, rachis width, feather asymmetry and barbule overlap) all were positively correlated with peak C(V):C(H). Throughout ontogeny, mallard primary feathers became stiffer and less transmissive to air at both macroscale (between individual feathers) and microscale (between barbs/barbules/barbicels) levels. Differences between species were manifest primarily as heterochrony of aerodynamic force development. Chukar wings generated measurable aerodynamic forces early (<8 days), and improved gradually throughout a 100 day ontogenetic period. Mallard wings exhibited delayed aerodynamic force production until just prior to fledging (day 60), and showed dramatic improvement within a condensed 2-week period. These differences in timing may be related to mechanisms of escape used by juveniles, with mallards swimming to safety and chukar flap-running up slopes to take refuge. Future comparative work should test whether the need for early onset of aerodynamic force production in the chukar, compared with delayed, but rapid, change in the mallard wing, leads to a limited repertoire of flight behavior in adult chukar compared with mallards.
Mathematical modeling of planar cell polarity signaling in the Drosophila melanogaster wing
NASA Astrophysics Data System (ADS)
Amonlirdviman, Keith
Planar cell polarity (PCP) signaling refers to the coordinated polarization of cells within the plane of various epithelial tissues to generate sub-cellular asymmetry along an axis orthogonal to their apical-basal axes. For example, in the Drosophila wing, PCP is seen in the parallel orientation of hairs that protrude from each of the approximately 30,000 epithelial cells to robustly point toward the wing tip. Through a poorly understood mechanism, cell clones mutant for some PCP signaling components, including some, but not all alleles of the receptor frizzled, cause polarity disruptions of neighboring, wild-type cells, a phenomenon referred to as domineering nonautonomy. Previous models have proposed diffusible factors to explain nonautonomy, but no such factors have yet been found. This dissertation describes the mathematical modeling of PCP in the Drosophila wing, based on a contact dependent signaling hypothesis derived from experimental results. Intuition alone is insufficient to deduce that this hypothesis, which relies on a local feedback loop acting at the cell membrane, underlies the complex patterns observed in large fields of cells containing mutant clones, and others have argued that it cannot account for observed phenotypes. Through reaction-diffusion, partial differential equation modeling and simulation, the feedback loop is shown to fully reproduce PCP phenotypes, including domineering nonautonomy. The sufficiency of this model and the experimental validation of model predictions argue that previously proposed diffusible factors need not be invoked to explain PCP signaling and reveal how specific protein-protein interactions lead to autonomy or domineering nonautonomy. Based on these results, an ordinary differential equation model is derived to study the relationship of the feedback loop with upstream signaling components. The cadherin Fat transduces a cue to the local feedback loop, biasing the polarity direction of each cell toward the wing tip. The feedback loop then amplifies and propagates PCP across the pupal wing, but polarity information does not always propagate correctly across cells lacking Fat function. Using the simplified model, the presence and severity of polarity defects in fat clones is shown to be an inherent consequence of the feedback loop when confronted with irregular variations in cell geometry.
The effect of butterfly-scale inspired patterning on leading-edge vortex growth
NASA Astrophysics Data System (ADS)
Wilroy, Jacob; Lang, Amy; Wahidi, Redha
2014-11-01
Leading edge vortices (LEVs) are important for generating thrust and lift in flapping flight, and the surface patterning (scales) on butterfly wings is hypothesized to play a role in the vortex formation of the LEV. To simplify this complex flow problem, we designed an experiment to focus on the alteration of 2-D vortex development with a variation in surface patterning. Specifically we are interested in the secondary vorticity generated by the LEV interacting at the patterned surface and how this can affect the growth rate of the circulation in the LEV. For this experiment we used rapid-prototyped longitudinal and transverse square grooves attached to a flat plate and compared the vortex formation as the plate moved vertically. The plate is impulsively started in quiescent water and flow fields at Re = 1500, 3000, and 6000 are examined using Digital Particle Image Velocimetry (DPIV). The vortex formation time is 0.6 and is based on the flat plate travel length and chord length. Support for this research came from NSF REU Grant 1358991 and CBET 1335848.
The effect of butterfly-scale inspired patterning on leading-edge vortex growth
NASA Astrophysics Data System (ADS)
Wilroy, Jacob; Lang, Amy
2015-11-01
Leading edge vortices (LEVs) are important for generating thrust and lift in flapping flight, and the surface patterning (scales) on butterfly wings is hypothesized to play a role in the vortex formation of the LEV. To simplify this complex flow problem, an experiment was designed to focus on the alteration of 2-D vortex development with a variation in surface patterning. Specifically, the secondary vorticity generated by the LEV interacting at the patterned surface was studied and the subsequent affect on the growth rate of the circulation in the LEV. For this experiment we used butterfly inspired grooves attached to a flat plate and compared the vortex formation to a smooth plate case as the plate moved vertically. The plate is impulsively started in quiescent water and flow fields at Re = 1500, 3000, and 6000 are examined using Digital Particle Image Velocimetry (DPIV). The vortex formation time is 3.0 and is based on the flat plate travel length and chord length. We would like to thank the National Science Foundation REU Site Award 1358991 for funding this research.
Structural analysis and testing of a carbon-composite wing using fiber Bragg gratings
NASA Astrophysics Data System (ADS)
Nicolas, Matthew James
The objective of this study was to determine the deflected wing shape and the out-of-plane loads of a large-scale carbon-composite wing of an ultralight aerial vehicle using Fiber Bragg Grating (FBG) technology. The composite wing was instrumented with an optical fiber on its top and bottom surfaces positioned over the main spar, resulting in approximately 780 strain sensors bonded to the wings. The strain data from the FBGs was compared to that obtained from four conventional strain gages, and was used to obtain the out-of-plane loads as well as the wing shape at various load levels using NASA-developed real-time load and displacement algorithms. The composite wing measured 5.5 meters and was fabricated from laminated carbon uniaxial and biaxial prepreg fabric with varying laminate ply patterns and wall thickness dimensions. A three-tier whiffletree system was used to load the wing in a manner consistent with an in-flight loading condition.
Iwata, Masaki; Ohno, Yoshikazu; Otaki, Joji M.
2014-01-01
Butterfly wings are covered with regularly arranged single-colored scales that are formed at the pupal stage. Understanding pupal wing development is therefore crucial to understand wing color pattern formation. Here, we successfully employed real-time in vivo imaging techniques to observe pupal hindwing development over time in the blue pansy butterfly, Junonia orithya. A transparent sheet of epithelial cells that were not yet regularly arranged was observed immediately after pupation. Bright-field imaging and autofluorescent imaging revealed free-moving hemocytes and tracheal branches of a crinoid-like structure underneath the epithelium. The wing tissue gradually became gray-white, epithelial cells were arranged regularly, and hemocytes disappeared, except in the bordering lacuna, after which scales grew. The dynamics of the epithelial cells and scale growth were also confirmed by fluorescent imaging. Fluorescent in vivo staining further revealed that these cells harbored many mitochondria at the surface of the epithelium. Organizing centers for the border symmetry system were apparent immediately after pupation, exhibiting a relatively dark optical character following treatment with fluorescent dyes, as well as in autofluorescent images. The wing tissue exhibited slow and low-frequency contraction pulses with a cycle of approximately 10 to 20 minutes, mainly occurring at 2 to 3 days postpupation. The pulses gradually became slower and weaker and eventually stopped. The wing tissue area became larger after contraction, which also coincided with an increase in the autofluorescence intensity that might have been caused by scale growth. Examination of the pattern of color development revealed that the black pigment was first deposited in patches in the central areas of an eyespot black ring and a parafocal element. These results of live in vivo imaging that covered wide wing area for a long time can serve as a foundation for studying the cellular dynamics of living wing tissues in butterflies. PMID:24586829
Iwata, Masaki; Ohno, Yoshikazu; Otaki, Joji M
2014-01-01
Butterfly wings are covered with regularly arranged single-colored scales that are formed at the pupal stage. Understanding pupal wing development is therefore crucial to understand wing color pattern formation. Here, we successfully employed real-time in vivo imaging techniques to observe pupal hindwing development over time in the blue pansy butterfly, Junonia orithya. A transparent sheet of epithelial cells that were not yet regularly arranged was observed immediately after pupation. Bright-field imaging and autofluorescent imaging revealed free-moving hemocytes and tracheal branches of a crinoid-like structure underneath the epithelium. The wing tissue gradually became gray-white, epithelial cells were arranged regularly, and hemocytes disappeared, except in the bordering lacuna, after which scales grew. The dynamics of the epithelial cells and scale growth were also confirmed by fluorescent imaging. Fluorescent in vivo staining further revealed that these cells harbored many mitochondria at the surface of the epithelium. Organizing centers for the border symmetry system were apparent immediately after pupation, exhibiting a relatively dark optical character following treatment with fluorescent dyes, as well as in autofluorescent images. The wing tissue exhibited slow and low-frequency contraction pulses with a cycle of approximately 10 to 20 minutes, mainly occurring at 2 to 3 days postpupation. The pulses gradually became slower and weaker and eventually stopped. The wing tissue area became larger after contraction, which also coincided with an increase in the autofluorescence intensity that might have been caused by scale growth. Examination of the pattern of color development revealed that the black pigment was first deposited in patches in the central areas of an eyespot black ring and a parafocal element. These results of live in vivo imaging that covered wide wing area for a long time can serve as a foundation for studying the cellular dynamics of living wing tissues in butterflies.
Hoyal Cuthill, Jennifer F; Charleston, Michael
2015-12-01
Examples of long-term coevolution are rare among free-living organisms. Müllerian mimicry in Heliconius butterflies had been suggested as a key example of coevolution by early genetic studies. However, research over the last two decades has been dominated by the idea that the best-studied comimics, H. erato and H. melpomene, did not coevolve at all. Recently sequenced genes associated with wing color pattern phenotype offer a new opportunity to resolve this controversy. Here, we test the hypothesis of coevolution between H. erato and H. melpomene using Bayesian multilocus analysis of five color pattern genes and five neutral genetic markers. We first explore the extent of phylogenetic agreement versus conflict between the different genes. Coevolution is then tested against three aspects of the mimicry diversifications: phylogenetic branching patterns, divergence times, and, for the first time, phylogeographic histories. We show that all three lines of evidence are compatible with strict coevolution of the diverse mimicry wing patterns, contrary to some recent suggestions. Instead, these findings tally with a coevolutionary diversification driven primarily by the ecological force of Müllerian mimicry. © 2015 The Author(s). Evolution © 2015 The Society for the Study of Evolution.
NASA Technical Reports Server (NTRS)
Lomax, Harvard; Heaslet, Max A
1956-01-01
For a given wing and supersonic Mach number, the problem of shaping an adjoining fuselage so that the combination will have a low wave drag is considered. Only fuselages that can be simulated by singularities (multipoles) distributed along the body axis are studied. However, the optimum variations of such singularities are completely specified in terms of the given wing geometry. An application is made to an elliptic wing having a biconvex section, a thickness-chord ratio equal to 0.05 at the root, and an aspect ratio equal to 3. A comparison of the theoretical results with a wind-tunnel experiment is also presented.
NASA Technical Reports Server (NTRS)
Perry, B., III
1981-01-01
Comparisons are presented analytically predicted and experimental turbulence responses of a wind tunnel model of a DC-10 derivative wing equipped with an active control system. The active control system was designed for the purpose of flutter suppression, but it had additional benefit of alleviating gust loads (wing bending moment) by about 25%. Comparisions of various wing responses are presented for variations in active control system parameters and tunnel speed. The analytical turbulence responses were obtained using DYLOFLEX, a computer program for dynamic loads analyses of flexible airplanes with active controls. In general, the analytical predictions agreed reasonably well with the experimental data.
The effect of wing flexibility on sound generation of flapping wings.
Geng, Biao; Xue, Qian; Zheng, Xudong; Liu, Geng; Ren, Yan; Dong, Haibo
2017-12-13
In this study, the unsteady flow and acoustic characteristics of a three-dimensional (3D) flapping wing model of a Tibicen linnei cicada in forward-flight are numerically investigated. A single cicada wing is modelled as a membrane with a prescribed motion reconstructed from high-speed videos of a live insect. The numerical solution takes a hydrodynamic/acoustic splitting approach: the flow field is solved with an incompressible Navier-Stokes flow solver based on an immersed boundary method, and the acoustic field is solved with linearized perturbed compressible equations. The 3D simulation allows for the examination of both the directivity and frequency compositions of the flapping wing sound in a full space. Along with the flexible wing model, a rigid wing model that is extracted from real motion is also simulated to investigate the effects of wing flexibility. The simulation results show that the flapping sound is directional; the dominant frequency varies around the wing. The first and second frequency harmonics show different radiation patterns in the rigid and flexible wing cases, which are demonstrated to be highly associated with wing kinematics and loadings. Furthermore, the rotation and deformation in the flexible wing is found to help lower the sound strength in all directions.
An introgressed wing pattern acts as a mating cue.
Sánchez, Angela P; Pardo-Diaz, Carolina; Enciso-Romero, Juan; Muñoz, Astrid; Jiggins, Chris D; Salazar, Camilo; Linares, Mauricio
2015-06-01
Heliconius butterflies provide good examples of both homoploid hybrid speciation and ecological speciation. In particular, examples of adaptive introgression have been detected among the subspecies of Heliconius timareta, which acquired red color pattern elements from H. melpomene. We tested whether the introgression of red wing pattern elements into H. timareta florencia might also be associated with incipient reproductive isolation (RI) from its close relative, H. timareta subsp. nov., found in the eastern Andes. No choice experiments show a 50% reduction in mating between females of H. t. subsp. nov. and males of H .t. florencia, but not in the reciprocal direction. In choice experiments using wing models, males of H. timareta subsp. nov. approach and court red phenotypes less than their own, whereas males of H. t. florencia prefer models with a red phenotype. Intrinsic postzygotic isolation was not detected in crosses between these H. timareta races. These results suggest that a color pattern trait gained by introgression is triggering RI between H. timareta subsp. nov. and H. t. florencia. © 2015 The Author(s). Evolution © 2015 The Society for the Study of Evolution.
The wing and the eye: a parsimonious theory for scaling and growth control?
Romanova-Michaelides, Maria; Aguilar-Hidalgo, Daniel; Jülicher, Frank; Gonzalez-Gaitan, Marcos
2015-01-01
How a developing organ grows and patterns to its final shape is an important question in developmental biology. Studies of growth and patterning in the Drosophila wing imaginal disc have identified a key player, the morphogen Decapentaplegic (Dpp). These studies provided insights into our understanding of growth control and scaling: expansion of the Dpp gradient correlated with the growth of the tissue. A recent report on growth of a Drosophila organ other than the wing, the eye imaginal disc, prompts a reconsideration of our models of growth control. Despite striking differences between the two, the Dpp gradient scales with the target tissues of both organs and the growth of both the wing and the eye is controlled by Dpp. The goal of this review is to discuss whether a parsimonious model of scaling and growth control can explain the relationship between the Dpp gradient and growth in these two different developmental systems. © 2015 Wiley Periodicals, Inc.
Water Tunnel Flow Visualization Study Through Poststall of 12 Novel Planform Shapes
NASA Technical Reports Server (NTRS)
Gatlin, Gregory M.; Neuhart, Dan H.
1996-01-01
To determine the flow field characteristics of 12 planform geometries, a flow visualization investigation was conducted in the Langley 16- by 24-Inch Water Tunnel. Concepts studied included flat plate representations of diamond wings, twin bodies, double wings, cutout wing configurations, and serrated forebodies. The off-surface flow patterns were identified by injecting colored dyes from the model surface into the free-stream flow. These dyes generally were injected so that the localized vortical flow patterns were visualized. Photographs were obtained for angles of attack ranging from 10' to 50', and all investigations were conducted at a test section speed of 0.25 ft per sec. Results from the investigation indicate that the formation of strong vortices on highly swept forebodies can improve poststall lift characteristics; however, the asymmetric bursting of these vortices could produce substantial control problems. A wing cutout was found to significantly alter the position of the forebody vortex on the wing by shifting the vortex inboard. Serrated forebodies were found to effectively generate multiple vortices over the configuration. Vortices from 65' swept forebody serrations tended to roll together, while vortices from 40' swept serrations were more effective in generating additional lift caused by their more independent nature.
Installation effects on propeller wake/vortex induced structure-borne noise transmission
NASA Technical Reports Server (NTRS)
Unruh, J. F.
1989-01-01
A laboratory-based test apparatus was employed to investigate the effects of power-plant placement, engine/nacelle mass installation, and wing-to-fuselage attachment methods on propeller-induced structure-borne noise (SBN) transmission levels and their effects on noise-control measures. Data are presented showing SBN transmission is insensitive to propeller spanwise placement, however some sensitivity is seen in propeller-to-wing spacing. Installation of an engine/nacelle mass and variation in wing-to-fuselage attachments have measurable influences on SBN transmission and control measures.
NASA Technical Reports Server (NTRS)
Abel, I.; Newsom, J. R.
1981-01-01
Two flutter suppression control laws were synthesized, implemented, and tested on a low speed aeroelastic wing model of a DC-10 derivative. The methodology used to design the control laws is described. Both control laws demonstrated increases in flutter speed in excess of 25 percent above the passive wing flutter speed. The effect of variations in gain and phase on the closed loop performance was measured and compared with analytical predictions. The analytical results are in good agreement with experimental data.
Aerodynamic consequences of wing morphing during emulated take-off and gliding in birds.
Klaassen van Oorschot, Brett; Mistick, Emily A; Tobalske, Bret W
2016-10-01
Birds morph their wings during a single wingbeat, across flight speeds and among flight modes. Such morphing may allow them to maximize aerodynamic performance, but this assumption remains largely untested. We tested the aerodynamic performance of swept and extended wing postures of 13 raptor species in three families (Accipitridae, Falconidae and Strigidae) using a propeller model to emulate mid-downstroke of flapping during take-off and a wind tunnel to emulate gliding. Based on previous research, we hypothesized that (1) during flapping, wing posture would not affect maximum ratios of vertical and horizontal force coefficients (C V :C H ), and that (2) extended wings would have higher maximum C V :C H when gliding. Contrary to each hypothesis, during flapping, extended wings had, on average, 31% higher maximum C V :C H ratios and 23% higher C V than swept wings across all biologically relevant attack angles (α), and, during gliding, maximum C V :C H ratios were similar for the two postures. Swept wings had 11% higher C V than extended wings in gliding flight, suggesting flow conditions around these flexed raptor wings may be different from those in previous studies of swifts (Apodidae). Phylogenetic affiliation was a poor predictor of wing performance, due in part to high intrafamilial variation. Mass was only significantly correlated with extended wing performance during gliding. We conclude that wing shape has a greater effect on force per unit wing area during flapping at low advance ratio, such as take-off, than during gliding. © 2016. Published by The Company of Biologists Ltd.
Utilization of Optimization for Design of Morphing Wing Structures for Enhanced Flight
NASA Astrophysics Data System (ADS)
Detrick, Matthew Scott
Conventional aircraft control surfaces constrain maneuverability. This work is a comprehensive study that looks at both smart material and conventional actuation methods to achieve wing twist to potentially improve flight capability using minimal actuation energy while allowing minimal wing deformation under aerodynamic loading. A continuous wing is used in order to reduce drag while allowing the aircraft to more closely approximate the wing deformation used by birds while loitering. The morphing wing for this work consists of a skin supported by an underlying truss structure whose goal is to achieve a given roll moment using less actuation energy than conventional control surfaces. A structural optimization code has been written in order to achieve minimal wing deformation under aerodynamic loading while allowing wing twist under actuation. The multi-objective cost function for the optimization consists of terms that ensure small deformation under aerodynamic loading, small change in airfoil shape during wing twist, a linear variation of wing twist along the length of the wing, small deviation from the desired wing twist, minimal number of truss members, minimal wing weight, and minimal actuation energy. Hydraulic cylinders and a two member linkage driven by a DC motor are tested separately to provide actuation. Since the goal of the current work is simply to provide a roll moment, only one actuator is implemented along the wing span. Optimization is also used to find the best location within the truss structure for the actuator. The active structure produced by optimization is then compared to simulated and experimental results from other researchers as well as characteristics of conventional aircraft.
Gradual and contingent evolutionary emergence of leaf mimicry in butterfly wing patterns.
Suzuki, Takao K; Tomita, Shuichiro; Sezutsu, Hideki
2014-11-25
Special resemblance of animals to natural objects such as leaves provides a representative example of evolutionary adaptation. The existence of such sophisticated features challenges our understanding of how complex adaptive phenotypes evolved. Leaf mimicry typically consists of several pattern elements, the spatial arrangement of which generates the leaf venation-like appearance. However, the process by which leaf patterns evolved remains unclear. In this study we show the evolutionary origin and process for the leaf pattern in Kallima (Nymphalidae) butterflies. Using comparative morphological analyses, we reveal that the wing patterns of Kallima and 45 closely related species share the same ground plan, suggesting that the pattern elements of leaf mimicry have been inherited across species with lineage-specific changes of their character states. On the basis of these analyses, phylogenetic comparative methods estimated past states of the pattern elements and enabled reconstruction of the wing patterns of the most recent common ancestor. This analysis shows that the leaf pattern has evolved through several intermediate patterns. Further, we use Bayesian statistical methods to estimate the temporal order of character-state changes in the pattern elements by which leaf mimesis evolved, and show that the pattern elements changed their spatial arrangement (e.g., from a curved line to a straight line) in a stepwise manner and finally establish a close resemblance to a leaf venation-like appearance. Our study provides the first evidence for stepwise and contingent evolution of leaf mimicry. Leaf mimicry patterns evolved in a gradual, rather than a sudden, manner from a non-mimetic ancestor. Through a lineage of Kallima butterflies, the leaf patterns evolutionarily originated through temporal accumulation of orchestrated changes in multiple pattern elements.
Marcus, Jeffrey M; Evans, Travis M
2008-09-01
The color patterns on the wings of butterflies have been an important model system in evolutionary developmental biology. A recent computational model tested genetic regulatory hierarchies hypothesized to underlie the formation of butterfly eyespot foci [Evans, T.M., Marcus, J.M., 2006. A simulation study of the genetic regulatory hierarchy for butterfly eyespot focus determination. Evol. Dev. 8, 273-283]. The computational model demonstrated that one proposed hierarchy was incapable of reproducing the known patterns of gene expression associated with eyespot focus determination in wild-type butterflies, but that two slightly modified alternative hierarchies were capable of reproducing all of the known gene expressions patterns. Here we extend the computational models previously implemented in Delphi 2.0 to two mutants derived from the squinting bush brown butterfly (Bicyclus anynana). These two mutants, comet and Cyclops, have aberrantly shaped eyespot foci that are produced by different mechanisms. The comet mutation appears to produce a modified interaction between the wing margin and the eyespot focus that results in a series of comet-shaped eyespot foci. The Cyclops mutation causes the failure of wing vein formation between two adjacent wing-cells and the fusion of two adjacent eyespot foci to form a single large elongated focus in their place. The computational approach to modeling pattern formation in these mutants allows us to make predictions about patterns of gene expression, which are largely unstudied in butterfly mutants. It also suggests a critical experiment that will allow us to distinguish between two hypothesized genetic regulatory hierarchies that may underlie all butterfly eyespot foci.
Beldade, P; McMillan, W O; Papanicolaou, A
2008-02-01
Technological and conceptual advances of the last decade have led to an explosion of genomic data and the emergence of new research avenues. Evolutionary and ecological functional genomics, with its focus on the genes that affect ecological success and adaptation in natural populations, benefits immensely from a phylogenetically widespread sampling of biological patterns and processes. Among those organisms outside established model systems, butterflies offer exceptional opportunities for multidisciplinary research on the processes generating and maintaining variation in ecologically relevant traits. Here we highlight research on wing color pattern variation in two groups of Nymphalid butterflies, the African species Bicyclus anynana (subfamily Satyrinae) and species of the South American genus Heliconius (subfamily Heliconiinae), which are emerging as important systems for studying the nature and origins of functional diversity. Growing genomic resources including genomic and cDNA libraries, dense genetic maps, high-density gene arrays, and genetic transformation techniques are extending current gene mapping and expression profiling analysis and enabling the next generation of research questions linking genes, development, form, and fitness. Efforts to develop such resources in Bicyclus and Heliconius underscore the general challenges facing the larger research community and highlight the need for a community-wide effort to extend ongoing functional genomic research on butterflies.
NASA Technical Reports Server (NTRS)
Phillips, W. H. (Inventor)
1983-01-01
A cruciform wing structure for a solar powered aircraft is disclosed. Solar cells are mounted on horizontal wing surfaces. Wing surfaces with spanwise axis perpendicular to surfaces maintain these surfaces normal to the Sun's rays by allowing aircraft to be flown in a controlled pattern at a large bank angle. The solar airplane may be of conventional design with respect to fuselage, propeller and tail, or may be constructed around a core and driven by propeller mechanisms attached near the tips of the airfoils.
Simulation of Cell Patterning Triggered by Cell Death and Differential Adhesion in Drosophila Wing.
Nagai, Tatsuzo; Honda, Hisao; Takemura, Masahiko
2018-02-27
The Drosophila wing exhibits a well-ordered cell pattern, especially along the posterior margin, where hair cells are arranged in a zigzag pattern in the lateral view. Based on an experimental result observed during metamorphosis of Drosophila, we considered that a pattern of initial cells autonomously develops to the zigzag pattern through cell differentiation, intercellular communication, and cell death (apoptosis) and performed computer simulations of a cell-based model of vertex dynamics for tissues. The model describes the epithelial tissue as a monolayer cell sheet of polyhedral cells. Their vertices move according to equations of motion, minimizing the sum total of the interfacial and elastic energies of cells. The interfacial energy densities between cells are introduced consistently with an ideal zigzag cell pattern, extracted from the experimental result. The apoptosis of cells is modeled by gradually reducing their equilibrium volume to zero and by assuming that the hair cells prohibit neighboring cells from undergoing apoptosis. Based on experimental observations, we also assumed wing elongation along the proximal-distal axis. Starting with an initial cell pattern similar to the micrograph experimentally obtained just before apoptosis, we carried out the simulations according to the model mentioned above and successfully reproduced the ideal zigzag cell pattern. This elucidates a physical mechanism of patterning triggered by cell apoptosis theoretically and exemplifies, to our knowledge, a new framework to study apoptosis-induced patterning. We conclude that the zigzag cell pattern is formed by an autonomous communicative process among the participant cells. Copyright © 2018 Biophysical Society. All rights reserved.
NASA Technical Reports Server (NTRS)
Alford, William J., Jr.
1956-01-01
The flow-field characteristics beneath swept and unswept wings as determined by potential-flow theory are compared with the experimentally determined flow fields beneath swept and unswept wing-fuselage combinations. The potential-flow theory utilized considered both spanwise and chordwise distributions of vorticity as well as the wing-thickness effects. The perturbation velocities induced by a unit horseshoe vortex are included in tabular form. The results indicated that significant chordwise flow gradients existed beneath both swept and unswept wings at zero lift and throughout the lift range. The theoretical predictions of the flow-field characteristics were qualitatively correct in all cases considered, although there were indications that the magnitudes of the downwash angles tended to be overpredicted as the tip of the swept wing was approached and that the sidewash angles ahead of the unswept wing were underpredicted. The calculated effects of compressibility indicated that significant increases in the chordwise variation of flow angles and dynamic-pressure ratios should be expected in going from low to high subsonic speeds.
Installation and airspeed effects on jet shock-associated noise
NASA Technical Reports Server (NTRS)
Vonglahn, U.; Goodykoontz, J.
1975-01-01
Experimental acoustic data are presented to illustrate, at model scale, the effect of varying the nozzle-wing installation on shock-associated noise, statically and with airspeed. The variation in installations included nozzle only, nozzle under-the-wing (with and without flaps deflected), and nozzle over-the-wing (unattached flow). The nozzles used were a conical and a 6-tube mixer nozzle with a cold-flow nozzle pressure ratio of 2.1. A 33-cm diameter free jet was used to simulate airspeed. With the nozzle only, shock wave noise dominated the spectra in the forward quadrant, while jet mixing noise dominated in the rearward quadrant. Similar trends were observed when a wing (flaps retracted) was included. Shock noise was attenuated with an over-the-wing configuration and increased with an under-the-wing configuration (due to reflection from the wing surface). With increasing flap deflection (under-the-wing configuration), the jet-flap interaction noise exceeded the shock noise and became dominant in both quadrants. The free jet results showed that airspeed had no effect on shock noise. The free jet noise data were corrected for convective amplification to approximate flight and comparisons between the various configurations are made.
Effect of pylon cross-sectional geometries on propulsion integration for a low-wing transport
NASA Technical Reports Server (NTRS)
Ingraldi, Anthony M.; Naik, Dinesh A.; Pendergraft, Odis C., Jr.
1993-01-01
An experimental program was conducted in the Langley 16-Foot Transonic Tunnel to evaluate the performance effects of various types of pylons on a 1/17th-scale, low-wing transport model. The model wing was designed for cruise at a Mach number of 0.77 and a lift coefficient of 0.55. The pylons were tested at two wing semispan locations over a range of toe-in angles. The effects of toe-in angle were found to be minimal, but the variation in geometry had a more pronounced effect on the lift characteristics of the model. A pylon whose maximum thickness occurred at the wing trailing edge, known as a compression pylon, proved to be the best choice in terms of retaining the flow characteristics of the wing without pylons. Practical considerations such as structural viability may necessitate modification of the compression pylon concept in order to take advantage of its apparent benefits.
NASA Technical Reports Server (NTRS)
Chapman, Rowe, Jr; Morrow, John D
1952-01-01
A modified triangular wing of aspect ratio 2.53 having an airfoil section 3.7 percent thick at the root and 5.98 percent thick at the tip was designed in an attempt to improve the lift and drag characteristics of triangular wings. Free-flight drag and stability tests were made using rocket-propelled models equipped with the modified wing. The Mach number range of the test was from 0.70 to 1.37. Test results indicated the following: The lift-curve slope of wing plus fuselage approaches the theoretical value of wing alone at supersonic Mach numbers. The drag coefficient, based on total wing area, for wing plus interference was approximately 0.0035 at subsonic Mach numbers and 0.0080 at supersonic Mach numbers. The maximum shift in aerodynamic center for the complete configuration was 14 percent in the rearward direction from the forward position of 51.5 percent of mean aerodynamic chord at subsonic Mach numbers. The variation of lift and moment with angle of attack was linear at supersonic Mach numbers for the range of coefficients covered in the test. The high value of lift-curve slope was considered to be a significant result attributable to the wing modifications.
Percin, M; van Oudheusden, B W; de Croon, G C H E; Remes, B
2016-05-19
The study investigates the aerodynamic performance and the relation between wing deformation and unsteady force generation of a flapping-wing micro air vehicle in hovering flight configuration. Different experiments were performed where fluid forces were acquired with a force sensor, while the three-dimensional wing deformation was measured with a stereo-vision system. In these measurements, time-resolved power consumption and flapping-wing kinematics were also obtained under both in-air and in-vacuum conditions. Comparison of the results for different flapping frequencies reveals different wing kinematics and deformation characteristics. The high flapping frequency case produces higher forces throughout the complete flapping cycle. Moreover, a phase difference occurs in the variation of the forces, such that the low flapping frequency case precedes the high frequency case. A similar phase lag is observed in the temporal evolution of the wing deformation characteristics, suggesting that there is a direct link between the two phenomena. A considerable camber formation occurs during stroke reversals, which is mainly determined by the stiffener orientation. The wing with the thinner surface membrane displays very similar characteristics to the baseline wing, which implies the dominance of the stiffeners in terms of providing rigidity to the wing. Wing span has a significant effect on the aerodynamic efficiency such that increasing the span length by 4 cm results in a 6% enhancement in the cycle-averaged X-force to power consumption ratio compared to the standard DelFly II wings with a span length of 28 cm.
Development of Bird-like Micro Aerial Vehicle with Flapping and Feathering Wing Motions
NASA Astrophysics Data System (ADS)
Maglasang, Jonathan; Goto, Norihiro; Isogai, Koji
To investigate the feasibility of a highly efficient flapping system capable of avian maneuvers, such as rapid takeoff, hover and gliding, a full scale bird-like (ornithopter) flapping-wing micro aerial vehicle (MAV) shaped and patterned after a typical pigeon (Columba livia) has been designed and constructed. Both numerical and experimental methods have been used in the development of this vehicle. This flapping-wing micro aerial vehicle utilizes both the flapping and feathering motions of an avian wing by employing a novel flapping-feathering mechanism, which has been synthesized and constructed so as to best describe the properly coordinated flapping and feathering wing motions at phase angle difference of 90° in a horizontal steady level flight condition. This design allows high flapping and feathering amplitudes and is configurable for asymmetric wing motions which are desirable in high-speed flapping flight and maneuvering. The preliminary results indicate its viability as a practical and an efficient flapping-wing micro aerial vehicle.
Ge, Dengteng; Wu, Gaoxiang; Yang, Lili; Kim, Hye-Na; Hallwachs, Winnie; Burns, John M; Janzen, Daniel H; Yang, Shu
2017-07-11
Whiteness, although frequently apparent on the wings, legs, antennae, or bodies of many species of moths and butterflies, along with other colors and shades, has often escaped our attention. Here, we investigate the nanostructure and microstructure of white spots on the wings of Carystoides escalantei , a dusk-active and shade-inhabiting Costa Rican rain forest butterfly (Hesperiidae). On both males and females, two types of whiteness occur: angle dependent (dull or bright) and angle independent, which differ in the microstructure, orientation, and associated properties of their scales. Some spots on the male wings are absent from the female wings. Whether the angle-dependent whiteness is bright or dull depends on the observation directions. The angle-dependent scales also show enhanced retro-reflection. We speculate that the biological functions and evolution of Carystoides spot patterns, scale structures, and their varying whiteness are adaptations to butterfly's low light habitat and to airflow experienced on the wing base vs. wing tip.
Cicada-Wing-Inspired Self-Cleaning Antireflection Coatings on Polymer Substrates.
Chen, Ying-Chu; Huang, Zhe-Sheng; Yang, Hongta
2015-11-18
The cicada has transparent wings with remarkable self-cleaning properties and high transmittance over the whole visible spectral range, which is derived from periodic conical structures covering the wing surface. Here we report a scalable self-assembly technique for fabricating multifunctional optical coatings that mimic cicada-wing structures. Spin-coated two-dimensional non-close-packed colloidal crystals are utilized as etching masks to pattern subwavelength-structured cone arrays directly on polymer substrates. The resulting gratings exhibit broadband antireflection performance and superhydrophobic properties after surface modification. The dependence of the cone shape and size on the antireflective and self-cleaning properties has also been investigated in this study.
Sri Lankan Villagers' Attitudes toward Winged Bean, New Third World Food Source.
ERIC Educational Resources Information Center
Hacklander, Effie H.
1984-01-01
Discusses a survey on the acquisition, preparation, and consumption patterns of the winged bean as a food source in various parts of Sri Lanka. It was found that expanding the knowledge and production of indigenous legumes can help satisfy food needs of Third World countries. (JOW)
Static Performance of a Wing-Mounted Thrust Reverser Concept
NASA Technical Reports Server (NTRS)
Asbury, Scott C.; Yetter, Jeffrey A.
1998-01-01
An experimental investigation was conducted in the Jet-Exit Test Facility at NASA Langley Research Center to study the static aerodynamic performance of a wing-mounted thrust reverser concept applicable to subsonic transport aircraft. This innovative engine powered thrust reverser system is designed to utilize wing-mounted flow deflectors to produce aircraft deceleration forces. Testing was conducted using a 7.9%-scale exhaust system model with a fan-to-core bypass ratio of approximately 9.0, a supercritical left-hand wing section attached via a pylon, and wing-mounted flow deflectors attached to the wing section. Geometric variations of key design parameters investigated for the wing-mounted thrust reverser concept included flow deflector angle and chord length, deflector edge fences, and the yaw mount angle of the deflector system (normal to the engine centerline or parallel to the wing trailing edge). All tests were conducted with no external flow and high pressure air was used to simulate core and fan engine exhaust flows. Test results indicate that the wing-mounted thrust reverser concept can achieve overall thrust reverser effectiveness levels competitive with (parallel mount), or better than (normal mount) a conventional cascade thrust reverser system. By removing the thrust reverser system from the nacelle, the wing-mounted concept offers the nacelle designer more options for improving nacelle aero dynamics and propulsion-airframe integration, simplifying nacelle structural designs, reducing nacelle weight, and improving engine maintenance access.
NASA Technical Reports Server (NTRS)
Nguyen, Nhan; Ting, Eric; Lebofsky, Sonia
2015-01-01
This paper presents data analysis of a flexible wing wind tunnel model with a variable camber continuous trailing edge flap (VCCTEF) design for drag minimization tested at the University of Washington Aeronautical Laboratory (UWAL). The wind tunnel test was designed to explore the relative merit of the VCCTEF concept for improved cruise efficiency through the use of low-cost aeroelastic model test techniques. The flexible wing model is a 10%-scale model of a typical transport wing and is constructed of woven fabric composites and foam core. The wing structural stiffness in bending is tailored to be half of the stiffness of a Boeing 757-era transport wing while the torsional stiffness is about the same. This stiffness reduction results in a wing tip deflection of about 10% of the wing semi-span. The VCCTEF is a multi-segment flap design having three chordwise camber segments and five spanwise flap sections for a total of 15 individual flap elements. The three chordwise camber segments can be positioned appropriately to create a desired trailing edge camber. Elastomeric material is used to cover the gaps in between the spanwise flap sections, thereby creating a continuous trailing edge. Wind tunnel data analysis conducted previously shows that the VCCTEF can achieve a drag reduction of up to 6.31% and an improvement in the lift-to-drag ratio (L=D) of up to 4.85%. A method for estimating the bending and torsional stiffnesses of the flexible wingUWAL wind tunnel model from static load test data is presented. The resulting estimation indicates that the stiffness of the flexible wing is significantly stiffer in torsion than in bending by as much as 9 to 1. The lift prediction for the flexible wing is computed by a coupled aerodynamic-structural model. The coupled model is developed by coupling a conceptual aerodynamic tool Vorlax with a finite-element model of the flexible wing via an automated geometry deformation tool. Based on the comparison of the lift curve slope, the lift prediction for the rigid wing is in good agreement with the estimated lift coefficients derived from the wind tunnel test data. Due to the movement of the VCCTEF during the wind tunnel test, uncertainty in the lift prediction due to the indicated variations of the VCCTEF deflection is studied. The results show a significant spread in the lift prediction which contradicts the consistency in the aerodynamic measurements, thus suggesting that the indicated variations as measured by the VICON system may not be reliable. The lift prediction of the flexible wing agrees very well with the measured lift curve for the baseline configuration. The computed bending deflection and wash-out twist of the flexible wing also match reasonably well with the aeroelastic deflection measurements. The results demonstrate the validity of the aerodynamic-structural tool for use to analyze aerodynamic performance of flexible wings.
Slender wing theory including regions of embedded total pressure loss
NASA Technical Reports Server (NTRS)
Mccune, James E.; Tavares, T. Sean; Lee, Norman K. W.; Weissbein, David
1988-01-01
An aerodynamic theory of the flow about slender delta wings is described. The theory includes a treatment of the self-consistent development of the vortex wake patterns above the wing necessary to maintain smooth flow at the wing edges. The paper focuses especially on the formation within the wake of vortex 'cores' as embedded regions of total pressure loss, fed and maintained by umbilical vortex sheets emanating from the wing edges. Criteria are developed for determining the growing size and location of these cores, as well as the distribution and strength of the vorticity within them. In this paper, however, the possibility of vortex breakup is omitted. The aerodynamic consequences of the presence and evolution of the cores and the associated wake structure are illustrated and discussed. It is noted that wake history effects can have substantial influence on the distribution of normal force on the wing as well as on its magnitude.
The effects of leading edge modifications on the post-stall characteristics of wings
NASA Technical Reports Server (NTRS)
Winkelmann, A. E.; Barlow, J. B.; Saini, J. K.; Anderson, J. D., Jr.; Jones, E.
1980-01-01
An investigation of the effects of leading edge modifications on the post-stall characteristics of two rectangular planform wings in a series of low speed wind tunnel tests is presented. Abrupt discontinuities in the leading edge shape of the wings were produced by placing a nose glove over a portion of the span or by deflecting sections of a segmented leading edge flap. Six component balance data, oil flow visualization photographs, and pressure distribution measurements were obtained, and tests made to study the development of flow separation at stall on small scale planform wing models. Results of oil flow visualization tests at and beyond stall showed the formation of counter-rotating swirl patterns on the upper surface of the '2-D' and '3-D' wings, and results of a numerical lifting line technique applied to wings with leading edge modifications are included.
Analysis of high-aspect-ratio jet-flap wings of arbitrary geometry
NASA Technical Reports Server (NTRS)
Lissaman, P. B. S.
1973-01-01
An analytical technique to compute the performance of an arbitrary jet-flapped wing is developed. The solution technique is based on the method of Maskell and Spence in which the well-known lifting-line approach is coupled with an auxiliary equation providing the extra function needed in jet-flap theory. The present method is generalized to handle straight, uncambered wings of arbitrary planform, twist, and blowing (including unsymmetrical cases). An analytical procedure is developed for continuous variations in the above geometric data with special functions to exactly treat discontinuities in any of the geometric and blowing data. A rational theory for the effect of finite wing thickness is introduced as well as simplified concepts of effective aspect ratio for rapid estimation of performance.
Flight of Sharovipteryx mirabilis: the world's first delta-winged glider.
Dyke, G J; Nudds, R L; Rayner, J M V
2006-07-01
The 225 million-year-old reptile Sharovipteryx mirabilis was the world's first delta-winged glider; this remarkable animal had a flight surface composed entirely of a hind-limb membrane. We use standard delta-wing aerodynamics to reconstruct the flight of S. mirabilis demonstrating that wing shape could have been controlled simply by protraction of the femora at the knees, and by variation in incidence of a small forelimb canard. Our method has allowed us to address the question of how identifying realistic glide performance can be used to set limits on aerodynamic design in this small animal. Our novel interpretation of the bizarre flight mode of S. mirabilis is the first based directly on interpretation of the fossil itself and the first grounded in aerodynamics.
NASA Technical Reports Server (NTRS)
Mann, M. J.; Mercer, C. E.
1986-01-01
A transonic computational analysis method and a transonic design procedure have been used to design the wing and the canard of a forward-swept-wing fighter configuration for good transonic maneuver performance. A model of this configuration was tested in the Langley 16-Foot Transonic Tunnel. Oil-flow photographs were obtained to examine the wind flow patterns at Mach numbers from 0.60 to 0.90. The transonic theory gave a reasonably good estimate of the wing pressure distributions at transonic maneuver conditions. Comparison of the forward-swept-wing configuration with an equivalent aft-swept-wing-configuration showed that, at a Mach number of 0.90 and a lift coefficient of 0.9, the two configurations have the same trimmed drag. The forward-swept wing configuration was also found to have trimmed drag levels at transonic maneuver conditions which are comparable to those of the HiMAT (highly maneuverable aircraft technology) configuration and the X-29 forward-swept-wing research configuration. The configuration of this study was also tested with a forebody strake.
Subsonic annular wing theory with application to flow about nacelles
NASA Technical Reports Server (NTRS)
Mann, M. J.
1974-01-01
A method has recently been developed for calculating the flow over a subsonic nacelle at zero angle of attack. The method makes use of annular wing theory and boundary-layer theory and has shown good agreement with both experimental data and more complex theoretical solutions. The method permits variation of the mass flow by changing the size of a center body.
de Souza, João Maria Gomes Alencar; Molina, Wagner Franco; de Almeida, Lúcia Maria; de Gouveia, Milson Bezerra; de Macêdo, Francisco Pepino; Laumann, Raul Alberto; Paranhos, Beatriz Aguiar Jordão
2015-01-01
The sterile insect technique (SIT) is widely utilized in the biological control of fruit flies of the family Tephritidae, particularly against the Mediterranean fruit fly. This study investigated the interaction between mating success and morphometric variation in the wings and the production of acoustic signals among three male groups of Ceratitis capitata (Wiedemann): (1) wild males, (2) irradiated with Co-60 (steriles), and (3) irradiated (steriles) and treated with ginger oil. The canonical variate analysis discriminated two groups (males irradiated and males wild), based on the morphological shape of the wings. Among males that emit buzz signals, wild males obtained copulation more frequently than males in Groups 2 and 3. The individuals of Group 3 achieved more matings than those in Group 2. Wild males displayed lower pulse duration, higher intervals between pulses, and higher dominant frequency. Regarding the reproductive success, the morphological differences in the wings' shape between accepted and nonaccepted males are higher in wild males than in the irradiated ones. The present results can be useful in programs using the sterile insect technique for biological control of C. capitata. PMID:26075293
The Influence of Second Harmonic Phase and Amplitude Variation in Cyclically Pitching Wings
NASA Astrophysics Data System (ADS)
Culler, Ethan; Farnsworth, John
2017-11-01
From wind tunnel testing of a cyber-physical wing model, it has been found that the pitch trajectory for stall flutter is described by an array of higher harmonic frequencies with decaying energy content. These frequencies distort the stall flutter motion from that of a pure sinusoidal oscillation in pitch and can have a significant effect on the resulting force production. In order to understand how these higher harmonic frequencies contribute to the overall pitching moment characteristics of a wing in stall flutter, a rigid finite span wing model, with aspect ratio four, was pitched in the wind tunnel. The prescribed motion of the pitch cycle was varied by changing the amplitude ratio and phase of the second harmonic of the oscillation frequency. The second harmonic represents the second highest energy mode in the pitching cycle spectra. Pitching moment and planar particle image velocimetry data was collected. From these pitching trajectories, a significant dependence of pitching moment on both the phase and amplitude of the prescribed waveforms was found. Specifically, for the same amplitude ratio, variations in the phase produced changes of approximately 30 percent in the phase averaged pitching moment.
Qualitative skeletal correlates of wing shape in extant birds (Aves: Neoaves).
Hieronymus, Tobin L
2015-02-27
Among living fliers (birds, bats, and insects), birds display relatively high aspect ratios, a dimensionless shape variable that distinguishes long and narrow vs. short and broad wings. Increasing aspect ratio results in a functional tradeoff between low induced drag (efficient cruise) and increased wing inertia (difficult takeoff). Given the wide scope of its functional effects, the pattern of aspect ratio evolution is an important factor that contributes to the substantial ecological and phylogenetic diversity of living birds. However, because the feathers that define the wingtip (and hence wingspan and aspect ratio) often do not fossilize, resolution in the pattern of avian wing shape evolution is obscured by missing information. Here I use a comparative approach to investigate the relationship between skeletal proxies of flight feather attachment and wing shape. An accessory lobe of the internal index process of digit II-1, a bony correlate of distal primary attachment, shows weak but statistically significant relationships to aspect ratio and mass independent of other skeletal morphology. The dorsal phalangeal fossae of digit II-1, which house distal primaries VIII and IX, also show a trend of increased prominence with higher aspect ratio. Quill knobs on the ulna are examined concurrently, but do not show consistent signal with respect to wing shape. Although quill knobs are cited as skeletal correlates of flight performance in birds, their relationship to wing shape is inconsistent among extant taxa, and may reflect diverging selection pressures acting on a conserved architecture. In contrast, correlates of distal primary feather attachment on the major digit show convergent responses to increasing aspect ratio. In light of the diversity of musculoskeletal and integumentary mophology that underlies wing shape in different avian clades, it is unlikely that a single skeletal feature will show consistent predictive power across Neoaves. Confident inference of wing shape in basal ornithurine birds will require multiple lines of evidence, together with an understanding of clade-specific evolutionary trends within the crown.
Control law system for X-Wing aircraft
NASA Technical Reports Server (NTRS)
Lawrence, Thomas H. (Inventor); Gold, Phillip J. (Inventor)
1990-01-01
Control law system for the collective axis, as well as pitch and roll axes, of an X-Wing aircraft and for the pneumatic valving controlling circulation control blowing for the rotor. As to the collective axis, the system gives the pilot single-lever direct lift control and insures that maximum cyclic blowing control power is available in transition. Angle-of-attach de-coupling is provided in rotary wing flight, and mechanical collective is used to augment pneumatic roll control when appropriate. Automatic gain variations with airspeed and rotor speed are provided, so a unitary set of control laws works in all three X-Wing flight modes. As to pitch and roll axes, the system produces essentially the same aircraft response regardless of flight mode or condition. Undesirable cross-couplings are compensated for in a manner unnoticeable to the pilot without requiring pilot action, as flight mode or condition is changed. A hub moment feedback scheme is implemented, utilizing a P+I controller, significantly improving bandwidth. Limits protect aircraft structure from inadvertent damage. As to pneumatic valving, the system automatically provides the pressure required at each valve azimuth location, as dictated by collective, cyclic and higher harmonic blowing commands. Variations in the required control phase angle are automatically introduced, and variations in plenum pressure are compensated for. The required switching for leading, trailing and dual edge blowing is automated, using a simple table look-up procedure. Non-linearities due to valve characteristics of circulation control lift are linearized by map look-ups.
2012-01-01
16.64 Figure 3. Venation map of Manduca sexta forewing [11]. 2.4. Venation Insect wings are formed from a complex makeup of polymer based chains, Chitin ...for coloration, but may subtly influence flow patterns and boundary layer structure over wings [4, 24]. There is significant understanding of chitin ...biological specimen to vary the bonding chains, assemblage of nanofibers and crystalline structure, the material properties of chitin can vary over a
Comparative Analysis of Uninhibited and Constrained Avian Wing Aerodynamics
NASA Astrophysics Data System (ADS)
Cox, Jordan A.
The flight of birds has intrigued and motivated man for many years. Bird flight served as the primary inspiration of flying machines developed by Leonardo Da Vinci, Otto Lilienthal, and even the Wright brothers. Avian flight has once again drawn the attention of the scientific community as unmanned aerial vehicles (UAV) are not only becoming more popular, but smaller. Birds are once again influencing the designs of aircraft. Small UAVs operating within flight conditions and low Reynolds numbers common to birds are not yet capable of the high levels of control and agility that birds display with ease. Many researchers believe the potential to improve small UAV performance can be obtained by applying features common to birds such as feathers and flapping flight to small UAVs. Although the effects of feathers on a wing have received some attention, the effects of localized transient feather motion and surface geometry on the flight performance of a wing have been largely overlooked. In this research, the effects of freely moving feathers on a preserved red tailed hawk wing were studied. A series of experiments were conducted to measure the aerodynamic forces on a hawk wing with varying levels of feather movement permitted. Angle of attack and air speed were varied within the natural flight envelope of the hawk. Subsequent identical tests were performed with the feather motion constrained through the use of externally-applied surface treatments. Additional tests involved the study of an absolutely fixed geometry mold-and-cast wing model of the original bird wing. Final tests were also performed after applying surface coatings to the cast wing. High speed videos taken during tests revealed the extent of the feather movement between wing models. Images of the microscopic surface structure of each wing model were analyzed to establish variations in surface geometry between models. Recorded aerodynamic forces were then compared to the known feather motion and surface geometry to correlate the performance to these two features. The results of this study revealed that the performance of the bird wing was directly affected by feather motion. It was also found that the motion of covert and secondary covert feathers had the greatest influence on the performance. Increased coefficients of lift and drag were found when higher frequencies of these feathers were observed. Noticeable reductions in the coefficient of drag were found to be associated with micron level variations in the depth of surface features on the wing.
NASA Technical Reports Server (NTRS)
Heald, R H; Strother, D H; Monish, B H
1931-01-01
This report presents the results of an extension to higher angles of attack of the investigation of the rolling and yawing moments due to ailerons of various chords and spans on two airfoils having the Clark Y and U. S. A. 27 wings. The measurements were made at various angles of pitch but at zero angle of roll and yaw, the wing chord being set at an angle of +4 degrees to the fuselage axis. In the case of the Clark Y airfoil the measurements have been extended to a pitch angle of 40 degrees, using ailerons of span equal to 67 per cent of the wing semispan and chord equal to 20 and 30 per cent of the wing chord. The work was conducted on wing models of 60-inch span and 10-inch chord.
Real-time monitoring system of composite aircraft wings utilizing Fibre Bragg Grating sensor
NASA Astrophysics Data System (ADS)
Vorathin, E.; Hafizi, Z. M.; Che Ghani, S. A.; Lim, K. S.
2016-10-01
Embedment of Fibre Bragg Grating (FBG) sensor in composite aircraft wings leads to the advancement of structural condition monitoring. The monitored aircraft wings have the capability to give real-time response under critical loading circumstances. The main objective of this paper is to develop a real-time FBG monitoring system for composite aircraft wings to view real-time changes when the structure undergoes some static loadings and dynamic impact. The implementation of matched edge filter FBG interrogation system to convert wavelength variations to strain readings shows that the structure is able to response instantly in real-time when undergoing few loadings and dynamic impact. This smart monitoring system is capable of updating the changes instantly in real-time and shows the weight induced on the composite aircraft wings instantly without any error. It also has a good agreement with acoustic emission (AE) sensor in the dynamic test.
Obara, Yoshiaki; Ozawa, Gaku; Fukano, Yuya
2008-11-01
The British and Japanese subspecies of the cabbage butterfly, Pieris rapae , differ in terms of the UV reflectance of their wings ( Obara and Majerus, 2000 ). We studied the biogeographical distribution of the female cabbage butterfly having wings with UV reflectance around the Eurasian continent, and between Britain and Japan. For the study, we collected specimens from various locations. A gradient in the UV reflectance of the wings appears to exist along the west-east axis; reflectance was higher toward the east and reached a peak in butterflies in Japan. The UV-reflecting Japanese subspecies Pieris rapae crucivora was found exclusively along the east coast of the Eurasian continent. This suggests that the Japanese subspecies has evolved from a continental ancestor, with females having UV-absorbing wings. We discuss the results of our study with regard to the evolution and adaptive significance of UV coloration in the Japanese subspecies.
NASA Technical Reports Server (NTRS)
Bamber, M J; House, R O
1937-01-01
An investigation was made to determine the spinning characteristics of Clark Y monoplane wings with different plan forms. A rectangular wing and a wing tapered 5:2, both with rounded tips, were tested on the N.A.C.A. spinning balance in the 5-foot vertical wind tunnel. The aerodynamic characteristics of the models and a prediction of the angles of sideslip for steady spins are given. Also included is an estimate of the yawning moment that must be furnished by the parts of the airplane to balance the inertia couples and wing yawing moment for spinning equilibrium. The effects on the spin of changes in plan form and of variations of some of the important parameters are discussed and the results are compared with those for a rectangular wing with square tips. It is concluded that for a conventional monoplane using Clark Y wing the sideslip will be algebraically larger for the wing with the rounded tip than for the wing with the square tip and will be largest for the tapered wing. The effect of plan form on the spin will vary with the type of airplane; and the provision of a yawing-moment coefficient of -0.025 (i.e., opposing the spin) by the tail, fuselage, and interference effects will insure against the attainment of equilibrium on a steady spin for any of the plan forms tested and for any of the parameters used in the analysis.
NASA Technical Reports Server (NTRS)
Suarez, Carlos J.; Smith, Brooke C.; Kramer, Brian R.; Ng, T. Terry; Ong, Lih-Yenn; Malcolm, Gerald N.
1993-01-01
Free-to-roll tests were conducted in water and wind tunnels in an effort to investigate the mechanisms of wing rock on a NASP-type vehicle. The configuration tested consisted of a highly-slender forebody and a 78 deg swept delta wing. In the water tunnel test, extensive flow visualization was performed and roll angle histories were obtained. In the wind tunnel test, the roll angle, forces and moments, and limited forebody and wing surface pressures were measured during the wing rock motion. A limit cycle oscillation was observed for angles of attack between 22 deg and 30 deg. In general, the experiments confirmed that the main flow phenomena responsible for the wing-body-tail wing rock are the interactions between the forebody and the wing vortices. The variation of roll acceleration (determined from the second derivative of the roll angle time history) with roll angle clearly slowed the energy balance necessary to sustain the limit cycle oscillation. Different means of suppressing wing rock by controlling the forebody vortices using small blowing jets were also explored. Steady blowing was found to be capable of suppressing wing rock, but significant vortex asymmetrices are created, causing the model to stop at a non-zero roll angle. On the other hand, alternating pulsed blowing on the left and right sides of the fore body was demonstrated to be a potentially effective means of suppressing wing rock and eliminating large asymmetric moments at high angles of attack.
Experimental and numerical studies of beetle-inspired flapping wing in hovering flight.
Van Truong, Tien; Le, Tuyen Quang; Park, Hoon Cheol; Byun, Doyoung
2017-05-17
In this paper, we measure unsteady forces and visualize 3D vortices around a beetle-like flapping wing model in hovering flight by experiment and numerical simulation. The measurement of unsteady forces and flow patterns around the wing were conducted using a dynamically scaled wing model in the mineral-oil tank. The wing kinematics were directly derived from the experiment of a real beetle. The 3D flow structures of the flapping wing were captured by using air bubble visualization while forces were measured by a sensor attached at the wing base. In comparison, the size and topology of spiral leading edge vortex, trailing edge vortex and tip vortex are well matched from experimental and numerical studies. In addition, the time history of forces calculated from numerical simulation is also similar to that from theforce measurement. A difference of average force is in order of 10 percent. The results indicate that the leading edge vortex due to rotational acceleration at the end of the stroke during flapping wing causes significant reduction of lift. The present study provides useful information on hover flight to develop a beetle-like flapping wing Micro Air Vehicle.
Global-Local Analysis and Optimization of a Composite Civil Tilt-Rotor Wing
NASA Technical Reports Server (NTRS)
Rais-Rohani, Masound
1999-01-01
This report gives highlights of an investigation on the design and optimization of a thin composite wing box structure for a civil tilt-rotor aircraft. Two different concepts are considered for the cantilever wing: (a) a thin monolithic skin design, and (b) a thick sandwich skin design. Each concept is examined with three different skin ply patterns based on various combinations of 0, +/-45, and 90 degree plies. The global-local technique is used in the analysis and optimization of the six design models. The global analysis is based on a finite element model of the wing-pylon configuration while the local analysis uses a uniformly supported plate representing a wing panel. Design allowables include those on vibration frequencies, panel buckling, and material strength. The design optimization problem is formulated as one of minimizing the structural weight subject to strength, stiffness, and d,vnamic constraints. Six different loading conditions based on three different flight modes are considered in the design optimization. The results of this investigation reveal that of all the loading conditions the one corresponding to the rolling pull-out in the airplane mode is the most stringent. Also the frequency constraints are found to drive the skin thickness limits, rendering the buckling constraints inactive. The optimum skin ply pattern for the monolithic skin concept is found to be (((0/+/-45/90/(0/90)(sub 2))(sub s))(sub s), while for the sandwich skin concept the optimal ply pattern is found to be ((0/+/-45/90)(sub 2s))(sub s).
Inhibition of Shh signalling in the chick wing gives insights into digit patterning and evolution.
Pickering, Joseph; Towers, Matthew
2016-10-01
In an influential model of pattern formation, a gradient of Sonic hedgehog (Shh) signalling in the chick wing bud specifies cells with three antero-posterior positional values, which give rise to three morphologically different digits by a self-organizing mechanism with Turing-like properties. However, as four of the five digits of the mouse limb are morphologically similar in terms of phalangeal pattern, it has been suggested that self-organization alone could be sufficient. Here, we show that inhibition of Shh signalling at a specific stage of chick wing development results in a pattern of four digits, three of which can have the same number of phalanges. These patterning changes are dependent on a posterior extension of the apical ectodermal ridge, and this also allows the additional digit to arise from the Shh-producing cells of the polarizing region - an ability lost in ancestral theropod dinosaurs. Our analyses reveal that, if the specification of antero-posterior positional values is curtailed, self-organization can then produce several digits with the same number of phalanges. We present a model that may give important insights into how the number of digits and phalanges has diverged during the evolution of avian and mammalian limbs. © 2016. Published by The Company of Biologists Ltd.
Zhao, Lü-Quan; Zhu, Dao-Hong
2014-01-01
The effects of environmental factors and appendage injury on the wing variation in Velarifictorus ornatus (Shiraki) (Orthoptera: Gryllidae) were investigated. The percentage of micropters was more than 95% when the nymphs were reared at constant photoperiods, and changing photoperiod did not affect wing variation in V. ornatus at 25 or 30°C. In the crowding experiment, the percentage of macropters was only 11.2% when the nymphs were reared separately at 25°C. In contrast, the percentage of macropters was significantly higher when the rearing density was increased to two nymphs per container and lower when the rearing density was increased to five or 10 nymphs per container. These results indicate that low and high rearing densities induce micropters, but intermediate rearing density stimulates the formation of macropters. Meanwhile, severance of appendages, such as antennae, femora, and tibiae, in the nymph stage exerted a micropterizing effect. The period sensitive to such stresses ranged from 35 to 60 days of nymph development. This is an open access paper. We use the Creative Commons Attribution 3.0 license that permits unrestricted use, provided that the paper is properly attributed.
NASA Technical Reports Server (NTRS)
Henderson, W. P.; Huffman, J. K.
1974-01-01
An investigation has been conducted to determine the effects of configuration variables on the lateral-directional stability characteristics of a wing-fuselage configuration. The variables under study included variations in the location of a single center-line vertical tail and twin vertical tails, wing height, fuselage strakes, and horizontal tails. The study was conducted in the Langley high-speed 7-by 10-foot tunnel at a Mach number of 0.30, at angles of attack up to 44 deg and at sideslip angles of 0 deg and plus or minus 5 deg.
Structureborne noise measurements on a small twin-engine aircraft
NASA Technical Reports Server (NTRS)
Cole, J. E., III; Martini, K. F.
1988-01-01
Structureborne noise measurements performed on a twin-engine aircraft (Beechcraft Baron) are reported. There are two overall objectives of the test program. The first is to obtain data to support the development of analytical models of the wing and fuselage, while the second is to evaluate effects of structural parameters on cabin noise. Measurements performed include structural and acoustic responses to impact excitation, structural and acoustic loss factors, and modal parameters of the wing. Path alterations include added mass to simulate fuel, variations in torque of bolts joining wing and fuselage, and increased acoustic absorption. Conclusions drawn regarding these measurements are presented.
Structureborne noise measurements on a small twin-engine aircraft
NASA Astrophysics Data System (ADS)
Cole, J. E., III; Martini, K. F.
1988-06-01
Structureborne noise measurements performed on a twin-engine aircraft (Beechcraft Baron) are reported. There are two overall objectives of the test program. The first is to obtain data to support the development of analytical models of the wing and fuselage, while the second is to evaluate effects of structural parameters on cabin noise. Measurements performed include structural and acoustic responses to impact excitation, structural and acoustic loss factors, and modal parameters of the wing. Path alterations include added mass to simulate fuel, variations in torque of bolts joining wing and fuselage, and increased acoustic absorption. Conclusions drawn regarding these measurements are presented.
Flow structure of vortex-wing interaction
NASA Astrophysics Data System (ADS)
McKenna, Christopher K.
Impingement of a streamwise-oriented vortex upon a fin, tail, blade or wing represents a fundamental class of flow-structure interaction that extends across a range of applications. This interaction can give rise to time-averaged loading, as well as unsteady loading known as buffeting. The loading is sensitive to parameters of the incident vortex as well as the location of vortex impingement on the downstream aerodynamic surface, generically designated as a wing. Particle image velocimetry is employed to determine patterns of velocity, vorticity, swirl ratio, and streamlines on successive cross-flow planes upstream of and along the wing, which lead to volume representations and thereby characterization of the interaction. At locations upstream of the leading edge of the wing, the evolution of the incident vortex is affected by the presence of the wing, and is highly dependent on the spanwise location of vortex impingement. Even at spanwise locations of impingement well outboard of the wing tip, a substantial influence on the structure of the incident vortex at locations significantly upstream of the leading edge of the wing was observed. For spanwise locations close to or intersecting the vortex core, the effects of upstream influence of the wing on the vortex are to: decrease the swirl ratio; increase the streamwise velocity deficit; decrease the streamwise vorticity; increase the azimuthal vorticity; increase the upwash; decrease the downwash; and increase the root-mean-square fluctuations of both streamwise velocity and vorticity. The interrelationship between these effects is addressed, including the rapid attenuation of axial vorticity in presence of an enhanced defect of axial velocity in the central region of the vortex. Moreover, when the incident vortex is aligned with, or inboard of, the tip of the wing, the swirl ratio decreases to values associated with instability of the vortex, giving rise to enhanced values of azimuthal vorticity relative to the streamwise (axial) vorticity, as well as relatively large root-mean-square values of streamwise velocity and vorticity. Along the chord of the wing, the vortex interaction gives rise to distinct modes, which may involve either enhancement or suppression of the vortex generated at the tip of the wing. These modes are classified and interpreted in conjunction with computed modes at the Air Force Research Laboratory. Occurrence of a given mode of interaction is predominantly determined by the dimensionless location of the incident vortex relative to the tip of the wing and is generally insensitive to the Reynolds number and dimensionless circulation of the incident vortex. The genesis of the basic modes of interaction is clarified using streamline topology with associated critical points. Whereas formation of an enhanced tip vortex involves a region of large upwash in conjunction with localized flow separation, complete suppression of the tip vortex is associated with a small-scale separation-attachment bubble bounded by downwash at the wing tip. Oscillation of the wing at an amplitude and velocity nearly two orders of magnitude smaller than the wing chord and free stream velocity respectively can give rise to distinctive patterns of upwash, downwash, and shed vorticity, which are dependent on the outboard displacement of the incident vortex relative to the wing tip. Moreover, these patterns are a strong function of the phase of the wing motion during its oscillation cycle. At a given value of phase, the wing oscillation induces upwash that is reinforced by the upwash of the incident vortex, giving a maximum value of net upwash. Conversely, when these two origins of upwash counteract, rather than reinforce, one another during the oscillation cycle, the net upwash has its minimum value. Analogous interpretations hold for regions of maximum and minimum net downwash located outboard of the regions of upwash. During the oscillation cycle of the wing, the magnitude and scale of the vorticity shed from the tip of the wing are directly correlated with the net upwash, which takes different forms related to the outboard displacement of the incident vortex. As the location of the incident vortex is displaced towards the wing tip, both the maximum upwash and the maximum vorticity of the tip vortex initially increase, then decrease. For the limiting case where the incident vortex impinges directly upon the tip of the wing, there is no tip vortex or induced region of upwash. Furthermore, at small values of vortex displacement from the wing tip, the position of the incident vortex varies significantly from its nominal position during the oscillation cycle. For all locations of the incident vortex, it is shown that, despite the small amplitude of the wing motion, the flow topology is fundamentally different at maximum positive and negative values of the wing velocity, that is, they are not symmetric.
Structural dynamics and aerodynamics measurements of biologically inspired flexible flapping wings.
Wu, P; Stanford, B K; Sällström, E; Ukeiley, L; Ifju, P G
2011-03-01
Flapping wing flight as seen in hummingbirds and insects poses an interesting unsteady aerodynamic problem: coupling of wing kinematics, structural dynamics and aerodynamics. There have been numerous studies on the kinematics and aerodynamics in both experimental and computational cases with both natural and artificial wings. These studies tend to ignore wing flexibility; however, observation in nature affirms that passive wing deformation is predominant and may be crucial to the aerodynamic performance. This paper presents a multidisciplinary experimental endeavor in correlating a flapping micro air vehicle wing's aeroelasticity and thrust production, by quantifying and comparing overall thrust, structural deformation and airflow of six pairs of hummingbird-shaped membrane wings of different properties. The results show that for a specific spatial distribution of flexibility, there is an effective frequency range in thrust production. The wing deformation at the thrust-productive frequencies indicates the importance of flexibility: both bending and twisting motion can interact with aerodynamic loads to enhance wing performance under certain conditions, such as the deformation phase and amplitude. By measuring structural deformations under the same aerodynamic conditions, beneficial effects of passive wing deformation can be observed from the visualized airflow and averaged thrust. The measurements and their presentation enable observation and understanding of the required structural properties for a thrust effective flapping wing. The intended passive responses of the different wings follow a particular pattern in correlation to their aerodynamic performance. Consequently, both the experimental technique and data analysis method can lead to further studies to determine the design principles for micro air vehicle flapping wings.
Localised JAK/STAT Pathway Activation Is Required for Drosophila Wing Hinge Development
Johnstone, Kirsty; Wells, Richard E.; Strutt, David; Zeidler, Martin P.
2013-01-01
Extensive morphogenetic remodelling takes place during metamorphosis from a larval to an adult insect body plan. These changes are particularly intricate in the generation of the dipteran wing hinge, a complex structure that is derived from an apparently simple region of the wing imaginal disc. Using the characterisation of original outstretched alleles of the unpaired locus as a starting point, we demonstrate the role of JAK/STAT pathway signalling in the process of wing hinge development. We show that differences in JAK/STAT signalling within the proximal most of three lateral folds present in the wing imaginal disc is required for fold morphology and the subsequent differentiation of the first and second auxiliary sclerites as well as the posterior notal wing process. Changes in these domains are consistent with the established fate map of the wing disc. We show that outstretched wing posture phenotypes arise from the loss of a region of Unpaired expression in the proximal wing fold and demonstrate that this results in a decrease in JAK/STAT pathway activity. Finally we show that reduction of JAK/STAT pathway activity within the proximal wing fold is sufficient to phenocopy the outstretched phenotype. Taken together, we suggest that localised Unpaired expression and hence JAK/STAT pathway activity, is required for the morphogenesis of the adult wing hinge, providing new insights into the link between signal transduction pathways, patterning and development. PMID:23741461
Husak, J F; Ribak, G; Baker, R H; Rivera, G; Wilkinson, G S; Swallow, J G
2013-06-01
Exaggerated male ornaments are predicted to be costly to their bearers, but these negative effects may be offset by the correlated evolution of compensatory traits. However, when locomotor systems, such as wings in flying species, evolve to decrease such costs, it remains unclear whether functional changes across related species are achieved via the same morphological route or via alternate changes that have similar function. We conducted a comparative analysis of wing shape in relation to eye-stalk elongation across 24 species of stalk-eyed flies, using geometric morphometrics to determine how species with increased eye span, a sexually selected trait, have modified wing morphology as a compensatory mechanism. Using traditional and phylogenetically informed multivariate analyses of shape in combination with phenotypic trajectory analysis, we found a strong phylogenetic signal in wing shape. However, dimorphic species possessed shifted wing veins with the result of lengthening and narrowing wings compared to monomorphic species. Dimorphic species also had changes that seem unrelated to wing size, but instead may govern wing flexion. Nevertheless, the lack of a uniform, compensatory pattern suggests that stalk-eyed flies used alternative modifications in wing structure to increase wing area and aspect ratio, thus taking divergent morphological routes to compensate for exaggerated eye stalks. © 2013 The Authors. Journal of Evolutionary Biology © 2013 European Society For Evolutionary Biology.
Nishigori, Hideaki; Ito, Masaaki; Nishizawa, Yuji
2017-04-01
We introduce a novel transanal tube (TAT), named the "WING DRAIN", designed to prevent anastomotic leakage after rectal cancer surgery, and report the fundamental experiments that led to its development. We performed the basic experiments to evaluate the effect of TATs on intestinal decompression, the changes they make in patterns of watery fluid drainage, the changes in their decompression effect when the extension tube connecting the TAT to the collection bag fills with watery drainage fluid, and the variations in intestinal contact and crushing pressure made by some types of TAT. Any type of TAT contributed to decompression in the intestinal tract. Watery drainage commenced from when the water level first rose to the hole in the tip of drain. The intestinal pressure increased with the length of the vertical twist in an extension tube. The crushing pressures of most types of TAT were high enough to cause injury to the intestine. We resolved the problems using an existing TAT for the purpose of intestinal decompression and by creating the first specialized TAT designed to prevent anastomotic leakage after rectal cancer surgery in Japan.
Lima, C B S; Nunes, L A; Carvalho, C A L; Ribeiro, M F; Souza, B A; Silva, C S B
2016-01-01
A geometric morphometrics approach was applied to evaluate differences in forewing patterns of the Jandaira bee (Melipona subnitida Ducke). For this, we studied the presence of fluctuating asymmetry (FA) in forewing shape and size of colonies kept in either rational hive boxes or natural tree trunks. We detected significant FA for wing size as well as wing shape independent of the type of housing (rational box or tree trunks), indicating the overall presence of stress during the development of the studied specimens. FA was also significant (p < 0.01) between rational boxes, possibly related to the use of various models of rational boxes used for keeping stingless bees. In addition, a Principal Component Analysis indicated morphometric variation between bee colonies kept in either rational hive boxes or in tree trunks, that may be related to the different origins of the bees: tree trunk colonies were relocated natural colonies while rational box colonies originated from multiplying other colonies. We conclude that adequate measures should be taken to reduce the amount of stress during bee handling by using standard models of rational boxes that cause the least disruption.
NASA Astrophysics Data System (ADS)
Hafiz, Shopan; Andrade, Nicolas; Monavarian, Morteza; Izyumskaya, Natalia; Das, Saikat; Zhang, Fan; Avrutin, Vitaliy; Morkoç, Hadis; Özgür, Ümit
2016-02-01
Near-field scanning optical microscopy was applied to investigate the spatial variations of extended defects and their effects on the optical quality for semi-polar (1-101) and (11-22) InGaN light emitting diodes (LEDs). (1-101) and (11-22) oriented InGaN LEDs emitting at 450-470 nm were grown on patterned Si (001) 7° offcut substrates and m-sapphire substrates by means of nano-epitaxial lateral overgrowth (ELO), respectively. For (1-101) structures, the photoluminescence (PL) at 85 K from the near surface c+ wings was found to be relatively uniform and strong across the sample. However, emission from the c- wings was substantially weaker due to the presence of high density of threading dislocations (TDs) and basal plane stacking faults (BSFs) as revealed from the local PL spectra. In case of (11-22) LED structures, near-field PL intensity correlated with the surface features and the striations along the direction parallel to the c-axis projection exposed facets where the Indium content was higher as deduced from shift in the PL peak energy.
Do digestive contents confound body mass as a measure of relative condition in nestling songbirds?
Streby, Henry M.; Peterson, Sean M.; Lehman, Justin A.; Kramer, Gunnar R.; Vernasco, Ben J.; Andersen, David E.
2014-01-01
Relative nestling condition, typically measured as nestling mass or as an index including nestling mass, is commonly purported to correlate with fledgling songbird survival. However, most studies directly investigating fledgling survival have found no such relationship. We weighed feces and stomach contents of nestling golden-winged warblers (Vermivora chrysoptera) to investigate the potential contribution of variation in digestive contents to differences in nestling mass. We estimated that the mass of a seventh-day (near fledging) nestling golden-winged warbler varies by 0.65 g (approx. 9% of mean nestling mass) depending on the contents of the nestling's digestive system at the time of weighing, and that digestive contents are dissimilar among nestlings at any moment the brood is removed from the nest for weighing. Our conservative estimate of within-individual variation in digestive contents equals 72% and 24% of the mean within-brood and population-wide range in nestling mass, respectively. Based on our results, a substantive but typically unknown amount of the variation in body mass among nestlings is confounded by differences in digestive contents. We conclude that short-term variation in digestive contents likely precludes the use of body mass, and therefore any mass-dependent index, as a measure of relative nestling condition or as a predictor of survival in golden-winged warblers and likely in many other songbirds of similar size.
Hummingbird wing efficacy depends on aspect ratio and compares with helicopter rotors
Kruyt, Jan W.; Quicazán-Rubio, Elsa M.; van Heijst, GertJan F.; Altshuler, Douglas L.; Lentink, David
2014-01-01
Hummingbirds are the only birds that can sustain hovering. This unique flight behaviour comes, however, at high energetic cost. Based on helicopter and aeroplane design theory, we expect that hummingbird wing aspect ratio (AR), which ranges from about 3.0 to 4.5, determines aerodynamic efficacy. Previous quasi-steady experiments with a wing spinner set-up provide no support for this prediction. To test this more carefully, we compare the quasi-steady hover performance of 26 wings, from 12 hummingbird taxa. We spun the wings at angular velocities and angles of attack that are representative for every species and measured lift and torque more precisely. The power (aerodynamic torque × angular velocity) required to lift weight depends on aerodynamic efficacy, which is measured by the power factor. Our comparative analysis shows that AR has a modest influence on lift and drag forces, as reported earlier, but interspecific differences in power factor are large. During the downstroke, the power required to hover decreases for larger AR wings at the angles of attack at which hummingbirds flap their wings (p < 0.05). Quantitative flow visualization demonstrates that variation in hover power among hummingbird wings is driven by similar stable leading edge vortices that delay stall during the down- and upstroke. A side-by-side aerodynamic performance comparison of hummingbird wings and an advanced micro helicopter rotor shows that they are remarkably similar. PMID:25079868
Ren, Jing; Bai, Ming; Yang, Xing-Ke; Zhang, Run-Zhi; Ge, Si-Qin
2017-01-01
The success of beetles is mainly attributed to the possibility to hide the hindwings under the sclerotised elytra. The acquisition of the transverse folding function of the hind wing is an important event in the evolutionary history of beetles. In this study, the morphological and functional variances in the hind wings of 94 leaf beetle species (Coleoptera: Chrysomelinae) is explored using geometric morphometrics based on 36 landmarks. Principal component analysis and Canonical variate analysis indicate that changes of apical area, anal area, and middle area are three useful phylogenetic features at a subtribe level of leaf beetles. Variances of the apical area are the most obvious, which strongly influence the entire venation variance. Partial least squares analysis indicates that the proximal and distal parts of hind wings are weakly associated. Modularity tests confirm that the proximal and distal compartments of hind wings are separate modules. It is deduced that for leaf beetles, or even other beetles, the hind wing possibly exhibits significant functional divergences that occurred during the evolution of transverse folding that resulted in the proximal and distal compartments of hind wings evolving into separate functional modules.
Dai, Mengyao; Wang, Yao; Fang, Lu; Irwin, David M; Zhu, Tengteng; Zhang, Junpeng; Zhang, Shuyi; Wang, Zhe
2014-01-01
Bats are the only mammals capable of self-powered flight using wings. Differing from mouse or human limbs, four elongated digits within a broad wing membrane support the bat wing, and the foot of the bat has evolved a long calcar that spread the interfemoral membrane. Our recent mRNA sequencing (mRNA-Seq) study found unique expression patterns for genes at the 5' end of the Hoxd gene cluster and for Tbx3 that are associated with digit elongation and wing membrane growth in bats. In this study, we focused on two additional genes, Meis2 and Mab21l2, identified from the mRNA-Seq data. Using whole-mount in situ hybridization (WISH) we validated the mRNA-Seq results for differences in the expression patterns of Meis2 and Mab21l2 between bat and mouse limbs, and further characterize the timing and location of the expression of these two genes. These analyses suggest that Meis2 may function in wing membrane growth and Mab21l2 may have a role in AP and DV axial patterning. In addition, we found that Tbx3 is uniquely expressed in the unique calcar structure found in the bat hindlimb, suggesting a role for this gene in calcar growth and elongation. Moreover, analysis of the coding sequences for Meis2, Mab21l2 and Tbx3 showed that Meis2 and Mab21l2 have high sequence identity, consistent with the functions of genes being conserved, but that Tbx3 showed accelerated evolution in bats. However, evidence for positive selection in Tbx3 was not found, which would suggest that the function of this gene has not been changed. Together, our findings support the hypothesis that the modulation of the spatiotemporal expression patterns of multiple functional conserved genes control limb morphology and drive morphological change in the diversification of mammalian limbs.
Fang, Lu; Irwin, David M.; Zhu, Tengteng; Zhang, Junpeng; Zhang, Shuyi; Wang, Zhe
2014-01-01
Bats are the only mammals capable of self-powered flight using wings. Differing from mouse or human limbs, four elongated digits within a broad wing membrane support the bat wing, and the foot of the bat has evolved a long calcar that spread the interfemoral membrane. Our recent mRNA sequencing (mRNA-Seq) study found unique expression patterns for genes at the 5′ end of the Hoxd gene cluster and for Tbx3 that are associated with digit elongation and wing membrane growth in bats. In this study, we focused on two additional genes, Meis2 and Mab21l2, identified from the mRNA-Seq data. Using whole-mount in situ hybridization (WISH) we validated the mRNA-Seq results for differences in the expression patterns of Meis2 and Mab21l2 between bat and mouse limbs, and further characterize the timing and location of the expression of these two genes. These analyses suggest that Meis2 may function in wing membrane growth and Mab21l2 may have a role in AP and DV axial patterning. In addition, we found that Tbx3 is uniquely expressed in the unique calcar structure found in the bat hindlimb, suggesting a role for this gene in calcar growth and elongation. Moreover, analysis of the coding sequences for Meis2, Mab21l2 and Tbx3 showed that Meis2 and Mab21l2 have high sequence identity, consistent with the functions of genes being conserved, but that Tbx3 showed accelerated evolution in bats. However, evidence for positive selection in Tbx3 was not found, which would suggest that the function of this gene has not been changed. Together, our findings support the hypothesis that the modulation of the spatiotemporal expression patterns of multiple functional conserved genes control limb morphology and drive morphological change in the diversification of mammalian limbs. PMID:25166052
Lehmann, Fritz-Olaf; Pick, Simon
2007-04-01
Flying insects may enhance their flight force production by contralateral wing interaction during dorsal stroke reversal ('clap-and-fling'). In this study, we explored the forces and moments due to clap-and-fling at various wing tip trajectories, employing a dynamically scaled electromechanical flapping device. The 17 tested bio-inspired kinematic patterns were identical in stroke amplitude, stroke frequency and angle of attack with respect to the horizontal stroke plane but varied in heaving motion. Clap-and-fling induced vertical force augmentation significantly decreased with increasing vertical force production averaged over the entire stroke cycle, whereas total force augmentation was independent from changes in force produced by a single wing. Vertical force augmentation was also largely independent of forces produced due to wing rotation at the stroke reversals, the sum of rotational circulation and wake capture force. We obtained maximum (17.4%) and minimum (1.4%) vertical force augmentation in two types of figure-eight stroke kinematics whereby rate and direction of heaving motion during fling may explain 58% of the variance in vertical force augmentation. This finding suggests that vertical wing motion distinctly alters the flow regime at the beginning of the downstroke. Using an analytical model, we determined pitching moments acting on an imaginary body of the flapping device from the measured time course of forces, the changes in length of the force vector's moment arm, the position of the centre of mass and body angle. The data show that pitching moments are largely independent from mean vertical force; however, clap-and-fling reinforces mean pitching moments by approximately 21%, compared to the moments produced by a single flapping wing. Pitching moments due to clap-and-fling significantly increase with increasing vertical force augmentation and produce nose-down moments in most of the tested patterns. The analytical model, however, shows that algebraic sign and magnitude of these moments may vary distinctly depending on both body angle and the distance between the wing hinge and the animal's centre of mass. Altogether, the data suggest that the benefit of clap-and-fling wing beat for vertical force enhancement and pitch balance may change with changing heaving motion and thus wing tip trajectory during manoeuvring flight. We hypothesize that these dependencies may have shaped the evolution of wing kinematics in insects that are limited by aerodynamic lift rather than by mechanical power of their flight musculature.
Lokemoen, John T.; Johnson, Douglas H.; Sharp, David E.
1990-01-01
During 1976-81 we weighed several thousands of wild Mallard, Gadwall, and Blue-winged Teal in central North Dakota to examine duckling growth patterns, adult weights, and the factors influencing them. One-day-old Mallard and Gadwall averaged 32.4 and 30.4 g, respectively, a reduction of 34% and 29% from fresh egg weights. In all three species, the logistic growth curve provided a good fit for duckling growth patterns. Except for the asymptote, there was no difference in growth curves between males and females of a species. Mallard and Gadwall ducklings were heavier in years when wetland area was extensive or had increased from the previous year. Weights of after-second-year females were greater than yearlings for Mallard but not for Gadwall or Blue-winged Teal. Adult Mallard females lost weight continuously from late March to early July. Gadwall and Blue-winged Teal females, which nest later than Mallard, gained weight after spring arrival, lost weight from the onset of nesting until early July, and then regained some weight. Females of all species captured on nests were lighter than those captured off nests at the same time. Male Mallard weights decreased from spring arrival until late May. Male Gadwall and Blue-winged Teal weights increased after spring arrival, then declined until early June. Males of all three species then gained weight until the end of June. Among adults, female Gadwall and male Mallard and Blue-winged Teal were heavier in years when wetland area had increased from the previous year; female Blue-winged Teal were heavier in years with more wetland area.
Nattero, Julieta; Dujardin, Jean-Pierre; Del Pilar Fernández, María; Gürtler, Ricardo E
2015-12-01
Fluctuating asymmetry (FA), a slight and random departure from bilateral symmetry that is normally distributed around a 0 mean, has been widely used to infer developmental instability. We investigated whether habitats (ecotopes) and host-feeding sources influenced wing FA of the hematophagous bug Triatoma infestans. Because bug populations occupying distinct habitats differed substantially and consistently in various aspects such as feeding rates, engorgement status and the proportion of gravid females, we predicted that bugs from more open peridomestic habitats (i.e., goat corrals) were more likely to exhibit higher FA than bugs from domiciles. We examined patterns of asymmetry and the amount of wing size and shape FA in 196 adult T. infestans collected across a gradient of habitat suitability and stability that decreased from domiciles, storerooms, kitchens, chicken coops, pig corrals, to goat corrals in a well-defined area of Figueroa, northwestern Argentina. The bugs had unmixed blood meals on human, chicken, pig and goat depending on the bug collection ecotope. We documented the occurrence of FA in wing shape for bugs fed on all host-feeding sources and in all ecotopes except for females from domiciles or fed on humans. FA indices for wing shape differed significantly among host-feeding sources, ecotopes and sexes. The patterns of wing asymmetry in females from domiciles and from goat corrals were significantly different; differences in male FA were congruent with evidence showing that they had higher mobility than females across habitats. The host-feeding sources and habitats of T. infestans affected wing developmental stability depending on sex. Copyright © 2015 Elsevier B.V. All rights reserved.
NASA Technical Reports Server (NTRS)
Collins, W.; Chang, S. H.; Kuo, J. T.
1984-01-01
Data from field surveys and biogeochemical tests conducted in Maine, Montana, and Washington strongly correlate with results obtained using high resolution airborne spectroradiometer which detects an anomalous spectral waveform that appears definitely associated with sulfide mineralization. The spectral region most affected by mineral stress is between 550 nm and 750 nm. Spectral variations observed in the field occur on the wings of the red chlorophyll band centered at about 690 nm. The metal-stress-induced variations on the absorption band wing are most successfully resolved in the high spectral resolution field data using a waveform analysis technique. The development of chlorophyll pigments was retarded in greenhouse plants doped with copper and zinc in the laboratory. The lowered chlorophyll production resulted in changes on the wings of the chlorophyll bands of reflectance spectra of the plants. The airborne spectroradiometer system and waveform analysis remains the most sensitive technique for biogeochemical surveys.
Urbanization disrupts latitude-size rule in 17-year cicadas.
Beasley, DeAnna E; Penick, Clint A; Boateng, Nana S; Menninger, Holly L; Dunn, Robert R
2018-03-01
Many ectotherms show a decrease in body size with increasing latitude due to changes in climate, a pattern termed converse Bergmann's rule. Urban conditions-particularly warmer temperatures and fragmented landscapes-may impose stresses on development that could disrupt these body size patterns. To test the impact of urbanization on development and latitudinal trends in body size, we launched a citizen science project to collect periodical cicadas ( Magicicada septendecim ) from across their latitudinal range during the 2013 emergence of Brood II. Periodical cicadas are long-lived insects whose distribution spans a broad latitudinal range covering both urban and rural habitats. We used a geometric morphometric approach to assess body size and developmental stress based on fluctuating asymmetry in wing shape. Body size of rural cicadas followed converse Bergmann's rule, but this pattern was disrupted in urban habitats. In the north, urban cicadas were larger than their rural counterparts, while southern populations showed little variation in body size between habitats. We detected no evidence of differences in developmental stress due to urbanization. To our knowledge, this is the first evidence that urbanization disrupts biogeographical trends in body size, and this pattern highlights how the effects of urbanization may differ over a species' range.
Sound radiation and wing mechanics in stridulating field crickets (Orthoptera: Gryllidae).
Montealegre-Z, Fernando; Jonsson, Thorin; Robert, Daniel
2011-06-15
Male field crickets emit pure-tone mating calls by rubbing their wings together. Acoustic radiation is produced by rapid oscillations of the wings, as the right wing (RW), bearing a file, is swept across the plectrum borne on the left wing (LW). Earlier work found the natural resonant frequency (f(o)) of individual wings to be different, but there is no consensus on the origin of these differences. Previous studies suggested that the frequency along the song pulse is controlled independently by each wing. It has also been argued that the stridulatory file has a variable f(o) and that the frequency modulation observed in most species is associated with this variability. To test these two hypotheses, a method was developed for the non-contact measurement of wing vibrations during singing in actively stridulating Gryllus bimaculatus. Using focal microinjection of the neuroactivator eserine into the cricket's brain to elicit stridulation and micro-scanning laser Doppler vibrometry, we monitored wing vibration in actively singing insects. The results show significantly lower f(o) in LWs compared with RWs, with the LW f(o) being identical to the sound carrier frequency (N=44). But during stridulation, the two wings resonate at one identical frequency, the song carrier frequency, with the LW dominating in amplitude response. These measurements also demonstrate that the stridulatory file is a constant resonator, as no variation was observed in f(o) along the file during sound radiation. Our findings show that, as they engage in stridulation, cricket wings work as coupled oscillators that together control the mechanical oscillations generating the remarkably pure species-specific song.
Murillo-Maldonado, Juan M; Zeineddine, Fouad Bou; Stock, Rachel; Thackeray, Justin; Riesgo-Escovar, Juan R
2011-01-01
Coordination between growth and patterning/differentiation is critical if appropriate final organ structure and size is to be achieved. Understanding how these two processes are regulated is therefore a fundamental and as yet incompletely answered question. Here we show through genetic analysis that the phospholipase C-γ (PLC-γ) encoded by small wing (sl) acts as such a link between growth and patterning/differentiation by modulating some MAPK outputs once activated by the insulin pathway; particularly, sl promotes growth and suppresses ectopic differentiation in the developing eye and wing, allowing cells to attain a normal size and differentiate properly. sl mutants have previously been shown to have a combination of both growth and patterning/differentiation phenotypes: small wings, ectopic wing veins, and extra R7 photoreceptor cells. We show here that PLC-γ activated by the insulin pathway participates broadly and positively during cell growth modulating EGF pathway activity, whereas in cell differentiation PLC-γ activated by the insulin receptor negatively regulates the EGF pathway. These roles require different SH2 domains of PLC-γ, and act via classic PLC-γ signaling and EGF ligand processing. By means of PLC-γ, the insulin receptor therefore modulates differentiation as well as growth. Overall, our results provide evidence that PLC-γ acts during development at a time when growth ends and differentiation begins, and is important for proper coordination of these two processes.
Transonic Drag Prediction on a DLR-F6 Transport Configuration Using Unstructured Grid Solvers
NASA Technical Reports Server (NTRS)
Lee-Rausch, E. M.; Frink, N. T.; Mavriplis, D. J.; Rausch, R. D.; Milholen, W. E.
2004-01-01
A second international AIAA Drag Prediction Workshop (DPW-II) was organized and held in Orlando Florida on June 21-22, 2003. The primary purpose was to inves- tigate the code-to-code uncertainty. address the sensitivity of the drag prediction to grid size and quantify the uncertainty in predicting nacelle/pylon drag increments at a transonic cruise condition. This paper presents an in-depth analysis of the DPW-II computational results from three state-of-the-art unstructured grid Navier-Stokes flow solvers exercised on similar families of tetrahedral grids. The flow solvers are USM3D - a tetrahedral cell-centered upwind solver. FUN3D - a tetrahedral node-centered upwind solver, and NSU3D - a general element node-centered central-differenced solver. For the wingbody, the total drag predicted for a constant-lift transonic cruise condition showed a decrease in code-to-code variation with grid refinement as expected. For the same flight condition, the wing/body/nacelle/pylon total drag and the nacelle/pylon drag increment predicted showed an increase in code-to-code variation with grid refinement. Although the range in total drag for the wingbody fine grids was only 5 counts, a code-to-code comparison of surface pressures and surface restricted streamlines indicated that the three solvers were not all converging to the same flow solutions- different shock locations and separation patterns were evident. Similarly, the wing/body/nacelle/pylon solutions did not appear to be converging to the same flow solutions. Overall, grid refinement did not consistently improve the correlation with experimental data for either the wingbody or the wing/body/nacelle pylon configuration. Although the absolute values of total drag predicted by two of the solvers for the medium and fine grids did not compare well with the experiment, the incremental drag predictions were within plus or minus 3 counts of the experimental data. The correlation with experimental incremental drag was not significantly changed by specifying transition. Although the sources of code-to-code variation in force and moment predictions for the three unstructured grid codes have not yet been identified, the current study reinforces the necessity of applying multiple codes to the same application to assess uncertainty.
NASA Technical Reports Server (NTRS)
Lamar, J. E.
1971-01-01
The development of a nonplanar lifting surface method having a continuous distribution of singularities and satisfying the tangent flow boundary condition on the mean camber surface is given. The method predicts some incompressible longitudinal aerodynamic coefficients of rectangular wings which have circular-arc camber. The solution method is of the integral-equation type and the resulting surface integrals are evaluated by either using numerical or analytical techniques, as are appropriate. Applications are made and the results compared with those from an exact two-dimensional circular-arc camber solution, a three-dimensional flat-wing solution which represents the camber by a projected slope onto the flat surface, and a flat-wing experiment. From these comparisons, the present method is found to predict well the flat-wing experiment and limiting values, in addition to the center of pressure variation at an angle of attack of zero for any camber. For wings having camber ratios larger than about 1.25% and moderate to high aspect ratios, the results deterioriate due to the inadequacy of lifting pressure modes employed.
Supersonic flow visualization of a nacelle in close proximity to a simulated wing
NASA Technical Reports Server (NTRS)
Biber, Kasim; Ellis, David R.
1993-01-01
A flow visualization study was made in the 9 x 9 inch supersonic wind tunnel at Wichita State University to examine shock and boundary layer flow interaction for a nacelle in close proximity to the lower surface of a simulated wing. The test matrix included variations of angle of attack from -2 degrees to +4 degrees, nacelle-wing gap from 0.5 to 3-nacelle inlet diameter (0.12 inch), and Reynolds number based on nacelle length (1.164 inch) from 1.16 x 10(exp 6) to 1.45 x 10(exp 6) at a nominal Mach number of 2. Schlieren pictures of wing and nacelle flowfield were recorded by a video camera during each tunnel run. Results show that the nacelle inlet shock wave remains attached to the inlet lip and its impingement does not significantly affect the wing boundary layer. At the nacelle trailing edge location, the wing boundary layer thickness is approximately one nacelle inlet diameter at alpha = 0 degrees and it decreases with increase of angle of attack.
Novel method for measuring a dense 3D strain map of robotic flapping wings
NASA Astrophysics Data System (ADS)
Li, Beiwen; Zhang, Song
2018-04-01
Measuring dense 3D strain maps of the inextensible membranous flapping wings of robots is of vital importance to the field of bio-inspired engineering. Conventional high-speed 3D videography methods typically reconstruct the wing geometries through measuring sparse points with fiducial markers, and thus cannot obtain the full-field mechanics of the wings in detail. In this research, we propose a novel system to measure a dense strain map of inextensible membranous flapping wings by developing a superfast 3D imaging system and a computational framework for strain analysis. Specifically, first we developed a 5000 Hz 3D imaging system based on the digital fringe projection technique using the defocused binary patterns to precisely measure the dynamic 3D geometries of rapidly flapping wings. Then, we developed a geometry-based algorithm to perform point tracking on the precisely measured 3D surface data. Finally, we developed a dense strain computational method using the Kirchhoff-Love shell theory. Experiments demonstrate that our method can effectively perform point tracking and measure a highly dense strain map of the wings without many fiducial markers.
Smith, N. Adam; Clarke, Julia A.
2012-01-01
Just as skeletal characteristics provide clues regarding behavior of extinct vertebrates, phylogenetically-informed evaluation of endocranial morphology facilitates comparisons among extinct taxa and extant taxa with known behavioral characteristics. Previous research has established that endocranial morphology varies across Aves; however, variation of those systems among closely related species remains largely unexplored. The Charadriiformes (shorebirds and allies) are an ecologically diverse clade with a comparatively rich fossil record, and therefore, are well suited for investigating interspecies variation, and potential links between endocranial morphology, phylogeny, ecology and other life history attributes. Endocranial endocasts were rendered from high resolution X-ray computed tomography data for 17 charadriiforms (15 extant and two flightless extinct species). Evaluation of endocranial character state changes on a phylogeny for Charadriiformes resulted in identification of characters that vary in taxa with distinct feeding and locomotor ecologies. In comparison with all other charadriiforms, stem and crown clade wing-propelled diving Pan-Alcidae displayed compressed semicircular canals, and indistinct occipital sinuses and cerebellar fissures. Flightless wing-propelled divers have relatively smaller brains for their body mass and smaller optic lobes than volant pan-alcids. Observed differences between volant and flightless wing-propelled sister taxa are striking given that flightless pan-alcids continue to rely on the flight stroke for underwater propulsion. Additionally, the brain of the Black Skimmer Rynchops niger, a taxon with a unique feeding ecology that involves continuous forward aerial motion and touch-based prey detection used both at day and night, is discovered to be unlike that of any other sampled charadriiform in having an extremely large wulst as well as a small optic lobe and distinct occipital sinus. Notably, the differences between the Black Skimmer and other charadriiforms are more pronounced than between wing-propelled divers and other charadriiforms. Finally, aspects of endosseous labyrinth morphology are remarkably similar between divers and non-divers, and may deserve further evaluation. PMID:23209585
Microscopic modulation of mechanical properties in transparent insect wings
DOE Office of Scientific and Technical Information (OSTI.GOV)
Arora, Ashima; Kumar, Pramod; Bhagavathi, Jithin
We report on the measurement of local friction and adhesion of transparent insect wings using an atomic force microscope cantilever down to nanometre length scales. We observe that the wing-surface is decorated with 10 μm long and 2 μm wide islands that have higher topographic height. The friction on the islands is two orders of magnitude higher than the back-ground while the adhesion on the islands is smaller. Furthermore, the high islands are decorated with ordered nano-wire-like structures while the background is full of randomly distributed granular nano-particles. Coherent optical diffraction through the wings produce a stable diffraction pattern revealing a quasi-periodicmore » organization of the high islands over the entire wing. This suggests a long-range order in the modulation of friction and adhesion which is directly correlated with the topography. The measurements unravel novel functional design of complex wing surface and could find application in miniature biomimetic devices.« less
Replication of cicada wing's nano-patterns by hot embossing and UV nanoimprinting.
Hong, Sung-Hoon; Hwang, Jaeyeon; Lee, Heon
2009-09-23
The hydrophobicity of the cicada wing originates from its naturally occurring, surface nano-structure. The nano-structure of the cicada wing consists of an array of nano-sized pillars, 100 nm in diameter and 300 nm in height. In this study, the nano-structure of the cicada wing was successfully duplicated by using hot embossing lithography and UV nanoimprint lithography (NIL). The diameter and pitch of replication were the same as those of the original cicada wing and the height was a little smaller than that of the original master. The transmittance of the hot embossed PVC film was increased by 2-6% compared with that of the bare PVC film. The hydrophobicity was measured by water contact angle measurements. The water contact angle of the replica, made of UV cured polymer, was 132 degrees +/- 2 degrees , which was slightly lower than that of the original cicada wing (138 degrees +/- 2 degrees ), but much higher than that of the UV cured polymer surface without any nano-sized pillars (86 degrees ).
NASA Technical Reports Server (NTRS)
Lomax, Harvard
1957-01-01
Several variational problems involving optimum wing and body combinations having minimum wave drag for different kinds of geometrical restraints are analyzed. Particular attention is paid to the effect on the wave drag of shortening the fuselage and, for slender axially symmetric bodies, the effect of fixing the fuselage diameter at several points or even of fixing whole portions of its shape.
Host plant affects morphometric variation of Diaphorina citri (Hemiptera: Liviidae).
Paris, Thomson M; Allan, Sandra A; Hall, David G; Hentz, Matthew G; Hetesy, Gabriella; Stansly, Philip A
2016-01-01
The Asian citrus psyllid (ACP), Diaphorina citri Kuwayama, is one of the most serious citrus pests worldwide due to its role as vector of huanglongbing or citrus greening disease. While some optimal plant species for ACP oviposition and development have been identified, little is known of the influence of host plants on ACP size and shape. Our goal was to determine how size and shape of ACP wing and body size varies when development occurs on different host plants in a controlled rearing environment. ACP were reared on six different rutaceous species; Bergera koenigii , Citrus aurantifolia , Citrus macrophylla , Citrus maxima , Citrus taiwanica and Murraya paniculata . Adults were examined for morphometric variation using traditional and geometric analysis based on 12 traits or landmarks. ACP reared on C. taiwanica were consistently smaller than those reared on the other plant species. Wing aspect ratio also differed between C. maxima and C. taiwanica . Significant differences in shape were detected with those reared on M. paniculata having narrower wings than those reared on C. macrophylla . This study provides evidence of wing size and shape differences of ACP based on host plant species which potentially may impact dispersal. Further study is needed to determine if behavioral and physiological differences are associated with the observed phenotypic differences.
Host plant affects morphometric variation of Diaphorina citri (Hemiptera: Liviidae)
Paris, Thomson M.; Hall, David G.; Hentz, Matthew G.; Hetesy, Gabriella; Stansly, Philip A.
2016-01-01
The Asian citrus psyllid (ACP), Diaphorina citri Kuwayama, is one of the most serious citrus pests worldwide due to its role as vector of huanglongbing or citrus greening disease. While some optimal plant species for ACP oviposition and development have been identified, little is known of the influence of host plants on ACP size and shape. Our goal was to determine how size and shape of ACP wing and body size varies when development occurs on different host plants in a controlled rearing environment. ACP were reared on six different rutaceous species; Bergera koenigii, Citrus aurantifolia, Citrus macrophylla, Citrus maxima, Citrus taiwanica and Murraya paniculata. Adults were examined for morphometric variation using traditional and geometric analysis based on 12 traits or landmarks. ACP reared on C. taiwanica were consistently smaller than those reared on the other plant species. Wing aspect ratio also differed between C. maxima and C. taiwanica. Significant differences in shape were detected with those reared on M. paniculata having narrower wings than those reared on C. macrophylla. This study provides evidence of wing size and shape differences of ACP based on host plant species which potentially may impact dispersal. Further study is needed to determine if behavioral and physiological differences are associated with the observed phenotypic differences. PMID:27833820
NASA Technical Reports Server (NTRS)
Abel, I.; Perry, B., III; Newsom, J. R.
1982-01-01
Two flutter suppression control laws wre designed and tested on a low speed aeroelastic model of a DC-10 derivative wing. Both control laws demontrated increases in flutter speed in excess of 25 percent above the passive wing flutter speed. In addition, one of the control laws was effective in reducing loads due to turbulence generated in the wind tunnel. The effect of variations in gain and phase on the closed-loop performance was measured and is compared with predictions. In general, both flutter and gust response predictions agree reasonably well with experimental data.
An experimental study of mushroom shaped stall cells. [on finite wings with separated flow
NASA Technical Reports Server (NTRS)
Winkelmann, A. E.
1982-01-01
Surface patterns characterized by a pair of counter-rotating swirls have been observed in connection with the conduction of surface flow visualization experiments involving test geometries with separated flows. An example of this phenomenon occurring on a finite wing with trailing edge stall has been referred to by Winkelmann and Barlow (1980) as 'mushroom shaped'. A description is presented of a collection of experimental results which show or suggest the occurrence of mushroom shaped stall cells on a variety of test geometries. Investigations conducted with finite wings, airfoil models, and flat plates are considered, and attention is given to studies involving the use of bluff models, investigations of shock induced boundary layer separation, and mushroom shaped patterns observed in a number of miscellaneous cases. It is concluded that the mushroom shaped stall cell appears commonly in separated flow regions.
Wingless is a positive regulator of eyespot color patterns in Bicyclus anynana butterflies.
Özsu, Nesibe; Chan, Qian Yi; Chen, Bin; Gupta, Mainak Das; Monteiro, Antónia
2017-09-01
Eyespot patterns of nymphalid butterflies are an example of a novel trait yet, the developmental origin of eyespots is still not well understood. Several genes have been associated with eyespot development but few have been tested for function. One of these genes is the signaling ligand, wingless, which is expressed in the eyespot centers during early pupation and may function in eyespot signaling and color ring differentiation. Here we tested the function of wingless in wing and eyespot development by down-regulating it in transgenic Bicyclus anynana butterflies via RNAi driven by an inducible heat-shock promoter. Heat-shocks applied during larval and early pupal development led to significant decreases in wingless mRNA levels and to decreases in eyespot size and wing size in adult butterflies. We conclude that wingless is a positive regulator of eyespot and wing development in B. anynana butterflies. Copyright © 2017 Elsevier Inc. All rights reserved.
NASA Technical Reports Server (NTRS)
Erickson, Gary E.
1991-01-01
The vortex dominated aerodynamic characteristics of a generic 65 degree cropped delta wing model were studied in a wind tunnel at subsonic through supersonic speeds. The lee-side flow fields over the wing-alone configuration and the wing with leading edge extension (LEX) added were observed at M (infinity) equals 0.40 to 1.60 using a laser vapor screen technique. These results were correlated with surface streamline patterns, upper surface static pressure distributions, and six-component forces and moments. The wing-alone exhibited vortex breakdown and asymmetry of the breakdown location at the subsonic and transonic speeds. An earlier onset of vortex breakdown over the wing occurred at transonic speeds due to the interaction of the leading edge vortex with the normal shock wave. The development of a shock wave between the vortex and wing surface caused an early separation of the secondary boundary layer. With the LEX installed, wing vortex breakdown asymmetry did not occur up to the maximum angle of attack in the present test of 24 degrees. The favorable interaction of the LEX vortex with the wing flow field reduced the effects of shock waves on the wing primary and secondary vortical flows. The direct interaction of the wing and LEX vortex cores diminished with increasing Mach number. The maximum attainable vortex-induced pressure signatures were constrained by the vacuum pressure limit at the transonic and supersonic speeds.
NASA Technical Reports Server (NTRS)
Benepe, D. B.; Cunningham, A. M., Jr.; Traylor, S., Jr.; Dunmyer, W. D.
1978-01-01
A detailed investigation of the flight buffeting response of the F-111A was performed in two phases. In Phase 1 stochastic analysis techniques were applied to wing and fuselage responses for maneuvers flown at subsonic speeds and wing leading edge sweep of 26 degrees. Power spectra and rms values were obtained. This report gives results of Phase 2 where the analyses were extended to include maneuvers flown at wing leading edge sweep values of 50 and 75.5 degrees at subsonic and supersonic speeds and the responses examined were expanded to include vertical shear, bending moment, and hingeline torque of the left and right horizontal tails. Power spectra, response time histories, variations of rms response with angle of attack and effects of wing sweep and Mach number are presented and discussed. Some Phase 1 results are given for comparison purposes.
NASA Astrophysics Data System (ADS)
Vigliotti, Andrea; Pasini, Damiano
2015-03-01
In the past few decades, several concepts for morphing wings have been proposed with the aim of improving the structural and aerodynamic performance of conventional aircraft wings. One of the most interesting challenges in the design of a morphing wing is represented by the skin, which needs to meet specific deformation requirements. In particular when morphing involves changes of cord or curvature, the skin is required to undergo large recoverable deformation in the actuation direction, while maintaining the desired shape and strength in the others. One promising material concept that can meet these specifications is represented by lattice materials. This paper examines the use of alternative planar lattices in the embodiment of a skin panel for cord and camber morphing of an aircraft wing. We use a structural homogenization scheme capable of capturing large geometric nonlinearity, to examine the structural performance of lattice skin concepts, as well as to tune their mechanical properties in desired directions.
An aerodynamic assessment of various supersonic fighter airplanes based on Soviet design concepts
NASA Technical Reports Server (NTRS)
Spearman, M. L.
1983-01-01
The aerodynamic, stability, and control characteristics of several supersonic fighter airplane concepts were assessed. The configurations include fixed-wing airplanes having delta wings, swept wings, and trapezoidal wings, and variable wing-sweep airplanes. Each concept employs aft tail controls. The concepts vary from lightweight, single engine, air superiority, point interceptor, or ground attack types to larger twin-engine interceptor and reconnaissance designs. Results indicate that careful application of the transonic or supersonic area rule can provide nearly optimum shaping for minimum drag for a specified Mach number requirement. Through the proper location of components and the exploitation of interference flow fields, the concepts provide linear pitching moment characteristics, high control effectiveness, and reasonably small variations in aerodynamic center location with a resulting high potential for maneuvering capability. By careful attention to component shaping and location and through the exploitation of local flow fields, favorable roll-to-yaw ratios may result and a high degree of directional stability can be achieved.
Effects of Inertial Power and Inertial Force on Bat Wings.
Yin, Dongfu; Zhang, Zhisheng; Dai, Min
2016-06-01
The inertial power and inertial force of wings are important factors in evaluating the flight performance of native bats. Based on measurement data of wing size and motions of Eptesicus fuscus, we present a new computational bat wing model with divided fragments of skeletons and membrane. The motions of the model were verified by comparing the joint and tip trajectories with native bats. The influences of flap, sweep, elbow, wrist and digits motions, the effects of different bones and membrane of bat wing, the components on vertical, spanwise and fore-aft directions of the inertial power and force were analyzed. Our results indicate that the flap, sweep, and elbow motions contribute the main inertial power and force; the membrane occupies an important proportion of the inertial power and force; inertial power on flap direction was larger, while variations of inertial forces on different directions were not evident. These methods and results offer insights into flight dynamics in other flying animals and may contribute to the design of future robotic bats.
In situ protocol for butterfly pupal wings using riboprobes.
Ramos, Diane; Monteiro, Antonia
2007-01-01
Here we present, in video format, a protocol for in situ hybridizations in pupal wings of the butterfly Bicyclus anynana using riboprobes. In situ hybridizations, a mainstay of developmental biology, are useful to study the spatial and temporal patterns of gene expression in developing tissues at the level of transcription. If antibodies that target the protein products of gene transcription have not yet been developed, and/or there are multiple gene copies of a particular protein in the genome that cannot be differentiated using available antibodies, in situs can be used instead. While an in situ technique for larval wing discs has been available to the butterfly community for several years, the current protocol has been optimized for the larger and more fragile pupal wings.
Boundary Dpp promotes growth of medial and lateral regions of the Drosophila wing.
Barrio, Lara; Milán, Marco
2017-07-04
The gradient of Decapentaplegic (Dpp) in the Drosophila wing has served as a paradigm to characterize the role of morphogens in regulating patterning. However, the role of this gradient in regulating tissue size is a topic of intense debate as proliferative growth is homogenous. Here, we combined the Gal4/UAS system and a temperature-sensitive Gal80 molecule to induce RNAi-mediated depletion of dpp and characterise the spatial and temporal requirement of Dpp in promoting growth. We show that Dpp emanating from the AP compartment boundary is required throughout development to promote growth by regulating cell proliferation and tissue size. Dpp regulates growth and proliferation rates equally in central and lateral regions of the developing wing appendage and reduced levels of Dpp affects similarly the width and length of the resulting wing. We also present evidence supporting the proposal that graded activity of Dpp is not an absolute requirement for wing growth.
An exploratory study of apex fence flaps on a 74 deg delta wing
NASA Technical Reports Server (NTRS)
Wahls, R. A.; Vess, R. J.
1985-01-01
An exploratory wind tunnel investigation was performed to observe the flow field effects produced by vertically deployed apex fences on a planar 74 degree delta wing. The delta shaped fences, each comprising approximately 3.375 percent of the wing area, were affixed along the first 25 percent of the wing leading edge in symmetric as well as asymmetric (i.e., fence on one side only) arrangements. The vortex flow field was visualized at angles of attack from 0 to 20 degrees using helium bubble and oil flow techniques; upper surface pressures were also measured along spanwise rows. The results were used to construct a preliminary description of the vortex patterns and induced pressures associated with vertical apex fence deployment. The objective was to obtain an initial evaluation of the potential of apex fences as vortex devices for subsonic lift modulation as well as lateral directional control of delta wing aircraft.
NASA Technical Reports Server (NTRS)
Rossow, V. J.; Corsiglia, V. R.; Schwind, R. G.; Frick, J. K. D.; Lemmer, O. J.
1975-01-01
Measurements were made in the wake of a swept wing model to study the structure of lift generated vortex wakes shed by conventional span loadings and by several span loadings designed to reduce wake velocities. Variations in the span loading on the swept wing generator were obtained by deflecting seven flap segments on each side by amounts determined by vortex lattice theory to approximate the desired span loadings. The resulting wakes were probed with a three component, hot wire probe to measure velocity, and with a wing to measure the rolling moment that would be induced on a following aircraft. The experimental techniques are described herein, and the measured velocity and rolling moments are presented, along with some comparisons with the applicable theories.
NASA Technical Reports Server (NTRS)
Kaufman, L. G., II; Johnson, C. B.
1977-01-01
Surface pressure distributions are presented for regions where three-dimensional separated flow effects are prominent on swept-wing-elevon-end-plate models of 0 degree, 50 degree, and 70 degree sweepback, and with 0 degree, 10 degree, 20 degree, and 30 degree elevon deflections. Surface-oil-flow photographs and pressure distributions on the flat-plate wing, elevon, and end-plate surfaces are presented for numerous geometric variations, including various spacings between the elevon and the end plate, with and without a tip fin. The data, for a free-stream Mach number of 6 and a wing-root-chord Reynolds number of 20 x 10 to the sixth power, reveal considerably larger regions of elevon induced loads on the adjacent end-plate surface than would be anticipated by using inviscid flow analyses.
Inviscid Analysis of Extended Formation Flight
NASA Technical Reports Server (NTRS)
Kless, James; Aftosmis, Michael J.; Ning, Simeon Andrew; Nemec, Marian
2012-01-01
Flying airplanes in extended formations, with separation distances of tens of wingspans, significantly improves safety while maintaining most of the fuel savings achieved in close formations. The present study investigates the impact of roll trim and compressibility at fixed lift coefficient on the benefits of extended formation flight. An Euler solver with adjoint-based mesh refinement combined with a wake propagation model is used to analyze a two-body echelon formation at a separation distance of 30 spans. Two geometries are examined: a simple wing and a wing-body geometry. Energy savings, quantified by both formation drag fraction and span efficiency factor, are investigated at subsonic and transonic speeds for a matrix of vortex locations. The results show that at fixed lift and trimmed for roll, the optimal location of vortex impingement is about 10% inboard of the trailing airplane s wing-tip. Interestingly, early results show the variation in drag fraction reduction is small in the neighborhood of the optimal position. Over 90% of energy benefits can be obtained with a 5% variation in transverse and 10% variation in crossflow directions. Early results suggest control surface deflections required to achieve trim reduce the benefits of formation flight by 3-5% at subsonic speeds. The final paper will include transonic effects and trim on extended formation flight drag benefits.
Neural basis of singing in crickets: central pattern generation in abdominal ganglia
NASA Astrophysics Data System (ADS)
Schöneich, Stefan; Hedwig, Berthold
2011-12-01
The neural mechanisms underlying cricket singing behavior have been the focus of several studies, but the central pattern generator (CPG) for singing has not been localized conclusively. To test if the abdominal ganglia contribute to the singing motor pattern and to analyze if parts of the singing CPG are located in these ganglia, we systematically truncated the abdominal nerve cord of fictively singing crickets while recording the singing motor pattern from a front-wing nerve. Severing the connectives anywhere between terminal ganglion and abdominal ganglion A3 did not preclude singing, although the motor pattern became more variable and failure-prone as more ganglia were disconnected. Singing terminated immediately and permanently after transecting the connectives between the metathoracic ganglion complex and the first unfused abdominal ganglion A3. The contribution of abdominal ganglia for singing pattern generation was confirmed by intracellular interneuron recordings and current injections. During fictive singing, an ascending interneuron with its soma and dendrite in A3 depolarized rhythmically. It spiked 10 ms before the wing-opener activity and hyperpolarized in phase with the wing-closer activity. Depolarizing current injection elicited rhythmic membrane potential oscillations and spike bursts that elicited additional syllables and reliably reset the ongoing chirp rhythm. Our results disclose that the abdominal ganglion A3 is directly involved in generating the singing motor pattern, whereas the more posterior ganglia seem to provide only stabilizing feedback to the CPG circuit. Localizing the singing CPG in the anterior abdominal neuromeres now allows analyzing its circuitry at the level of identified interneurons in subsequent studies.
On the aerodynamic forces of flapping finite-wings in forward flight: a numerical study
NASA Astrophysics Data System (ADS)
Gonzalo, Alejandro; Uhlmann, Markus; Garcia-Villalba, Manuel; Flores, Oscar
2017-11-01
We study the flow around two flapping wings in forward flight at a low Reynolds number, Re = 500 , with 3D direct numerical simulations. The flow solver used is TUCAN, an in-house code which solves the Navier-Stokes equations for incompressible flow using an immersed boundary method to model the presence of the wings. The wings are rectangular with a NACA0012 airfoil of chord c as a cross-section. They are located side by side at a distance 0.5 c between their inboard tips. The wings flap with respect to an axis parallel to the streamwise velocity, without pitching. The angle of rotation is defined using a sinusoidal function with a reduced frequency k = 1 and an amplitude such that the maximum height of the outboard tips is c in all cases. We perform several simulations varying the aspect ratio of the wings (AR = 2 and 4) and the distance between the inboard tip of the wings and the axis of rotation (R = 0 , 2 and ∞), the latter case corresponding to wings in heaving motion. In this way we can study the variation of the fictitious forces on the wings and the induced spanwise flows, and their relation to the vortical structures on the wing (i.e. leading edge vortex, trailing edge votex, tip vortices) and the resulting aerodynamic forces. This work was funded by project TRA2013-41103-P (Mineco/Feder UE). The simulations were partially performed at the Steinbuch Centre for Computing, Karlsruhe, whose support is thankfully acknowledged.
NASA Technical Reports Server (NTRS)
Toll, T. A.
1980-01-01
A parametric analysis was made to investigate the relationship between current cargo airplanes and possible future designs that may differ greatly in both size and configuration. The method makes use of empirical scaling laws developed from statistical studies of data from current and advanced airplanes and, in addition, accounts for payload density, effects of span distributed load, and variations in tail area ratio. The method is believed to be particularly useful for exploratory studies of design and technology options for large airplanes. The analysis predicts somewhat more favorable variations of the ratios of payload to gross weight and block fuel to payload as the airplane size is increased than has been generally understood from interpretations of the cube-square law. In terms of these same ratios, large all wing (spanloader) designs show an advantage over wing-fuselage designs.
The sail wing windmill and its adaptation for use in rural India
NASA Technical Reports Server (NTRS)
Sherman, M. M.
1973-01-01
An 8 meter-diameter prototype sail wing windmill is reported that uses a one meter-diameter bullock cartwheel to which three bamboo poles are latched in a triangular pattern with overlapping ends, to form the airframe for cloth sails. This device lifts 300 pounds to a height of 20 feet in one minute in a 10 mph wind.
Waliwitiya, Ranil; Belton, Peter; Nicholson, Russell A; Lowenberger, Carl A
2010-03-01
The effects were evaluated of the plant terpenoid thymol and eight other neuroactive compounds on flight muscle impulses (FMIs) and wing beat frequency (WBF) of tethered blowflies (Phaenicia sericata Meig.). The electrical activity of the dorsolongitudinal flight muscles was closely linked to the WBF of control insects. Topically applied thymol inhibited WBF within 15-30 min and reduced FMI frequency. Octopamine and chlordimeform caused a similar, early-onset bursting pattern that decreased in amplitude with time. Desmethylchlordimeform blocked wing beating within 60 min and generated a profile of continuous but lower-frequency FMIs. Fipronil suppressed wing beating and induced a pattern of continuous, variable-frequency spiking that diminished gradually over 6 h. Cypermethrin- and rotenone-treated flies had initial strong FMIs that declined with time. In flies injected with GABA, the FMIs were generally unidirectional and frequency was reduced, as was seen with thymol. Thymol readily penetrates the cuticle and interferes with flight muscle and central nervous function in the blowfly. The similarity of the action of thymol and GABA suggests that this terpenoid acts centrally in blowflies by mimicking or facilitating GABA action.
NASA Astrophysics Data System (ADS)
Zhao, Guang-yin; Li, Ying-hong; Liang, Hua; Han, Meng-hu; Hua, Wei-zhuo
2015-01-01
Wind tunnel experiments are conducted for improving the aerodynamic performance of delta wing using a leading-edge pulsed nanosecond dielectric barrier discharge (NS-DBD). The whole effects of pulsed NS-DBD on the aerodynamic performance of the delta wing are studied by balanced force measurements. Pressure measurements and particle image velocimetry (PIV) measurements are conducted to investigate the formation of leading-edge vortices affected by the pulsed NS-DBD, compared to completely stalled flow without actuation. Various pulsed actuation frequencies of the plasma actuator are examined with the freestream velocity up to 50 m/s. Stall has been delayed substantially and significant shifts in the aerodynamic forces can be achieved at the post-stall regions when the actuator works at the optimum reduced frequency of F + = 2. The upper surface pressure measurements show that the largest change of static pressure occurs at the forward part of the wing at the stall region. The time-averaged flow pattern obtained from the PIV measurement shows that flow reattachment is promoted with excitation, and a vortex flow pattern develops. The time-averaged locations of the secondary separation line and the center of the vortical region both move outboard with excitation.
Sawada, H; Nakagoshi, M; Reinhardt, R K; Ziegler, I; Koch, P B
2002-06-01
Color patterns of butterfly wings are composed of single color points represented by each scale. In the case of Precis coenia, at the end of pupal development, different types of pigments are synthesized sequentially in the differently colored scales beginning with white (pterins) followed by red (ommatins) and then black (melanin). In order to explain how formation of these different colors is regulated, we examined the expression of an mRNA-encoding guanosine triphosphate-cyclohydrolase I (GTP-CH I; EC 3.5.4.16), the first key enzyme in the biosynthesis of pteridines, during pigment formation in the wings of P. coenia. The strongest positive signal was recognized around pigment formation one day before butterfly emergence. This GTP-CH I gene expression is paralleled by GTP-CH I enzyme activity measured in wing extracts. We also investigated the effect of 20-hydroxyecdysone on the expression of GTP-CH I mRNA and the enzyme activity during color formation. The results strongly suggest that the onset and duration of the expression of a GTP-CH I mRNA is triggered by a declining ecdysteroid hormone titer during late pupal development.
NASA Technical Reports Server (NTRS)
Helms, V. T., III; Bradley, P. F.
1984-01-01
Results are presented for oil flow and phase change paint heat transfer tests conducted on a 0.006 scale model of a proposed single stage to orbit control configured vehicle. The data were taken at angles of attack up to 40 deg at a free stream Mach number of 10 for Reynolds numbers based on model length of 0.5 x 10 to the 6th power, 1.0 x 10 to the 6th power and 2.0 x 10 to the 6th power. The magnitude and distribution of heating are characterized in terms of angle of attack and Reynolds number aided by an analysis of the flow data which are used to suggest the presence of various three dimensional flow structures that produce the observed heating patterns. Of particular interest are streak heating patterns that result in high localized heat transfer rates on the wing windward surface at low to moderate angles of attack. These streaks are caused by the bow-shock/wing-shock interaction and formation of the wing-shock. Embedded vorticity was found to be associated with these interactions.
Estimation of wing nonlinear aerodynamic characteristics at supersonic speeds
NASA Technical Reports Server (NTRS)
Carlson, H. W.; Mack, R. J.
1980-01-01
A computational system for estimation of nonlinear aerodynamic characteristics of wings at supersonic speeds was developed and was incorporated in a computer program. This corrected linearized theory method accounts for nonlinearities in the variation of basic pressure loadings with local surface slopes, predicts the degree of attainment of theoretical leading edge thrust, and provides an estimate of detached leading edge vortex loadings that result when the theoretical thrust forces are not fully realized.
NASA Technical Reports Server (NTRS)
Mugler, John P., Jr.
1958-01-01
Pressure distributions are presented for a thin highly tapered untwisted 45 deg sweptback wing in combination with a body. These tests were made in the Langley 8-foot transonic pressure tunnel at both 1.0 and 0.5 atmosphere stagnation pressures at Mach numbers from 0.800 to 1.200 through an angle-of-attack range of -4 deg to 12 deg.
Effects of spoiler surfaces on the aeroelastic behavior of a low-aspect-ratio rectangular wing
NASA Technical Reports Server (NTRS)
Cole, Stanley R.
1990-01-01
An experimental research study to determine the effectiveness of spoiler surfaces in suppressing flutter onset for a low-aspect-ratio, rectangular wing was conducted in the Langley Transonic Dynamics Tunnel (TDT). The wing model used in this flutter test consisted of a rigid wing mounted to the wind-tunnel wall by a flexible, rectangular beam. The flexible beam was connected to the wing root and cantilever mounted to the wind-tunnel wall. The wing had a 1.5 aspect ratio based on wing semispan and a NACA 64A010 airfoil shape. The spoiler surfaces consisted of thin, rectangular aluminum plates that were vertically mounted to the wing surface. The spoiler surface geometry and location on the wing surface were varied to determine the effects of these parameters on the classical flutter of the wing model. Subsonically, the experiment showed that spoiler surfaces increased the flutter dynamic pressure with each successive increase in spoiler height or width. This subsonic increase in flutter dynamic pressure was approximately 15 percent for the maximum height spoiler configuration and for the maximum width spoiler configuration. At transonic Mach numbers, the flutter dynamic pressure conditions were increased even more substantially than at subsonic Mach numbers for some of the smaller spoiler surfaces. But greater than a certain spoiler size (in terms of either height or width) the spoilers forced a torsional instability in the transonic regime that was highly Mach number dependent. This detrimental torsional instability was found at dynamic pressures well below the expected flutter conditions. Variations in the spanwise location of the spoiler surfaces on the wing showed little effect on flutter. Flutter analysis was conducted for the basic configuration (clean wing with all spoiler surface mass properties included). The analysis correlated well with the clean wing experimental flutter results.
Open Rotor Noise Shielding by Blended-Wing-Body Aircraft
NASA Technical Reports Server (NTRS)
Guo, Yueping; Czech, Michael J.; Thomas, Russell H.
2015-01-01
This paper presents an analysis of open rotor noise shielding by Blended Wing Body (BWB) aircraft by using model scale test data acquired in the Boeing Low Speed Aeroacoustic Facility (LSAF) with a legacy F7/A7 rotor model and a simplified BWB platform. The objective of the analysis is the understanding of the shielding features of the BWB and the method of application of the shielding data for noise studies of BWB aircraft with open rotor propulsion. By studying the directivity patterns of individual tones, it is shown that though the tonal energy distribution and the spectral content of the wind tunnel test model, and thus its total noise, may differ from those of more advanced rotor designs, the individual tones follow directivity patterns that characterize far field radiations of modern open rotors, ensuring the validity of the use of this shielding data. Thus, open rotor tonal noise shielding should be categorized into front rotor tones, aft rotor tones and interaction tones, not only because of the different directivities of the three groups of tones, but also due to the differences in their source locations and coherence features, which make the respective shielding characteristics of the three groups of tones distinctly different from each other. To reveal the parametric trends of the BWB shielding effects, results are presented with variations in frequency, far field emission angle, rotor operational condition, engine installation geometry, and local airframe features. These results prepare the way for the development of parametric models for the shielding effects in prediction tools.
Willkommen, Jana; Hörnschemeyer, Thomas
2007-06-01
The ability to fly is the decisive factor for the evolutionary success of winged insects (Pterygota). Despite this, very little is known about the ground-pattern and evolution of the functionally very important wing base. Here we use the Ephemeroptera, usually regarded as the most ancient flying insects, as a model for the analysis of the flight musculature and the sclerites of the wing base. Morphology and anatomy of the pterothorax of 13 species of Ephemeroptera and five species of Plecoptera were examined and a detailed description of Habroleptoides confusa (Ephemeroptera: Leptophlebiidae) is given. A new homology of the wing base sclerites in Ephemeroptera is proposed. The wing base of Ephemeroptera possesses three axillary sclerites that are homologous to the first axillary, the second axillary and the third axillary of Neoptera. For example, the third axillary possesses the axillary-pleural muscle that mostly is considered as a characteristic feature of the Neoptera. Many of the muscles and sclerites of the flight system of the Ephemeroptera and Neoptera can be readily homologised. In fact, there are indications that a foldable wing base may be a ground plan feature of pterygote insects and that the non-foldable wing base of the Ephemeroptera is a derived state.
Insect Wing Displacement Measurement Using Digital Holography
DOE Office of Scientific and Technical Information (OSTI.GOV)
Aguayo, Daniel D.; Mendoza Santoyo, Fernando; Torre I, Manuel H. de la
2008-04-15
Insects in flight have been studied with optical non destructive techniques with the purpose of using meaningful results in aerodynamics. With the availability of high resolution and large dynamic range CCD sensors the so called interferometric digital holographic technique was used to measure the surface displacement of in flight insect wings, such as butterflies. The wings were illuminated with a continuous wave Verdi laser at 532 nm, and observed with a CCD Pixelfly camera that acquire images at a rate of 11.5 frames per second at a resolution of 1392x1024 pixels and 12 Bit dynamic range. At this frame ratemore » digital holograms of the wings were captured and processed in the usual manner, namely, each individual hologram is Fourier processed in order to find the amplitude and phase corresponding to the digital hologram. The wings displacement is obtained when subtraction between two digital holograms is performed for two different wings position, a feature applied to all consecutive frames recorded. The result of subtracting is seen as a wrapped phase fringe pattern directly related to the wing displacement. The experimental data for different butterfly flying conditions and exposure times are shown as wire mesh plots in a movie of the wings displacement.« less
Aerodynamic Interaction between Delta Wing and Hemisphere-Cylinder in Supersonic Flow
NASA Astrophysics Data System (ADS)
Nishino, Atsuhiro; Ishikawa, Takahumi; Nakamura, Yoshiaki
As future space vehicles, Reusable Launch Vehicle (RLV) needs to be developed, where there are two kinds of RLV: Single Stage To Orbit (SSTO) and Two Stage To Orbit (TSTO). In the latter case, the shock/shock interaction and shock/boundary layer interaction play a key role. In the present study, we focus on the supersonic flow field with aerodynamic interaction between a delta wing and a hemisphere-cylinder, which imitate a TSTO, where the clearance, h, between the delta wing and hemisphere-cylinder is a key parameter. As a result, complicated flow patterns were made clear, including separation bubbles.
Propeller swirl effect on single-engine general-aviation aircraft stall-spin tendencies
NASA Technical Reports Server (NTRS)
Katz, Joseph; Feistel, Terry W.
1987-01-01
An investigation is conducted of the effect of a single engine, untapered low wing general aviation aircraft propeller's swirl on the craft's stall pattern. The asymmetrical character of the propeller's swirl can trigger an early stall of one of the wings, aggravating the spin-entry condition. It is shown that the combination of this propeller-induced effect with adverse sideslip can result in large and abrupt changes in the rolling moment, in such conditions as uncoordinated low speed turning maneuvers where the pilot yaws the aircraft with wings level, rather than rolling it.
Genome of Drosophila suzukii, the Spotted Wing Drosophila
Chiu, Joanna C.; Jiang, Xuanting; Zhao, Li; Hamm, Christopher A.; Cridland, Julie M.; Saelao, Perot; Hamby, Kelly A.; Lee, Ernest K.; Kwok, Rosanna S.; Zhang, Guojie; Zalom, Frank G.; Walton, Vaughn M.; Begun, David J.
2013-01-01
Drosophila suzukii Matsumura (spotted wing drosophila) has recently become a serious pest of a wide variety of fruit crops in the United States as well as in Europe, leading to substantial yearly crop losses. To enable basic and applied research of this important pest, we sequenced the D. suzukii genome to obtain a high-quality reference sequence. Here, we discuss the basic properties of the genome and transcriptome and describe patterns of genome evolution in D. suzukii and its close relatives. Our analyses and genome annotations are presented in a web portal, SpottedWingFlyBase, to facilitate public access. PMID:24142924
In-Flight Wing Pressure Distributions for the NASA F/A-18A High Alpha Research Vehicle
NASA Technical Reports Server (NTRS)
Davis, Mark C.; Saltzman, John A.
2000-01-01
Pressure distributions on the wings of the F/A-18A High Alpha Research Vehicle (HARV) were obtained using both flush-mounted pressure orifices and surface-mounted pressure tubing. During quasi-stabilized 1-g flight, data were gathered at ranges for angle of attack from 5 deg to 70 deg, for angle of sideslip from -12 deg to +12 deg, and for Mach from 0.23 to 0.64, at various engine settings, and with and without the leading edge extension fence installed. Angle of attack strongly influenced the wing pressure distribution, as demonstrated by a distinct flow separation pattern that occurred between the range from 15 deg to 30 deg. Influence by the leading edge extension fence was evident on the inboard wing pressure distribution, but little influence was seen on the outboard portion of the wing. Angle-of-sideslip influence on wing pressure distribution was strongest at low angle of attack. Influence of Mach number was observed in the regions of local supersonic flow, diminishing as angle of attack was increased. Engine throttle setting had little influence on the wing pressure distribution.
Evolution of female-specific wingless forms in bagworm moths.
Niitsu, Shuhei; Sugawara, Hirotaka; Hayashi, Fumio
2017-01-01
The evolution of winglessness in insects has been typically interpreted as a consequence of developmental and other adaptations to various environments that are secondarily derived from a winged morph. Several species of bagworm moths (Insecta: Lepidoptera, Psychidae) exhibit a case-dwelling larval life style along with one of the most extreme cases of sexual dimorphism: wingless female adults. While the developmental process that led to these wingless females is well known, the origins and evolutionary transitions are not yet understood. To examine the evolutionary patterns of wing reduction in bagworm females, we reconstruct the molecular phylogeny of over 30 Asian species based on both mitochondrial (cytochrome c oxidase subunit I) and nuclear (28S rRNA) DNA sequences. Under a parsimonious assumption, the molecular phylogeny implies that: (i) the evolutionary wing reduction towards wingless females consisted of two steps: (Step I) from functional wings to vestigial wings (nonfunctional) and (Step II) from vestigial wings to the most specialized vermiform adults (lacking wings and legs); and (ii) vermiform morphs evolved independently at least twice. Based on the results of our study, we suggest that the evolutionary changes in the developmental system are essential for the establishment of different wingless forms in insects. © 2016 Wiley Periodicals, Inc.
Linz, David M; Tomoyasu, Yoshinori
2015-01-01
The amazing array of diversity among insect wings offers a powerful opportunity to study the mechanisms guiding morphological evolution. Studies in Drosophila (the fruit fly) have identified dozens of genes important for wing development. These genes are often called candidate genes, serving as an ideal starting point to study wing development in other insects. However, we also need to explore beyond the candidate genes to gain a more comprehensive view of insect wing evolution. As a first step away from the traditional candidate genes, we utilized Tribolium (the red flour beetle) as a model and assessed the potential involvement of a group of developmental toolkit genes (embryonic patterning genes) in beetle wing development. We hypothesized that the highly pleiotropic nature of these developmental genes would increase the likelihood of finding novel wing genes in Tribolium. Through the RNA interference screening, we found that Tc-cactus has a less characterized (but potentially evolutionarily conserved) role in wing development. We also found that the odd-skipped family genes are essential for the formation of the thoracic pleural plates, including the recently discovered wing serial homologs in Tribolium. In addition, we obtained several novel insights into the function of these developmental genes, such as the involvement of mille-pattes and Tc-odd-paired in metamorphosis. Despite these findings, no gene we examined was found to have novel wing-related roles unique in Tribolium. These results suggest a relatively conserved nature of developmental toolkit genes and highlight the limited degree to which these genes are co-opted during insect wing evolution.
Fiedler, Wolfgang
2005-06-01
An analysis of the external flight apparatus of 700 blackcaps from eight different populations (sedentary to long-distance migrators) is presented. With increasing migration distances of populations, (1) wing length, aspect ratio, and wing pointedness increase; (2) wing load decreases; (3) slots on the wing tips become relatively shorter; (4) the alula tends to be shorter in relation to wing length; and (5) the tail is shorter in relation to wing length. Although body mass increases from southern to northern populations, changes in wing length and wing area are two to three times larger than expected for simple isometric relationships. Regarding the aerodynamic background of these changes, it can be stated that traits for energy-effective flight are more strongly developed and traits for maneuverability are less developed in birds traveling longer distances, presumably as a consequence of trade-offs. Nonmigratory blackcaps from Madeira and the Cape Verde islands do not always show the traits we would expect in view of their sedentary behavior. This can be seen as a result of recent colonization of these islands by migrants or of selection by factors other than migration behavior. In migratory populations, changes between the first and the second set of primaries during first complete molt show almost the same pattern as the changes from nonmigratory to migratory populations. During molt of the primaries, blackcaps of nonmigratory populations do not show these changes. Hybrids between migrating and nonmigrating blackcap populations (Moscow and Madeira) showed intermediate values between parent populations in wing length, wing shape, and wing area; in the other variables they resembled either parent population.
NASA Technical Reports Server (NTRS)
Jones, Kenneth M.; Biedron, Robert T.; Whitlock, Mark
1995-01-01
A computational study was performed to determine the predictive capability of a Reynolds averaged Navier-Stokes code (CFL3D) for two-dimensional and three-dimensional multielement high-lift systems. Three configurations were analyzed: a three-element airfoil, a wing with a full span flap and a wing with a partial span flap. In order to accurately model these complex geometries, two different multizonal structured grid techniques were employed. For the airfoil and full span wing configurations, a chimera or overset grid technique was used. The results of the airfoil analysis illustrated that although the absolute values of lift were somewhat in error, the code was able to predict reasonably well the variation with Reynolds number and flap position. The full span flap analysis demonstrated good agreement with experimental surface pressure data over the wing and flap. Multiblock patched grids were used to model the partial span flap wing. A modification to an existing patched- grid algorithm was required to analyze the configuration as modeled. Comparisons with experimental data were very good, indicating the applicability of the patched-grid technique to analyses of these complex geometries.
Vortex wake alleviation studies with a variable twist wing
NASA Technical Reports Server (NTRS)
Holbrook, G. T.; Dunham, D. M.; Greene, G. C.
1985-01-01
Vortex wake alleviation studies were conducted in a wind tunnel and a water towing tank using a multisegmented wing model which provided controlled and measured variations in span load. Fourteen model configurations are tested at a Reynolds number of one million and a lift coefficient of 0.6 in the Langley 4- by 7-Meter Tunnel and the Hydronautics Ship Model Basin water tank at Hydronautics, Inc., Laurel, Md. Detailed measurements of span load and wake velocities at one semispan downstream correlate well with each other, with inviscid predictions of span load and wake roll up, and with peak trailing-wing rolling moments measured in the far wake. Average trailing-wing rolling moments are found to be an unreliable indicator of vortex wake intensity because vortex meander does not scale between test facilities and free-air conditions. A tapered-span-load configuration, which exhibits little or no drag penalty, is shown to offer significant downstream wake alleviation to a small trailing wing. The greater downstream wake alleviation achieved with the addition of spoilers to a flapped-wing configuration is shown to result directly from the high incremental drag and turbulence associated with the spoilers and not from the span load alteration they cause.
Hummingbirds generate bilateral vortex loops during hovering: evidence from flow visualization
NASA Astrophysics Data System (ADS)
Pournazeri, Sam; Segre, Paolo S.; Princevac, Marko; Altshuler, Douglas L.
2012-12-01
Visualization of the vortex wake of a flying animal provides understanding of how wingbeat kinematics are translated into the aerodynamic forces for powering and controlling flight. Two general vortex flow patterns have been proposed for the wake of hovering hummingbirds: (1) The two wings form a single, merged vortex ring during each wing stroke; and (2) the two wings form bilateral vortex loops during each wing stroke. The second pattern was proposed after a study with particle image velocimetry that demonstrated bilateral source flows in a horizontal measurement plane underneath hovering Anna's hummingbirds ( Calypte anna). Proof of this hypothesis requires a clear perspective of bilateral pairs of vortices. Here, we used high-speed image sequences (500 frames per second) of C. anna hover feeding within a white plume to visualize the vortex wake from multiple perspectives. The films revealed two key structural features: (1) Two distinct jets of downwards airflow are present under each wing; and (2) vortex loops around each jet are shed during each upstroke and downstroke. To aid in the interpretation of the flow visualization data, we analyzed high-speed kinematic data (1,000 frames per second) of wing tips and wing roots as C. anna hovered in normal air. These data were used to refine several simplified models of vortex topology. The observed flow patterns can be explained by either a single loop model with an hourglass shape or a bilateral model, with the latter being more likely. When hovering in normal air, hummingbirds used an average stroke amplitude of 153.6° (range 148.9°-164.4°) and a wingbeat frequency of 38.5 Hz (range 38.1-39.1 Hz). When hovering in the white plume, hummingbirds used shallower stroke amplitudes ( bar{x} = 129.8°, range 116.3°-154.1°) and faster wingbeat frequencies ( bar{x} = 41.1 Hz, range 38.5-44.7 Hz), although the bilateral jets and associated vortices were observed across the full kinematic range. The plume did not significantly alter the air density or constrain the sustained muscle contractile frequency. Instead, higher wingbeat frequencies likely incurred a higher metabolic cost with the possible benefit of allowing the birds to more rapidly escape from the visually disruptive plume.
Hummingbirds generate bilateral vortex loops during hovering: evidence from flow visualization
NASA Astrophysics Data System (ADS)
Pournazeri, Sam; Segre, Paolo S.; Princevac, Marko; Altshuler, Douglas L.
2013-01-01
Visualization of the vortex wake of a flying animal provides understanding of how wingbeat kinematics are translated into the aerodynamic forces for powering and controlling flight. Two general vortex flow patterns have been proposed for the wake of hovering hummingbirds: (1) The two wings form a single, merged vortex ring during each wing stroke; and (2) the two wings form bilateral vortex loops during each wing stroke. The second pattern was proposed after a study with particle image velocimetry that demonstrated bilateral source flows in a horizontal measurement plane underneath hovering Anna's hummingbirds ( Calypte anna). Proof of this hypothesis requires a clear perspective of bilateral pairs of vortices. Here, we used high-speed image sequences (500 frames per second) of C. anna hover feeding within a white plume to visualize the vortex wake from multiple perspectives. The films revealed two key structural features: (1) Two distinct jets of downwards airflow are present under each wing; and (2) vortex loops around each jet are shed during each upstroke and downstroke. To aid in the interpretation of the flow visualization data, we analyzed high-speed kinematic data (1,000 frames per second) of wing tips and wing roots as C. anna hovered in normal air. These data were used to refine several simplified models of vortex topology. The observed flow patterns can be explained by either a single loop model with an hourglass shape or a bilateral model, with the latter being more likely. When hovering in normal air, hummingbirds used an average stroke amplitude of 153.6° (range 148.9°-164.4°) and a wingbeat frequency of 38.5 Hz (range 38.1-39.1 Hz). When hovering in the white plume, hummingbirds used shallower stroke amplitudes ( bar{x} = 129.8°, range 116.3°-154.1°) and faster wingbeat frequencies ( bar{x} = 41.1 Hz, range 38.5-44.7 Hz), although the bilateral jets and associated vortices were observed across the full kinematic range. The plume did not significantly alter the air density or constrain the sustained muscle contractile frequency. Instead, higher wingbeat frequencies likely incurred a higher metabolic cost with the possible benefit of allowing the birds to more rapidly escape from the visually disruptive plume.
Subsonic Reynolds Number Effects on a Diamond Wing Configuration
NASA Technical Reports Server (NTRS)
Luckring, J. M.; Ghee, T. A.
2001-01-01
An advanced diamond-wing configuration was tested at low speeds in the National Transonic Facility (NTF) in air at chord Reynolds numbers from 4.4 million (typical wind-tunnel conditions) to 24 million (nominal flight value). Extensive variations on high-lift rigging were explored as part of a broad multinational program. The analysis for this study is focused on the cruise and landing settings of the wing high-lift systems. Three flow domains were identified from the data and provide a context for the ensuing data analysis. Reynolds number effects were examined in incremental form based upon attached-flow theory. A similar approach showed very little effect of low-speed compressibility.
The aerodynamic characteristics of seven frequently used wing sections at full Reynolds number
NASA Technical Reports Server (NTRS)
Munk, Max M; Miller, Elton W
1927-01-01
This report contains the aerodynamic properties of the wing sections U.S.A. 5, U.S.A. 27, U.S.A. 35 A, U.S.A. 35 B, Clark Y, R.A.F. 15, and Gottingen 387, as determined at various Reynolds numbers up to an approximately full scale value in the variable density wind tunnel of the National Advisory Committee for Aeronautics. It is shown that the characteristics of the wings investigated are affected greatly and in a somewhat erratic manner by variation of the Reynolds number. In general there is a small increase in maximum lift and an appreciable decrease in drag at all lifts.
NASA Technical Reports Server (NTRS)
Gloss, B. B.
1978-01-01
A close-coupled canard-wing configuration was tested in the Langely high-speed 7 by 10 foot tunnel at a Mach number of 0.30 to determine the effect of changing wing camber on the trimmed lift capability. Trimmed lift coefficients of near 2.0 were attained; however, the data indicated that the highest buffet-free trimmed lift coefficient attainable was approximately 1.30. The buffet used in this investigation were qualitative in nature and gave no indication of buffet intensity. Thus, the trimmed lift coefficient of near 2.0 might be attainable if the buffet intensity was not too high. The data showed that there was approximately a 10 percent variation in drag coefficient, for different model configurations, at a given trimmed lift coefficient. Large increases in wing lift had only small effects on canard lift.
Piloted simulation study of two tilt-wing control concepts
NASA Technical Reports Server (NTRS)
Birckelbaw, Lourdes G.; Corliss, Lloyd D.
1994-01-01
A two-phase piloted simulation study was conducted to investigate alternative wing and flap controls for tilt-wing aircraft. The initial phase of the study compared the flying qualities of both a conventional (programmed) flap and an innovative geared flap. The second phase of the study introduced an alternate method of pilot control for the geared flap and further studied the flying qualities of the programmed flap, and two geared flap configurations. In general, the pilot rating showed little variation between the programmed flap and the geared flap control concepts. Some differences between the two concepts were noticed and are discussed in this paper. The addition of pitch attitude stabilization in the second phase of the study greatly enhanced the aircraft flying qualities. This paper describes the simulated tilt-wing aircraft and the flap control concepts and presents the results of both phases of the simulation study.
Replication of cicada wing's nano-patterns by hot embossing and UV nanoimprinting
NASA Astrophysics Data System (ADS)
Hong, Sung-Hoon; Hwang, Jaeyeon; Lee, Heon
2009-09-01
The hydrophobicity of the cicada wing originates from its naturally occurring, surface nano-structure. The nano-structure of the cicada wing consists of an array of nano-sized pillars, 100 nm in diameter and 300 nm in height. In this study, the nano-structure of the cicada wing was successfully duplicated by using hot embossing lithography and UV nanoimprint lithography (NIL). The diameter and pitch of replication were the same as those of the original cicada wing and the height was a little smaller than that of the original master. The transmittance of the hot embossed PVC film was increased by 2-6% compared with that of the bare PVC film. The hydrophobicity was measured by water contact angle measurements. The water contact angle of the replica, made of UV cured polymer, was 132° ± 2°, which was slightly lower than that of the original cicada wing (138° ± 2°), but much higher than that of the UV cured polymer surface without any nano-sized pillars (86°).
Kaulfuss, Uwe; Moulds, Max
2015-01-01
Abstract A new genus and species of primitive cicada (Hemiptera: Tettigarctidae) is described from the early Miocene of southern New Zealand. Paratettigarcta zealandica gen. et sp. n. is the first cicada (Cicadoidea) fossil from New Zealand and exhibits wing venation patterns typical for the subfamily Tettigarctinae. It differs from other fossil taxa and the extant genus Tettigarcta in the early divergence of CuA2 from the nodal line in the forewing, its parallel-sided subcostal cell, the early bifurcation of vein M and long apical cells of the hindwing, and in wing pigmentation patterns. PMID:25829843
Kaulfuss, Uwe; Moulds, Max
2015-01-01
A new genus and species of primitive cicada (Hemiptera: Tettigarctidae) is described from the early Miocene of southern New Zealand. Paratettigarctazealandica gen. et sp. n. is the first cicada (Cicadoidea) fossil from New Zealand and exhibits wing venation patterns typical for the subfamily Tettigarctinae. It differs from other fossil taxa and the extant genus Tettigarcta in the early divergence of CuA2 from the nodal line in the forewing, its parallel-sided subcostal cell, the early bifurcation of vein M and long apical cells of the hindwing, and in wing pigmentation patterns.
Seasonal variation of columnar aerosol optical properties and radiative forcing over Beijing, China
NASA Astrophysics Data System (ADS)
Yu, Xingna; Lü, Rui; Liu, Chao; Yuan, Liang; Shao, Yixing; Zhu, Bin; Lei, Lu
2017-10-01
Long-term seasonal characteristics of aerosol optical properties and radiative forcing at Beijing (during March 2001-March 2015) were investigated using a combination of ground-based Sun/sky radiometer retrievals from the AERONET and a radiative transfer model. Aerosol optical depth (AOD) showed a distinct seasonal variation with higher values in spring and summer, and relatively lower values in fall and winter. Average Angstrom exponent (AE) in spring was lower than other seasons, implying the significant impact of dust episodes on aerosol size distribution. AE mainly distributed between 1.0 and 1.4 with an obvious uni-peak pattern in each season. The observation data showed that high AODs (>1.0) were clustered in the fine mode growth wing and the coarse mode. Compared to AOD, seasonal variation in single scattering albedo (SSA) showed an opposite pattern with larger values in summer and spring, and smaller ones in winter and fall. The highest volume size distribution and median radius of fine mode particles occurred in summer, while those of coarse mode particles in spring. The averaged aerosol radiative forcing (ARF) at the top of the atmosphere (TOA) in spring, summer, fall and winter were -33 ± 22 W m-2, -35 ± 22 W m-2, -28 ± 20 W m-2, and -24 ± 23 W m-2 respectively, and these differences were mainly due to the SSA seasonal variation. The largest positive ARF within atmosphere occurred in spring, implying strong warming in the atmosphere. The low heating ratio in summer was caused by the increase in water vapor content, which enhanced light scattering capacity (i.e., increased SSA).
2016-09-07
approach in co simulation with fluid-dynamics solvers is used. An original variational formulation is developed for the inverse problem of...by the inverse solution meshing. The same approach is used to map the structural and fluid interface kinematics and loads during the fluid structure...co-simulation. The inverse analysis is verified by reconstructing the deformed solution obtained with a corresponding direct formulation, based on
Bennema-Broos, M; Groenewegen, P P; Westert, G P
2001-06-01
In this paper, the hypothesis that the spatial distribution of hospital beds is more even in countries with socialist or social democratic governments than in countries with conservative or Christian democratic governments was tested. To avoid the confounding influences of historical and institutional differences between countries, we used the Federal Republic of Germany as a case study. The German federal states have their own governments who play an important role in creating structures for the planning of hospital facilities. The test of the hypothesis was largely quantitative. At the level of federal states the rank correlation was computed between the weighted number of years of left-wing government participation and the coefficient of variation in the number of hospital beds per 1000 inhabitants. In addition to this, the hospital plans of two federal states were studied. The hypothesis was supported by the data, showing a positive association between the number of years of left-wing government participation and regional variation in the number of hospital beds. A comparison of the hospital plans of two contrasting federal states showed less government interference in hospital planning in the state with a tradition of right-wing government. There seems to be a relation between left-wing government participation in West German states and a more equal distribution of the number of hospital beds per 1,000 inhabitants.
Bolstad, Geir H.; Cassara, Jason A.; Márquez, Eladio; Hansen, Thomas F.; van der Linde, Kim; Houle, David; Pélabon, Christophe
2015-01-01
Precise exponential scaling with size is a fundamental aspect of phenotypic variation. These allometric power laws are often invariant across taxa and have long been hypothesized to reflect developmental constraints. Here we test this hypothesis by investigating the evolutionary potential of an allometric scaling relationship in drosophilid wing shape that is nearly invariant across 111 species separated by at least 50 million years of evolution. In only 26 generations of artificial selection in a population of Drosophila melanogaster, we were able to drive the allometric slope to the outer range of those found among the 111 sampled species. This response was rapidly lost when selection was suspended. Only a small proportion of this reversal could be explained by breakup of linkage disequilibrium, and direct selection on wing shape is also unlikely to explain the reversal, because the more divergent wing shapes produced by selection on the allometric intercept did not revert. We hypothesize that the reversal was instead caused by internal selection arising from pleiotropic links to unknown traits. Our results also suggest that the observed selection response in the allometric slope was due to a component expressed late in larval development and that variation in earlier development did not respond to selection. Together, these results are consistent with a role for pleiotropic constraints in explaining the remarkable evolutionary stability of allometric scaling. PMID:26371319
The generation and diversification of butterfly eyespot color patterns.
Brunetti, C R; Selegue, J E; Monteiro, A; French, V; Brakefield, P M; Carroll, S B
2001-10-16
A fundamental challenge of evolutionary and developmental biology is understanding how new characters arise and change. The recently derived eyespots on butterfly wings vary extensively in number and pattern between species and play important roles in predator avoidance. Eyespots form through the activity of inductive organizers (foci) at the center of developing eyespot fields. Foci are the proposed source of a morphogen, the levels of which determine the color of surrounding wing scale cells. However, it is unknown how reception of the focal signal translates into rings of different-colored scales, nor how different color schemes arise in different species. We have identified several transcription factors, including butterfly homologs of the Drosophila Engrailed/Invected and Spalt proteins, that are deployed in concentric territories corresponding to the future rings of pigmented scales that compose the adult eyespot. We have isolated a new Bicyclus anynana wing pattern mutant, Goldeneye, in which the scales of one inner color ring become the color of a different ring. These changes correlate with shifts in transcription factor expression, suggesting that Goldeneye affects an early regulatory step in eyespot color patterning. In different butterfly species, the same transcription factors are expressed in eyespot fields, but in different relative spatial domains that correlate with divergent eyespot color schemes. Our results suggest that signaling from the focus induces nested rings of regulatory gene expression that subsequently control the final color pattern. Furthermore, the remarkably plastic regulatory interactions downstream of focal signaling have facilitated the evolution of eyespot diversity.
Hindlimb motion during steady flight of the lesser dog-faced fruit bat, Cynopterus brachyotis.
Cheney, Jorn A; Ton, Daniel; Konow, Nicolai; Riskin, Daniel K; Breuer, Kenneth S; Swartz, Sharon M
2014-01-01
In bats, the wing membrane is anchored not only to the body and forelimb, but also to the hindlimb. This attachment configuration gives bats the potential to modulate wing shape by moving the hindlimb, such as by joint movement at the hip or knee. Such movements could modulate lift, drag, or the pitching moment. In this study we address: 1) how the ankle translates through space during the wingbeat cycle; 2) whether amplitude of ankle motion is dependent upon flight speed; 3) how tension in the wing membrane pulls the ankle; and 4) whether wing membrane tension is responsible for driving ankle motion. We flew five individuals of the lesser dog-faced fruit bat, Cynopterus brachyotis (Family: Pteropodidae), in a wind tunnel and documented kinematics of the forelimb, hip, ankle, and trailing edge of the wing membrane. Based on kinematic analysis of hindlimb and forelimb movements, we found that: 1) during downstroke, the ankle moved ventrally and during upstroke the ankle moved dorsally; 2) there was considerable variation in amplitude of ankle motion, but amplitude did not correlate significantly with flight speed; 3) during downstroke, tension generated by the wing membrane acted to pull the ankle dorsally, and during upstroke, the wing membrane pulled laterally when taut and dorsally when relatively slack; and 4) wing membrane tension generally opposed dorsoventral ankle motion. We conclude that during forward flight in C. brachyotis, wing membrane tension does not power hindlimb motion; instead, we propose that hindlimb movements arise from muscle activity and/or inertial effects.
Mikó, István; Copeland, Robert S.; Balhoff, James P.; Yoder, Matthew J.; Deans, Andrew R.
2014-01-01
We revise two relatively rare ensign wasp genera, whose species are restricted to Sub-Saharan Africa: Afrevania and Trissevania. Afrevania longipetiolata sp. nov., Trissevania heatherae sp. nov., T. hugoi sp. nov., T. mrimaensis sp. nov. and T. slideri sp. nov. are described, males and females of T. anemotis and Afrevania leroyi are redescribed, and an identification key for Trissevaniini is provided. We argue that Trissevania mrimaensis sp. nov. and T. heatherae sp. nov. populations are vulnerable, given their limited distributions and threats from mining activities in Kenya. We hypothesize that these taxa together comprise a monophyletic lineage, Trissevaniini, tr. nov., the members of which share the ability to fold their fore wings along two intersecting fold lines. Although wing folding of this type has been described for the hind wing of some insects four-plane wing folding of the fore wing has never been documented. The wing folding mechanism and the pattern of wing folds of Trissevaniini is shared only with some cockroach species (Blattodea). It is an interesting coincidence that all evaniids are predators of cockroach eggs. The major wing fold lines of Trissevaniini likely are not homologous to any known longitudinal anatomical structures on the wings of other Evaniidae. Members of the new tribe share the presence of a coupling mechanism between the fore wing and the mesosoma that is composed of a setal patch on the mesosoma and the retinaculum of the fore wing. While the setal patch is an evolutionary novelty, the retinaculum, which originally evolved to facilitate fore and hind wing coupling in Hymenoptera, exemplifies morphological exaptation. We also refine and clarify the Semantic Phenotype approach used in previous taxonomic revisions and explore the consequences of merging new with existing data. The way that semantic statements are formulated can evolve in parallel, alongside improvements to the ontologies themselves. PMID:24787704
Rajabi, H.; Ghoroubi, N.; Malaki, M.; Darvizeh, A.; Gorb, S. N.
2016-01-01
Dragonflies and damselflies, belonging to the order Odonata, are known to be excellent fliers with versatile flight capabilities. The ability to fly over a wide range of speeds, high manoeuvrability and great agility are a few characteristics of their flight. The architecture of the wings and their structural elements have been found to play a major role in this regard. However, the precise influence of individual wing components on the flight performance of these insects remains unknown. The design of the wing basis (so called basal complex) and the venation of this part are responsible for particular deformability and specific shape of the wing blade. However, the wing bases are rather different in representatives of different odonate groups. This presumably reflects the dimensions of the wings on one hand, and different flight characteristics on the other hand. In this article, we develop the first three-dimensional (3D) finite element (FE) models of the proximal part of the wings of typical representatives of five dragonflies and damselflies families. Using a combination of the basic material properties of insect cuticle, a linear elastic material model and a nonlinear geometric analysis, we simulate the mechanical behaviour of the wing bases. The results reveal that although both the basal venation and the basal complex influence the structural stiffness of the wings, it is only the latter which significantly affects their deformation patterns. The use of numerical simulations enabled us to address the role of various wing components such as the arculus, discoidal cell and triangle on the camber formation in flight. Our study further provides a detailed representation of the stress concentration in the models. The numerical analysis presented in this study is not only of importance for understanding structure-function relationship of insect wings, but also might help to improve the design of the wings for biomimetic micro-air vehicles (MAVs). PMID:27513753
Diversity in the organization of elastin bundles and intramembranous muscles in bat wings.
Cheney, Jorn A; Allen, Justine J; Swartz, Sharon M
2017-04-01
Unlike birds and insects, bats fly with wings composed of thin skin that envelops the bones of the forelimb and spans the area between the limbs, digits, and sometimes the tail. This skin is complex and unusual; it is thinner than typical mammalian skin and contains organized bundles of elastin and embedded skeletal muscles. These elements are likely responsible for controlling the shape of the wing during flight and contributing to the aerodynamic capabilities of bats. We examined the arrangement of two macroscopic architectural elements in bat wings, elastin bundles and wing membrane muscles, to assess the diversity in bat wing skin morphology. We characterized the plagiopatagium and dactylopatagium of 130 species from 17 families of bats using cross-polarized light imaging. This method revealed structures with distinctive relative birefringence, heterogeneity of birefringence, variation in size, and degree of branching. We used previously published anatomical studies and tissue histology to identify birefringent structures, and we analyzed their architecture across taxa. Elastin bundles, muscles, neurovasculature, and collagenous fibers are present in all species. Elastin bundles are oriented in a predominantly spanwise or proximodistal direction, and there are five characteristic muscle arrays that occur within the plagiopatagium, far more muscle than typically recognized. These results inform recent functional studies of wing membrane architecture, support the functional hypothesis that elastin bundles aid wing folding and unfolding, and further suggest that all bats may use these architectural elements for flight. All species also possess numerous muscles within the wing membrane, but the architecture of muscle arrays within the plagiopatagium varies among families. To facilitate present and future discussion of these muscle arrays, we refine wing membrane muscle nomenclature in a manner that reflects this morphological diversity. The architecture of the constituents of the skin of the wing likely plays a key role in shaping wings during flight. © 2017 Anatomical Society.
Taira, Wataru; Otaki, Joji M
2016-01-01
Butterfly wing color patterns often contain eyespots, which are developmentally determined at the late larval and early pupal stages by organizing activities of focal cells that can later form eyespot foci. In the pupal stage, the focal position of a future eyespot is often marked by a focal spot, one of the pupal cuticle spots, on the pupal surface. Here, we examined the possible relationships of the pupal focal spots with the underneath pupal wing tissues and with the adult wing eyespots using Junonia butterflies. Large pupal focal spots were found in two species with large adult eyespots, J. orithya and J. almana, whereas only small pupal focal spots were found in a species with small adult eyespots, J. hedonia. The size of five pupal focal spots on a single wing was correlated with the size of the corresponding adult eyespots in J. orithya. A pupal focal spot was a three-dimensional bulge of cuticle surface, and the underside of the major pupal focal spot exhibited a hollowed cuticle in a pupal case. Cross sections of a pupal wing revealed that the cuticle layer shows a curvature at a focal spot, and a positional correlation was observed between the cuticle layer thickness and its corresponding cell layer thickness. Adult major eyespots of J. orithya and J. almana exhibited surface elevations and depressions that approximately correspond to the coloration within an eyespot. Our results suggest that a pupal focal spot is produced by the organizing activity of focal cells underneath the focal spot. Probably because the focal cell layer immediately underneath a focal spot is thicker than that of its surrounding areas, eyespots of adult butterfly wings are three-dimensionally constructed. The color-height relationship in adult eyespots might have an implication in the developmental signaling for determining the eyespot color patterns.
Taira, Wataru; Otaki, Joji M.
2016-01-01
Butterfly wing color patterns often contain eyespots, which are developmentally determined at the late larval and early pupal stages by organizing activities of focal cells that can later form eyespot foci. In the pupal stage, the focal position of a future eyespot is often marked by a focal spot, one of the pupal cuticle spots, on the pupal surface. Here, we examined the possible relationships of the pupal focal spots with the underneath pupal wing tissues and with the adult wing eyespots using Junonia butterflies. Large pupal focal spots were found in two species with large adult eyespots, J. orithya and J. almana, whereas only small pupal focal spots were found in a species with small adult eyespots, J. hedonia. The size of five pupal focal spots on a single wing was correlated with the size of the corresponding adult eyespots in J. orithya. A pupal focal spot was a three-dimensional bulge of cuticle surface, and the underside of the major pupal focal spot exhibited a hollowed cuticle in a pupal case. Cross sections of a pupal wing revealed that the cuticle layer shows a curvature at a focal spot, and a positional correlation was observed between the cuticle layer thickness and its corresponding cell layer thickness. Adult major eyespots of J. orithya and J. almana exhibited surface elevations and depressions that approximately correspond to the coloration within an eyespot. Our results suggest that a pupal focal spot is produced by the organizing activity of focal cells underneath the focal spot. Probably because the focal cell layer immediately underneath a focal spot is thicker than that of its surrounding areas, eyespots of adult butterfly wings are three-dimensionally constructed. The color-height relationship in adult eyespots might have an implication in the developmental signaling for determining the eyespot color patterns. PMID:26731532
Genotoxicity testing of two lead-compounds in somatic cells of Drosophila melanogaster.
Carmona, Erico R; Creus, Amadeu; Marcos, Ricard
2011-09-18
The in vivo genotoxic activity of two inorganic lead compounds was studied in Drosophila melanogaster by measurement of two different genetic endpoints. We used the wing-spot test and the comet assay. The comet assay was conducted with larval haemocytes. The results from the wing-spot test showed that neither lead chloride, PbCl(2), nor lead nitrate, Pb(NO(3))(2), were able to induce significant increases in the frequency of mutant spots. In addition, the combined treatments with gamma-radiation and PbCl(2) or Pb(NO(3))(2) did not show significant variations in the frequency of the three categories of mutant spots recorded, compared with the frequency induced by gamma-radiation alone. This seems to indicate that the lead compounds tested do not interact with the repair of the genetic damage induced by ionizing radiation. When the lead compounds were evaluated in the in vivo comet assay with haemocytes, Pb(NO(3))(2) was effective in inducing significant increases of DNA damage with a direct dose-response pattern. These results confirm the usefulness of the comet assay with haemocytes as an in vivo model and support the assumption that there is a genotoxic risk associated with lead exposure. Copyright © 2011 Elsevier B.V. All rights reserved.
Theoretical stability and control characteristics of wings with various amounts of taper and twist
NASA Technical Reports Server (NTRS)
Pearson, Henry A; Jones, Robert T
1938-01-01
Stability derivatives have been computed for twisted wings of different plan forms that include variations in both the wing taper and the aspect ratio. Taper ratios of 1.0, 0,50, and 0.25 are considered for each of three aspect ratios: 6, 10, and 16. The specific derivatives for which results are given are the rolling-moment and the yawing-moment derivatives with respect to (a) rolling velocity, (b) yawing velocity, and (c) angle of sideslip. These results are given in such a form that the effect of any initial symmetrical wing twist (such as may be produced by flaps) on the derivatives may easily be taken into account. In addition to the stability derivatives, results are included for determining the theoretical rolling moment due to aileron deflection and a series of influence lines is given by which the loading across the span may be determined for any angle-of-attack distribution that may occur on the wing plan forms considered. The report also includes incidental references to the application of the results.
Formation and Development of the Dynamic Stall Vortex on a Wing with Leading Edge Tubercles
NASA Astrophysics Data System (ADS)
Hrynuk, John; Bohl, Douglas
2015-11-01
Humpback whales are unique in that their flippers have leading edge ``bumps'' or tubercles. Past work on airfoils inspired by whale flippers has centered on the static aerodynamic characteristics of these airfoils. The current study uses Molecular Tagging Velocimetry (MTV) to investigate the effects of tubercles on dynamically pitching NACA 0012 airfoils. A baseline (i.e. straight leading edge) wing and one modified with leading edge tubercles are investigated. Tracking of the Dynamic Stall Vortex (DSV) is performed to quantitatively compare the DSV formation location, path, and convective velocity for tubercled and baseline wings. The results show that there is a spanwise variation in the initial formation location and motion of the DSV on the modified wing. Once formed, the DSV aligns into a more uniform spanwise structure. As the pitching motion progresses, the DSV on the modified wing convects away from the airfoil surface later and slower than is observed for the baseline airfoil. The results indicate that the tubercles may delay stall when compared to the baseline airfoil. This work was supported by NSF Grant # 0845882.
Aeroelastic character of a National Aerospace Plane demonstrator concept
NASA Technical Reports Server (NTRS)
Spain, Charles V.; Zeiler, Thomas A.; Gibbons, Michael D.; Soistmann, David L.; Pozefsky, Peter; Dejesus, Rafael O.; Brannon, Cyprian P.
1993-01-01
The paper provides an analytical assessment of the flutter character of an unclassified National Aerospace Plane configuration known as the demonstrator. Linear subsonic, supersonic, and hypersonic analysis indicate that the vehicle is prone to body-freedom flutter resulting from the decrease in vibration frequency of the all-moveable wing at high flight dynamic pressures. As the wing-pivot frequency decreases, it couples with the vehicle short-period mode resulting in dynamic instability. A similar instability sometimes occurs when the pivot mode couples with the fuselage-bending mode. Also assessed, for supersonic flight conditions, are configuration variations that include relocation of the wing further aft on the lifting-body fuselage, and the addition of body flaps to the rear of the vehicle. These changes are destabilizing because they result in severe wing-pivot/fuselage-bending instabilities at dynamic pressures lower than the instabilities indicated for the original demonstrator. Finally, a two-point wing support and actuation system concept is proposed for the National Aerospace Plane, which if developed may (according to cursory analysis) enhance overall stability.
The Effect of Large Angles of Yaw on the Accuracy of Wing-Tip Yawmeters
NASA Technical Reports Server (NTRS)
Golden, Jacob
1942-01-01
The present method used by the NACA for the measurement of sideslip angles in flight involves the use of a device called the yawmeter. The operation of this instrument depends on the motion of a free-swinging vane which, mounted ahead of the wing tip, alines itself with the local wind direction. Because of the flow pattern about the airplane, the local wind direction at the yaw vane may be slightly different from the direction of the relative wind and the yaw-vane readings may be in error. This error is corrected by using half the difference between the readings of two vanes, one on each wing, for unyawed flight as a calibration constant. It is possible, however, that, because of the change in location of the vane with respect to the flow pattern at large angles of yaw, the constant obtained for unyawed flight may not apply. The present report covers power-off tests made in the free-flight tunnel to check the validity of this method.
Rizzi, Samantha Karlla Lopes de Almeida; Haddad, Cinira Assad Simão; Giron, Patricia Santolia; Pinheiro, Thaís Lúcia; Nazário, Afonso Celso Pinto; Facina, Gil
2016-06-01
The aim of this study was to determine the incidence of winged scapula after breast cancer surgery, its impact on shoulder morbidity and difference in incidence according to surgery type. Patients with breast cancer and surgical indication for axillary dissection were included. A total of 112 patients were surveyed with one physical evaluation before the surgery and others 15, 30, 90, and 180 days after. Winged scapula was assessed with test proposed by Hoppenfeld. Shoulder range of motion (ROM) was assessed with goniometer for flexion, extension, adduction, abduction, internal rotation, and external rotation. A verbal scale from 0 to 10 was used to assess pain. Winged scapula incidence was 8.0 % 15 days after surgery. Two patients recovered from winged scapula 90 days after surgery and four more 180 days after surgery, while three patients still had winged scapula at this time. The incidence after 15 days from surgery was 20.9 and 22.6 % among patients submitted to sentinel node biopsy or axillary lymphadenectomy (AL), respectively (p < 0.01). There was no statistical difference of incidence according to breast surgery type. Operated side shoulder flexion, adduction, and abduction ROM changes were statistically different in patients with or without winged scapula. The mean reduction was higher in patients with winged scapula. Both groups showed the same pattern over time in pain. Winged scapula incidence was 8.0 % and was higher in AL, and prevalence decreased during 6 months after surgery. Patients who developed winged scapula had more shoulder flexion, adduction, and abduction limitation.
NASA Astrophysics Data System (ADS)
Krishnan, Krishnamoorthy; Naqavi, Iftekhar Z.; Gurka, Roi
2017-11-01
Understanding the physics of flapping wings at moderate Reynolds number flows takes on greater importance in the context of avian aerodynamics as well as in the design of miniature-aerial-vehicles. Analyzing the characteristics of wake vortices generated downstream of flapping wings can help to explain the unsteady contribution to the aerodynamics loads. In this study, numerical simulations of flow over a bio-inspired pseudo-2D flapping wing model was conducted to characterize the evolution of unsteady flow structures in the downstream wake of flapping wing. The wing model was based on a European starling's wing and wingbeat kinematics were incorporated to simulate a free-forward flight. The starling's wingbeat kinematics were extracted from experiments conducted in a wind tunnel where freely flying starling was measured using high-speed PIV as well as high-speed imaging yielding a series of kinematic images sampled at 500 Hz. The average chord of the wing section was 6 cm and simulations were carried out at a Reynolds number of 54,000, reduced frequency of 0.17, and Strouhal number of 0.16. Large eddy simulation was performed using a second order, finite difference code ParLES. Characteristics of wake vortex structures during the different phases of the wing strokes were examined. The role of wingbeat kinematics in the configuration of downstream vortex patterns is discussed. Evaluated wake topology and lift-drag characteristics are compared with the starling's wind tunnel results.
Cooper, R G; El Doumani, H A A
2006-03-01
Quill mites (Gabucinia bicaudata) and lice (Struthiolipeurus struthionis) may infest ostrich feathers, resulting in skin damage, pruritus and excessive feather preening and loss. Four different feather types (prime white, femina extra wide, femina class 1, and femina short; n = 10) were collected. The quill mites and lice were removed with fine forceps, studied using a photographic optical microscope and counted microscopically at x 100 magnification following collection by sedimentation. They were placed in separate Petri dishes containing lactophenol solution and examined (x40 magnification). Anatomical features are described. The density of quill mites in all feather types of both wings was higher than that of the lice. There was no significant difference between the counts of both arthropods on the left wing and the right wing, respectively, except for the femina class 1 quill mites (P = 0.01). The femina extra wide feathers were a preferred habitat in both wings. Large standard deviations (quill mites left wing: 73 +/- 8; quill mites right wing: 69 +/- 7) suggested variations in the degree of migration between feather shafts or as a response to escape preening. It is recommended that ostriches be treated with an oral preparation of Ivermectin administered per os at a dosage rate of 0.2 mg/kg at 30-day intervals for quill mites, and with a 1-5 % Malathion dust at 14-day intervals for lice.
Hahn, Steffen; Korner-Nievergelt, Fränzi; Emmenegger, Tamara; Amrhein, Valentin; Csörgő, Tibor; Gursoy, Arzu; Ilieva, Mihaela; Kverek, Pavel; Pérez-Tris, Javier; Pirrello, Simone; Zehtindjiev, Pavel; Salewski, Volker
2016-01-01
In migratory birds, morphological adaptations for efficient migratory flight often oppose morphological adaptations for efficient behavior during resident periods. This includes adaptations in wing shape for either flying long distances or foraging in the vegetation and in climate-driven variation of body size. In addition, the timing of migratory flights and particularly the timely arrival at local breeding sites is crucial because fitness prospects depend on site-specific phenology. Thus, adaptations for efficient long-distance flights might be also related to conditions at destination areas. For an obligatory long-distance migrant, the common nightingale, we verified that wing length as the aerodynamically important trait, but not structural body size increased from the western to the eastern parts of the species range. In contrast with expectation from aerodynamic theory, however, wing length did not increase with increasing migration distances. Instead, wing length was associated with the phenology at breeding destinations, namely the speed of local spring green-up. We argue that longer wings are beneficial for adjusting migration speed to local conditions for birds breeding in habitats with fast spring green-up and thus short optimal arrival periods. We suggest that the speed of spring green-up at breeding sites is a fundamental variable determining the timing of migration that fine tune phenotypes in migrants across their range.
Lin, Xinda; Yao, Yun; Wang, Bo; Emlen, Douglas J; Lavine, Laura Corley
2016-01-01
Crowding and changes in food availability are two critical environmental conditions that impact an animal's trajectory toward either migration or reproduction. Many insects facing this challenge have evolved wing polyphenisms. When conditions favor reproduction, wing polyphenic species produce adults that either have no wings or short, non-functional wings. Facultative wing growth reflects a physiological and evolutionary trade-off between migration and reproduction, triggered by environmental conditions. How environmental cues are transduced to produce these alternative forms, and their associated ecological shift from migration to reproduction, remains an important unsolved problem in evolutionary ecology. The brown planthopper, a wing polymorphic insect exhibiting strong trade-offs in investment between migration and reproduction, is one of the most serious rice pests in Asia. In this study, we investigated the function of four genes in the insulin-signaling pathway known to couple nutrition with growth, PI3 Kinase (PI3K), PDK1, Akt (Protein Kinase B), and the forkhead gene FOXO. Using a combination of RNA interference and pharmacological inhibitor treatment, we show that all four genes contribute to tissue level regulation of wing polymorphic development in this insect. As predicted, silencing of the NlPI3K, NlAkt and NlPDK1 through dsRNA and with the pharmacological inhibitor Perifosine resulted in short-winged brown planthoppers, whereas knockdown of NlFOXO resulted in long-winged planthoppers. Morphometric analyses confirm that phenotypes from our manipulations mimic what would be found in nature, i.e., major parameters such as bristle number, wing area and body weight are not significantly different from non-experimental animals. Taken together, these data implicate the insulin-signaling pathway in the transduction of environmental factors into condition-dependent patterns of wing growth in insects.
Morphogenesis in bat wings: linking development, evolution and ecology.
Adams, Rick A
2008-01-01
The evolution of powered flight in mammals required specific developmental shifts from an ancestral limb morphology to one adapted for flight. Through studies of comparative morphogenesis, investigators have quantified points and rates of divergence providing important insights into how wings evolved in mammals. Herein I compare growth,development and skeletogenesis of forelimbs between bats and the more ancestral state provided by the rat (Rattus norvegicus)and quantify growth trajectories that illustrate morphological divergence both developmentally and evolutionarily. In addition, I discuss how wing shape is controlled during morphogenesis by applying multivariate analyses of wing bones and wing membranes and discuss how flight dynamics are stabilized during flight ontogeny. Further, I discuss the development of flight in bats in relation to the ontogenetic niche and how juveniles effect populational foraging patterns. In addition, I provide a hypothetical ontogenetic landscape model that predicts how and when selection is most intense during juvenile morphogenesis and test this model with data from a population of the little brown bat, Myotis lucifugus. (c) 2007 S. Karger AG, Basel
Boundary Dpp promotes growth of medial and lateral regions of the Drosophila wing
Barrio, Lara; Milán, Marco
2017-01-01
The gradient of Decapentaplegic (Dpp) in the Drosophila wing has served as a paradigm to characterize the role of morphogens in regulating patterning. However, the role of this gradient in regulating tissue size is a topic of intense debate as proliferative growth is homogenous. Here, we combined the Gal4/UAS system and a temperature-sensitive Gal80 molecule to induce RNAi-mediated depletion of dpp and characterise the spatial and temporal requirement of Dpp in promoting growth. We show that Dpp emanating from the AP compartment boundary is required throughout development to promote growth by regulating cell proliferation and tissue size. Dpp regulates growth and proliferation rates equally in central and lateral regions of the developing wing appendage and reduced levels of Dpp affects similarly the width and length of the resulting wing. We also present evidence supporting the proposal that graded activity of Dpp is not an absolute requirement for wing growth. DOI: http://dx.doi.org/10.7554/eLife.22013.001 PMID:28675372
An experimental study of separated flow on a finite wing
NASA Technical Reports Server (NTRS)
Winkelmann, A. E.
1981-01-01
The flow field associated with the formation of a mushroom shaped trailing edge stall cell on a low-aspect-ratio (AR = 4.0) wing was investigated in a series of low speed wind tunnel tests (Reynolds number based on 15.2 cm chord = 480,000). Flow field surveys of the separation bubble and wake of a partially stalled and fully stalled wing were completed using a hot-wire probe, a split-film probe, and a directional sensitive pressure probe. A new color video display technique was developed to display the flow field survey data. Photographs were obtained of surface oil flow patterns and smoke flow visualization
Exploring bird aerodynamics using radio-controlled models.
Hoey, Robert G
2010-12-01
A series of radio-controlled glider models was constructed by duplicating the aerodynamic shape of soaring birds (raven, turkey vulture, seagull and pelican). Controlled tests were conducted to determine the level of longitudinal and lateral-directional static stability, and to identify the characteristics that allowed flight without a vertical tail. The use of tail-tilt for controlling small bank-angle changes, as observed in soaring birds, was verified. Subsequent tests, using wing-tip ailerons, inferred that birds use a three-dimensional flow pattern around the wing tip (wing tip vortices) to control adverse yaw and to create a small amount of forward thrust in gliding flight.
Wilts, Bodo D.; Michielsen, Kristel; De Raedt, Hans; Stavenga, Doekele G.
2012-01-01
The cover scales on the wing of the Emerald-patched Cattleheart butterfly, Parides sesostris, contain gyroid-type biological photonic crystals that brightly reflect green light. A pigment, which absorbs maximally at approximately 395 nm, is immersed predominantly throughout the elaborate upper lamina. This pigment acts as a long-pass filter shaping the reflectance spectrum of the underlying photonic crystals. The additional effect of the filtering is that the spatial distribution of the scale reflectance is approximately angle-independent, leading to a stable wing pattern contrast. The spectral tuning of the original reflectance is verified by photonic band structure modelling. PMID:24098853
Saito, Kazuya; Nomura, Shuhei; Yamamoto, Shuhei; Niiyama, Ryuma; Okabe, Yoji
2017-05-30
Ladybird beetles are high-mobility insects and explore broad areas by switching between walking and flying. Their excellent wing transformation systems enabling this lifestyle are expected to provide large potential for engineering applications. However, the mechanism behind the folding of their hindwings remains unclear. The reason is that ladybird beetles close the elytra ahead of wing folding, preventing the observation of detailed processes occurring under the elytra. In the present study, artificial transparent elytra were transplanted on living ladybird beetles, thereby enabling us to observe the detailed wing-folding processes. The result revealed that in addition to the abdominal movements mentioned in previous studies, the edge and ventral surface of the elytra, as well as characteristic shaped veins, play important roles in wing folding. The structures of the wing frames enabling this folding process and detailed 3D shape of the hindwing were investigated using microcomputed tomography. The results showed that the tape spring-like elastic frame plays an important role in the wing transformation mechanism. Compared with other beetles, hindwings in ladybird beetles are characterized by two seemingly incompatible properties: ( i ) the wing rigidity with relatively thick veins and ( ii ) the compactness in stored shapes with complex crease patterns. The detailed wing-folding process revealed in this study is expected to facilitate understanding of the naturally optimized system in this excellent deployable structure.
Nomura, Shuhei; Yamamoto, Shuhei; Niiyama, Ryuma; Okabe, Yoji
2017-01-01
Ladybird beetles are high-mobility insects and explore broad areas by switching between walking and flying. Their excellent wing transformation systems enabling this lifestyle are expected to provide large potential for engineering applications. However, the mechanism behind the folding of their hindwings remains unclear. The reason is that ladybird beetles close the elytra ahead of wing folding, preventing the observation of detailed processes occurring under the elytra. In the present study, artificial transparent elytra were transplanted on living ladybird beetles, thereby enabling us to observe the detailed wing-folding processes. The result revealed that in addition to the abdominal movements mentioned in previous studies, the edge and ventral surface of the elytra, as well as characteristic shaped veins, play important roles in wing folding. The structures of the wing frames enabling this folding process and detailed 3D shape of the hindwing were investigated using microcomputed tomography. The results showed that the tape spring-like elastic frame plays an important role in the wing transformation mechanism. Compared with other beetles, hindwings in ladybird beetles are characterized by two seemingly incompatible properties: (i) the wing rigidity with relatively thick veins and (ii) the compactness in stored shapes with complex crease patterns. The detailed wing-folding process revealed in this study is expected to facilitate understanding of the naturally optimized system in this excellent deployable structure. PMID:28507159
NASA Astrophysics Data System (ADS)
Tarulescu, R.; Tarulescu, S.; Leahu, C.
2017-10-01
The conventional downforce devices (with fixed geometry) of high speed vehicles have parameters such as area, angle of incidence and head resistance coefficients, all with constant values. The downforce is proportional with the square of movement speed and the power consumed for the neutralization of aerodynamic road resistance is proportional with the cube of speed. The authors carried out an analytical study of downforce, adjustable/monitored by optimum incidence (modification of incidence angle of rear wing for performance improvement).
NASA Technical Reports Server (NTRS)
Braden, J. A.; Hancock, J. P.; Burdges, K. P.; Hackett, J. E.
1979-01-01
The work to develop a wing-nacelle arrangement to accommodate a wide range of upper surface blown configuration is reported. Pertinent model and installation details are described. Data of the effects of a wide range of nozzle geometric variations are presented. Nozzle aspect ratio, boattail angle, and chordwise position are among the parameters investigated. Straight and swept wing configurations were tested across a range of nozzle pressure ratios, lift coefficients, and Mach numbers.
Zhou, Shuang-Shuang; Sun, Ze; Ma, Weihua; Chen, Wei; Wang, Man-Qun
2014-03-01
We sequenced the antenna transcriptome of the brown planthopper (BPH), Nilaparvata lugens (Stål), a global rice pest, and performed transcriptome analysis on BPH antenna. We obtained about 40million 90bp reads that were assembled into 75,874 unigenes with a mean size of 456bp. Among the antenna transcripts, 32,856 (43%) showed significant similarity (E-value <1e(-5)) to known proteins in the NCBI database. Gene ontology and Kyoto Encyclopedia of Genes and Genomes (KEGG) analyses were used to classify functions of BPH antenna genes. We identified 10 odorant-binding proteins (OBPs), including 7 previously unidentified, and 11 chemosensory proteins (CSPs), including two new members. The expression profiles of 4 OBPs and 2 CSPs were determined by q-PCR for antenna, abdomen, leg and wing of insects of different age, gender, and mating status including two BPH adult wing-morphology types. NlugCSP10 and 4 OBPs appeared to be antenna-specific because they were highly and differentially expressed in male and female antennae. NlugCSP11 was expressed ubiquitously, with particularly high expression in wings. The transcript levels of several olfactory genes depended on adult wing form, age, gender, and mating status, although no clear expression patterns were determined. Copyright © 2013 Elsevier Inc. All rights reserved.
Preserving and vouchering butterflies and moths for large-scale museum-based molecular research
Epstein, Samantha W.; Mitter, Kim; Hamilton, Chris A.; Plotkin, David; Mitter, Charles
2016-01-01
Butterflies and moths (Lepidoptera) comprise significant portions of the world’s natural history collections, but a standardized tissue preservation protocol for molecular research is largely lacking. Lepidoptera have traditionally been spread on mounting boards to display wing patterns and colors, which are often important for species identification. Many molecular phylogenetic studies have used legs from pinned specimens as the primary source for DNA in order to preserve a morphological voucher, but the amount of available tissue is often limited. Preserving an entire specimen in a cryogenic freezer is ideal for DNA preservation, but without an easily accessible voucher it can make specimen identification, verification, and morphological work difficult. Here we present a procedure that creates accessible and easily visualized “wing vouchers” of individual Lepidoptera specimens, and preserves the remainder of the insect in a cryogenic freezer for molecular research. Wings are preserved in protective holders so that both dorsal and ventral patterns and colors can be easily viewed without further damage. Our wing vouchering system has been implemented at the University of Maryland (AToL Lep Collection) and the University of Florida (Florida Museum of Natural History, McGuire Center of Lepidoptera and Biodiversity), which are among two of the largest Lepidoptera molecular collections in the world. PMID:27366654
Inter-species variation in yolk steroid levels and a cowbird-host comparison
Hahn, D. Caldwell; Hatfield, Jeffrey S.; Abdelnabi, Mahmoud A.; Wu, Julie M.; Igl, Lawrence D.; Ottinger, Mary A.
2005-01-01
We examined variability in yolk hormone levels among songbird species and the role of yolk steroids as a mechanism for enhanced exploitation of hosts by the parasitic Brown-headed Cowbird Molothrus ater. Within-clutch variation in yolk steroids has been found in several avian species in single species studies, but few comparisons have been made among species. We found a large range of differences in yolk testosterone among the seven passerine species examined, with significant differences between those at the high end (Song Sparrow Melospiza melodia , Red-winged Blackbird Agelaius phoeniceus, and House Sparrow, Passer domesticus ) and those at the low end (Eastern Phoebe Sayornis phoebe, and House Finch Carpodacus mexicanus ). We also found that the testosterone level in cowbird eggs was intermediate in relation to host species levels and was significantly lower than that in three common cowbird hosts (Song Sparrow, Red-winged Blackbird, and House Sparrow), but not significantly different from three others. Geographical comparisons of yolk testosterone levels in all cowbird subspecies and populations from several regions showed no significant differences, though a trend that deserves further exploration was the pattern of lowest level in the ancestral population of cowbirds in the central prairies and of highest level in the northwestern population where range invasion occurred approximately 40 years ago. The levels of 17 betaestradiol were similar in the seven songbird species examined, which is consistent with current hypotheses that this hormone plays a role in embryonic sexual differentiation. Further investigation is needed to determine whether the large differences observed among species in absolute level of yolk testosterone are the relevant focal point or whether target tissue sensitivity differences mediate the effects of this yolk steroid, particularly between parasitic and non-parasitic species.
Shape matters: improved flight in tapered auto-rotating wings
NASA Astrophysics Data System (ADS)
Liu, Yucen; Vincent, Lionel; Kanso, Eva
2017-11-01
Many plants use gravity and wind to disperse their seeds. The shape of seed pods influence their aerodynamics. For example, Liana seeds form aerodynamic gliders and Sycamore trees release airborne ``helicopters.'' Here, we use carefully-controlled experiments and high-speed photography to examine dispersion by tumbling (auto-rotation) and we focus on the effect of geometry on flight characteristics. We consider four families of shapes: rectangular, elliptic, tapered, and sharp-tip wings, and we vary the span-to-chord ratio. We find that tapered wings exhibit extended flight time and range, that is, better performance. A quasi-steady two-dimensional model is used to highlight the mechanisms by which shape affects flight performance. These findings could have significant implications on linking seedpod designs to seed dispersion patterns as well as on optimizing wing design in active flight problems.
Design and development of flapping wing micro air vehicle
NASA Astrophysics Data System (ADS)
Hynes, N. Rajesh Jesudoss; Solomon, A. Jeffey Markus; Kathiresh, E.; Brighton, D.; Velu, P. Shenbaga
2018-05-01
Birds and insects have different methods of producing lift and thrust for hovering and forward flight. Most birds, however, cannot hover. Wing tips of birds follow simple paths in flight, whereas insects have very complicated wing tip paths, for hovering and forward flight, which vary with each species. FMAV based on avian flight. Development of Flapping Wing Air Vehicle (FWAV) is an on-going quest to master the natural flyers by mechanical means. It is characterized by unsteady aerodynamics, whose knowledge is still developing. The present work aims at include being capable of manoeuvring around and over obstacles by adjusting pitch, yaw, and roll, able to glide for five seconds under its own power, skilful at alternating between flapping and gliding with minimal disruption of flight pattern and being durable enough to withstand impacts with minimal to no damage.
Waliwitiya, Ranil; Belton, Peter; Nicholson, Russell A; Lowenberger, Carl A
2012-02-01
We evaluated the acute toxicities and the physiological effects of plant monoterpenoids (eugenol, pulegone, citronellal and alpha-terpineol) and neuroactive insecticides (malathion, dieldrin and RH3421) on flight muscle impulses (FMI) and wing beat signals (WBS) of the blow fly (Phaenicia sericata). Topically-applied eugenol, pulegone, citronellal, and alpha-terpineol produced neurotoxic symptoms, but were less toxic than malathion, dieldrin, or RH3421. Topical application of eugenol, pulegone, and citronellal reduced spike amplitude in one of the two banks of blow fly dorsolongitudinal flight muscles within 6-8 min, but with citronellal, the amplitude of FMIs reverted to a normal pattern within 1 hr. In contrast to pulegone and citronellal, where impulse frequency remained relatively constant, eugenol caused a gradual increase, then a decline in the frequency of spikes in each muscle bank. Wing beating was blocked permanently within 6-7 min of administering pulegone or citronellal and within 16 mins with eugenol. alpha-Terpineol-treated blow flies could not beat their wings despite normal FMI patterns. The actions of these monoterpenoids on blow fly flight motor patterns are discussed and compared with those of dieldrin, malathion, RH3421, and a variety of other neuroactive substances we have previously investigated in this system. Eugenol, pulegone and citronellal readily penetrate blow fly cuticle and interfere with flight muscle and/or central nervous function. Although there were differences in the effects of these compounds, they mainly depressed flight-associated responses, and acted similarly to compounds that block sodium channels and facilitate GABA action.
NASA Technical Reports Server (NTRS)
Krenkel, A. R.
1978-01-01
The finite-step method was programmed for computing the span loading and stability derivatives of trapezoidal shaped wings in symmetric, yawed, and rotary flight. Calculations were made for a series of different wing planforms and the results compared with several available methods for estimating these derivatives in the linear angle of attack range. The agreement shown was generally good except in a few cases. An attempt was made to estimate the nonlinear variation of lift with angle of attack in the high alpha range by introducing the measured airfoil section data into the finite-step method. The numerical procedure was found to be stable only at low angles of attack.
Experimental parametric studies of transonic T-tail flutter. [wind tunnel tests
NASA Technical Reports Server (NTRS)
Ruhlin, C. L.; Sandford, M. C.
1975-01-01
Wind-tunnel tests of the T-tail of a wide-body jet airplane were made at Mach numbers up to 1.02. The model consisted of a 1/13-size scaled version of the T-tail, fuselage, and inboard wing of the airplane. Two interchangeable T-tails were tested, one with design stiffness for flutter-clearance studies and one with reduced stiffness for flutter-trend studies. Transonic antisymmetric-flutter boundaries were determined for the models with variations in: (1) fin-spar stiffness, (2) stabilizer dihedral angle (-5 deg and 0 deg), (3) wing and forward-fuselage shape, and (4) nose shape of the fin-stabilizer juncture. A transonic symmetric-flutter boundary and flutter trends were established for variations in stabilizer pitch stiffness. Photographs of the test configurations are shown.
Rolling Moments Due to Rolling and Yaw for Four Wing Models in Rotation
NASA Technical Reports Server (NTRS)
Knight, Montgomery; Wenzinger, Carl J
1932-01-01
This report presents the results of a series of autorotation and torque tests on four different rotating wing systems at various rates of roll and at several angles of yaw. The investigation covered an angle of attack range up to 90 degrees and angles of yaw of 0 degree, 5 degrees, 10 degrees, and 20 degrees. The tests were made in a 5-foot, closed-throat atmospheric wind tunnel. The object of the tests was primarily to determine the effects of various angles of yaw on the rolling moments of the rotating wings up to large angles of attack. It was found that at angles of attack above that of maximum lift the rolling moments on the wings due to yaw (or side slip) from 5 degrees to 20 degrees were roughly of the same magnitude as those due to rolling. There was a wide variation in magnitude of the rolling moment due to yaw angle. The rates and ranges of stable autorotation for the monoplane models were considerably increased by yaw, whereas for an unstaggered biplane they were little affected. The immediate cause of the rolling moment due to yaw is apparently the building up of large loads on the forward wing tip and the reduction of loads on the rearward wing tip.
NASA Technical Reports Server (NTRS)
Chawla, Kalpana
1993-01-01
Attached as appendices to this report are documents describing work performed on the simulation of a landing powered-lift delta wing, the tracking of flow features using overset grids, and the simulation of flaps on the Wright Patterson Lab's fighter-lift-and-control (FLAC) wing. Numerical simulation of a powered-lift landing includes the computation of flow about a delta wing at four fixed heights as well as a simulated landing, in which the delta wing descends toward the ground. Comparison of computed and experimental lift coefficients indicates that the simulations capture the qualitative trends in lift-loss encountered by thrust-vectoring aircraft operating in ground effect. Power spectra of temporal variations of pressure indicate computed vortex shedding frequencies close to the jet exit are in the experimentally observed frequency range; the power spectra of pressure also provide insights into the mechanisms of lift oscillations. Also, a method for using overset grids to track dynamic flow features is described and the method is validated by tracking a moving shock and vortices shed behind a circular cylinder. Finally, Chimera gridding strategies were used to develop pressure coefficient contours for the FLAC wing for a Mach no. of 0.18 and Reynolds no. of 2.5 million.
Experiments on tip vortices interacting with downstream wings
NASA Astrophysics Data System (ADS)
Chen, C.; Wang, Z.; Gursul, I.
2018-05-01
The interaction of meandering tip vortices shed from a leading wing with a downstream wing was investigated experimentally in a water tunnel using flow visualization, particle image velocimetry measurements, and volumetric velocity measurements. Counter-rotating upstream vortices may exhibit sudden variations of the vortex core location when the wing-tip separation is within approximately twice the vortex core radius. This is caused by the formation of vortex dipoles near the wing tip. In contrast, co-rotating upstream vortices do not exhibit such sensitivity. Large spanwise displacement of the trajectory due to the image vortex is possible when the incident vortex is further inboard. For both co-rotating and counter-rotating vortices, as long as there is no direct impingement upon the wing, there is a little change in the structure of the time-averaged vortex past the wing, even though the tip vortex shed from the downstream wing may be substantially weakened or strengthened. In the absence of the downstream wing, as well as for weak interactions, the most energetic unsteady modes represent the first helical mode | m| = 1, which is estimated from the three-dimensional Proper Orthogonal Decomposition modes and has a very large wavelength, on the order of 102 times the vortex core radius, λ/ a = O(102). Instantaneous vorticity measurements as well as flow visualization suggest the existence of a smaller wavelength, λ/ a = 5-6, which is not among the most energetic modes. These two-orders of magnitude different wavelengths are in agreement with the previous measurements of tip vortices and also exhibit qualitative agreement with the transient energy growth analysis. The very long wavelength mode in the upstream vortex may persist during the interaction, and reveal coupling with the trailing vortex as well as increased meandering.
Confocal imaging of butterfly tissue.
Brunetti, Craig R
2014-01-01
To understand the molecular events responsible for morphological change requires the ability to examine gene expression in a wide range of organisms in addition to model systems to determine how the differences in gene expression correlate with phenotypic differences. There are approximately 12,000 species of butterflies, most, with distinct patterns on their wings. The most important tool for studying gene expression in butterflies is confocal imaging of butterfly tissue by indirect immunofluorescence using either cross-reactive antibodies from closely related species such as Drosophila or developing butterfly-specific antibodies. In this report, we describe how indirect immunofluorescence protocols can be used to visualize protein expression patterns on the butterfly wing imaginal disc and butterfly embryo.
Transcriptomic insights into the genetic basis of mammalian limb diversity.
Maier, Jennifer A; Rivas-Astroza, Marcelo; Deng, Jenny; Dowling, Anna; Oboikovitz, Paige; Cao, Xiaoyi; Behringer, Richard R; Cretekos, Chris J; Rasweiler, John J; Zhong, Sheng; Sears, Karen E
2017-03-23
From bat wings to whale flippers, limb diversification has been crucial to the evolutionary success of mammals. We performed the first transcriptome-wide study of limb development in multiple species to explore the hypothesis that mammalian limb diversification has proceeded through the differential expression of conserved shared genes, rather than by major changes to limb patterning. Specifically, we investigated the manner in which the expression of shared genes has evolved within and among mammalian species. We assembled and compared transcriptomes of bat, mouse, opossum, and pig fore- and hind limbs at the ridge, bud, and paddle stages of development. Results suggest that gene expression patterns exhibit larger variation among species during later than earlier stages of limb development, while within species results are more mixed. Consistent with the former, results also suggest that genes expressed at later developmental stages tend to have a younger evolutionary age than genes expressed at earlier stages. A suite of key limb-patterning genes was identified as being differentially expressed among the homologous limbs of all species. However, only a small subset of shared genes is differentially expressed in the fore- and hind limbs of all examined species. Similarly, a small subset of shared genes is differentially expressed within the fore- and hind limb of a single species and among the forelimbs of different species. Taken together, results of this study do not support the existence of a phylotypic period of limb development ending at chondrogenesis, but do support the hypothesis that the hierarchical nature of development translates into increasing variation among species as development progresses.
Otaki, Joji M
2011-06-01
Butterfly wing color patterns consist of many color-pattern elements such as eyespots. It is believed that eyespot patterns are determined by a concentration gradient of a single morphogen species released by diffusion from the prospective eyespot focus in conjunction with multiple thresholds in signal-receiving cells. As alternatives to this single-morphogen model, more flexible multiple-morphogen model and induction model can be proposed. However, the relevance of these conceptual models to actual eyespots has not been examined systematically. Here, representative eyespots from nymphalid butterflies were analyzed morphologically to determine if they are consistent with these models. Measurement of ring widths of serial eyespots from a single wing surface showed that the proportion of each ring in an eyespot is quite different among homologous rings of serial eyespots of different sizes. In asymmetric eyespots, each ring is distorted to varying degrees. In extreme cases, only a portion of rings is expressed remotely from the focus. Similarly, there are many eyespots where only certain rings are deleted, added, or expanded. In an unusual case, the central area of an eyespot is composed of multiple "miniature eyespots," but the overall macroscopic eyespot structure is maintained. These results indicate that each eyespot ring has independence and flexibility to a certain degree, which is less consistent with the single-morphogen model. Considering a "periodic eyespot", which has repeats of a set of rings, damage-induced eyespots in mutants, and a scale-size distribution pattern in an eyespot, the induction model is the least incompatible with the actual eyespot diversity.
Burger, Joanna; Gochfeld, Michael; Jeitner, Christian; Donio, Mark; Pittfield, Taryn
2014-01-01
A number of factors affect the consumption risk from mercury in fish, including mercury levels, seasonal patterns of mercury concentrations, human consumption patterns, and sensitive populations (e.g. pregnant women, fetuses, young children, and yet unknown genetic factors). Recently the protective effects of selenium on methylmercury toxicity have been publicized, particularly for saltwater fish. We examine levels of mercury and selenium in several species of fish and seabirds from the Aleutians (Alaska), determine selenium:mercury molar ratios, and examine species-specific and individual variation in the ratios as a means of exploring the use of the ratio in risk assessment and risk management. Variation among species was similar for mercury and selenium. There was significant inter-specific and intraspecific variation in selenium:mercury molar ratios for fish, and for birds. The mean selenium:mercury molar ratios for all fish and bird species were above 1, meaning there was an excess of selenium relative to mercury. It has been suggested that an excess of selenium confers some protective advantage for salt water fish, although the degree of excess necessary is unclear. The selenium:mercury molar ratio was significantly correlated negatively with total length for most fish species, but not for dolly varden. Some individuals of Pacific cod, yellow irish lord, rock greenling, Pacific halibut, dolly varden, and to a lesser extent, flathead sole, had selenium:mercury ratios below 1. No bird muscle had an excess of mercury (ratio below 1), and only glaucous-winged gull and pigeon guillemot had ratios between 1 and 5. There was a great deal of variation in selenium:mercury molar ratios within fish species, and within bird species, making it difficult and impractical to use these ratios in risk assessment or management, for fish advisories, or for consumers, particularly given the difficulty of interpreting the ratios. PMID:22664537
Infrared Imaging Of Flows Seeded With SF6
NASA Technical Reports Server (NTRS)
Manuel, Gregory S.; Daryabeigi, Kamran; Alderfer, David W.; Obara, Clifford J.
1993-01-01
Novel technique enables repeated measurements of flow patterns during flight. Wing-tip vorticity studied in flight by observing infrared emissions from SF6 gas entrained in wing-tip flow. System makes vortical flows visible throughout all altitude and speed ranges of all subsonic aircraft. Also useful for transonic and supersonic speeds. Primary application is testing of aircraft in flight, also proves useful in testing fast land vehicles and structures or devices subject to strong winds.
Measurements of Flow Rate and Trajectory of Aircraft Tire-Generated Water Spray
NASA Technical Reports Server (NTRS)
Daugherty, Robert H.; Stubbs, Sandy M.
1987-01-01
An experimental investigation was conducted at the NASA Langley Research Center to measure the flow rate and trajectory of water spray generated by an aircraft tire operating on a flooded runway. Tests were conducted in the Hydrodynamics Research Facility and made use of a partial airframe and a nose tire from a general aviation aircraft. Nose tires from a commercial transport aircraft were also used. The effects of forward speed, tire load, and water depth on water spray patterns were evaluated by measuring the amount and location of water captured by an array of tubes mounted behind the test tire. Water ejected from the side of the tire footprint had the most significant potential for ingestion into engine inlets. A lateral wake created on the water surface by the rolling tire can dominate the shape of the spray pattern as the distance aft of the tire is increased. Forward speed increased flow rates and moved the spray pattern inboard. Increased tire load caused the spray to become less dense. Near the tire, increased water depths caused flow rates to increase. Tests using a fuselage and partial wing along with the nose gear showed that for certain configurations, wing aerodynamics can cause a concentration of spray above the wing.
Evaluation of structural design concepts for an arrow-wing supersonic cruise aircraft
NASA Technical Reports Server (NTRS)
Sakata, I. F.; Davis, G. W.
1977-01-01
An analytical study was performed to determine the best structural approach for design of primary wing and fuselage structure of a Mach 2.7 arrow wing supersonic cruise aircraft. Concepts were evaluated considering near term start of design. Emphasis was placed on the complex interactions between thermal stress, static aeroelasticity, flutter, fatigue and fail safe design, static and dynamic loads, and the effects of variations in structural arrangements, concepts and materials on these interactions. Results indicate that a hybrid wing structure incorporating low profile convex beaded and honeycomb sandwich surface panels of titanium alloy 6Al-4V were the most efficient. The substructure includes titanium alloy spar caps reinforced with boron polyimide composites. The fuselage shell consists of hat stiffened skin and frame construction of titanium alloy 6Al-4V. A summary of the study effort is presented, and a discussion of the overall logic, design philosophy and interaction between the analytical methods for supersonic cruise aircraft design are included.
NASA Technical Reports Server (NTRS)
Bradley, D.; Buchholz, R. E.
1971-01-01
A 0.015 scale model of a modified version of the MDAC space shuttle booster was tested in the Naval Ship Research and Development Center 7 x 10 foot transonic wind tunnel, to obtain force, static stability, and control effectiveness data. Data were obtained for a cruise Mach Number of 0.38, altitude of 10,000 ft, and Reynolds Number per foot of approximately 2 x one million. The model was tested through an angle of attack range of -4 deg to 15 deg at zero degree angle of sideslip, and at an angle of sideslip range of -6 deg to 6 deg at fixed angles of attack of 0 deg, 6 deg, and 15 deg. Other test variables were elevon deflections, canard deflections, aileron deflections, rudder deflections, wing dihedral angle, canard incidence angle, wing incidence angle, canard position, wing position, wing and canard control flap size and dorsal fin size.
NASA Technical Reports Server (NTRS)
Parsons-Wingerter, Patricia A.; Weitzel, Alexander; Vyas, Ruchi J.; Murray, Matthew C.; Wyatt, Sarah E.
2016-01-01
One fundamental requirement shared by humans with all higher terrestrial life forms, including insect wings, higher land plants and other vertebrates, is a complex, fractally branching vascular system. NASA's VESsel GENeration Analysis (VESGEN) software maps and quantifies vascular trees, networks, and tree-network composites according to weighted physiological rules such as vessel connectivity, tapering and bifurcational branching. According to fluid dynamics, successful vascular transport requires a complex distributed system of highly regulated laminar flow. Microvascular branching rules within vertebrates, dicot leaves and the other organisms therefore display many similarities. One unifying perspective is that vascular patterning offers a useful readout that necessarily integrates complex molecular signaling pathways. VESGEN has elucidated changes in vascular pattern resulting from inflammatory, stress response, developmental and other signaling within numerous tissues and major model organisms studied for Space Biology. For a new VESGEN systems approach, we analyzed differential gene expression in leaves of Arabidopsis thaliana reported by GeneLab (GLDS-7) for spaceflight. Vascular-related changes in leaf gene expression were identified that can potentially be phenocopied by mutants in ground-based experiments. To link transcriptional, protein and other molecular change with phenotype, alterations in the Euclidean and dynamic dimensions (x,y,t) of vascular patterns for Arabidopsis leaves and other model species are being co-localized with signaling patterns of single molecular expression analyzed as information dimensions (i,j,k,...). Previously, Drosophila microarray data returned from space suggested significant changes in genes related to wing venation development that include EGF, Notch, Hedghog, Wingless and Dpp signaling. Phenotypes of increasingly abnormal ectopic wing venation in the (non-spaceflight) Drosophila wing generated by overexpression of a Notch antagonist were analyzed by VESGEN. Other VESGEN research applications include the mouse retina, GI and coronary vessels, avian placental analogs and translational studies in the astronaut retina related to health challenges for long-duration missions.
Imperfect chemical female mimicry in males of the ant Cardiocondyla obscurior
NASA Astrophysics Data System (ADS)
Cremer, Sylvia; D'Ettorre, Patrizia; Drijfhout, Falko P.; Sledge, Matthew F.; Turillazzi, Stefano; Heinze, Jürgen
2008-11-01
Winged and wingless males coexist in the ant Cardiocondyla obscurior. Wingless (“ergatoid”) males never leave their maternal colony and fight remorselessly among each other for the access to emerging females. The peaceful winged males disperse after about 10 days, but beforehand also mate in the nest. In the first 5 days of their life, winged males perform a chemical female mimicry that protects them against attack and even makes them sexually attractive to ergatoid males. When older, the chemical profile of winged males no longer matches that of virgin females; nevertheless, they are still tolerated, which so far has been puzzling. Contrasting this general pattern, we have identified a single aberrant colony in which all winged males were attacked and killed by the ergatoid males. A comparative analysis of the morphology and chemical profile of these untypical attacked winged males and the tolerated males from several normal colonies revealed that normal old males are still performing some chemical mimicry to the virgin queens, though less perfect than in their young ages. The anomalous attacked winged males, on the other hand, had a very different odour to the females. Our study thus exemplifies that the analysis of rare malfunctioning can add valuable insight on functioning under normal conditions and allows the conclusion that older winged males from normal colonies of the ant C. obscurior are guarded through an imperfect chemical female mimicry, still close enough to protect against attacks by the wingless fighters yet dissimilar enough not to elicit their sexual interest.
Thermo-orientation and the movement of feather-feeding lice on hosts.
Harbison, Christopher W; Boughton, Rachel M
2014-08-01
Temperature variation on the host is known to influence ectoparasite distributions. Ectoparasites may also use temperature gradients between host regions when moving on the host; however, tests are rare. Feather-feeding wing lice (Phthiraptera: Ischnocera) spend the majority of their time on the flight feathers of their avian hosts where they insert their bodies between feather barbs to escape host preening. However, because wing lice feed on downy abdominal feathers, they must repeatedly migrate between the flight feathers and body regions of their hosts. We performed a series of experiments that tested thermo-orientation in wing lice and evaluated its potential use during louse migrations between host regions. We found that wing lice can rapidly and accurately locate nearby heat targets that approximate host temperatures (37 C), demonstrating a capacity for directed thermo-orientation. We next tested the preference of wing lice for temperatures found along migration routes between bird flight feathers and their body regions. Wing lice could distinguish between temperatures found within distinct bird regions, and lice that had recently fed preferred the cooler temperatures (32 C), similar to those within bird flight feathers where they typically reside. However, when starved for 18-20 hr, wing lice shifted their preferences toward temperatures typical of bird body regions where they feed (36 C), demonstrating an ability to use thermal cues when moving between bird regions. We discuss the use of thermal cues during louse migration and microhabitat selection, as well as other potential impacts of thermo-orientation on host-parasite interactions.
Walker, Simon M.; Thomas, Adrian L. R.; Taylor, Graham K.
2009-01-01
Here, we present a detailed analysis of the wing kinematics and wing deformations of desert locusts (Schistocerca gregaria, Forskål) flying tethered in a wind tunnel. We filmed them using four high-speed digital video cameras, and used photogrammetry to reconstruct the motion of more than 100 identified points. Whereas the hindwing motions were highly stereotyped, the forewing motions showed considerable variation, consistent with a role in flight control. Both wings were positively cambered on the downstroke. The hindwing was cambered through an ‘umbrella effect’ whereby the trailing edge tension compressed the radial veins during the downstroke. Hindwing camber was reversed on the upstroke as the wing fan corrugated, reducing the projected area by 30 per cent, and releasing the tension in the trailing edge. Both the wings were strongly twisted from the root to the tip. The linear decrease in incidence along the hindwing on the downstroke precisely counteracts the linear increase in the angle of attack that would otherwise occur in root flapping for an untwisted wing. The consequent near-constant angle of attack is reminiscent of the optimum for a propeller of constant aerofoil section, wherein a linear twist distribution allows each section to operate at the unique angle of attack maximizing the lift to drag ratio. This implies tuning of the structural, morphological and kinematic parameters of the hindwing for efficient aerodynamic force production. PMID:19091683
Genome-wide introgression among distantly related Heliconius butterfly species.
Zhang, Wei; Dasmahapatra, Kanchon K; Mallet, James; Moreira, Gilson R P; Kronforst, Marcus R
2016-02-27
Although hybridization is thought to be relatively rare in animals, the raw genetic material introduced via introgression may play an important role in fueling adaptation and adaptive radiation. The butterfly genus Heliconius is an excellent system to study hybridization and introgression but most studies have focused on closely related species such as H. cydno and H. melpomene. Here we characterize genome-wide patterns of introgression between H. besckei, the only species with a red and yellow banded 'postman' wing pattern in the tiger-striped silvaniform clade, and co-mimetic H. melpomene nanna. We find a pronounced signature of putative introgression from H. melpomene into H. besckei in the genomic region upstream of the gene optix, known to control red wing patterning, suggesting adaptive introgression of wing pattern mimicry between these two distantly related species. At least 39 additional genomic regions show signals of introgression as strong or stronger than this mimicry locus. Gene flow has been on-going, with evidence of gene exchange at multiple time points, and bidirectional, moving from the melpomene to the silvaniform clade and vice versa. The history of gene exchange has also been complex, with contributions from multiple silvaniform species in addition to H. besckei. We also detect a signature of ancient introgression of the entire Z chromosome between the silvaniform and melpomene/cydno clades. Our study provides a genome-wide portrait of introgression between distantly related butterfly species. We further propose a comprehensive and efficient workflow for gene flow identification in genomic data sets.
The gust-mitigating potential of flapping wings.
Fisher, Alex; Ravi, Sridhar; Watkins, Simon; Watmuff, Jon; Wang, Chun; Liu, Hao; Petersen, Phred
2016-08-02
Nature's flapping-wing flyers are adept at negotiating highly turbulent flows across a wide range of scales. This is in part due to their ability to quickly detect and counterract disturbances to their flight path, but may also be assisted by an inherent aerodynamic property of flapping wings. In this study, we subject a mechanical flapping wing to replicated atmospheric turbulence across a range of flapping frequencies and turbulence intensities. By means of flow visualization and surface pressure measurements, we determine the salient effects of large-scale freestream turbulence on the flow field, and on the phase-average and fluctuating components of pressure and lift. It is shown that at lower flapping frequencies, turbulence dominates the instantaneous flow field, and the random fluctuating component of lift contributes significantly to the total lift. At higher flapping frequencies, kinematic forcing begins to dominate and the flow field becomes more consistent from cycle to cycle. Turbulence still modulates the flapping-induced flow field, as evidenced in particular by a variation in the timing and extent of leading edge vortex formation during the early downstroke. The random fluctuating component of lift contributes less to the total lift at these frequencies, providing evidence that flapping wings do indeed provide some inherent gust mitigation.
Wake structure and wing motion in bat flight
NASA Astrophysics Data System (ADS)
Hubel, Tatjana; Breuer, Kenneth; Swartz, Sharon
2008-11-01
We report on experiments concerning the wake structure and kinematics of bat flight, conducted in a low-speed wind tunnel using time-resolved PIV (200Hz) and 4 high-speed cameras to capture wake and wing motion simultaneously. 16 Lesser dog-faced fruit bats (C. brachyotis) were trained to fly in the wind tunnel at 3-6.5m/s. The PIV recordings perpendicular to the flow stream allowed observing the development of the tip vortex and circulation over the wing beat cycle. Each PIV acquisition sequence is correlated with the respective kinematic history. Circulation within wing beat cycles were often quite repeatable, however variations due to maneuvering of the bat are clearly visible. While no distinct vortex structure was observed at the upper reversal point (defined according the vertical motion of the wrist) a tip vortex was observed to develop in the first third of the downstroke, growing in strength, and persisting during much of the upstroke. Correlated to the presence of a strong tip vortex the circulation has almost constant strength over the middle half of the wing beat. At relatively low flight speeds (3.4 m/s), a closed vortex structure behind the bat is postulated.
Anderson, Christopher N; Grether, Gregory F
2010-02-22
In zones of sympatry between closely related species, species recognition errors in a competitive context can cause character displacement in agonistic signals and competitor recognition functions, just as species recognition errors in a mating context can cause character displacement in mating signals and mate recognition. These two processes are difficult to distinguish because the same traits can serve as both agonistic and mating signals. One solution is to test for sympatric shifts in recognition functions. We studied competitor recognition in Hetaerina damselflies by challenging territory holders with live tethered conspecific and heterospecific intruders. Heterospecific intruders elicited less aggression than conspecific intruders in species pairs with dissimilar wing coloration (H. occisa/H. titia, H. americana/H. titia) but not in species pairs with similar wing coloration (H. occisa/H. cruentata, H. americana/H. cruentata). Natural variation in the area of black wing pigmentation on H. titia intruders correlated negatively with heterospecific aggression. To directly examine the role of wing coloration, we blackened the wings of H. occisa or H. americana intruders and measured responses of conspecific territory holders. This treatment reduced territorial aggression at multiple sites where H. titia is present, but not at allopatric sites. These results provide strong evidence for agonistic character displacement.
Interactive flutter analysis and parametric study for conceptual wing design
NASA Technical Reports Server (NTRS)
Mukhopadhyay, Vivek
1995-01-01
An interactive computer program was developed for wing flutter analysis in the conceptual design stage. The objective was to estimate the flutter instability boundary of a flexible cantilever wing, when well defined structural and aerodynamic data are not available, and then study the effect of change in Mach number, dynamic pressure, torsional frequency, sweep, mass ratio, aspect ratio, taper ratio, center of gravity, and pitch inertia, to guide the development of the concept. The software was developed on MathCad (trademark) platform for Macintosh, with integrated documentation, graphics, database and symbolic mathematics. The analysis method was based on nondimensional parametric plots of two primary flutter parameters, namely Regier number and Flutter number, with normalization factors based on torsional stiffness, sweep, mass ratio, aspect ratio, center of gravity location and pitch inertia radius of gyration. The plots were compiled in a Vaught Corporation report from a vast database of past experiments and wind tunnel tests. The computer program was utilized for flutter analysis of the outer wing of a Blended Wing Body concept, proposed by McDonnell Douglas Corporation. Using a set of assumed data, preliminary flutter boundary and flutter dynamic pressure variation with altitude, Mach number and torsional stiffness were determined.
Theory of an airfoil equipped with a jet flap under low-speed flight conditions
NASA Technical Reports Server (NTRS)
Addessio, F. L.; Skifstad, J. G.
1975-01-01
A theory is developed, for the inviscid, incompressible flow past a thin airfoil equipped with a thin, part-span jet flap, by treating the induced flowfields of the jet and the wing separately and by obtaining the fully coupled solution in an iterative manner. Spanwise variation of the jet vortex strength is assumed to be elliptical in the analysis. Since the method considers the vorticity associated with the jet to be positioned on the computed locus of the jet, the downwash aft of the wing is evaluated as well as forces and moments on the wing. A lifting-surface theory is incorporated for the aerodynamics of the wing. Computational results are presented for a rectangular wing at momentum coefficients above 2.0 and compared with existing linear theories and experimental data. Good agreement is found for small angles of attack, jet-deflection angles, and jet-momentum coefficients where the linear theories and experimental data are applicable. Downwash data at a point in the vicinity of a control surface, the load distribution on the airfoil, and the jet, and the jet location are also presented for representative flight conditons.
Over-the-wing model thrust reverser noise tests
NASA Technical Reports Server (NTRS)
Goodykoontz, J.; Gutierrez, O.
1977-01-01
Static acoustic tests were conducted on a 1/12 scale model over-the-wing target type thrust reverser. The model configuration simulates a design that is applicable to the over-the-wing short-haul advanced technology engine. Aerodynamic screening tests of a variety of reverser designs identified configurations that satisfied a reverse thrust requirement of 35 percent of forward thrust at a nozzle pressure ratio of 1.29. The variations in the reverser configuration included, blocker door angle, blocker door lip angle and shape, and side skirt shape. Acoustic data are presented and compared for the various configurations. The model data scaled to a single full size engine show that peak free field perceived noise (PN) levels at a 152.4 meter sideline distance range from 98 to 104 PNdb.
Smart wing rotation and trailing-edge vortices enable high frequency mosquito flight
Bomphrey, Richard J.; Nakata, Toshiyuki; Phillips, Nathan; Walker, Simon M.
2017-01-01
Summary Mosquitoes exhibit unique wing kinematics; their long, slender wings flap at remarkably high frequencies for their size (>800 Hz) and with lower stroke amplitudes than any other insect group1. This shifts weight support away from the translation-dominated, aerodynamic mechanisms used by most insects2, as well as by helicopters and aeroplanes, towards poorly understood rotational mechanisms that occur when pitching at the end of each half-stroke. Here we report wing kinematics and solve the full Navier-Stokes equations using computational fluid dynamics with overset grids and validate our results with in vivo flow measurements. We show that, while familiar separated flow patterns are used by mosquitoes, much of the aerodynamic force that supports their weight is generated in a manner unlike any previously described flying animal. In total, there are three key features: leading-edge vortices (a well-known mechanism that appears to be almost ubiquitous in insect flight), trailing-edge vortices caused by a novel form of wake capture at stroke reversal, and rotational drag. The two new elements are largely independent of the wing velocity, instead relying on rapid changes in the pitch angle (wing rotation) at the end of each half stroke, and are therefore relatively immune to the shallow flapping amplitude. Moreover, these mechanisms are particularly well-suited to high-aspect ratio mosquito wings. PMID:28355184
Smart wing rotation and trailing-edge vortices enable high frequency mosquito flight
NASA Astrophysics Data System (ADS)
Bomphrey, Richard J.; Nakata, Toshiyuki; Phillips, Nathan; Walker, Simon M.
2017-03-01
Mosquitoes exhibit unusual wing kinematics; their long, slender wings flap at remarkably high frequencies for their size (>800 Hz)and with lower stroke amplitudes than any other insect group. This shifts weight support away from the translation-dominated, aerodynamic mechanisms used by most insects, as well as by helicopters and aeroplanes, towards poorly understood rotational mechanisms that occur when pitching at the end of each half-stroke. Here we report free-flight mosquito wing kinematics, solve the full Navier-Stokes equations using computational fluid dynamics with overset grids, and validate our results with in vivo flow measurements. We show that, although mosquitoes use familiar separated flow patterns, much of the aerodynamic force that supports their weight is generated in a manner unlike any previously described for a flying animal. There are three key features: leading-edge vortices (a well-known mechanism that appears to be almost ubiquitous in insect flight), trailing-edge vortices caused by a form of wake capture at stroke reversal, and rotational drag. The two new elements are largely independent of the wing velocity, instead relying on rapid changes in the pitch angle (wing rotation) at the end of each half-stroke, and they are therefore relatively immune to the shallow flapping amplitude. Moreover, these mechanisms are particularly well suited to high aspect ratio mosquito wings.
NASA Technical Reports Server (NTRS)
Mineck, Raymond E.; Vijgen, Paul M. H. W.
1993-01-01
Three planar, untwisted wings with the same elliptical chord distribution but with different curvatures of the quarter-chord line were tested in the Langley 8-Foot Transonic Pressure Tunnel (8-ft TPT) and the Langley 7- by 10-Foot High-Speed Tunnel (7 x 10 HST). A fourth wing with a rectangular planform and the same projected area and span was also tested. Force and moment measurements from the 8-ft TPT tests are presented for Mach numbers from 0.3 to 0.5 and angles of attack from -4 degrees to 7 degrees. Sketches of the oil-flow patterns on the upper surfaces of the wings and some force and moment measurements from the 7 x 10 HST tests are presented at a Mach number of 0.5. Increasing the curvature of the quarter-chord line makes the angle of zero lift more negative but has little effect on the drag coefficient at zero lift. The changes in lift-curve slope and in the Oswald efficiency factor with the change in curvature of the quarter-chord line (wingtip location) indicate that the elliptical wing with the unswept quarter-chord line has the lowest lifting efficiency and the elliptical wing with the unswept trailing edge has the highest lifting efficiency; the crescent-shaped planform wing has an efficiency in between.
Shang, Feng; Ding, Bi-Yue; Xiong, Ying; Dou, Wei; Wei, Dong; Jiang, Hong-Bo; Wei, Dan-Dan; Wang, Jin-Jun
2016-01-01
Winged and wingless morphs in insects represent a trade-off between dispersal ability and reproduction. We studied key genes associated with apterous and alate morphs in Toxoptera citricida (Kirkaldy) using RNAseq, digital gene expression (DGE) profiling, and RNA interference. The de novo assembly of the transcriptome was obtained through Illumina short-read sequencing technology. A total of 44,199 unigenes were generated and 27,640 were annotated. The transcriptomic differences between alate and apterous adults indicated that 279 unigenes were highly expressed in alate adults, whereas 5,470 were expressed at low levels. Expression patterns of the top 10 highly expressed genes in alate adults agreed with wing bud development trends. Silencing of the lipid synthesis and degradation gene (3-ketoacyl-CoA thiolase, mitochondrial-like) and glycogen genes (Phosphoenolpyruvate carboxykinase [GTP]-like and Glycogen phosphorylase-like isoform 2) resulted in underdeveloped wings. This suggests that both lipid and glycogen metabolism provide energy for aphid wing development. The large number of sequences and expression data produced from the transcriptome and DGE sequencing, respectively, increases our understanding of wing development mechanisms. PMID:27577531
ERIC Educational Resources Information Center
Malone, Molly
2012-01-01
Most middle school students comprehend that organisms have adaptations that enable their survival and that successful adaptations prevail in a population over time. Yet they often miss that those bird beaks, moth-wing colors, or whatever traits are the result of random, normal genetic variations that just happen to confer a negative, neutral, or…
NASA Technical Reports Server (NTRS)
Kukreja, Sunil L.; Vio, Gareth A.; Andrianne, Thomas; azak, Norizham Abudl; Dimitriadis, Grigorios
2012-01-01
The stall flutter response of a rectangular wing in a low speed wind tunnel is modelled using a nonlinear difference equation description. Static and dynamic tests are used to select a suitable model structure and basis function. Bifurcation criteria such as the Hopf condition and vibration amplitude variation with airspeed were used to ensure the model was representative of experimentally measured stall flutter phenomena. Dynamic test data were used to estimate model parameters and estimate an approximate basis function.
Transonic Reynolds Number and Leading-Edge Bluntness Effects on a 65 deg Delta Wing
NASA Technical Reports Server (NTRS)
Luckring, J. M.
2003-01-01
A 65 degree delta wing has been tested in the National Transonic Facility (NTF) at mean aerodynamic chord Reynolds numbers from 6 million to 120 million at subsonic and transonic speeds. The configuration incorporated a systematic variation of the leading edge bluntness. The analysis for this paper is focused on the Reynolds number and bluntness effects at transonic speeds (M = 0.85) from this data set. The results show significant effects of both these parameters on the onset and progression of leading edge vortex separation.
Reynolds Number and Leading-Edge Bluntness Effects on a 65 Deg Delta Wing
NASA Technical Reports Server (NTRS)
Luckring, J. M.
2002-01-01
A 65 deg delta wing has been tested in the National Transonic Facility (NTF) at mean aerodynamic chord Reynolds numbers from 6 million to 120 million at subsonic and transonic speeds. The configuration incorporated systematic variation of the leading edge bluntness. The analysis for this paper is focused on the Reynolds number and bluntness effects at subsonic speeds (M = 0.4) from this data set. The results show significant effects of both these parameters on the onset and progression of leading-edge vortex separation.
Transonic Reynolds Number and Leading-Edge Bluntness Effects on a 65 deg Delta Wing
NASA Technical Reports Server (NTRS)
Luckring, J. M.
2003-01-01
A 65 deg delta wing has been tested in the National Transonic Facility (NTF) at mean aerodynamic chord Reynolds numbers from 6 million to 120 million at subsonic and transonic speeds. The configuration incorporated a systematic variation of the leading edge bluntness. The analysis for this paper is focused on the Reynolds number and bluntness effects at transonic speeds (M = 0.85) from this data set. The results show significant effects of both these parameters on the onset and progression of leading- edge vortex separation.
Reynolds Number and Leading-Edge Bluntness Effects on a 65 deg Delta Wing
NASA Technical Reports Server (NTRS)
Luckring, J. M.
2002-01-01
A 65 degree delta wing has been tested in the National Transonic Facility (NTF) at mean aerodynamic chord Reynolds numbers from 6 million to 120 million at subsonic and transonic speeds. The configuration incorporated systematic variation of the leading edge bluntness. The analysis for this paper is focused on the Reynolds number and bluntness effects at subsonic speeds (M = 0.4) from this data set. The results show significant effects of both these parameters on the onset and progression of leading-edge vortex separation.
Transonic Reynolds Number and Leading-Edge Bluntness Effects on a 65 deg Delta Wing
NASA Technical Reports Server (NTRS)
Luckring, J. M.
2003-01-01
A 65 deg delta wing has been tested in the National Transonic Facility (NTF) at mean aerodynamic chord Reynolds numbers from 6 million to 120 million at subsonic and transonic speeds. The configuration incorporated a systematic variation of the leading edge bluntness. The analysis for this paper is focused on the Reynolds number and bluntness effects at transonic speeds (M=0.85) from this data set. The results show significant effects of both these parameters on the onset and progression of leading-edge vortex separation.
The Iconography and Symbolism of Sun God in Urartian Art
NASA Astrophysics Data System (ADS)
Poghosyan, Gayane
2016-12-01
The predominating symbol of the winged sun disc in Urartian religious iconography testifies the significant role and importance of the sun in worship. The stylistic variation and peculiar iconographic features of the winged discs, sacred animals and divine images associated with solar deity shows the relationship between the cult of the sun god, sequence of the different phases of the year and constellations in Urartian culture. Such kind of iconography is possibly formed and stylized in result of interaction of ancient human imaginations, influence of rock paintings and religious beliefs.
A Wingless and Notch double-repression mechanism regulates G1–S transition in the Drosophila wing
Herranz, Héctor; Pérez, Lidia; Martín, Francisco A; Milán, Marco
2008-01-01
The control of tissue growth and patterning is orchestrated in various multicellular tissues by the coordinated activity of the signalling molecules Wnt/Wingless (Wg) and Notch, and mutations in these pathways can cause cancer. The role of these molecules in the control of cell proliferation and the crosstalk between their corresponding pathways remain poorly understood. Crosstalk between Notch and Wg has been proposed to organize pattern and growth in the Drosophila wing primordium. Here we report that Wg and Notch act in a surprisingly linear pathway to control G1–S progression. We present evidence that these molecules exert their function by regulating the expression of the dmyc proto-oncogene and the bantam micro-RNA, which positively modulated the activity of the E2F transcription factor. Our results demonstrate that Notch acts in this cellular context as a repressor of cell-cycle progression and Wg has a permissive role in alleviating Notch-mediated repression of G1–S progression in wing cells. PMID:18451803
Environmental heterogeneity, dispersal mode, and co-occurrence in stream macroinvertebrates
Heino, Jani
2013-01-01
Both environmental heterogeneity and mode of dispersal may affect species co-occurrence in metacommunities. Aquatic invertebrates were sampled in 20–30 streams in each of three drainage basins, differing considerably in environmental heterogeneity. Each drainage basin was further divided into two equally sized sets of sites, again differing profoundly in environmental heterogeneity. Benthic invertebrate data were divided into three groups of taxa based on overland dispersal modes: passive dispersers with aquatic adults, passive dispersers with terrestrial winged adults, and active dispersers with terrestrial winged adults. The co-occurrence of taxa in each dispersal mode group, drainage basin, and heterogeneity site subset was measured using the C-score and its standardized effect size. The probability of finding high levels of species segregation tended to increase with environmental heterogeneity across the drainage basins. These patterns were, however, contingent on both dispersal mode and drainage basin. It thus appears that environmental heterogeneity and dispersal mode interact in affecting co-occurrence in metacommunities, with passive dispersers with aquatic adults showing random patterns irrespective of environmental heterogeneity, and active dispersers with terrestrial winged adults showing increasing segregation with increasing environmental heterogeneity. PMID:23467653
Saison, Tamar; Peroz, Christophe; Chauveau, Vanessa; Berthier, Serge; Sondergard, Elin; Arribart, Hervé
2008-12-01
An original and low cost method for the fabrication of patterned surfaces bioinspired from butterfly wings and lotus leaves is presented. Silica-based sol-gel films are thermally imprinted from elastomeric molds to produce stable structures with superhydrophobicity values as high as 160 degrees water contact angle. The biomimetic surfaces are demonstrated to be tuned from superhydrophobic to superhydrophilic by annealing between 200 degrees C and 500 degrees C.
Carr, David M; Ellsworth, Ashley A; Fisher, Gregory L; Valeriano, Wescley W; Vasco, Juan P; Guimarães, Paulo S S; de Andrade, Rodrigo R; da Silva, Elizabeth R; Rodrigues, Wagner N
2018-02-05
The iridescent wings of the Chalcopterix rutilans damselfly (Rambur) (Odonata, Polythoridae) are investigated with focused ion beam/scanning electron microscopy, transmission electron microscopy, and time-of-flight secondary ion mass spectrometry. The electron microscopy images reveal a natural photonic crystal as the source of the varying colors. The photonic crystal has a consistent number and thickness (∼195 nm) of the repeat units on the ventral side of the wing, which is consistent with the red color visible from the bottom side of the wing in all regions. The dorsal side of the wing shows strong color variations ranging from red to blue depending on the region. In the electron microscopy images, the dorsal side of the wing exhibits varied number and thicknesses of the repeat units. The repeat unit spacings for the red, yellow/green, and blue regions are approximately 195, 180, and 145 nm, respectively. Three-dimensional analysis of the natural photonic crystals by time-of-flight secondary ion mass spectrometry reveals that changes in the relative levels of Na, K, and eumelanin are responsible for the varying dielectric constant needed to generate the photonic crystal. The photonic crystal also appears to be assembled with a chemical tricomponent layer structure due to the enhancement of the CH 6 N 3 + species at every other interface between the high/low dielectric constant layers.
Habitat assessment, Missouri River at Hermann, Missouri
Jacobson, Robert B.; Laustrup, Mark S.; Reuter, Joanna M.
2002-01-01
This report documents methods and results of aquatic habitat assessment in the Missouri River near Hermann, Missouri. The assessment is intended to improve understanding of spatial and temporal variability of aquatic habitat, including habitats thought to be critical for the endangered pallid sturgeon (Scaphirhynchus albus). Physical aquatic habitat - depth, velocity, and substrate - was assessed around 9 wing dikes and adjacent to the U.S. Route 19 bridge, at discharges varying from 44,000 cubic feet per second (cfs) to 146, 000 cfs during August 2000-May, 2001. For the river as a whole, velocities are bi-modally distributed with distinct peaks relating to navigation channel and wing-dike environments. Velocities predictably showed an increasing trend with increasing discharge. Substrate within wing dikes was dominated by mud at low discharges, whereas the navigation channel had patches of transporting sand, rippled sand, and coarse sand. Discharges that overtopped the wing dikes (about 93,000 cfs, March 2001) were associated with increases of patchy sand, rippled sand, and coarse sand within the wing dikes. When flows were substantially over the wing dikes (146,000 cfs, May 2001) substrates within most wing dikes showed substantial reorganization and coarsening. The habitat assessment provides a geospatial database that can be used to query wing dikes for distributions of depth, velocity, and substrate for comparison with fish samples collected by US Fish and Wildlife Service biologists (Grady and others, 2001). In addition, the assessment documented spatial and temporal variation in habitat within the Hermann reach and over a range of discharges. Measurable geomorphic change--alteration of substrate conditions plus substantial erosion and deposition--was associated with flows equaled or exceeded 12-40% of the time (40-140 days per year). Documented geomorphic change associated with high-frequency flows underscores the natural temporal variability of physical habitat in the Lower Missouri River.
Román, Jessica K; Walsh, Callee M; Oh, Junho; Dana, Catherine E; Hong, Sungmin; Jo, Kyoo D; Alleyne, Marianne; Miljkovic, Nenad; Cropek, Donald M
2018-03-01
Laser-ablation electrospray ionization (LAESI) imaging mass spectrometry (IMS) is an emerging bioanalytical tool for direct imaging and analysis of biological tissues. Performing ionization in an ambient environment, this technique requires little sample preparation and no additional matrix, and can be performed on natural, uneven surfaces. When combined with optical microscopy, the investigation of biological samples by LAESI allows for spatially resolved compositional analysis. We demonstrate here the applicability of LAESI-IMS for the chemical analysis of thin, desiccated biological samples, specifically Neotibicen pruinosus cicada wings. Positive-ion LAESI-IMS accurate ion-map data was acquired from several wing cells and superimposed onto optical images allowing for compositional comparisons across areas of the wing. Various putative chemical identifications were made indicating the presence of hydrocarbons, lipids/esters, amines/amides, and sulfonated/phosphorylated compounds. With the spatial resolution capability, surprising chemical distribution patterns were observed across the cicada wing, which may assist in correlating trends in surface properties with chemical distribution. Observed ions were either (1) equally dispersed across the wing, (2) more concentrated closer to the body of the insect (proximal end), or (3) more concentrated toward the tip of the wing (distal end). These findings demonstrate LAESI-IMS as a tool for the acquisition of spatially resolved chemical information from fragile, dried insect wings. This LAESI-IMS technique has important implications for the study of functional biomaterials, where understanding the correlation between chemical composition, physical structure, and biological function is critical. Graphical abstract Positive-ion laser-ablation electrospray ionization mass spectrometry coupled with optical imaging provides a powerful tool for the spatially resolved chemical analysis of cicada wings.
NASA Astrophysics Data System (ADS)
Khan, Mahasin Ali; Bera, Subir
2017-07-01
The occurrences of Pinus L. (family Pinaceae) megafossils (cones and leaf remains) have been abundantly documented from the Cenozoic sediments of eastern Asia (Japan and China), but none has been confirmed from the Indian Cenozoic till date. Here, we describe Pinus arunachalensis Khan and Bera, sp. nov. on the basis of seed remains from the middle to late Miocene Siwalik sediments of the Dafla Formation exposed around West Kameng district in Arunachal Pradesh, eastern Himalaya. Seeds are winged, broadly oblong to oval in outline, 1.3-1.5 cm long and 0.4-0.6 cm broad (in the middle part), located basipetally and symmetrically to wing, cellular pattern of wing is seemingly undulatory and parallel with the long axis of the wing. So far, this report provides the first ever fossil record of Pinus winged seeds from India. This record suggests that Pinus was an important component of tropical-subtropical evergreen forest in the area during the Miocene and this group subsequently declined from the local vegetation probably because of the gradual intensification of MSI (monsoon index) from the Miocene to the present. We also review the historical phytogeography and highlight the phytogeographic implication of this genus.
Quinn, Lucy R; Meharg, Andrew A; van Franeker, Jan A; Graham, Isla M; Thompson, Paul M
Many wildlife studies use chemical analyses to explore spatio-temporal variation in diet, migratory patterns and contaminant exposure. Intrinsic markers are particularly valuable for studying non-breeding marine predators, when direct methods of investigation are rarely feasible. However, any inferences regarding foraging ecology are dependent upon the time scale over which tissues such as feathers are formed. In this study, we validate the use of body feathers for studying non-breeding foraging patterns in a pelagic seabird, the northern fulmar. Analysis of carcasses of successfully breeding adult fulmars indicated that body feathers moulted between September and March, whereas analyses of carcasses and activity patterns suggested that wing feather and tail feather moult occurred during more restricted periods (September to October and September to January, respectively). By randomly sampling relevant body feathers, average values for individual birds were shown to be consistent. We also integrated chemical analyses of body feather with geolocation tracking data to demonstrate that analyses of δ 13 C and δ 15 N values successfully assigned 88 % of birds to one of two broad wintering regions used by breeding adult fulmars from a Scottish study colony. These data provide strong support for the use of body feathers as a tool for exploring non-breeding foraging patterns and diet in wide-ranging, pelagic seabirds.
Bédard, Emilie; Lévesque, Simon; Martin, Philippe; Pinsonneault, Linda; Paranjape, Kiran; Lalancette, Cindy; Dolcé, Charles-Éric; Villion, Manuela; Valiquette, Louis; Faucher, Sébastien P; Prévost, Michèle
2016-12-01
OBJECTIVE To determine the source of a Legionella pneumophila serogroup 5 nosocomial outbreak and the role of the heat exchanger installed on the hot water system within the previous year. SETTING A 400-bed tertiary care university hospital in Sherbrooke, Canada. METHODS Hot water samples were collected and cultured for L. pneumophila from 25 taps (baths and sinks) within wing A and 9 taps in wing B. Biofilm (5) and 2 L water samples (3) were collected within the heat exchangers for L. pneumophila culture and detection of protists. Sequence-based typing was performed on strain DNA extracts and pulsed-field gel electrophoresis patterns were analyzed. RESULTS Following 2 cases of hospital-acquired legionellosis, the hot water system investigation revealed a large proportion of L. pneumophila serogroup 5 positive taps (22/25 in wing A and 5/9 in wing B). High positivity was also detected in the heat exchanger of wing A in water samples (3/3) and swabs from the heat exchanger (4/5). The outbreak genotyping investigation identified the hot water system as the source of infections. Genotyping results revealed that all isolated environmental strains harbored the same related pulsed-field gel electrophoresis pattern and sequence-based type. CONCLUSIONS Two cases of hospital-acquired legionellosis occurred in the year following the installation of a heat exchanger to preheat hospital hot water. No cases were reported previously, although the same L. pneumophila strain was isolated from the hot water system in 1995. The heat exchanger promoted L. pneumophila growth and may have contributed to confirmed clinical cases. Infect. Control Hosp. Epidemiol. 2016;1475-1480.
Flow structures around a flapping wing considering ground effect
NASA Astrophysics Data System (ADS)
Van Truong, Tien; Kim, Jihoon; Kim, Min Jun; Park, Hoon Cheol; Yoon, Kwang Joon; Byun, Doyoung
2013-07-01
Over the past several decades, there has been great interest in understanding the aerodynamics of flapping flight, namely the two flight modes of hovering and forward flight. However, there has been little focus on the aerodynamic characteristics during takeoff of insects. In a previous study we found that the Rhinoceros Beetle ( Trypoxylusdichotomus) takes off without jumping, which is uncommon for other insects. In this study we built a scaled-up electromechanical model of a flapping wing and investigated fluid flow around the beetle's wing model. In particular, the present dynamically scaled mechanical model has the wing kinematics pattern achieved from the real beetle's wing kinematics during takeoff. In addition, we could systematically change the three-dimensional inclined motion of the flapping model through each stroke. We used digital particle image velocimetry with high spatial resolution, and were able to qualitatively and quantitatively study the flow field around the wing at a Reynolds number of approximately 10,000. The present results provide insight into the aerodynamics and the evolution of vortical structures, as well as the ground effect experienced by a beetle's wing during takeoff. The main unsteady mechanisms of beetles have been identified and intensively analyzed as the stability of the leading edge vortex (LEV) during strokes, the delayed stall during upstroke, the rotational circulation in pronation periods, and wake capture in supination periods. Due to the ground effect, the LEV was enhanced during half downstroke, and the lift force could thus be increased to lift the beetle during takeoff. This is useful for researchers in developing a micro air vehicle that has a beetle-like flapping wing motion.
Simons, Erin L R; O'connor, Patrick M
2012-03-01
Wing bone histology in three species of birds was characterized in order to test hypotheses related to the relationship between skeletal microstructure and inferred wing loading during flight. Data on the degree of laminarity (the proportion of circular vascular canals) and the occurrence of secondary osteons were obtained from three species that utilize different primary flight modes: the Double-crested cormorant, a continuous flapper; the Brown pelican, a static soarer; and the Laysan albatross, a dynamic soarer. Laminarity indices were calculated for four quadrants for each of the three main wing elements. Ulnae and carpometacarpi were predicted to exhibit quadrant specific patterns of laminarity due to hypothesized differences in locally applied loads related to the attachment of flight feathers. However, few differences among the quadrants were identified. No significant differences were identified among the three elements, which is notable as different bones are likely experiencing different loading conditions. These results do not support the concept of bone functional adaptation in the primary structure of the wing elements. Significant differences in laminarity were found among the three primary flight modes. The dynamic soaring birds exhibited significantly lower laminarity than the flapping and static soaring birds. These results support the proposed hypothesis that laminarity is an adaptation for resisting torsional loading. This may be explained by overall wing shape: whereas dynamic soaring birds have long slender wings, flappers and static soaring birds have broader wings with a larger wing chord that would necessarily impart a higher torsional moment on the feather-bearing bones. Copyright © 2012 Wiley Periodicals, Inc.
Vortex Flap Technology: a Stability and Control Assessment
NASA Technical Reports Server (NTRS)
Carey, K. M.; Erickson, G. E.
1984-01-01
A comprehensive low-speed wind tunnel investigation was performed of leading edge vortex flaps applied to representative aircraft configurations. A determination was made of the effects of analytically- and empirically-designed vortex flaps on the static longitudinal and lateral-directional aerodynamics, stability, and control characteristics of fighter wings having leading-edge sweep angles of 45 to 76.5 degrees. The sensitivity to several configuration modifications was assessed, which included the effects of flap planform, leading- and trailing-edge flap deflection angles, wing location on the fuselage, forebody strakes, canards, and centerline and outboard vertical tails. Six-component forces and moments, wing surface static pressure distributions, and surface flow patterns were obtained using the Northrop 21- by 30-inch low-speed wind tunnel.
NASA Astrophysics Data System (ADS)
Faber, Jakob A.; Arrieta, Andres F.; Studart, André R.
2018-03-01
Origami enables folding of objects into a variety of shapes in arts, engineering, and biological systems. In contrast to well-known paper-folded objects, the wing of the earwig has an exquisite natural folding system that cannot be sufficiently described by current origami models. Such an unusual biological system displays incompatible folding patterns, remains open by a bistable locking mechanism during flight, and self-folds rapidly without muscular actuation. We show that these notable functionalities arise from the protein-rich joints of the earwig wing, which work as extensional and rotational springs between facets. Inspired by this biological wing, we establish a spring origami model that broadens the folding design space of traditional origami and allows for the fabrication of precisely tunable, four-dimensional–printed objects with programmable bioinspired morphing functionalities.
On the quasi-steady aerodynamics of normal hovering flight part I: the induced power factor
Nabawy, Mostafa R. A.; Crowther, William J.
2014-01-01
An analytical treatment to quantify the losses captured in the induced power factor, k, is provided for flapping wings in normal hover, including the effects of non-uniform downwash, tip losses and finite flapping amplitude. The method is based on a novel combination of actuator disc and lifting line blade theories that also takes into account the effect of advance ratio. The model has been evaluated against experimental results from the literature and qualitative agreement obtained for the effect of advance ratio on the lift coefficient of revolving wings. Comparison with quantitative experimental data for the circulation as a function of span for a fruitfly wing shows that the model is able to correctly predict the circulation shape of variation, including both the magnitude of the peak circulation and the rate of decay in circulation towards zero. An evaluation of the contributions to induced power factor in normal hover for eight insects is provided. It is also shown how Reynolds number can be accounted for in the induced power factor, and good agreement is obtained between predicted span efficiency as a function of Reynolds number and numerical results from the literature. Lastly, it is shown that for a flapping wing in hover k owing to the non-uniform downwash effect can be reduced to 1.02 using an arcsech chord distribution. For morphologically realistic wing shapes based on beta distributions, it is shown that a value of 1.07 can be achieved for a radius of first moment of wing area at 40% of wing length. PMID:24522785
On the quasi-steady aerodynamics of normal hovering flight part I: the induced power factor.
Nabawy, Mostafa R A; Crowther, William J
2014-04-06
An analytical treatment to quantify the losses captured in the induced power factor, k, is provided for flapping wings in normal hover, including the effects of non-uniform downwash, tip losses and finite flapping amplitude. The method is based on a novel combination of actuator disc and lifting line blade theories that also takes into account the effect of advance ratio. The model has been evaluated against experimental results from the literature and qualitative agreement obtained for the effect of advance ratio on the lift coefficient of revolving wings. Comparison with quantitative experimental data for the circulation as a function of span for a fruitfly wing shows that the model is able to correctly predict the circulation shape of variation, including both the magnitude of the peak circulation and the rate of decay in circulation towards zero. An evaluation of the contributions to induced power factor in normal hover for eight insects is provided. It is also shown how Reynolds number can be accounted for in the induced power factor, and good agreement is obtained between predicted span efficiency as a function of Reynolds number and numerical results from the literature. Lastly, it is shown that for a flapping wing in hover k owing to the non-uniform downwash effect can be reduced to 1.02 using an arcsech chord distribution. For morphologically realistic wing shapes based on beta distributions, it is shown that a value of 1.07 can be achieved for a radius of first moment of wing area at 40% of wing length.
Stephens, C. R.; Juliano, S. A.
2012-01-01
Estimating a mosquito’s vector competence, or likelihood of transmitting disease, if it takes an infectious blood meal, is an important aspect of predicting when and where outbreaks of infectious diseases will occur. Vector competence can be affected by rearing temperature and inter- and intraspecific competition experienced by the individual mosquito during its larval development. This research investigates whether a new morphological indicator of larval rearing conditions, wing shape, can be used to distinguish reliably temperature and competitive conditions experienced during larval stages. Aedes albopictus and Aedes aegypti larvae were reared in low intra-specific, high intra-specific, or high inter-specific competition treatments at either 22°C or 32°C. The right wing of each dried female was removed and photographed. Nineteen landmarks and twenty semilandmarks were digitized on each wing. Shape variables were calculated using geometric morphometric software. Canonical variate analysis, randomization multivariate analysis of variance, and visualization of landmark movement using deformation grids provided evidence that although semilandmark position was significantly affected by larval competition and temperature for both species, the differences in position did not translate into differences in wing shape, as shown in deformation grids. Two classification procedures yielded success rates of 26–49%. Accounting for wing size produced no increase in classification success. There appeared to be a significant relationship between shape and size. These results, particularly the low success rate of classification based on wing shape, show that shape is unlikely to be a reliable indicator of larval rearing competition and temperature conditions for Aedes albopictus and Aedes aegypti. PMID:22897054
Body molt of male long-tailed ducks in the nearshore waters of the north slope, Alaska
Howell, M.D.; Grand, J.B.; Flint, Paul L.
2003-01-01
We examined the timing and intensity of body molt in relation to stage of remige growth for postbreeding adult male Long-tailed Ducks (Clangula hyemalis) off the coast of northern Alaska. During this period, remige and rectrix feathers are molted simultaneously with body feathers during the prebasic molt, which results in a period of increased energetic and nutritional demands. We collected birds from late July through mid-August and recorded intensity of molt in eight regions: head and neck, back and rump, greater coverts, lesser coverts, flank and sides, breast, belly, and tail. Using nonlinear regression, we estimated the peak intensity and variation for each region in relation to ninth primary length. We found little evidence of molt in the head and neck region. The greater and lesser coverts, and back and rump reached peak molt intensities earliest and were followed by tail, breast, and belly. Molt intensity in the flank and side region was highly variable and indicated a more prolonged molting pattern in relation to other regions. While body molt occurs simultaneously with wing molt, we found that molt among regions occurred in a staggered pattern. Long-tailed Ducks may employ this staggered molting pattern to minimize the energetic and nutritional requirements of molt.
Proprioceptive input patterns elevator activity in the locust flight system.
Wolf, H; Pearson, K G
1988-06-01
1. In the locust, Locusta migratoria, the roles of two groups of wing sense organs, hind wing tegulae and wing-hinge stretch receptors, in the generation of the flight motor pattern were investigated. A preparation was employed that allowed the intracellular recording of neural activity in almost intact tethered flying locusts or after selective manipulations of sensory input. The functions of the two sets of receptors were assessed 1) by studying the phases of their discharges in the wingbeat cycle (Fig. 3), 2) by the selective ablation of input from the receptors (Figs. 4-7), and 3) by the selective stimulation of the receptor afferents (Figs. 8-12). 2. Input from the tegulae was found to be responsible for the initiation of elevator activity (Figs. 9 and 10) and for the generation of a distinct initial rapid depolarization (Figs. 4, 5, and 8) characteristic of elevator motor neuron activity in intact locusts (Figs. 1 and 16). 3. Input from the wing-hinge stretch receptors was found to control the duration of elevator depolarizations by the graded suppression of a second late component of the elevator depolarizations as wingbeat frequency increased (Figs. 6, 7, 11, and 12). The characteristics of this late component of elevator activity suggested that it is generated by the same (central nervous) mechanism that produces the elevator depolarizations recorded in deafferented animals (Fig. 2). Apparently this late component contributes to the intact pattern of elevator depolarizations only at lower wingbeat frequencies and is abolished by the action of stretch-receptor input at frequencies above approximately 15 Hz (Figs. 1, 2, and 4). At these high wingbeat frequencies elevator activity is dominated by the rapid depolarizations generated as a result of tegula input. 4. The present study demonstrates 1) that the timing of elevator motor neuron activity is determined by phasic afferent input from tegulae and stretch receptors and 2) that input from the stretch receptors controls the duration of elevator activity in the wingbeat cycle following the wing movement that was responsible for the generation of the receptor discharge.
Mimicking the colourful wing scale structure of the Papilio blumei butterfly.
Kolle, Mathias; Salgard-Cunha, Pedro M; Scherer, Maik R J; Huang, Fumin; Vukusic, Pete; Mahajan, Sumeet; Baumberg, Jeremy J; Steiner, Ullrich
2010-07-01
The brightest and most vivid colours in nature arise from the interaction of light with surfaces that exhibit periodic structure on the micro- and nanoscale. In the wings of butterflies, for example, a combination of multilayer interference, optical gratings, photonic crystals and other optical structures gives rise to complex colour mixing. Although the physics of structural colours is well understood, it remains a challenge to create artificial replicas of natural photonic structures. Here we use a combination of layer deposition techniques, including colloidal self-assembly, sputtering and atomic layer deposition, to fabricate photonic structures that mimic the colour mixing effect found on the wings of the Indonesian butterfly Papilio blumei. We also show that a conceptual variation to the natural structure leads to enhanced optical properties. Our approach offers improved efficiency, versatility and scalability compared with previous approaches.
Pressure-Distribution Measurements on O-2H Airplane in Flight
NASA Technical Reports Server (NTRS)
Pearson, H A
1937-01-01
Results are given of pressure-distribution measurements made over two different horizontal tail surfaces and the right wing cellule, including the slipstream area, of an observation-type biplane. Measurements were also taken of air speed, control-surface positions, control-stick forces, angular velocities, and accelerations during various abrupt maneuvers. These maneuvers consisted of push-downs and pull-ups from level flight, dive pull-outs, and aileron rolls with various thrust conditions. The results from the pressure-distribution measurements over the wing cellule are given on charts showing the variation of individual rib coefficients with wing coefficients; the data from the tail-surface pressure-distribution measurements are given mainly as total loads and moments. These data are supplemented by time histories of the measured quantities and isometric views of the rib pressure distributions occurring in abrupt maneuvers.
NASA Technical Reports Server (NTRS)
Taylor, A. B.
1984-01-01
Work done under the Energy Efficient Transport project in the field of advanced aerodynamics and active controls is summarized. The project task selections focused on the following: the investigation of long-duct nacelle shape variation on interference drag; the investigation of the adequacy of a simple control law for the elastic modes of a wing; the development of the aerodynamic technology at cruise and low speed of high-aspect-ratio supercritical wings of high performance; and the development of winglets for a second-generation jet transport. All the tasks involved analysis and substantial wind tunnel testing. The winglet program also included flight evaluation. It is considered that the technology base has been built for the application of high-aspect-ratio supercritical wings and for the use of winglets on second-generation transports.
NASA Technical Reports Server (NTRS)
Stimpert, D. L.
1978-01-01
An acoustic and aerodynamic test program was conducted on a 1/6.25 scale model of the Quiet, Clean, Short-Haul Experimental Engine (QCSEE) forward thrust over-the-wing (OTW) nozzle and OTW thrust reverser. In reverse thrust, the effect of reverser geometry was studied by parametric variations in blocker spacing, blocker height, lip angle, and lip length. Forward thrust nozzle tests determined the jet noise levels of the cruise and takeoff nozzles, the effect of opening side doors to achieve takeoff thrust, and scrubbing noise of the cruise and takeoff jet on a simulated wing surface. Velocity profiles are presented for both forward and reverse thrust nozzles. An estimate of the reverse thrust was made utilizing the measured centerline turning angle.
NASA Technical Reports Server (NTRS)
Choo, Yung K.; Slater, John W.; Vickerman, Mary B.; VanZante, Judith F.; Wadel, Mary F. (Technical Monitor)
2002-01-01
Issues associated with analysis of 'icing effects' on airfoil and wing performances are discussed, along with accomplishments and efforts to overcome difficulties with ice. Because of infinite variations of ice shapes and their high degree of complexity, computational 'icing effects' studies using available software tools must address many difficulties in geometry acquisition and modeling, grid generation, and flow simulation. The value of each technology component needs to be weighed from the perspective of the entire analysis process, from geometry to flow simulation. Even though CFD codes are yet to be validated for flows over iced airfoils and wings, numerical simulation, when considered together with wind tunnel tests, can provide valuable insights into 'icing effects' and advance our understanding of the relationship between ice characteristics and their effects on performance degradation.
Automatic identification of bird targets with radar via patterns produced by wing flapping.
Zaugg, Serge; Saporta, Gilbert; van Loon, Emiel; Schmaljohann, Heiko; Liechti, Felix
2008-09-06
Bird identification with radar is important for bird migration research, environmental impact assessments (e.g. wind farms), aircraft security and radar meteorology. In a study on bird migration, radar signals from birds, insects and ground clutter were recorded. Signals from birds show a typical pattern due to wing flapping. The data were labelled by experts into the four classes BIRD, INSECT, CLUTTER and UFO (unidentifiable signals). We present a classification algorithm aimed at automatic recognition of bird targets. Variables related to signal intensity and wing flapping pattern were extracted (via continuous wavelet transform). We used support vector classifiers to build predictive models. We estimated classification performance via cross validation on four datasets. When data from the same dataset were used for training and testing the classifier, the classification performance was extremely to moderately high. When data from one dataset were used for training and the three remaining datasets were used as test sets, the performance was lower but still extremely to moderately high. This shows that the method generalizes well across different locations or times. Our method provides a substantial gain of time when birds must be identified in large collections of radar signals and it represents the first substantial step in developing a real time bird identification radar system. We provide some guidelines and ideas for future research.
Ectopic KNOX Expression Affects Plant Development by Altering Tissue Cell Polarity and Identity[OPEN
Rebocho, Alexandra B.
2016-01-01
Plant development involves two polarity types: tissue cell (asymmetries within cells are coordinated across tissues) and regional (identities vary spatially across tissues) polarity. Both appear altered in the barley (Hordeum vulgare) Hooded mutant, in which ectopic expression of the KNOTTED1-like Homeobox (KNOX) gene, BKn3, causes inverted polarity of differentiated hairs and ectopic flowers, in addition to wing-shaped outgrowths. These lemma-specific effects allow the spatiotemporal analysis of events following ectopic BKn3 expression, determining the relationship between KNOXs, polarity, and shape. We show that tissue cell polarity, based on localization of the auxin transporter SISTER OF PINFORMED1 (SoPIN1), dynamically reorients as ectopic BKn3 expression increases. Concurrently, ectopic expression of the auxin importer LIKE AUX1 and boundary gene NO APICAL MERISTEM is activated. The polarity of hairs reflects SoPIN1 patterns, suggesting that tissue cell polarity underpins oriented cell differentiation. Wing cell files reveal an anisotropic growth pattern, and computational modeling shows how polarity guiding growth can account for this pattern and wing emergence. The inverted ectopic flower orientation does not correlate with SoPIN1, suggesting that this form of regional polarity is not controlled by tissue cell polarity. Overall, the results suggest that KNOXs trigger different morphogenetic effects through interplay between tissue cell polarity, identity, and growth. PMID:27553356
Cruising the rain forest floor: butterfly wing shape evolution and gliding in ground effect.
Cespedes, Ann; Penz, Carla M; DeVries, Philip J
2015-05-01
Flight is a key innovation in the evolutionary success of insects and essential to dispersal, territoriality, courtship and oviposition. Wing shape influences flight performance and selection likely acts to maximize performance for conducting essential behaviours that in turn results in the evolution of wing shape. As wing shape also contributes to fitness, optimal shapes for particular flight behaviours can be assessed with aerodynamic predictions and placed in an ecomorphological context. Butterflies in the tribe Haeterini (Nymphalidae) are conspicuous members of understorey faunas in lowland Neotropical forests. Field observations indicate that the five genera in this clade differ in flight height and behaviour: four use gliding flight at the forest floor level, and one utilizes flapping flight above the forest floor. Nonetheless, the association of ground level gliding flight behaviour and wing shape has never been investigated in this or any other butterfly group. We used landmark-based geometric morphometrics to test whether wing shapes in Haeterini and their close relatives reflected observed flight behaviours. Four genera of Haeterini and some distantly related Satyrinae showed significant correspondence between wing shape and theoretical expectations in performance trade-offs that we attribute to selection for gliding in ground effect. Forewing shape differed between sexes for all taxa, and male wing shapes were aerodynamically more efficient for gliding flight than corresponding females. This suggests selection acts differentially on male and female wing shapes, reinforcing the idea that sex-specific flight behaviours contribute to the evolution of sexual dimorphism. Our study indicates that wing shapes in Haeterini butterflies evolved in response to habitat-specific flight behaviours, namely gliding in ground effect along the forest floor, resulting in ecomorphological partitions of taxa in morphospace. The convergent flight behaviour and wing morphology between tribes of Satyrinae suggest that the flight environment may offset phylogenetic constraints. Overall, this study provides a basis for exploring similar patterns of wing shape evolution in other taxa that glide in ground effect. © 2014 The Authors. Journal of Animal Ecology © 2014 British Ecological Society.
Structural efficiency study of composite wing rib structures
NASA Technical Reports Server (NTRS)
Swanson, Gary D.; Gurdal, Zafer; Starnes, James H., Jr.
1988-01-01
A series of short stiffened panel designs which may be applied to a preliminary design assessment of an aircraft wing rib is presented. The computer program PASCO is used as the primary design and analysis tool to assess the structural efficiency and geometry of a tailored corrugated panel, a corrugated panel with a continuous laminate, a hat stiffened panel, a blade stiffened panel, and an unstiffened flat plate. To correct some of the shortcomings in the PASCO analysis when shear is present, a two step iterative process using the computer program VICON is used. The loadings considered include combinations of axial compression, shear, and lateral pressure. The loading ranges considered are broad enough such that the designs presented may be applied to other stiffened panel applications. An assessment is made of laminate variations, increased spacing, and nonoptimum geometric variations, including a beaded panel, on the design of the panels.
Mechanical performance of aquatic rowing and flying.
Walker, J A; Westneat, M W
2000-09-22
Aquatic flight, performed by rowing or flapping fins, wings or limbs, is a primary locomotor mechanism for many animals. We used a computer simulation to compare the mechanical performance of rowing and flapping appendages across a range of speeds. Flapping appendages proved to be more mechanically efficient than rowing appendages at all swimming speeds, suggesting that animals that frequently engage in locomotor behaviours that require energy conservation should employ a flapping stroke. The lower efficiency of rowing appendages across all speeds begs the question of why rowing occurs at all. One answer lies in the ability of rowing fins to generate more thrust than flapping fins during the power stroke. Large forces are necessary for manoeuvring behaviours such as accelerations, turning and braking, which suggests that rowing should be found in slow-swimming animals that frequently manoeuvre. The predictions of the model are supported by observed patterns of behavioural variation among rowing and flapping vertebrates.
Schertel, Claus; Albarca, Monica; Rockel-Bauer, Claudia; Kelley, Nicholas W.; Bischof, Johannes; Hens, Korneel
2015-01-01
Transcription factors (TFs) are key regulators of cell fate. The estimated 755 genes that encode DNA binding domain-containing proteins comprise ∼5% of all Drosophila genes. However, the majority has remained uncharacterized so far due to the lack of proper genetic tools. We generated 594 site-directed transgenic Drosophila lines that contain integrations of individual UAS-TF constructs to facilitate spatiotemporally controlled misexpression in vivo. All transgenes were expressed in the developing wing, and two-thirds induced specific phenotypic defects. In vivo knockdown of the same genes yielded a phenotype for 50%, with both methods indicating a great potential for misexpression to characterize novel functions in wing growth, patterning, and development. Thus, our UAS-TF library provides an important addition to the genetic toolbox of Drosophila research, enabling the identification of several novel wing development-related TFs. In parallel, we established the chromatin landscape of wing imaginal discs by ChIP-seq analyses of five chromatin marks and RNA Pol II. Subsequent clustering revealed six distinct chromatin states, with two clusters showing enrichment for both active and repressive marks. TFs that carry such “bivalent” chromatin are highly enriched for causing misexpression phenotypes in the wing, and analysis of existing expression data shows that these TFs tend to be differentially expressed across the wing disc. Thus, bivalently marked chromatin can be used as a marker for spatially regulated TFs that are functionally relevant in a developing tissue. PMID:25568052
Wilts, Bodo D; Vey, Aidan J M; Briscoe, Adriana D; Stavenga, Doekele G
2017-11-21
Longwing butterflies, Heliconius sp., also called heliconians, are striking examples of diversity and mimicry in butterflies. Heliconians feature strongly colored patterns on their wings, arising from wing scales colored by pigments and/or nanostructures, which serve as an aposematic signal. Here, we investigate the coloration mechanisms among several species of Heliconius by applying scanning electron microscopy, (micro)spectrophotometry, and imaging scatterometry. We identify seven kinds of colored scales within Heliconius whose coloration is derived from pigments, nanostructures or both. In yellow-, orange- and red-colored wing patches, both cover and ground scales contain wavelength-selective absorbing pigments, 3-OH-kynurenine, xanthommatin and/or dihydroxanthommatin. In blue wing patches, the cover scales are blue either due to interference of light in the thin-film lower lamina (e.g., H. doris) or in the multilayered lamellae in the scale ridges (so-called ridge reflectors, e.g., H. sara and H. erato); the underlying ground scales are black. In the white wing patches, both cover and ground scales are blue due to their thin-film lower lamina, but because they are stacked upon each other and at the wing substrate, a faint bluish to white color results. Lastly, green wing patches (H. doris) have cover scales with blue-reflecting thin films and short-wavelength absorbing 3-OH-kynurenine, together causing a green color. The pigmentary and structural traits are discussed in relation to their phylogenetic distribution and the evolution of vision in this highly interesting clade of butterflies.
Efficiency of lift production in flapping and gliding flight of swifts.
Henningsson, Per; Hedenström, Anders; Bomphrey, Richard J
2014-01-01
Many flying animals use both flapping and gliding flight as part of their routine behaviour. These two kinematic patterns impose conflicting requirements on wing design for aerodynamic efficiency and, in the absence of extreme morphing, wings cannot be optimised for both flight modes. In gliding flight, the wing experiences uniform incident flow and the optimal shape is a high aspect ratio wing with an elliptical planform. In flapping flight, on the other hand, the wing tip travels faster than the root, creating a spanwise velocity gradient. To compensate, the optimal wing shape should taper towards the tip (reducing the local chord) and/or twist from root to tip (reducing local angle of attack). We hypothesised that, if a bird is limited in its ability to morph its wings and adapt its wing shape to suit both flight modes, then a preference towards flapping flight optimization will be expected since this is the most energetically demanding flight mode. We tested this by studying a well-known flap-gliding species, the common swift, by measuring the wakes generated by two birds, one in gliding and one in flapping flight in a wind tunnel. We calculated span efficiency, the efficiency of lift production, and found that the flapping swift had consistently higher span efficiency than the gliding swift. This supports our hypothesis and suggests that even though swifts have been shown previously to increase their lift-to-drag ratio substantially when gliding, the wing morphology is tuned to be more aerodynamically efficient in generating lift during flapping. Since body drag can be assumed to be similar for both flapping and gliding, it follows that the higher total drag in flapping flight compared with gliding flight is primarily a consequence of an increase in wing profile drag due to the flapping motion, exceeding the reduction in induced drag.
Efficiency of Lift Production in Flapping and Gliding Flight of Swifts
Henningsson, Per; Hedenström, Anders; Bomphrey, Richard J.
2014-01-01
Many flying animals use both flapping and gliding flight as part of their routine behaviour. These two kinematic patterns impose conflicting requirements on wing design for aerodynamic efficiency and, in the absence of extreme morphing, wings cannot be optimised for both flight modes. In gliding flight, the wing experiences uniform incident flow and the optimal shape is a high aspect ratio wing with an elliptical planform. In flapping flight, on the other hand, the wing tip travels faster than the root, creating a spanwise velocity gradient. To compensate, the optimal wing shape should taper towards the tip (reducing the local chord) and/or twist from root to tip (reducing local angle of attack). We hypothesised that, if a bird is limited in its ability to morph its wings and adapt its wing shape to suit both flight modes, then a preference towards flapping flight optimization will be expected since this is the most energetically demanding flight mode. We tested this by studying a well-known flap-gliding species, the common swift, by measuring the wakes generated by two birds, one in gliding and one in flapping flight in a wind tunnel. We calculated span efficiency, the efficiency of lift production, and found that the flapping swift had consistently higher span efficiency than the gliding swift. This supports our hypothesis and suggests that even though swifts have been shown previously to increase their lift-to-drag ratio substantially when gliding, the wing morphology is tuned to be more aerodynamically efficient in generating lift during flapping. Since body drag can be assumed to be similar for both flapping and gliding, it follows that the higher total drag in flapping flight compared with gliding flight is primarily a consequence of an increase in wing profile drag due to the flapping motion, exceeding the reduction in induced drag. PMID:24587260