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.
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
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.
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
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.
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.
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.
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.
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.
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.
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
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.
[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.
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
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.
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.
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
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.
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
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.
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.
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
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.
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.
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.
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
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
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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?
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
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.
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
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
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.
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.
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
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
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.
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.
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.
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.
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
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
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.
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.
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.
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.
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.
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
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.
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.
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
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
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.
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.
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.
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.
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.
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.
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.
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
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
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.
T-Box Genes in Drosophila Limb Development.
Pflugfelder, G O; Eichinger, F; Shen, J
2017-01-01
T-box genes are essential for limb development in vertebrates and arthropods. The Drosophila genome encodes eight T-box genes, six of which are expressed in limb ontogenesis. The Tbx20-related gene pair midline and H15 is essential for dorso-ventral patterning of the Drosophila legs. The three Tbx6-related Dorsocross genes are required for epithelial remodeling during wing development. The Drosophila gene optomotor-blind (omb) is the only member of the Tbx2 subfamily in the fly and is predominantly involved in wing development. Omb is essential for wing development and is sufficient to promote the development of a second wing pair. Targeted manipulations of omb expression have shown that the bulk omb requirement for wing development can be deconstructed into a number of individual functions. Even though omb expression in the wing disc is symmetrical with regard to the anterior/posterior (A/P) compartment boundary, anterior and posterior knockdowns have distinct consequences: Anterior Omb is required for the maintenance of a straight A/P lineage restriction boundary. Posterior Omb suppresses formation of an apical epithelial fold along the A/P boundary. Drosophila T-box gene expression is not confined to the ectoderm-derived epithelia of the imaginal discs. Both Doc and Omb are prominently expressed in leg disc muscle precursor cells. Omb is also strongly expressed in a tracheal branch that invades the extracellular matrix of the wing disc. The function of Doc and Omb in the latter tissues is not known, indicative of the many questions still open in the field. © 2017 Elsevier Inc. All rights reserved.
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
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.
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.
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.
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...
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.
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.
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.
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.
Evolution of insect wings and development - new details from Palaeozoic nymphs.
Haug, Joachim T; Haug, Carolin; Garwood, Russell J
2016-02-01
The nymphal stages of Palaeozoic insects differ significantly in morphology from those of their modern counterparts. Morphological details for some previously reported species have recently been called into question. Palaeozoic insect nymphs are important, however - their study could provide key insights into the evolution of wings, and complete metamorphosis. Here we review past work on these topics and juvenile insects in the fossil record, and then present both novel and previously described nymphs, documented using new imaging methods. Our results demonstrate that some Carboniferous nymphs - those of Palaeodictyopteroidea - possessed movable wing pads and appear to have been able to perform simple flapping flight. It remains unclear whether this feature is ancestral for Pterygota or an autapomorphy of Palaeodictyopteroidea. Further characters of nymphal development which were probably in the ground pattern of Pterygota can be reconstructed. Wing development was very gradual (archimetaboly). Wing pads did not protrude from the tergum postero-laterally as in most modern nymphs, but laterally, and had well-developed venation. The modern orientation of wing pads and the delay of wing development into later developmental stages (condensation) appears to have evolved several times independently within Pterygota: in Ephemeroptera, Odonatoptera, Eumetabola, and probably several times within Polyneoptera. Selective pressure appears to have favoured a more pronounced metamorphosis between the last nymphal and adult stage, ultimately reducing exploitation competition between the two. We caution, however, that the results presented herein remain preliminary, and the reconstructed evolutionary scenario contains gaps and uncertainties. Additional comparative data need to be collected. The present study is thus seen as a starting point for this enterprise. © 2014 Cambridge Philosophical Society.
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.
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.
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.
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.
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
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.
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.
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
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.
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.
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
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.
Dahal, Giri Raj; Pradhan, Sarala Joshi; Bates, Emily Anne
2017-08-01
Loss of embryonic ion channel function leads to morphological defects, but the underlying reason for these defects remains elusive. Here, we show that inwardly rectifying potassium (Irk) channels regulate release of the Drosophila bone morphogenetic protein Dpp in the developing fly wing and that this is necessary for developmental signaling. Inhibition of Irk channels decreases the incidence of distinct Dpp-GFP release events above baseline fluorescence while leading to a broader distribution of Dpp-GFP. Work by others in different cell types has shown that Irk channels regulate peptide release by modulating membrane potential and calcium levels. We found calcium transients in the developing wing, and inhibition of Irk channels reduces the duration and amplitude of calcium transients. Depolarization with high extracellular potassium evokes Dpp release. Taken together, our data implicate Irk channels as a requirement for regulated release of Dpp, highlighting the importance of the temporal pattern of Dpp presentation for morphogenesis of the wing. © 2017. Published by The Company of Biologists Ltd.
Transcriptomic and epigenomic characterization of the developing bat wing
Eckalbar, Walter L.; Schlebusch, Stephen A.; Mason, Mandy K.; Gill, Zoe; Parker, Ash V.; Booker, Betty M.; Nishizaki, Sierra; Muswamba-Nday, Christiane; Terhune, Elizabeth; Nevonen, Kimberly; Makki, Nadja; Friedrich, Tara; VanderMeer, Julia E.; Pollard, Katherine S.; Carbone, Lucia; Wall, Jeff D.; Illing, Nicola; Ahituv, Nadav
2016-01-01
Bats are the only mammals capable of powered flight, but little is known about the genetic determinants that shape their wings. Here, we generated a genome for Miniopterus natalensis and performed RNA-seq and ChIP-seq (H3K27ac, H3K27me3) on its developing forelimb and hindlimb autopods at sequential embryonic stages to decipher the molecular events that underlie bat wing development. Over 7,000 genes and several lncRNAs, including Tbx5-as1 and Hottip, were differentially expressed between forelimb, hindlimb and different stages. ChIP-seq identified thousands of regions that are differentially modified in forelimb versus hindlimb. Comparative genomics found 2,796 bat-accelerated regions within H3K27ac peaks, several of which cluster near limb-associated genes. Pathway analyses revealed multiple ribosomal proteins and known limb patterning signaling pathways as differentially regulated, and implicated increased forelimb mesenchymal condensations with differential growth. Combined, our work outlines multiple genetic components that contribute to bat wing formation, providing a genomic blueprint for this morphological innovation. PMID:27019111
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.
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
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.
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.
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).
Critical Point in Self-Organized Tissue Growth
NASA Astrophysics Data System (ADS)
Aguilar-Hidalgo, Daniel; Werner, Steffen; Wartlick, Ortrud; González-Gaitán, Marcos; Friedrich, Benjamin M.; Jülicher, Frank
2018-05-01
We present a theory of pattern formation in growing domains inspired by biological examples of tissue development. Gradients of signaling molecules regulate growth, while growth changes these graded chemical patterns by dilution and advection. We identify a critical point of this feedback dynamics, which is characterized by spatially homogeneous growth and proportional scaling of patterns with tissue length. We apply this theory to the biological model system of the developing wing of the fruit fly Drosophila melanogaster and quantitatively identify signatures of the critical point.
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.
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.
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.
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 ...
NASA Technical Reports Server (NTRS)
Amiet, R. K.
1991-01-01
A unified theory for aerodynamics and noise of advanced turboprops is presented. The theory and a computer code developed for evaluation at the shielding benefits that might be expected by an aircraft wing in a wing-mounted propeller installation are presented. Several computed directivity patterns are presented to demonstrate the theory. Recently with the advent of the concept of using the wing of an aircraft for noise shielding, the case of diffraction by a surface in a flow has been given attention. The present analysis is based on the case of diffraction of no flow. By combining a Galilean and a Lorentz transform, the wave equation with a mean flow can be reduced to the ordinary equation. Allowance is also made in the analysis for the case of a swept wing. The same combination of Galilean and Lorentz transforms lead to a problem with no flow but a different sweep. The solution procedures for the cases of leading and trailing edges are basically the same. Two normalizations of the solution are given by the computer program. FORTRAN computer programs are presented with detailed documentation. The output from these programs compares favorably with the results of other investigators.
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.
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
Velentzas, Panagiotis D; Velentzas, Athanassios D; Pantazi, Asimina D; Mpakou, Vassiliki E; Zervas, Christos G; Papassideri, Issidora S; Stravopodis, Dimitrios J
2013-01-01
Proteasome-dependent and autophagy-mediated degradation of eukaryotic cellular proteins represent the two major proteostatic mechanisms that are critically implicated in a number of signaling pathways and cellular processes. Deregulation of functions engaged in protein elimination frequently leads to development of morbid states and diseases. In this context, and through the utilization of GAL4/UAS genetic tool, we herein examined the in vivo contribution of proteasome and autophagy systems in Drosophila eye and wing morphogenesis. By exploiting the ability of GAL4-ninaE. GMR and P{GawB}Bx(MS1096) genetic drivers to be strongly and preferentially expressed in the eye and wing discs, respectively, we proved that proteasomal integrity and ubiquitination proficiency essentially control fly's eye and wing development. Indeed, subunit- and regulator-specific patterns of severe organ dysmorphia were obtained after the RNAi-induced downregulation of critical proteasome components (Rpn1, Rpn2, α5, β5 and β6) or distinct protein-ubiquitin conjugators (UbcD6, but not UbcD1 and UbcD4). Proteasome deficient eyes presented with either rough phenotypes or strongly dysmorphic shapes, while transgenic mutant wings were severely folded and carried blistered structures together with loss of vein differentiation. Moreover, transgenic fly eyes overexpressing the UBP2-yeast deubiquitinase enzyme were characterized by an eyeless-like phenotype. Therefore, the proteasome/ubiquitin proteolytic activities are undoubtedly required for the normal course of eye and wing development. In contrast, the RNAi-mediated downregulation of critical Atg (1, 4, 7, 9 and 18) autophagic proteins revealed their non-essential, or redundant, functional roles in Drosophila eye and wing formation under physiological growth conditions, since their reduced expression levels could only marginally disturb wing's, but not eye's, morphogenetic organization and architecture. However, Atg9 proved indispensable for the maintenance of structural integrity of adult wings in aged flies. In toto, our findings clearly demonstrate the gene-specific fundamental contribution of proteasome, but not autophagy, in invertebrate eye and wing organ development.
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
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.
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.
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
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.
Genomic Sequence around Butterfly Wing Development Genes: Annotation and Comparative Analysis
Conceição, Inês C.; Long, Anthony D.; Gruber, Jonathan D.; Beldade, Patrícia
2011-01-01
Background Analysis of genomic sequence allows characterization of genome content and organization, and access beyond gene-coding regions for identification of functional elements. BAC libraries, where relatively large genomic regions are made readily available, are especially useful for species without a fully sequenced genome and can increase genomic coverage of phylogenetic and biological diversity. For example, no butterfly genome is yet available despite the unique genetic and biological properties of this group, such as diversified wing color patterns. The evolution and development of these patterns is being studied in a few target species, including Bicyclus anynana, where a whole-genome BAC library allows targeted access to large genomic regions. Methodology/Principal Findings We characterize ∼1.3 Mb of genomic sequence around 11 selected genes expressed in B. anynana developing wings. Extensive manual curation of in silico predictions, also making use of a large dataset of expressed genes for this species, identified repetitive elements and protein coding sequence, and highlighted an expansion of Alcohol dehydrogenase genes. Comparative analysis with orthologous regions of the lepidopteran reference genome allowed assessment of conservation of fine-scale synteny (with detection of new inversions and translocations) and of DNA sequence (with detection of high levels of conservation of non-coding regions around some, but not all, developmental genes). Conclusions The general properties and organization of the available B. anynana genomic sequence are similar to the lepidopteran reference, despite the more than 140 MY divergence. Our results lay the groundwork for further studies of new interesting findings in relation to both coding and non-coding sequence: 1) the Alcohol dehydrogenase expansion with higher similarity between the five tandemly-repeated B. anynana paralogs than with the corresponding B. mori orthologs, and 2) the high conservation of non-coding sequence around the genes wingless and Ecdysone receptor, both involved in multiple developmental processes including wing pattern formation. PMID:21909358
2011-01-01
Background Mitogen-activated protein kinase (MAPK) cascades (p38, JNK, ERK pathways) are involved in cell fate acquisition during development. These kinase modules are associated with scaffold proteins that control their activity. In Drosophila, dMP1, that encodes an ERK scaffold protein, regulates ERK signaling during wing development and contributes to intervein and vein cell differentiation. Functional relationships during wing development between a chromatin regulator, the Enhancer of Trithorax and Polycomb Corto, ERK and its scaffold protein dMP1, are examined here. Results Genetic interactions show that corto and dMP1 act together to antagonize rolled (which encodes ERK) in the future intervein cells, thus promoting intervein fate. Although Corto, ERK and dMP1 are present in both cytoplasmic and nucleus compartments, they interact exclusively in nucleus extracts. Furthermore, Corto, ERK and dMP1 co-localize on several sites on polytene chromosomes, suggesting that they regulate gene expression directly on chromatin. Finally, Corto is phosphorylated. Interestingly, its phosphorylation pattern differs between cytoplasm and nucleus and changes upon ERK activation. Conclusions Our data therefore suggest that the Enhancer of Trithorax and Polycomb Corto could participate in regulating vein and intervein genes during wing tissue development in response to ERK signaling. PMID:21401930
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.
Mason, Mandy K; Hockman, Dorit; Curry, Lyle; Cunningham, Thomas J; Duester, Gregg; Logan, Malcolm; Jacobs, David S; Illing, Nicola
2015-01-01
The bat has strikingly divergent forelimbs (long digits supporting wing membranes) and hindlimbs (short, typically free digits) due to the distinct requirements of both aerial and terrestrial locomotion. During embryonic development, the morphology of the bat forelimb deviates dramatically from the mouse and chick, offering an alternative paradigm for identifying genes that play an important role in limb patterning. Using transcriptome analysis of developing Natal long-fingered bat (Miniopterus natalensis) fore- and hindlimbs, we demonstrate that the transcription factor Meis2 has a significantly higher expression in bat forelimb autopods compared to hindlimbs. Validation by reverse transcriptase and quantitative polymerase chain reaction (RT-qPCR) and whole mount in situ hybridisation shows that Meis2, conventionally known as a marker of the early proximal limb bud, is upregulated in the bat forelimb autopod from CS16. Meis2 expression is localised to the expanding interdigital webbing and the membranes linking the wing to the hindlimb and tail. In mice, Meis2 is also expressed in the interdigital region prior to tissue regression. This interdigital Meis2 expression is not activated by retinoic acid (RA) signalling as it is present in the retained interdigital tissue of Rdh10 (trex/trex) mice, which lack RA. Additionally, genes encoding RA-synthesising enzymes, Rdh10 and Aldh1a2, and the RA nuclear receptor Rarβ are robustly expressed in bat fore- and hindlimb interdigital tissues indicating that the mechanism that retains interdigital tissue in bats also occurs independently of RA signalling. Mammalian interdigital Meis2 expression, and upregulation in the interdigital webbing of bat wings, suggests an important role for Meis2 in autopod development. Interdigital Meis2 expression is RA-independent, and retention of interdigital webbing in bat wings is not due to the suppression of RA-induced cell death. Rather, RA signalling may play a role in the thinning (rather than complete loss) of the interdigital tissue in the bat forelimb, while Meis2 may interact with other factors during both bat and mouse autopod development to maintain a pool of interdigital cells that contribute to digit patterning and growth.
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
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
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.
A tool for developing an automatic insect identification system based on wing outlines
Yang, He-Ping; Ma, Chun-Sen; Wen, Hui; Zhan, Qing-Bin; Wang, Xin-Li
2015-01-01
For some insect groups, wing outline is an important character for species identification. We have constructed a program as the integral part of an automated system to identify insects based on wing outlines (DAIIS). This program includes two main functions: (1) outline digitization and Elliptic Fourier transformation and (2) classifier model training by pattern recognition of support vector machines and model validation. To demonstrate the utility of this program, a sample of 120 owlflies (Neuroptera: Ascalaphidae) was split into training and validation sets. After training, the sample was sorted into seven species using this tool. In five repeated experiments, the mean accuracy for identification of each species ranged from 90% to 98%. The accuracy increased to 99% when the samples were first divided into two groups based on features of their compound eyes. DAIIS can therefore be a useful tool for developing a system of automated insect identification. PMID:26251292
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.
Pantazi, Asimina D.; Mpakou, Vassiliki E.; Zervas, Christos G.; Papassideri, Issidora S.; Stravopodis, Dimitrios J.
2013-01-01
Proteasome-dependent and autophagy-mediated degradation of eukaryotic cellular proteins represent the two major proteostatic mechanisms that are critically implicated in a number of signaling pathways and cellular processes. Deregulation of functions engaged in protein elimination frequently leads to development of morbid states and diseases. In this context, and through the utilization of GAL4/UAS genetic tool, we herein examined the in vivo contribution of proteasome and autophagy systems in Drosophila eye and wing morphogenesis. By exploiting the ability of GAL4-ninaE. GMR and P{GawB}BxMS1096 genetic drivers to be strongly and preferentially expressed in the eye and wing discs, respectively, we proved that proteasomal integrity and ubiquitination proficiency essentially control fly’s eye and wing development. Indeed, subunit- and regulator-specific patterns of severe organ dysmorphia were obtained after the RNAi-induced downregulation of critical proteasome components (Rpn1, Rpn2, α5, β5 and β6) or distinct protein-ubiquitin conjugators (UbcD6, but not UbcD1 and UbcD4). Proteasome deficient eyes presented with either rough phenotypes or strongly dysmorphic shapes, while transgenic mutant wings were severely folded and carried blistered structures together with loss of vein differentiation. Moreover, transgenic fly eyes overexpressing the UBP2-yeast deubiquitinase enzyme were characterized by an eyeless-like phenotype. Therefore, the proteasome/ubiquitin proteolytic activities are undoubtedly required for the normal course of eye and wing development. In contrast, the RNAi-mediated downregulation of critical Atg (1, 4, 7, 9 and 18) autophagic proteins revealed their non-essential, or redundant, functional roles in Drosophila eye and wing formation under physiological growth conditions, since their reduced expression levels could only marginally disturb wing’s, but not eye’s, morphogenetic organization and architecture. However, Atg9 proved indispensable for the maintenance of structural integrity of adult wings in aged flies. In toto, our findings clearly demonstrate the gene-specific fundamental contribution of proteasome, but not autophagy, in invertebrate eye and wing organ development. PMID:24282550
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.
Distal-less regulates eyespot patterns and melanization in Bicyclus butterflies.
Monteiro, Antónia; Chen, Bin; Ramos, Diane M; Oliver, Jeffrey C; Tong, Xiaoling; Guo, Min; Wang, Wen-Kai; Fazzino, Lisa; Kamal, Firdous
2013-07-01
Butterfly eyespots represent novel complex traits that display substantial diversity in number and size within and across species. Correlative gene expression studies have implicated a large suite of transcription factors, including Distal-less (Dll), Engrailed (En), and Spalt (Sal), in eyespot development in butterflies, but direct evidence testing the function of any of these proteins is still missing. Here we show that the characteristic two-eyespot pattern of wildtype Bicyclus anynana forewings is correlated with dynamic progression of Dll, En, and Sal expression in larval wings from four spots to two spots, whereas no such decline in gene expression ensues in a four-eyespot mutant. We then conduct transgenic experiments testing whether over-expression of any of these genes in a wild-type genetic background is sufficient to induce eyespot differentiation in these pre-patterned wing compartments. We also produce a Dll-RNAi transgenic line to test how Dll down-regulation affects eyespot development. Finally we test how ectopic expression of these genes during the pupal stages of development alters adults color patters. We show that over-expressing Dll in larvae is sufficient to induce the differentiation of additional eyespots and increase the size of eyespots, whereas down-regulating Dll leads to a decrease in eyespot size. Furthermore, ectopic expression of Dll in the early pupal wing led to the appearance of ectopic patches of black scales. We conclude that Dll is a positive regulator of focal differentiation and eyespot signaling and that this gene is also a possible selector gene for scale melanization in butterflies. Copyright © 2013 Wiley Periodicals, Inc.
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.
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.
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.
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.
FijiWingsPolarity: An open source toolkit for semi-automated detection of cell polarity.
Dobens, Leonard L; Shipman, Anna; Axelrod, Jeffrey D
2018-01-02
Epithelial cells are defined by apical-basal and planar cell polarity (PCP) signaling, the latter of which establishes an orthogonal plane of polarity in the epithelial sheet. PCP signaling is required for normal cell migration, differentiation, stem cell generation and tissue repair, and defects in PCP have been associated with developmental abnormalities, neuropathologies and cancers. While the molecular mechanism of PCP is incompletely understood, the deepest insights have come from Drosophila, where PCP is manifest in hairs and bristles across the adult cuticle and organization of the ommatidia in the eye. Fly wing cells are marked by actin-rich trichome structures produced at the distal edge of each cell in the developing wing epithelium and in a mature wing the trichomes orient collectively in the distal direction. Genetic screens have identified key PCP signaling pathway components that disrupt trichome orientation, which has been measured manually in a tedious and error prone process. Here we describe a set of image processing and pattern-recognition macros that can quantify trichome arrangements in micrographs and mark these directly by color, arrow or colored arrow to indicate trichome location, length and orientation. Nearest neighbor calculations are made to exploit local differences in orientation to better and more reliably detect and highlight local defects in trichome polarity. We demonstrate the use of these tools on trichomes in adult wing preps and on actin-rich developing trichomes in pupal wing epithelia stained with phalloidin. FijiWingsPolarity is freely available and will be of interest to a broad community of fly geneticists studying the effect of gene function on PCP.
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.
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.
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
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.
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.
Selective chromatid segregation mechanism for Bruchus wings piebald color.
Klar, Amar J S
2015-01-01
The mechanisms of asymmetric organ development have been under intensive investigation for years, yet the proposed mechanisms remain controversial (1-3). The female Bruchus quadrimaculatus beetle insect develops two black-colored spots bilaterally located on each upper elytra wing by an unknown mechanism. Fifty percent of the P (for piebald, two colors) gene homozygous mutant insects, described in 1925, had a normal left elytrum (with two black spots) and an abnormal right elytrum (with two red spots) and the balance supported the converse lateralized pigment arrangement (4). Rather than supporting the conventional morphogen model for the wings pigmentation development, their biological origin is explained here with the somatic strand-specific epigenetic imprinting and selective sister chromatid segregation (SSIS) mechanism (5). We propose that the P gene product performs the selective sister chromatid segregation function to produce symmetric cell division of a specific cell during embryogenesis to result in the bilateral symmetric development of elytra black color spots and that the altered chromatid segregation pattern of the mutant causes asymmetric cell division to confer the piebald phenotype.
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
Comparative analysis of the development of wing-flapping and flight in the fowl.
Provine, R R; Strawbridge, C L; Harrison, B J
1984-01-01
The development of wing-flapping rate, lateral flight, wing area, and the ratio of wing area to body weight are described in the Japanese quail (Coturnix coturnix japonica) and three chickens (Gallus gallus) to determine common developmental phenomena and to assess the effects of domestication. The chickens were the White Leghorn (a commercial egg producer), the Cornish X Rock (a commercial meat producer), and the Red Jungle fowl (the probable ancestor of domestic chickens). All birds performed drop-evoked wing-flapping on the day of hatching, at least 1 week before lateral flight was possible. Flapping rate of chickens doubled between hatching (approximately 4-6 Hz) and 13 days (approximately 9-12 Hz), after which it leveled off. Japanese quail (JQ) maintained a high flapping rate (approximately 11-13 Hz) during the 21 days after hatching. The Jungle fowl (JF) and JQ flapped the fastest and the White Leghorn (WL) and Cornish X Rock (CR) chickens flapped the slowest. The JF, WL, and JQ developed lateral flight at 7-9 days. The CR first flew 1-2 weeks later but subsequently became flightless. The WL, JF, and JQ had similar ratios of wing area to body weight; the ratios increased to a peak at 11-15 days and later declined. The ratio of the very heavy, essentially flightless, CR was approximately one-half that of the flighted JQ, WL, and JF. The wing-flapping frequencies of the domestic WL and CR chickens approximated that of the JF, suggesting that domestication did not affect the motor pattern generator for flight. The artificial selection of the CR for high body weight drastically diminished its flight performance by producing an unfavorable ratio of wing area to body weight. The JF and the domestic WL both flew well and had similar ratios. Domestication affected flight performance but not the neural circuitry producing wing-flapping. The central nervous system is much more conservative in its response to selection than the peripheral effector structures that it drives.
Mateus, Ana Rita A; Marques-Pita, Manuel; Oostra, Vicencio; Lafuente, Elvira; Brakefield, Paul M; Zwaan, Bas J; Beldade, Patrícia
2014-11-21
The environmental regulation of development can result in the production of distinct phenotypes from the same genotype and provide the means for organisms to cope with environmental heterogeneity. The effect of the environment on developmental outcomes is typically mediated by hormonal signals which convey information about external cues to the developing tissues. While such plasticity is a wide-spread property of development, not all developing tissues are equally plastic. To understand how organisms integrate environmental input into coherent adult phenotypes, we must know how different body parts respond, independently or in concert, to external cues and to the corresponding internal signals. We quantified the effect of temperature and ecdysone hormone manipulations on post-growth tissue patterning in an experimental model of adaptive developmental plasticity, the butterfly Bicyclus anynana. Following a suite of traits evolving by natural or sexual selection, we found that different groups of cells within the same tissue have sensitivities and patterns of response that are surprisingly distinct for the external environmental cue and for the internal hormonal signal. All but those wing traits presumably involved in mate choice responded to developmental temperature and, of those, all but the wing traits not exposed to predators responded to hormone manipulations. On the other hand, while patterns of significant response to temperature contrasted traits on autonomously-developing wings, significant response to hormone manipulations contrasted neighboring groups of cells with distinct color fates. We also showed that the spatial compartmentalization of these responses cannot be explained by the spatial or temporal compartmentalization of the hormone receptor protein. Our results unravel the integration of different aspects of the adult phenotype into developmental and functional units which both reflect and impact evolutionary change. Importantly, our findings underscore the complexity of the interactions between environment and physiology in shaping the development of different body parts.
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.
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.
Özsu, Nesibe; Monteiro, Antónia
2017-10-16
One hypothesis surrounding the origin of novel traits is that they originate from the co-option of pre-existing genes or larger gene regulatory networks into novel developmental contexts. Insights into a trait's evolutionary origins can, thus, be gained via identification of the genes underlying trait development, and exploring whether those genes also function in other developmental contexts. Here we investigate the set of genes associated with the development of eyespot color patterns, a trait that originated once within the Nymphalid family of butterflies. Although several genes associated with eyespot development have been identified, the eyespot gene regulatory network remains largely unknown. In this study, next-generation sequencing and transcriptome analyses were used to identify a large set of genes associated with eyespot development of Bicyclus anynana butterflies, at 3-6 h after pupation, prior to the differentiation of the color rings. Eyespot-associated genes were identified by comparing the transcriptomes of homologous micro-dissected wing tissues that either develop or do not develop eyespots in wild-type and a mutant line of butterflies, Spotty, with extra eyespots. Overall, 186 genes were significantly up and down-regulated in wing tissues that develop eyespots compared to wing tissues that do not. Many of the differentially expressed genes have yet to be annotated. New signaling pathways, including the Toll, Fibroblast Growth Factor (FGF), extracellular signal-regulated kinase (ERK) and/or Jun N-terminal kinase (JNK) signaling pathways are associated for the first time with eyespot development. In addition, several genes involved in wound healing and calcium signaling were also found to be associated with eyespots. Overall, this study provides the identity of many new genes and signaling pathways associated with eyespots, and suggests that the ancient wound healing gene regulatory network may have been co-opted to cells at the center of the pattern to aid in eyespot origins. New transcription factors that may be providing different identities to distinct wing sectors, and genes with sexually dimorphic expression in the eyespots were also identified.
NASA Technical Reports Server (NTRS)
Teske, M. E.
1984-01-01
This is a user manual for the computer code ""AGDISP'' (AGricultural DISPersal) which has been developed to predict the deposition of material released from fixed and rotary wing aircraft in a single-pass, computationally efficient manner. The formulation of the code is novel in that the mean particle trajectory and the variance about the mean resulting from turbulent fluid fluctuations are simultaneously predicted. The code presently includes the capability of assessing the influence of neutral atmospheric conditions, inviscid wake vortices, particle evaporation, plant canopy and terrain on the deposition pattern.
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.
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
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
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.
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.
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.
Rodriguez, Isabel
2011-01-01
Acquisition of a final shape and size during organ development requires a regulated program of growth and patterning controlled by a complex genetic network of signalling molecules that must be coordinated to provide positional information to each cell within the corresponding organ or tissue. The mechanism by which all these signals are coordinated to yield a final response is not well understood. Here, I have characterized the Drosophila ortholog of the human TGF-β Inducible Early Gene 1 (dTIEG). TIEG are zinc-finger proteins that belong to the Krüppel-like factor (KLF) family and were initially identified in human osteoblasts and pancreatic tumor cells for the ability to enhance TGF-β response. Using the developing wing of Drosophila as “in vivo” model, the dTIEG function has been studied in the control of cell proliferation and patterning. These results show that dTIEG can modulate Dpp signalling. Furthermore, dTIEG also regulates the activity of JAK/STAT pathway suggesting a conserved role of TIEG proteins as positive regulators of TGF-β signalling and as mediators of the crosstalk between signalling pathways acting in a same cellular context. PMID:21494610
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.
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...
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
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.
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.
Numerical studies of the deposition of material released from fixed and rotary wing aircraft
NASA Technical Reports Server (NTRS)
Bilanin, A. J.; Teske, M. E.
1984-01-01
The computer code AGDISP (AGricultural DISPersal) has been developed to predict the deposition of material released from fixed and rotary wing aircraft in a single-pass, computationally efficient manner. The formulation of the code is novel in that the mean particle trajectory and the variance about the mean resulting from turbulent fluid fluctuations are simultaneously predicted. The code presently includes the capability of assessing the influence of neutral atmospheric conditions, inviscid wake vortices, particle evaporation, plant canopy and terrain on the deposition pattern. In this report, the equations governing the motion of aerially released particles are developed, including a description of the evaporation model used. A series of case studies, using AGDISP, are included.
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
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.
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
Kayal, Arivudainambi; Mohan, Viswanathan; Malanda, Belma; Anjana, Ranjit Mohan; Bhavadharini, Balaji; Mahalakshmi, Manni Mohanraj; Maheswari, Kumar; Uma, Ram; Unnikrishnan, Ranjit; Kalaiyarasi, Gunasekaran; Ninov, Lyudmil; Belton, Anne
2016-01-01
Aim: The Women In India with GDM Strategy (WINGS) project was conducted with the aim of developing a model of care (MOC) suitable for women with gestational diabetes mellitus (GDM) in low- and middle-income countries. Methodology: The WINGS project was carried out in Chennai, Southern India, in two phases. In Phase I, a situational analysis was conducted to understand the practice patterns of health-care professionals and to determine the best screening criteria through a pilot screening study. Results: Phase II involved developing a MOC-based on findings from the situational analysis and evaluating its effectiveness. The model focused on diagnosis, management, and follow-up of women with GDM who were followed prospectively throughout their pregnancy. An educational booklet was provided to all women with GDM, offering guidance on self-management of GDM including sample meal plans and physical activity tips. A pedometer was provided to all women to monitor step count. Medical nutrition therapy (MNT) was the first line of treatment given to women with GDM. Women were advised to undergo fasting blood glucose and postprandial blood glucose testing every fortnight. Insulin was indicated when the target blood glucose levels were not achieved with MNT. Women were evaluated for pregnancy outcomes and postpartum glucose tolerance status. Conclusions: The WINGS MOC offers a comprehensive package at every level of care for women with GDM. If successful, this MOC will be scaled up to other resource-constrained settings with the hope of improving lives of women with GDM. PMID:27730085
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.
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.
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.
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).
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).
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
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.
Hox genes, digit identities and the theropod/bird transition.
Galis, Frietson; Kundrát, Martin; Metz, Johan A J
2005-05-15
Vargas and Fallon (2005. J Exp Zool (Mol Dev Evol) 304B:86-90) propose that Hox gene expression patterns indicate that the most anterior digit in bird wings is homologous to digit 1 rather than to digit 2 in other amniotes. This interpretation is based on the presence of Hoxd13 expression in combination with the absence of Hoxd12 expression in the second digit condensation from which this digit develops (the first condensation is transiently present). This is a pattern that is similar to that in the developing digit 1 of the chicken foot and the mouse hand and foot. They have tested this new hypothesis by analysing Hoxd12 and Hoxd13 expression patterns in two polydactylous chicken mutants, Silkie and talpid2. They conclude that the data support the notion that the most anterior remaining digit of the bird wing is homologous to digit 1 in other amniotes either in a standard phylogenetic sense, or alternatively in a (limited) developmental sense in agreement with the Frameshift Hypothesis of Wagner and Gautier (1999, i.e., that the developmental pathway is homologous to the one that leads to a digit 1 identity in other amniotes, although it occurs in the second instead of the first digit condensation). We argue that the Hoxd12 and Hoxd13 expression patterns found for these and other limb mutants do not allow distinguishing between the hypothesis of Vargas and Fallon (2005. J Exp Zool (Mol Dev Evol) 304B:86-90) and the alternative one, i.e., the most anterior digit in bird wings is homologous to digit 2 in other amniotes, in a phylogenetic or developmental sense. Therefore, at the moment the data on limb mutants does not present a challenge to the hypothesis, based on other developmental data (Holmgren, 1955. Acta Zool 36:243-328; Hinchliffe, 1984. In: Hecht M, Ostrom JH, Viohl G, Wellnhofer P, editors. The beginnings of birds. Eichstätt: Freunde des Jura-Museum. p 141-147; Burke and Feduccia, 1997. Science 278:666-668; Kundrát et al., 2002. J Exp Zool (Mol Dev Evol) 294B:151-159; Larsson and Wagner, 2002. J Exp Zool (Mol Dev Evol) 294B:146-151; Feduccia and Nowicki, 2002. Naturwissenschaften 89:391-393), that the digits of bird wings are homologous to digits 2,3,4 in amniotes. We recommend further testing of the hypothesis by comparing Hoxd expression patterns in different taxa. Copyright 2005 Wiley-Liss, Inc
Sheeba, Caroline J; Andrade, Raquel P; Duprez, Delphine; Palmeirim, Isabel
2010-01-01
Specific interactions between fibroblast growth factors (Fgf1-22) and their tyrosine kinase receptors (FgfR1-4) activate different signalling pathways that are responsible for the biological processes in which Fgf signalling is implicated during embryonic development. In the chick, several Fgf ligands (Fgf2, 4, 8, 9, 10, 12, 13 and 18) and the four FgfRs (FgfR 1, 2, 3 and 4) have been reported to be expressed in the developing limb. The precise spatial and temporal expression of these transcripts is important to guide the limb bud to develop into a wing/leg. In this paper, we present a detailed and systematic analysis of the expression patterns of FgfR1, 2, 3 and 4 throughout chick wing development, by in situ hybridisation on whole mounts and sections. Moreover, we characterize for the first time the different isoforms of FGFR1-3 by analysing their differential expression in limb ectoderm and mesodermal tissues, using RT-PCR and in situ hybridisation on sections. Finally, isoform-specific sequences for FgfR1IIIb, FgfR1IIIc, FgfR3IIIb and FgfR3IIIc were determined and deposited in GenBank with the following accession numbers: GU053725, GU065444, GU053726, GU065445, respectively.
High-Speed Boundary-Layer Transition: Study of Stationary Crossflow Using Spectral Analysis
NASA Astrophysics Data System (ADS)
McGuire, Patrick Joseph
Crossflow instability is primary cause of boundary-layer transition on swept wings used in high-speed applications. Delaying the downstream location of transition would drastically reduce the viscous drag over the wing surface, and subsequently improves the overall aircraft efficiency. By studying the development of instability growth rates and how they interact with the surroundings, researchers can control the crossflow transition location. Experiments on the 35° swept-wing model were performed in the NASA Langley 20-Inch Supersonic Wind Tunnel with Mach 2.0 flow conditions and 20 μm tall discrete roughness elements (DRE) with varying spacing placed along the leading edge. Fluorene was used as the sublimating chemical in the surface flow visualization technique to observe the transition front and stationary crossflow vortex patterns in the laminar flow region. Spatial spectral decomposition was completed on high-resolution images of sublimating chemical runs using a newly developed image processing technique. Streamwise evolution of the vortex track wavelengths within the laminar boundary-layer region was observed. The spectral information was averaged to produce dominant modes present throughout the laminar region.
Flow field of flexible flapping wings
NASA Astrophysics Data System (ADS)
Sallstrom, Erik
The agility and maneuverability of natural fliers would be desirable to incorporate into engineered micro air vehicles (MAVs). However, there is still much for engineers to learn about flapping flight in order to understand how such vehicles can be built for efficient flying. The goal of this study is to develop a methodology for capturing high quality flow field data around flexible flapping wings in a hover environment and to interpret it to gain a better understanding of how aerodynamic forces are generated. The flow field data was captured using particle image velocimetry (PIV) and required that measurements be taken around a repeatable flapping motion to obtain phase-averaged data that could be studied throughout the flapping cycle. Therefore, the study includes the development of flapping devices with a simple repeatable single degree of freedom flapping motion. The acquired flow field data has been examined qualitatively and quantitatively to investigate the mechanisms behind force production in hovering flight and to relate it to observations in previous research. Specifically, the flow fields have been investigated around a rigid wing and several carbon fiber reinforced flexible membrane wings. Throughout the whole study the wings were actuated with either a sinusoidal or a semi-linear flapping motion. The semi-linear flapping motion holds the commanded angular velocity nearly constant through half of each half-stroke while the sinusoidal motion is always either accelerating or decelerating. The flow fields were investigated by examining vorticity and vortex structures, using the Q criterion as the definition for the latter, in two and three dimensions. The measurements were combined with wing deflection measurements to demonstrate some of the key links in how the fluid-structure interactions generated aerodynamic forces. The flow fields were also used to calculate the forces generated by the flapping wings using momentum balance methods which yielded details of where along the wing the forces were generated. As expected, these results indicated that the spanwise location of where the forces were generated depended upon the wings membrane material and reinforcement pattern, but in general it was in the outer third of the wing. (Full text of this dissertation may be available via the University of Florida Libraries web site. Please check http://www.uflib.ufl.edu/etd.html)
Magneto-optical Effects in the Scattering Polarization Wings of the Ca I 4227 Å Resonance Line
NASA Astrophysics Data System (ADS)
Alsina Ballester, E.; Belluzzi, L.; Trujillo Bueno, J.
2018-02-01
The linear polarization pattern produced by scattering processes in the Ca I 4227 Å resonance line is a valuable observable for probing the solar atmosphere. Via the Hanle effect, the very significant Q/I and U/I line-center signals are sensitive to the presence of magnetic fields in the lower chromosphere with strengths between 5 and 125 G, approximately. On the other hand, partial frequency redistribution (PRD) produces sizable signals in the wings of the Q/I profile, which have always been thought to be insensitive to the presence of magnetic fields. Interestingly, novel observations of this line revealed a surprising behavior: fully unexpected signals in the wings of the U/I profile and spatial variability in the wings of both Q/I and U/I. We show that the magneto-optical (MO) terms of the Stokes-vector transfer equation produce sizable signals in the wings of U/I and a clear sensitivity of the Q/I and U/I wings to the presence of photospheric magnetic fields with strengths similar to those that produce the Hanle effect in the line core. This radiative transfer investigation on the joint action of scattering processes and the Hanle and Zeeman effects in the Ca I 4227 Å line should facilitate the development of more reliable techniques for exploring the magnetism of stellar atmospheres. To this end, we can now exploit the circular polarization produced by the Zeeman effect, the magnetic sensitivity caused by the above-mentioned MO effects in the Q/I and U/I wings, and the Hanle effect in the line core.
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
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.
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.
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.
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.
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.
Predicting Aircraft Spray Patterns on Crops
NASA Technical Reports Server (NTRS)
Teske, M. E.; Bilanin, A. J.
1986-01-01
Agricultural Dispersion Prediction (AGDISP) system developed to predict deposition of agricultural material released from rotary- and fixed-wing aircraft. AGDISP computes ensemble average mean motion resulting from turbulent fluid fluctuations. Used to examine ways of making dispersal process more efficient by insuring uniformity, reducing waste, and saving money. Programs in AGDISP system written in FORTRAN IV for interactive execution.
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.
Physics-based Morphology Analysis and Adjoint Optimization of Flexible Flapping Wings
2016-08-30
understand the underlying physics of flexible wings in flying insects and birds towards the bio -inspired wing designs with superior aerodynamic...flapping flights have been developed to understand the underlying physics of flexible wings in flying insects and birds towards the bio -inspired wing...been developed to understand the underlying physics of flexible wings in flying insects and birds towards the bio -inspired wing designs with superior
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)
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.
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.
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
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
Fiber Optic Wing Shape Sensing on NASA's Ikhana UAV
NASA Technical Reports Server (NTRS)
Richards, Lance; Parker, Allen R.; Ko, William L.; Piazza, Anthony
2008-01-01
This document discusses the development of fiber optic wing shape sensing on NASA's Ikhana vehicle. The Dryden Flight Research Center's Aerostructures Branch initiated fiber-optic instrumentation development efforts in the mid-1990s. Motivated by a failure to control wing dihedral resulting in a mishap with the Helios aircraft, new wing displacement techniques were developed. Research objectives for Ikhana included validating fiber optic sensor measurements and real-time wing shape sensing predictions; the validation of fiber optic mathematical models and design tools; assessing technical viability and, if applicable, developing methodology and approaches to incorporate wing shape measurements within the vehicle flight control system; and, developing and flight validating approaches to perform active wing shape control using conventional control surfaces and active material concepts.
Wind Tunnel Test of a Risk-Reduction Wing/Fuselage Model to Examine Juncture-Flow Phenomena
NASA Technical Reports Server (NTRS)
Kegerise, Michael A.; Neuhart, Dan H.
2016-01-01
A wing/fuselage wind-tunnel model was tested in the Langley 14- by 22-foot Subsonic Wind Tunnel in preparation for a highly-instrumented Juncture Flow Experiment to be conducted in the same facility. This test, which was sponsored by the NASA Transformational Tool and Technologies Project, is part of a comprehensive set of experimental and computational research activities to develop revolutionary, physics-based aeronautics analysis and design capability. The objectives of this particular test were to examine the surface and off-body flow on a generic wing/body combination to: 1) choose a final wing for a future, highly instrumented model, 2) use the results to facilitate unsteady pressure sensor placement on the model, 3) determine the area to be surveyed with an embedded laser-doppler velocimetry (LDV) system, 4) investigate the primary juncture corner- flow separation region using particle image velocimetry (PIV) to see if the particle seeding is adequately entrained and to examine the structure in the separated region, and 5) to determine the similarity of observed flow features with those predicted by computational fluid dynamics (CFD). This report documents the results of the above experiment that specifically address the first three goals. Multiple wing configurations were tested at a chord Reynolds number of 2.4 million. Flow patterns on the surface of the wings and in the region of the wing/fuselage juncture were examined using oil- flow visualization and infrared thermography. A limited number of unsteady pressure sensors on the fuselage around the wing leading and trailing edges were used to identify any dynamic effects of the horseshoe vortex on the flow field. The area of separated flow in the wing/fuselage juncture near the wing trailing edge was observed for all wing configurations at various angles of attack. All of the test objectives were met. The staff of the 14- by 22-foot Subsonic Wind Tunnel provided outstanding support and delivered exceptional value to the experiment, which exceeded expectations. The results of this test will directly inform the planning for the first of a series of instrumented-model tests at the same Reynolds number. These tests will be performed on a slightly larger-scale model with the selected wing, and will include off-body measurements with LDV and PIV, steady and unsteady pressure measurements, and the flow-visualization techniques that are discussed in this report.
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).
Rhen, Turk; Simmons, Rebecca B.
2016-01-01
Butterfly eyespots are complex morphological traits that can vary in size, shape and color composition even on the same wing surface. Homology among eyespots suggests they share a common developmental basis and function as an integrated unit in response to selection. Despite strong evidence of genetic integration, eyespots can also exhibit modularity or plasticity, indicating an underlying flexibility in pattern development. The extent to which particular eyespots or eyespot color elements exhibit modularity or integration is poorly understood, particularly following exposure to novel conditions. We used perturbation experiments to explore phenotypic correlations among different eyespots and their color elements on the ventral hindwing of V. cardui. Specifically, we identified which eyespots and eyespot features are most sensitive to perturbation by heat shock and injection of heparin—a cold shock mimic. For both treatments, the two central eyespots (3 + 4) were most affected by the experimental perturbations, whereas the outer eyespot border was more resistant to modification than the interior color elements. Overall, the individual color elements displayed a similar response to heat shock across all eyespots, but varied in their response to each other. Graphical modeling also revealed that although eyespots differ morphologically, regulation of eyespot size and colored elements appear to be largely integrated across the wing. Patterns of integration, however, were disrupted following heat shock, revealing that the strength of integration varies across the wing and is strongest between the two central eyespots. These findings support previous observations that document coupling between eyespots 3 + 4 in other nymphalid butterflies. PMID:27560365
Connahs, Heidi; Rhen, Turk; Simmons, Rebecca B
2016-01-01
Butterfly eyespots are complex morphological traits that can vary in size, shape and color composition even on the same wing surface. Homology among eyespots suggests they share a common developmental basis and function as an integrated unit in response to selection. Despite strong evidence of genetic integration, eyespots can also exhibit modularity or plasticity, indicating an underlying flexibility in pattern development. The extent to which particular eyespots or eyespot color elements exhibit modularity or integration is poorly understood, particularly following exposure to novel conditions. We used perturbation experiments to explore phenotypic correlations among different eyespots and their color elements on the ventral hindwing of V. cardui. Specifically, we identified which eyespots and eyespot features are most sensitive to perturbation by heat shock and injection of heparin-a cold shock mimic. For both treatments, the two central eyespots (3 + 4) were most affected by the experimental perturbations, whereas the outer eyespot border was more resistant to modification than the interior color elements. Overall, the individual color elements displayed a similar response to heat shock across all eyespots, but varied in their response to each other. Graphical modeling also revealed that although eyespots differ morphologically, regulation of eyespot size and colored elements appear to be largely integrated across the wing. Patterns of integration, however, were disrupted following heat shock, revealing that the strength of integration varies across the wing and is strongest between the two central eyespots. These findings support previous observations that document coupling between eyespots 3 + 4 in other nymphalid butterflies.
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.
NASA Technical Reports Server (NTRS)
Chaderjian, N. M.
1986-01-01
A computer code is under development whereby the thin-layer Reynolds-averaged Navier-Stokes equations are to be applied to realistic fighter-aircraft configurations. This transonic Navier-Stokes code (TNS) utilizes a zonal approach in order to treat complex geometries and satisfy in-core computer memory constraints. The zonal approach has been applied to isolated wing geometries in order to facilitate code development. Part 1 of this paper addresses the TNS finite-difference algorithm, zonal methodology, and code validation with experimental data. Part 2 of this paper addresses some numerical issues such as code robustness, efficiency, and accuracy at high angles of attack. Special free-stream-preserving metrics proved an effective way to treat H-mesh singularities over a large range of severe flow conditions, including strong leading-edge flow gradients, massive shock-induced separation, and stall. Furthermore, lift and drag coefficients have been computed for a wing up through CLmax. Numerical oil flow patterns and particle trajectories are presented both for subcritical and transonic flow. These flow simulations are rich with complex separated flow physics and demonstrate the efficiency and robustness of the zonal approach.
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.
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.
Peterson, Aidan J.; O'Connor, Michael B.
2013-01-01
Imaginal disc development in Drosophila requires coordinated cellular proliferation and tissue patterning. In our studies of TGFβ superfamily signaling components, we found that a protein null mutation of Smad2, the only Activin subfamily R-Smad in the fruit fly, produces overgrown wing discs that resemble gain of function for BMP subfamily signaling. The wing discs are expanded specifically along the anterior-posterior axis, with increased proliferation in lateral regions. The morphological defect is not observed in mutants for the TGFβ receptor baboon, and epistasis tests showed that baboon is epistatic to Smad2 for disc overgrowth. Rescue experiments indicate that Baboon binding, but not canonical transcription factor activity, of Smad2 is required for normal disc growth. Smad2 mutant discs generate a P-Mad stripe that is narrower and sharper than the normal gradient, and activation targets are correspondingly expressed in narrowed domains. Repression targets of P-Mad are profoundly mis-regulated, with brinker and pentagone reporter expression eliminated in Smad2 mutants. Loss of expression requires a silencer element previously shown to be controlled by BMP signaling. Epistasis experiments show that Baboon, Mad and Schnurri are required to mediate the ectopic silencer output in the absence of Smad2. Taken together, our results show that loss of Smad2 permits promiscuous Baboon activity, which represses genes subject to control by Mad-dependent silencer elements. The absence of Brinker and Pentagone in Smad2 mutants explains the compound wing disc phenotype. Our results highlight the physiological relevance of substrate inhibition of a kinase, and reveal a novel interplay between the Activin and BMP pathways. PMID:23293296
A Wind-Tunnel Investigation of the Development of Lift on Wings in Accelerated Longitudinal Motion
NASA Technical Reports Server (NTRS)
Turner, Thomas R.
1960-01-01
An investigation was made in the Langley 300 MPH 7- by 10-foot tunnel to determine the development of lift on a wing during a simulated constant-acceleration catapult take-off. The investigation included models of a two-dimensional wing, an unswept wing having an aspect ratio of 6, a 35 deg. swept wing having an aspect ratio of 3.05, and a 60 deg. delta wing having an aspect ratio of 2.31. All the wings investigated developed at least 90 percent of their steady-state lift in the first 7 chord lengths of travel. The development of lift was essentially independent of the acceleration when based on chord lengths traveled, and was in qualitative agreement with theory.
Wang, Xiaoyi; Wei, Ke; Yang, Zhongqi; Jennings, David E; Duan, Jian J
2016-05-19
Wing phenotype polymorphism is commonly observed in insects, yet little is known about the influence of environmental cues on the development or expression of the alternative phenotypes. Here, we report how both biotic and abiotic factors affect the wing morph differentiation of a bethylid parasitoid Sclerodermus pupariae. The percentage of winged female parasitoid progeny increased exponentially with temperature between 20 °C to 30 °C. Low intensity light and short-day photoperiod conditions also significantly induced the development of winged morphs. Interestingly, wingless maternal parasitoids produced more winged progeny. Furthermore, the degree of wing dimorphism was significantly influenced by the interactions between light intensity and maternal wing morphs. The percentage of winged female progeny was not significantly influenced by foundress densities, but increased significantly with parasitoid brood sizes. However, the percentage of male progeny increased significantly with the densities of maternal parasitoids. Our findings highlight the phenotypic partitioning of wing morphology and development in the parasitoid S. pupariae under varied environmental cues, and reveal the most favourable conditions for the production of winged females in this bethylid wasp. It is thus possible to increase winged female parasitoid production for the purposes of biological control by manipulation of biotic and abiotic conditions.
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.
2014-01-01
Background The homology of the digits in the bird wing is a high-profile controversy in developmental and evolutionary biology. The embryonic position of the digits cartilages with respect to the primary axis (ulnare and ulna) corresponds to 2, 3, 4, but comparative-evolutionary morphology supports 1, 2, 3. A homeotic frameshift of digit identity in evolution could explain how cells in embryonic positions 2, 3, 4 began developing morphologies 1, 2, 3. Another alternative is that no re-patterning of cell fates occurred, and the primary axis shifted its position by some other mechanism. In the wing, only the anterior digit lacks expression of HoxD10 and HoxD12, resembling digit 1 of other limbs, as predicted by 1, 2, 3. However, upon loss of digit 1 in evolution, the most anterior digit 2 could have lost their expression, deceitfully resembling a digit 1. To test this notion, we observed HoxD10 and HoxD12 in a limb where digit 2 is the most anterior digit: The rabbit foot. We also explored whether early inhibition of Shh signalling in the embryonic wing bud induces an experimental homeotic frameshift, or an experimental axis shift. We tested these hypotheses using DiI injections to study the fate of cells in these experimental wings. Results We found strong transcription of HoxD10 and HoxD12 was present in the most anterior digit 2 of the rabbit foot. Thus, we found no evidence to question the use of HoxD expression as support for 1, 2, 3. When Shh signalling in early wing buds is inhibited, our fate maps demonstrate that an experimental homeotic frameshift is induced. Conclusion Along with comparative morphology, HoxD expression provides strong support for 1, 2, 3 identity of wing digits. As an explanation for the offset 2, 3, 4 embryological position, the homeotic frameshift hypothesis is consistent with known mechanisms of limb development, and further proven to be experimentally possible. In contrast, the underlying mechanisms and experimental plausibility of an axis shift remain unclear. PMID:24725625
Salinas-Saavedra, Miguel; Gonzalez-Cabrera, Cristian; Ossa-Fuentes, Luis; Botelho, Joao F; Ruiz-Flores, Macarena; Vargas, Alexander O
2014-04-12
The homology of the digits in the bird wing is a high-profile controversy in developmental and evolutionary biology. The embryonic position of the digits cartilages with respect to the primary axis (ulnare and ulna) corresponds to 2, 3, 4, but comparative-evolutionary morphology supports 1, 2, 3. A homeotic frameshift of digit identity in evolution could explain how cells in embryonic positions 2, 3, 4 began developing morphologies 1, 2, 3. Another alternative is that no re-patterning of cell fates occurred, and the primary axis shifted its position by some other mechanism. In the wing, only the anterior digit lacks expression of HoxD10 and HoxD12, resembling digit 1 of other limbs, as predicted by 1, 2, 3. However, upon loss of digit 1 in evolution, the most anterior digit 2 could have lost their expression, deceitfully resembling a digit 1. To test this notion, we observed HoxD10 and HoxD12 in a limb where digit 2 is the most anterior digit: The rabbit foot. We also explored whether early inhibition of Shh signalling in the embryonic wing bud induces an experimental homeotic frameshift, or an experimental axis shift. We tested these hypotheses using DiI injections to study the fate of cells in these experimental wings. We found strong transcription of HoxD10 and HoxD12 was present in the most anterior digit 2 of the rabbit foot. Thus, we found no evidence to question the use of HoxD expression as support for 1, 2, 3. When Shh signalling in early wing buds is inhibited, our fate maps demonstrate that an experimental homeotic frameshift is induced. Along with comparative morphology, HoxD expression provides strong support for 1, 2, 3 identity of wing digits. As an explanation for the offset 2, 3, 4 embryological position, the homeotic frameshift hypothesis is consistent with known mechanisms of limb development, and further proven to be experimentally possible. In contrast, the underlying mechanisms and experimental plausibility of an axis shift remain unclear.
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
Development of direct-inverse 3-D methods for applied transonic aerodynamic wing design and analysis
NASA Technical Reports Server (NTRS)
Carlson, Leland A.
1989-01-01
An inverse wing design method was developed around an existing transonic wing analysis code. The original analysis code, TAWFIVE, has as its core the numerical potential flow solver, FLO30, developed by Jameson and Caughey. Features of the analysis code include a finite-volume formulation; wing and fuselage fitted, curvilinear grid mesh; and a viscous boundary layer correction that also accounts for viscous wake thickness and curvature. The development of the inverse methods as an extension of previous methods existing for design in Cartesian coordinates is presented. Results are shown for inviscid wing design cases in super-critical flow regimes. The test cases selected also demonstrate the versatility of the design method in designing an entire wing or discontinuous sections of a wing.
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 ).
Wingless promotes proliferative growth in a gradient-independent manner.
Baena-Lopez, Luis Alberto; Franch-Marro, Xavier; Vincent, Jean-Paul
2009-10-06
Morphogens form concentration gradients that organize patterns of cells and control growth. It has been suggested that, rather than the intensity of morphogen signaling, it is its gradation that is the relevant modulator of cell proliferation. According to this view, the ability of morphogens to regulate growth during development depends on their graded distributions. Here, we describe an experimental test of this model for Wingless, one of the key organizers of wing development in Drosophila. Maximal Wingless signaling suppresses cellular proliferation. In contrast, we found that moderate and uniform amounts of exogenous Wingless, even in the absence of endogenous Wingless, stimulated proliferative growth. Beyond a few cell diameters from the source, Wingless was relatively constant in abundance and thus provided a homogeneous growth-promoting signal. Although morphogen signaling may act in combination with as yet uncharacterized graded growth-promoting pathways, we suggest that the graded nature of morphogen signaling is not required for proliferation, at least in the developing Drosophila wing, during the main period of growth.
NASA Astrophysics Data System (ADS)
Perkins, David A.; Reed, John L., Jr.; Havens, Ernie
2004-07-01
Cornerstone Research Group, Inc. (CRG), with specific no-cost guidance and support from Lockheed Martin, proposed to significantly increase the capability of loitering Unmanned Air Vehicles (UAVs) by developing a unique adaptive wing structure. This technology will offer significant operational benefit to air vehicles of this type currently under development. The development of this adaptive wing structure will enable such aircraft to adapt their wing configuration to maximize efficiency in each flight regime experienced during their mission. Additionally, the benefits of this development program will enhance the agility and maneuverability of the vehicle; therefore increasing its mission capability. The specific morphing ability CRG proposed to develop was a controlled expansion and contraction of the wing chord, which increases the wing planform area and therefore the lift produced. CRG proved feasibility of this concept and developed a sub-scale prototype integrating smart materials developed at CRG.
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.
A wing expressed sequence tag resource for Bicyclus anynana butterflies, an evo-devo model
Beldade, Patrícia; Rudd, Stephen; Gruber, Jonathan D; Long, Anthony D
2006-01-01
Background Butterfly wing color patterns are a key model for integrating evolutionary developmental biology and the study of adaptive morphological evolution. Yet, despite the biological, economical and educational value of butterflies they are still relatively under-represented in terms of available genomic resources. Here, we describe an Expression Sequence Tag (EST) project for Bicyclus anynana that has identified the largest available collection to date of expressed genes for any butterfly. Results By targeting cDNAs from developing wings at the stages when pattern is specified, we biased gene discovery towards genes potentially involved in pattern formation. Assembly of 9,903 ESTs from a subtracted library allowed us to identify 4,251 genes of which 2,461 were annotated based on BLAST analyses against relevant gene collections. Gene prediction software identified 2,202 peptides, of which 215 longer than 100 amino acids had no homology to any known proteins and, thus, potentially represent novel or highly diverged butterfly genes. We combined gene and Single Nucleotide Polymorphism (SNP) identification by constructing cDNA libraries from pools of outbred individuals, and by sequencing clones from the 3' end to maximize alignment depth. Alignments of multi-member contigs allowed us to identify over 14,000 putative SNPs, with 316 genes having at least one high confidence double-hit SNP. We furthermore identified 320 microsatellites in transcribed genes that can potentially be used as genetic markers. Conclusion Our project was designed to combine gene and sequence polymorphism discovery and has generated the largest gene collection available for any butterfly and many potential markers in expressed genes. These resources will be invaluable for exploring the potential of B. anynana in particular, and butterflies in general, as models in ecological, evolutionary, and developmental genetics. PMID:16737530
NASA Technical Reports Server (NTRS)
Hsu, C.-H.; Lan, C. E.
1985-01-01
Wing rock is one type of lateral-directional instabilities at high angles of attack. To predict wing rock characteristics and to design airplanes to avoid wing rock, parameters affecting wing rock characteristics must be known. A new nonlinear aerodynamic model is developed to investigate the main aerodynamic nonlinearities causing wing rock. In the present theory, the Beecham-Titchener asymptotic method is used to derive expressions for the limit-cycle amplitude and frequency of wing rock from nonlinear flight dynamics equations. The resulting expressions are capable of explaining the existence of wing rock for all types of aircraft. Wing rock is developed by negative or weakly positive roll damping, and sustained by nonlinear aerodynamic roll damping. Good agreement between theoretical and experimental results is obtained.
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.
Muscle function in avian flight: achieving power and control
Biewener, Andrew A.
2011-01-01
Flapping flight places strenuous requirements on the physiological performance of an animal. Bird flight muscles, particularly at smaller body sizes, generally contract at high frequencies and do substantial work in order to produce the aerodynamic power needed to support the animal's weight in the air and to overcome drag. This is in contrast to terrestrial locomotion, which offers mechanisms for minimizing energy losses associated with body movement combined with elastic energy savings to reduce the skeletal muscles' work requirements. Muscles also produce substantial power during swimming, but this is mainly to overcome body drag rather than to support the animal's weight. Here, I review the function and architecture of key flight muscles related to how these muscles contribute to producing the power required for flapping flight, how the muscles are recruited to control wing motion and how they are used in manoeuvring. An emergent property of the primary flight muscles, consistent with their need to produce considerable work by moving the wings through large excursions during each wing stroke, is that the pectoralis and supracoracoideus muscles shorten over a large fraction of their resting fibre length (33–42%). Both muscles are activated while being lengthened or undergoing nearly isometric force development, enhancing the work they perform during subsequent shortening. Two smaller muscles, the triceps and biceps, operate over a smaller range of contractile strains (12–23%), reflecting their role in controlling wing shape through elbow flexion and extension. Remarkably, pigeons adjust their wing stroke plane mainly via changes in whole-body pitch during take-off and landing, relative to level flight, allowing their wing muscles to operate with little change in activation timing, strain magnitude and pattern. PMID:21502121
NASA Technical Reports Server (NTRS)
Bertin, J. J.; Graumann, B. W.
1973-01-01
Numerical codes were developed to calculate the two dimensional flow field which results when supersonic flow encounters double wedge configurations whose angles are such that a type 4 pattern occurs. The flow field model included the shock interaction phenomena for a delta wing orbiter. Two numerical codes were developed, one which used the perfect gas relations and a second which incorporated a Mollier table to define equilibrium air properties. The two codes were used to generate theoretical surface pressure and heat transfer distributions for velocities from 3,821 feet per second to an entry condition of 25,000 feet per second.
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.
NASA Technical Reports Server (NTRS)
Parsons-Wingerter, P.; Weitzel, Alexander; Vyas, R. J.; Murray, M. C.; Vickerman, M. B.; Bhattacharya, S.; Wyatt, S. E.
2016-01-01
One fundamental requirement shared by humans with all higher terrestrial life forms, including other vertebrates, insects, and higher land plants, 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. A unifying perspective is that vascular patterning offers a useful readout of molecular signaling that necessarily integrates these complex pathways. VESGEN has elucidated changes in vascular pattern resulting from inflammatory, 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. Vascularrelated 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 spatial and dynamic dimensions 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. 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.
NASA Technical Reports Server (NTRS)
Gern, Frank H.; Naghshineh, Amir H.; Sulaeman, Erwin; Kapania, Rakesh K.; Haftka, Raphael T.
2000-01-01
This paper describes a structural and aeroelastic model for wing sizing and weight calculation of a strut-braced wing. The wing weight is calculated using a newly developed structural weight analysis module considering the special nature of strut-braced wings. A specially developed aeroelastic model enables one to consider wing flexibility and spanload redistribution during in-flight maneuvers. The structural model uses a hexagonal wing-box featuring skin panels, stringers, and spar caps, whereas the aerodynamics part employs a linearized transonic vortex lattice method. Thus, the wing weight may be calculated from the rigid or flexible wing spanload. The calculations reveal the significant influence of the strut on the bending material weight of the wing. The use of a strut enables one to design a wing with thin airfoils without weight penalty. The strut also influences wing spanload and deformations. Weight savings are not only possible by calculation and iterative resizing of the wing structure according to the actual design loads. Moreover, as an advantage over the cantilever wing, employment of the strut twist moment for further load alleviation leads to increased savings in structural weight.
Patterning of wound-induced intercellular Ca2+ flashes in a developing epithelium
NASA Astrophysics Data System (ADS)
Narciso, Cody; Wu, Qinfeng; Brodskiy, Pavel; Garston, George; Baker, Ruth; Fletcher, Alexander; Zartman, Jeremiah
2015-10-01
Differential mechanical force distributions are increasingly recognized to provide important feedback into the control of an organ’s final size and shape. As a second messenger that integrates and relays mechanical information to the cell, calcium ions (Ca2+) are a prime candidate for providing important information on both the overall mechanical state of the tissue and resulting behavior at the individual-cell level during development. Still, how the spatiotemporal properties of Ca2+ transients reflect the underlying mechanical characteristics of tissues is still poorly understood. Here we use an established model system of an epithelial tissue, the Drosophila wing imaginal disc, to investigate how tissue properties impact the propagation of Ca2+ transients induced by laser ablation. The resulting intercellular Ca2+ flash is found to be mediated by inositol 1,4,5-trisphosphate and depends on gap junction communication. Further, we find that intercellular Ca2+ transients show spatially non-uniform characteristics across the proximal-distal axis of the larval wing imaginal disc, which exhibit a gradient in cell size and anisotropy. A computational model of Ca2+ transients is employed to identify the principle factors explaining the spatiotemporal patterning dynamics of intercellular Ca2+ flashes. The relative Ca2+ flash anisotropy is principally explained by local cell shape anisotropy. Further, Ca2+ velocities are relatively uniform throughout the wing disc, irrespective of cell size or anisotropy. This can be explained by the opposing effects of cell diameter and cell elongation on intercellular Ca2+ propagation. Thus, intercellular Ca2+ transients follow lines of mechanical tension at velocities that are largely independent of tissue heterogeneity and reflect the mechanical state of the underlying tissue.
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 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.
Flow structure and aerodynamic performance of a hovering bristled wing in low Re
NASA Astrophysics Data System (ADS)
Lee, Seunghun; Lahooti, Mohsen; Kim, Daegyoum
2017-11-01
Previous studies on a bristled wing have mainly focused on simple kinematics of the wing such as translation or rotation. The aerodynamic performance of a bristled wing in a quasi-steady phase is known to be comparable to that of a smooth wing without a gap because shear layers in the gaps of the bristled wing are sufficiently developed to block the gaps. However, we point out that, in the starting transient phase where the shear layers are not fully developed, the force generation of a bristled wing is not as efficient as that of a quasi-steady state. The performance in the transient phase is important to understand the aerodynamics of a bristled wing in an unsteady motion. In the hovering motion, due to repeated stroke reversals, the formation and development of shear layers inside the gaps is repeated in each stroke. In this study, a bristled wing in hovering is numerically investigated in the low Reynolds number of O(10). We especially focus on the development of shear layers during a stroke reversal and its effect on the overall propulsive performance. Although the aerodynamic force generation is slightly reduced due to the gap vortices, the asymmetric behavior of vortices in a gap between bristles during a stroke reversal makes the bristled wing show higher lift to drag ratio than a smooth wing.
Overview of the ARPA/WL Smart Structures and Materials Development-Smart Wing contract
NASA Astrophysics Data System (ADS)
Kudva, Jayanth N.; Jardine, A. Peter; Martin, Christopher A.; Appa, Kari
1996-05-01
While the concept of an adaptive aircraft wing, i.e., a wing whose shape parameters such as camber, wing twist, and thickness can be varied to optimize the wing shape for various flight conditions, has been extensively studied, the complexity and weight penalty of the actuation mechanisms have precluded their practical implementation. Recent development of sensors and actuators using smart materials could potentially alleviate the shortcomings of prior designs, paving the way for a practical, `smart' adaptive wing which responds to changes in flight and environmental conditions by modifying its shape to provide optimal performance. This paper presents a summary of recent work done on adaptive wing designs under an on-going ARPA/WL contract entitled `Smart Structures and Materials Development--Smart Wing.' Specifically, the design, development and planned wind tunnel testing of a 16% model representative of a fighter aircraft wing and incorporating the following features, are discussed: (1) a composite wing torque box whose span-wise twist can be varied by activating built-in shape memory alloy (SMA) torque tubes to provide increased lift and enhanced maneuverability at multiple flight conditions, (2) trailing edge control surfaces deployed using composite SMA actuators to provide smooth, hingeless aerodynamic surfaces, and (3) a suite of fiber optic sensors integrated into the wing skin which provide real-time strain and pressure data to a feedback control system.
Nonplanar wing load-line and slender wing theory
NASA Technical Reports Server (NTRS)
Deyoung, J.
1977-01-01
Nonplanar load line, slender wing, elliptic wing, and infinite aspect ratio limit loading theories are developed. These are quasi two dimensional theories but satisfy wing boundary conditions at all points along the nonplanar spanwise extent of the wing. These methods are applicable for generalized configurations such as the laterally nonplanar wing, multiple nonplanar wings, or wing with multiple winglets of arbitrary shape. Two dimensional theory infers simplicity which is practical when analyzing complicated configurations. The lateral spanwise distribution of angle of attack can be that due to winglet or control surface deflection, wing twist, or induced angles due to multiwings, multiwinglets, ground, walls, jet or fuselage. In quasi two dimensional theory the induced angles due to these extra conditions are likewise determined for two dimensional flow. Equations are developed for the normal to surface induced velocity due to a nonplanar trailing vorticity distribution. Application examples are made using these methods.
Nonlinear Aerodynamics and the Design of Wing Tips
NASA Technical Reports Server (NTRS)
Kroo, Ilan
1991-01-01
The analysis and design of wing tips for fixed wing and rotary wing aircraft still remains part art, part science. Although the design of airfoil sections and basic planform geometry is well developed, the tip regions require more detailed consideration. This is important because of the strong impact of wing tip flow on wing drag; although the tip region constitutes a small portion of the wing, its effect on the drag can be significant. The induced drag of a wing is, for a given lift and speed, inversely proportional to the square of the wing span. Concepts are proposed as a means of reducing drag. Modern computational methods provide a tool for studying these issues in greater detail. The purpose of the current research program is to improve the understanding of the fundamental issues involved in the design of wing tips and to develop the range of computational and experimental tools needed for further study of these ideas.
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
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
Study on utilization of advanced composites in commercial aircraft wing structures, volume 2
NASA Technical Reports Server (NTRS)
Sakata, I. F.; Ostrom, R. B.
1978-01-01
A plan is defined for a composite wing development effort which will assist commercial transport manufacturers in reaching a level of technology readiness where the utilization of composite wing structure is a cost competitive option for a new aircraft production plan. The recommended development effort consists of two programs: a joint government/industry material development program and a wing structure development program. Both programs are described in detail.
Roughness Effects on the Formation of a Leading Edge Vortex
NASA Astrophysics Data System (ADS)
Elliott, Cassidy; Lang, Amy; Wahidi, Redha; Wilroy, Jacob
2016-11-01
Microscopic scales cover the wings of Monarch butterflies, creating a patterned surface. This patterning is an important natural flow control mechanism that is thought to delay the growth of the leading edge vortex (LEV) produced by the flapping motion of a wing. The increased skin friction caused by the scales leads to a weaker LEV being shed into the butterfly's wake, lessening drag and increasing flight efficiency. To test this theory, a plate of random roughness was designed in SolidWorks and printed on the Objet 30 Pro 3D printer. A 2x3x5 cubic foot tow tank was used to test the rough plate at Reynold's numbers of 1500, 3000, and 6000 (velocities of 8, 16, and 32 mm/s) at an angle of attack of 45 degrees. Particle Image Velocimetry (PIV) captured images of the LEV generated by the plate when towed upwards through the particle-seeded flow. Codes written in MatLab were used to automatically track and determine the strength of the LEV. Circulation values for the randomly-rough plate were then compared to the same values generated in a previous experiment that used a smooth plate and a grooved plate to determine the effect of the patterning on vortex development. Funding provided by NSF REU site Grant EEC 1358991 and CBET 1628600.
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.
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.
Optimal flapping wing for maximum vertical aerodynamic force in hover: twisted or flat?
Phan, Hoang Vu; Truong, Quang Tri; Au, Thi Kim Loan; Park, Hoon Cheol
2016-07-08
This work presents a parametric study, using the unsteady blade element theory, to investigate the role of twist in a hovering flapping wing. For the investigation, a flapping-wing system was developed to create a wing motion of large flapping amplitude. Three-dimensional kinematics of a passively twisted wing, which is capable of creating a linearly variable geometric angle of attack (AoA) along the wingspan, was measured during the flapping motion and used for the analysis. Several negative twist or wash-out configurations with different values of twist angle, which is defined as the difference in the average geometric AoAs at the wing root and the wing tip, were obtained from the measured wing kinematics through linear interpolation and extrapolation. The aerodynamic force generation and aerodynamic power consumption of these twisted wings were obtained and compared with those of flat wings. For the same aerodynamic power consumption, the vertical aerodynamic forces produced by the negatively twisted wings are approximately 10%-20% less than those produced by the flat wings. However, these twisted wings require approximately 1%-6% more power than flat wings to produce the same vertical force. In addition, the maximum-force-producing twisted wing, which was found to be the positive twist or wash-in configuration, was used for comparison with the maximum-force-producing flat wing. The results revealed that the vertical aerodynamic force and aerodynamic power consumption of the two types of wings are almost identical for the hovering condition. The power loading of the positively twisted wing is only approximately 2% higher than that of the maximum-force-producing flat wing. Thus, the flat wing with proper wing kinematics (or wing rotation) can be regarded as a simple and efficient candidate for the development of hovering flapping-wing micro air vehicle.
NASA Astrophysics Data System (ADS)
Perez-Rosado, Ariel; Gehlhar, Rachel D.; Nolen, Savannah; Gupta, Satyandra K.; Bruck, Hugh A.
2015-06-01
Currently, flapping wing unmanned aerial vehicles (a.k.a., ornithopters or robotic birds) sustain very short duration flight due to limited on-board energy storage capacity. Therefore, energy harvesting elements, such as flexible solar cells, need to be used as materials in critical components, such as wing structures, to increase operational performance. In this paper, we describe a layered fabrication method that was developed for realizing multifunctional composite wings for a unique robotic bird we developed, known as Robo Raven, by creating compliant wing structure from flexible solar cells. The deformed wing shape and aerodynamic lift/thrust loads were characterized throughout the flapping cycle to understand wing mechanics. A multifunctional performance analysis was developed to understand how integration of solar cells into the wings influences flight performance under two different operating conditions: (1) directly powering wings to increase operation time, and (2) recharging batteries to eliminate need for external charging sources. The experimental data is then used in the analysis to identify a performance index for assessing benefits of multifunctional compliant wing structures. The resulting platform, Robo Raven III, was the first demonstration of a robotic bird that flew using energy harvested from solar cells. We developed three different versions of the wing design to validate the multifunctional performance analysis. It was also determined that residual thrust correlated to shear deformation of the wing induced by torsional twist, while biaxial strain related to change in aerodynamic shape correlated to lift. It was also found that shear deformation of the solar cells induced changes in power output directly correlating to thrust generation associated with torsional deformation. Thus, it was determined that multifunctional solar cell wings may be capable of three functions: (1) lightweight and flexible structure to generate aerodynamic forces, (2) energy harvesting to extend operational time and autonomy, and (3) sensing of an aerodynamic force associated with wing deformation.
NASA Astrophysics Data System (ADS)
Mainwaring, David E.; Nguyen, Song Ha; Webb, Hayden; Jakubov, Timur; Tobin, Mark; Lamb, Robert N.; Wu, Alex H.-F.; Marchant, Richard; Crawford, Russell J.; Ivanova, Elena P.
2016-03-01
While insect wings are widely recognised as multi-functional, recent work showed that this extends to extensive bactericidal activity brought about by cell deformation and lysis on the wing nanotopology. We now quantitatively show that subtle changes to this topography result in substantial changes in bactericidal activity that are able to span an order of magnitude. Notably, the chemical composition of the lipid nanopillars was seen by XPS and synchrotron FTIR microspectroscopy to be similar across these activity differences. Modelling the interaction between bacterial cells and the wing surface lipids of 3 species of dragonflies, that inhabit similar environments, but with distinctly different behavioural repertoires, provided the relationship between surface structure and antibacterial functionality. In doing so, these principal behavioural patterns correlated with the demands for antimicrobial efficiency dictated by differences in their foraging strategies. This work now reveals a new feature in the design elegance of natural multi-functional surfaces as well providing insights into the bactericidal mechanism underlying inherently antimicrobial materials, while suggesting that nanotopology is related to the evolutionary development of a species through the demands of its behavioural repertoire. The underlying relationship between the processes of wetting, adhesion and capillarity of the lipid nanopillars and bactericidal efficiency suggests new prospects for purely mechano-responsive antibacterial surfaces.While insect wings are widely recognised as multi-functional, recent work showed that this extends to extensive bactericidal activity brought about by cell deformation and lysis on the wing nanotopology. We now quantitatively show that subtle changes to this topography result in substantial changes in bactericidal activity that are able to span an order of magnitude. Notably, the chemical composition of the lipid nanopillars was seen by XPS and synchrotron FTIR microspectroscopy to be similar across these activity differences. Modelling the interaction between bacterial cells and the wing surface lipids of 3 species of dragonflies, that inhabit similar environments, but with distinctly different behavioural repertoires, provided the relationship between surface structure and antibacterial functionality. In doing so, these principal behavioural patterns correlated with the demands for antimicrobial efficiency dictated by differences in their foraging strategies. This work now reveals a new feature in the design elegance of natural multi-functional surfaces as well providing insights into the bactericidal mechanism underlying inherently antimicrobial materials, while suggesting that nanotopology is related to the evolutionary development of a species through the demands of its behavioural repertoire. The underlying relationship between the processes of wetting, adhesion and capillarity of the lipid nanopillars and bactericidal efficiency suggests new prospects for purely mechano-responsive antibacterial surfaces. Electronic supplementary information (ESI) available. See DOI: 10.1039/c5nr08542j
NASA Technical Reports Server (NTRS)
Kulfan, R. M.; Neumann, F. D.; Nisbet, J. W.; Mulally, A. R.; Murakami, J. K.; Noble, E. C.; Mcbarron, J. P.; Stalter, J. L.; Gimmestad, D. W.; Sussman, M. B.
1973-01-01
An initial design study of high-transonic-speed transport aircraft has been completed. Five different design concepts were developed. These included fixed swept wing, variable-sweep wing, delta wing, double-fuselage yawed-wing, and single-fuselage yawed-wing aircraft. The boomless supersonic design objectives of range=5560 Km (3000 nmi), payload-18 143 kg (40 000lb), Mach=1.2, and FAR Part 36 aircraft noise levels were achieved by the single-fuselage yawed-wing configuration with a gross weight of 211 828 Kg (467 000 lb). A noise level of 15 EPNdB below FAR Part 36 requirements was obtained with a gross weight increase to 226 796 Kg (500 000 lb). Although wing aeroelastic divergence was a primary design consideration for the yawed-wing concepts, the graphite-epoxy wings of this study were designed by critical gust and maneuver loads rather than by divergence requirements. The transonic nacelle drag is shown to be very sensitive to the nacelle installation. A six-degree-of-freedom dynamic stability analysis indicated that the control coordination and stability augmentation system would require more development than for a symmetrical airplane but is entirely feasible. A three-phase development plan is recommended to establish the full potential of the yawed-wing concept.
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
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.
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.
The Aerodynamic Optimization of Wings at Subsonic Speeds and the Influence of Wingtip Design. Thesis
NASA Technical Reports Server (NTRS)
Zimmer, H.
1987-01-01
Some of the objectives of modern aircraft development are related to the achievement of reduced fuel consumption and aircraft noise. This investigation is mainly concerned with the aerodynamic aspects of aircraft development, i.e., reduction of induced drag. New studies of wing design, and in particular wing tips, are considered. Induced drag is important since, in cruising flight, it accounts for approximately one-third of the entire drag for the aircraft, and one-half while climbing. A survey is presented for the wing geometries and wing tip designs studied, and theoretical investigations of different planar wings with systematically varied wing tip forms are conducted. Attention is also paid to a theoretical study of some planar and nonplanar wings and their comparison with experimental data.
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
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
Higashi, C H V; Bressan, A
2013-10-01
We examined the population dynamics of the corn planthopper Peregrinus maidis (Ashmead) (Hemiptera: Delphacidae) throughout a cycle of corn (Zea mays L.) production on plants with or without symptoms of maize mosaic virus (MMV) (Rhabdoviridae: Nucleorhabdovirus) infection. Our results indicate that the timing of MMV plant infection greatly influenced the planthopper's host plant colonization patterns. Corn plants that expressed symptoms of MMV infection early in the crop cycle (28 d after planting) harbored, on average, 40 and 48% fewer planthoppers than plants that expressed symptoms of MMV infection later in the crop cycle (49 d after planting) and asymptomatic plants, respectively. We also observed a change in the number of brachypterous (short-wing type) and macropterous (long-wing type) winged forms produced; plants expressing early symptoms of MMV infection harbored, on average, 41 and 47% more of the brachypterous form than plants with late infections of MMV and plants with no symptoms of MMV, respectively. Furthermore, we determined the rates of MMV-infected planthoppers relative to their wing morphology (macropterous or brachypterous) and gender. MMV infection was 5 and 12% higher in females than in males in field and greenhouse experiments, respectively; however, these differences were not significantly different. This research provides evidence that MMV similarly infects P. maidis planthoppers regardless of the gender and wing morphotype. These results also suggest that the timing of symptom development greatly affects the population dynamics of the planthopper vector, and likely has important consequences for the dynamics of the disease in the field.
USDA-ARS?s Scientific Manuscript database
The sand cricket, Gryllus firmus, is a wing-dimorphic species with long-wing (LW) and short wing (LW) morphs. The LW forms have very well developed wings and flight muscles and their SW counterparts have reduced wings and flight muscles, coupled with greater resource allocations to reproduction. Thi...
Application of slender wing benefits to military aircraft
NASA Technical Reports Server (NTRS)
Polhamus, E. C.
1983-01-01
A review is provided of aerodynamic research conducted at the Langley Research Center with respect to the application of slender wing benefits in the design of high-speed military aircraft, taking into account the supersonic performance and leading-edge vortex flow associated with very highly sweptback wings. The beginning of the development of modern classical swept wing jet aircraft is related to the German Me 262 project during World War II. In the U.S., a theoretical study conducted by Jones (1945) pointed out the advantages of the sweptback wing concept. Developments with respect to variable sweep wings are discussed, taking into account early research in 1946, a joint program of the U.S. with the United Kingdom, the tactical aircraft concept, and the important part which the Langley variable-sweep research program played in the development of the F-111, F-14, and B-1. Attention is also given to hybrid wings, vortex flow theory development, and examples of flow design technology.
Butterflies regulate wing temperatures using radiative cooling
NASA Astrophysics Data System (ADS)
Tsai, Cheng-Chia; Shi, Norman Nan; Ren, Crystal; Pelaez, Julianne; Bernard, Gary D.; Yu, Nanfang; Pierce, Naomi
2017-09-01
Butterfly wings are live organs embedded with multiple sensory neurons and, in some species, with pheromoneproducing cells. The proper function of butterfly wings demands a suitable temperature range, but the wings can overheat quickly in the sun due to their small thermal capacity. We developed an infrared technique to map butterfly wing temperatures and discovered that despite the wings' diverse visible colors, regions of wings that contain live cells are the coolest, resulting from the thickness of the wings and scale nanostructures. We also demonstrated that butterflies use behavioral traits to prevent overheating of their 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.
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.
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.
SMA actuators for morphing wings
NASA Astrophysics Data System (ADS)
Brailovski, V.; Terriault, P.; Georges, T.; Coutu, D.
An experimental morphing laminar wing was developed to prove the feasibility of aircraft fuel consumption reduction through enhancement of the laminar flow regime over the wing extrados. The morphing wing prototype designed for subsonic cruise flight conditions (Mach 0.2 … 0.3; angle of attack - 1 … +2∘), combines three principal subsystems: (1) flexible extrados, (2) rigid intrados and (3) an actuator group located inside the wing box. The morphing capability of the wing relies on controlled deformation of the wing extrados under the action of shape memory alloys (SMA) actuators. A coupled fluid-structure model of the morphing wing was used to evaluate its mechanical and aerodynamic performances in different flight conditions. A 0.5 m chord and 1 m span prototype of the morphing wing was tested in a subsonic wind tunnel. In this work, SMA actuators for morphing wings were modeled using a coupled thermo-mechanical finite element model and they were windtunnel validated. If the thermo-mechanical model of SMA actuators presented in this work is coupled with the previously developed structureaerodynamic model of the morphing wing, it could serve for the optimization of the entire morphing wing system.
NASA Technical Reports Server (NTRS)
Runnels, J. N.; Gupfa, A.
1973-01-01
Augmentor wing ducting system studies conducted on a valveless system configuration that provides cruise thrust from the augmentor nozzles have shown that most of the duct system pressure loss would occur in the strut-wing duct y-junction and the wing duct-augmentor lobe nozzles. These components were selected for development testing over a range of duct Mach numbers and pressure ratios to provide a technical basis for predicting installed wing thrust loading and for evaluating design wing loading of a particular wing aspect ratios. The flow characteristics of ducting components with relatively high pressure loss coefficients were investigated. The turbulent pressure fluctuations associated with flows at high Mach numbers were analyzed to evaluate potential duct fatigue problems.
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
Zhang, Jing; Blessing, Danso; Wu, Chenyu; Liu, Na; Li, Juan; Qin, Sheng
2017-01-01
Wings of Bombyx mori (B. mori) develop from the primordium, and different B. mori strains have different wing types. In order to identify the key factors influencing B. mori wing development, we chose strains P50 and U11, which are typical for normal wing and minute wing phenotypes, respectively. We dissected the wing disc on the 1st-day of wandering stage (P50D1 and U11D1), 2nd-day of wandering stage (P50D2 and U11D2), and 3rd-day of wandering stage (P50D3 and U11D3). Subsequently, RNA-sequencing (RNA-Seq) was performed on both strains in order to construct their gene expression profiles. P50 exhibited 628 genes differentially expressed to U11, 324 up-regulated genes, and 304 down-regulated genes. Five enriched gene ontology (GO) terms were identified by GO enrichment analysis based on these differentially expressed genes (DEGs). KEGG enrichment analysis results showed that the DEGs were enriched in five pathways; of these, we identified three pathways related to the development of wings. The three pathways include amino sugar and nucleotide sugar metabolism pathway, proteasome signaling pathway, and the Hippo signaling pathway. The representative genes in the enrichment pathways were further verified by quantitative real-time reverse transcription polymerase chain reaction (qRT-PCR). The RNA-Seq and qRT-PCR results were largely consistent with each other. Our results also revealed that the significantly different genes obtained in our study might be involved in the development of the size of B. mori wings. In addition, several KEGG enriched pathways might be involved in the regulation of the pathways of wing formation. These results provide a basis for further research of wing development in B. mori. PMID:28617839
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
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.
Fiber Optic Wing Shape Sensing on NASA's Ikhana UAV
NASA Technical Reports Server (NTRS)
Richards, Lance; Parker, Allen R.; Ko, William L.; Piazza, Anthony
2008-01-01
Fiber Optic Wing Shape Sensing on Ikhana involves five major areas 1) Algorithm development: Local-strain-to-displacement algorithms have been developed for complex wing shapes for real-time implementation (NASA TP-2007-214612, patent application submitted) 2) FBG system development: Dryden advancements to fiber optic sensing technology have increased data sampling rates to levels suitable for monitoring structures in flight (patent application submitted) 3) Instrumentation: 2880 FBG strain sensors have been successfully installed on the Ikhana wings 4) Ground Testing: Fiber optic wing shape sensing methods for high aspect ratio UAVs have been validated through extensive ground testing in Dryden s Flight Loads Laboratory 5) Flight Testing: Real time fiber Bragg strain measurements successfully acquired and validated in flight (4/28/2008) Real-time fiber optic wing shape sensing successfully demonstrated in flight
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)
Sakata, I. F.; Ostrom, R. B.; Cardinale, S. V.
1978-01-01
The effort required by commercial transport manufacturers to accomplish the transition from current construction materials and practices to extensive use of composites in aircraft wings was investigated. The engineering and manufacturing disciplines which normally participate in the design, development, and production of an aircraft were employed to ensure that all of the factors that would enter a decision to commit to production of a composite wing structure were addressed. A conceptual design of an advanced technology reduced energy aircraft provided the framework for identifying and investigating unique design aspects. A plan development effort defined the essential technology needs and formulated approaches for effecting the required wing development. The wing development program plans, resource needs, and recommendations are summarized.
Insect-inspired wing actuation structures based on ring-type resonators
NASA Astrophysics Data System (ADS)
Bolsman, Caspar T.; Goosen, Johannes F. L.; van Keulen, Fred
2008-03-01
In this paper, we illustrate and study the opportunities of resonant ring type structures as wing actuation mechanisms for a flapping wing Micro Air Vehicle (MAV). Various design alternatives are presented and studied based on computational and physical models. Insects provide an excellent source of inspiration for the development of the wing actuation mechanisms for flapping wing MAVs. The insect thorax is a structure which in essence provides a mechanism to couple the wing muscles to the wings while offering weight reduction through application of resonance, using tailored elasticity. The resonant properties of the thorax are a very effective way to reducing the power expenditure of wing movement. The wing movement itself is fairly complex and is guided by a set of control muscles and thoracic structures which are present in proximity of the wing root. The development of flapping wing MAVs requires a move away from classical structures and actuators. The use of gears and rotational electric motors is hard to justify at the small scale. Resonant structures provide a large design freedom whilst also providing various options for actuation. The move away from deterministic mechanisms offers possibilities for mass reduction.
Development of multi-element active aerodynamics for the formula sae car
NASA Astrophysics Data System (ADS)
Merkel, James Patrick
This thesis focuses on the design, development, and implementation of an active aerodynamics system on 2013 Formula SAE car. The aerodynamics package itself consists of five element front and rear wings as well as an under body diffuser. Five element wings produce significant amounts of drag which is a compromise between the cornering ability of the car and the acceleration capability on straights. The active aerodynamics system allows for the wing angle of attack to dynamically change their configuration on track based on sensory data to optimize the wings for any given scenario. The wings are studied using computational fluid dynamics both in their maximum lift configuration as well as a minimum drag configuration. A control system is then developed using an electro mechanical actuation system to articulate the wings between these two states.
Notch signalling coordinates tissue growth and wing fate specification in Drosophila.
Rafel, Neus; Milán, Marco
2008-12-01
During the development of a given organ, tissue growth and fate specification are simultaneously controlled by the activity of a discrete number of signalling molecules. Here, we report that these two processes are extraordinarily coordinated in the Drosophila wing primordium, which extensively proliferates during larval development to give rise to the dorsal thoracic body wall and the adult wing. The developmental decision between wing and body wall is defined by the opposing activities of two secreted signalling molecules, Wingless and the EGF receptor ligand Vein. Notch signalling is involved in the determination of a variety of cell fates, including growth and cell survival. We present evidence that growth of the wing primordium mediated by the activity of Notch is required for wing fate specification. Our data indicate that tissue size modulates the activity range of the signalling molecules Wingless and Vein. These results highlight a crucial role of Notch in linking proliferation and fate specification in the developing wing primordium.
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.
Loads Model Development and Analysis for the F/A-18 Active Aeroelastic Wing Airplane
NASA Technical Reports Server (NTRS)
Allen, Michael J.; Lizotte, Andrew M.; Dibley, Ryan P.; Clarke, Robert
2005-01-01
The Active Aeroelastic Wing airplane was successfully flight-tested in March 2005. During phase 1 of the two-phase program, an onboard excitation system provided independent control surface movements that were used to develop a loads model for the wing structure and wing control surfaces. The resulting loads model, which was used to develop the control laws for phase 2, is described. The loads model was developed from flight data through the use of a multiple linear regression technique. The loads model input consisted of aircraft states and control surface positions, in addition to nonlinear inputs that were calculated from flight-measured parameters. The loads model output for each wing consisted of wing-root bending moment and torque, wing-fold bending moment and torque, inboard and outboard leading-edge flap hinge moment, trailing-edge flap hinge moment, and aileron hinge moment. The development of the Active Aeroelastic Wing loads model is described, and the ability of the model to predict loads during phase 2 research maneuvers is demonstrated. Results show a good match to phase 2 flight data for all loads except inboard and outboard leading-edge flap hinge moments at certain flight conditions. The average load prediction errors for all loads at all flight conditions are 9.1 percent for maximum stick-deflection rolls, 4.4 percent for 5-g windup turns, and 7.7 percent for 4-g rolling pullouts.
NASA Astrophysics Data System (ADS)
Bakunowicz, Jerzy; Święch, Łukasz; Meyer, Ralf
2016-12-01
In aeronautical research experimental data sets of high quality are essential to verify and improve simulation algorithms. For this reason the experimental techniques need to be constantly refined. The shape, movement or deformation of structural aircraft elements can be measured implicitly in multiple ways; however, only optical, correlation-based techniques are able to deliver direct high-order and spatial results. In this paper two different optical metrologies are used for on-ground preparation and the actual execution of in-flight wing deformation measurements on a PW-6U glider. Firstly, the commercial PONTOS system is used for static tests on the ground and for wind tunnel investigations to successfully certify an experimental sensor pod mounted on top of the test bed fuselage. Secondly, a modification of the glider is necessary to implement the optical method named image pattern correlation technique (IPCT), which has been developed by the German Aerospace Center DLR. This scientific technology uses a stereoscopic camera set-up placed inside the experimental pod and a stochastic dot matrix applied to the area of interest on the glider wing to measure the deformation of the upper wing surface in-flight. The flight test installation, including the preparation, is described and results are presented briefly. Focussing on the compensation for typical error sources, the paper concludes with a recommended procedure to enhance the data processing for better results. Within the presented project IPCT has been developed and optimized for a new type of test bed. Adapted to the special requirements of the glider, the IPCT measurements were able to deliver a valuable wing deformation data base which now can be used to improve corresponding numerical models and simulations.
Experimental Study of the Structure of a Wingtip Vortex
NASA Technical Reports Server (NTRS)
Anderson, Elgin A.; Wright, Christopher T.
2000-01-01
A complete look at the near-field development and subsequent role-up of a wingtip vortex from a NACA 0015 wing section is investigated. Two separate but equally important surveys of the vortex structure in the region adjacent to the wingtip and approximately one chord length downstream of the trailing edge are performed. The two surveys provide qualitative flow-visualization an quantitative velocity measurement data. The near-field development and subsequent role-up of the vortex structures is strongly influenced by the angle-of-attack and the end-cap treatment of the wing section. The velocity field near the wingtip of the NACA 0015 wing section was measured with a triple-sensor hot wire probe and compared to flow visualization images produced with titanium tetrachloride smoke injection and laser illumination. The flat end-cap results indicate the formation of multiple, relatively strong vortex structures as opposed to the formation of a single vortex produced with the round end-cap. The multiple vortices generated by the flat end-cap are seen to rotate around a common ce te in a helical pattern until they eventually merge into a single vortex. Compared to a non-dimensional loading parameter, the results of the velocity and flow visualization data shows a "jetlike" axial velocity profile for loading parameter values on the order of 0.1 and a "wakelike" profile for much lower loading parameter values.
NASA Technical Reports Server (NTRS)
Hall, G. F.; Shamroth, S. J.; Mcdonald, H.; Briley, W. R.
1976-01-01
A method was developed for determining the aerodynamic loads on the tip of an infinitely thin, swept, cambered semi-infinite wing at an angle of attack which is operating subsonically in an inviscid medium and is subjected to a sinusoidal gust. Under the assumption of linearized aerodynamics, the loads on the tip are obtained by superposition of the steady aerodynamic results for angle of attack and camber, and the unsteady results for the response to the sinusoidal gust. The near field disturbance pressures in the fluid surrounding the tip are obtained by assuming a dipole representation for the loading on the tip and calculating the pressures accordingly. The near field pressures are used to drive a reduced form of the Navier-Stokes equations which yield the tip vortex formation. The combined viscid-inviscid analysis is applied to determining the pressures and examining the vortex rollup in the vicinity of an unswept, uncambered wing moving steadily at a Mach number of 0.2 at an angle of attack of 0.1 rad. The viscous tip flow calculation shows features expected in the tip flow such as the qualitatively proper development of boundary layers on both the upper and lower airfoil surfaces. In addition, application of the viscous solution leads to the generation of a circular type flow pattern above the airfoil suction surface.
ROBUSTNESS OF SIGNALING GRADIENT IN DROSOPHILA WING IMAGINAL DISC
Lei, Jinzhi; Wan, Frederic Y. M.; Lander, Arthur D.; Nie, Qing
2012-01-01
Quasi-stable gradients of signaling protein molecules (known as morphogens or ligands) bound to cell receptors are known to be responsible for differential cell signaling and gene expressions. From these follow different stable cell fates and visually patterned tissues in biological development. Recent studies have shown that the relevant basic biological processes yield gradients that are sensitive to small changes in system characteristics (such as expression level of morphogens or receptors) or environmental conditions (such as temperature changes). Additional biological activities must play an important role in the high level of robustness observed in embryonic patterning for example. It is natural to attribute observed robustness to various type of feedback control mechanisms. However, our own simulation studies have shown that feedback control is neither necessary nor sufficient for robustness of the morphogen decapentaplegic (Dpp) gradient in wing imaginal disc of Drosophilas. Furthermore, robustness can be achieved by substantial binding of the signaling morphogen Dpp with nonsignaling cell surface bound molecules (such as heparan sulfate proteoglygans) and degrading the resulting complexes at a sufficiently rapid rate. The present work provides a theoretical basis for the results of our numerical simulation studies. PMID:24098092
Heers, Ashley M; Dial, Kenneth P
2015-02-01
Wings have long been regarded as a hallmark of evolutionary innovation, allowing insects, birds, and bats to radiate into aerial environments. For many groups, our intuitive and colloquial perspective is that wings function for aerial activities, and legs for terrestrial, in a relatively independent manner. However, insects and birds often engage their wings and legs cooperatively. In addition, the degree of autonomy between wings and legs may be constrained by tradeoffs, between allocating resources to wings versus legs during development, or between wing versus leg investment and performance (because legs must be carried as baggage by wings during flight and vice versa). Such tradeoffs would profoundly affect the development and evolution of locomotor strategies, and many related aspects of animal ecology. Here, we provide the first evaluation of wing versus leg investment, performance and relative use, in birds-both across species, and during ontogeny in three precocial species with different ecologies. Our results suggest that tradeoffs between wing and leg modules help shape ontogenetic and evolutionary trajectories, but can be offset by recruiting modules cooperatively. These findings offer a new paradigm for exploring locomotor strategies of flying organisms and their extinct precursors, and thereby elucidating some of the most spectacular diversity in animal history. © 2014 The Author(s). Evolution © 2014 The Society for the Study of Evolution.
Kim, Sabrina Y; Renihan, Maia K; Boulianne, Gabrielle L
2006-06-01
PDZ (PSD-95, Discs-large, ZO-1) domain proteins often function as scaffolding proteins and have been shown to play important roles in diverse cellular processes such as the establishment and maintenance of cell polarity, and signal transduction. Here, we report the identification and cloning of a novel Drosophila melanogaster gene that is predicted to produce several different PDZ domain-containing proteins through alternative promoter usage and alternative splicing. This gene, that we have named big bang (bbg), was first identified as C96-GAL4, a GAL4 enhancer trap line that was generated in our lab. To further characterize bbg, its expression pattern was examined in ovaries, embryos, and late third instar larvae using UAS reporter gene constructs, in situ hybridization, or immunocytochemistry. In addition, the expression of alternatively spliced transcripts was examined in more detail using in situ hybridization. We find that during embryogenesis bbg is predominantly expressed in the developing gut, but it is also expressed in external sensory organs found in the epidermis. In the late third instar larva, bbg is expressed along the presumptive wing margin in the wing disc, broadly in the eye disc, and in other imaginal discs as well as in the brain. The expression patterns observed are dynamic and specific during development, suggesting that like other genes that encode for several different PDZ domain protein isoforms, bbg likely plays important roles in multiple developmental processes.
Small Engine Technology (SET) Task 23 ANOPP Noise Prediction for Small Engines, Wing Reflection Code
NASA Technical Reports Server (NTRS)
Lieber, Lysbeth; Brown, Daniel; Golub, Robert A. (Technical Monitor)
2000-01-01
The work performed under Task 23 consisted of the development and demonstration of improvements for the NASA Aircraft Noise Prediction Program (ANOPP), specifically targeted to the modeling of engine noise enhancement due to wing reflection. This report focuses on development of the model and procedure to predict the effects of wing reflection, and the demonstration of the procedure, using a representative wing/engine configuration.
Induced drag ideal efficiency factor of arbitrary lateral-vertical wing forms
NASA Technical Reports Server (NTRS)
Deyoung, J.
1980-01-01
A relatively simple equation is presented for estimating the induced drag ideal efficiency factor e for arbitrary cross sectional wing forms. This equation is based on eight basic but varied wing configurations which have exact solutions. The e function which relates the basic wings is developed statistically and is a continuous function of configuration geometry. The basic wing configurations include boxwings shaped as a rectangle, ellipse, and diamond; the V-wing; end-plate wing; 90 degree cruciform; circle dumbbell; and biplane. Example applications of the e equations are made to many wing forms such as wings with struts which form partial span rectangle dumbbell wings; bowtie, cruciform, winglet, and fan wings; and multiwings. Derivations are presented in the appendices of exact closed form solutions found of e for the V-wing and 90 degree cruciform wing and for an asymptotic solution for multiwings.
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.
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.
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.
Measurement of shape and deformation of insect wing
NASA Astrophysics Data System (ADS)
Yin, Duo; Wei, Zhen; Wang, Zeyu; Zhou, Changqiu
2018-01-01
To measure the shape and deformation of an insect wing, a scanning setup adopting laser triangulation and image matching was developed. Only one industry camera with two light sources was employed to scan the transparent insect wings. 3D shape and point to point full field deformation of the wings could be obtained even when the wingspan is less than 3 mm. The venation and corrugation could be significantly identified from the results. The deformation of the wing under pin loading could be seen clearly from the results as well. Calibration shows that the shape and deformation measurement accuracies are no lower than 0.01 mm. Laser triangulation and image matching were combined dexterously to adapt wings' complex shape, size, and transparency. It is suitable for insect flight research or flapping wing micro-air vehicle development.
Elastically Shaped Wing Optimization and Aircraft Concept for Improved Cruise Efficiency
NASA Technical Reports Server (NTRS)
Nguyen, Nhan; Trinh, Khanh; Reynolds, Kevin; Kless, James; Aftosmis, Michael; Urnes, James, Sr.; Ippolito, Corey
2013-01-01
This paper presents the findings of a study conducted tn 2010 by the NASA Innovation Fund Award project entitled "Elastically Shaped Future Air Vehicle Concept". The study presents three themes in support of meeting national and global aviation challenges of reducing fuel burn for present and future aviation systems. The first theme addresses the drag reduction goal through innovative vehicle configurations via non-planar wing optimization. Two wing candidate concepts have been identified from the wing optimization: a drooped wing shape and an inflected wing shape. The drooped wing shape is a truly biologically inspired wing concept that mimics a seagull wing and could achieve about 5% to 6% drag reduction, which is aerodynamically significant. From a practical perspective, this concept would require new radical changes to the current aircraft development capabilities for new vehicles with futuristic-looking wings such as this concept. The inflected wing concepts could achieve between 3% to 4% drag reduction. While the drag reduction benefit may be less, the inflected-wing concept could have a near-term impact since this concept could be developed within the current aircraft development capabilities. The second theme addresses the drag reduction goal through a new concept of elastic wing shaping control. By aeroelastically tailoring the wing shape with active control to maintain optimal aerodynamics, a significant drag reduction benefit could be realized. A significant reduction in fuel burn for long-range cruise from elastic wing shaping control could be realized. To realize the potential of the elastic wing shaping control concept, the third theme emerges that addresses the drag reduction goal through a new aerodynamic control effector called a variable camber continuous trailing edge flap. Conventional aerodynamic control surfaces are discrete independent surfaces that cause geometric discontinuities at the trailing edge region. These discontinuities promote vorticities which result in drag rises as well as noise sources. The variable camber trailing edge flap concept could provide a substantial drag reduction benefit over a conventional discrete flap system. Aerodynamic simulations show a drag reduction of over 50% could be achieved with the flap concept over a conventional discrete flap system.
NASA Technical Reports Server (NTRS)
Adams, Gaynor J; DUGAN DUANE W
1952-01-01
A method of analysis based on slender-wing theory is developed to investigate the characteristics in roll of slender cruciform wings and wing-body combinations. The method makes use of the conformal mapping processes of classical hydrodynamics which transform the region outside a circle and the region outside an arbitrary arrangement of line segments intersecting at the origin. The method of analysis may be utilized to solve other slender cruciform wing-body problems involving arbitrarily assigned boundary conditions. (author)
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.
Zhang, Xubo; Luo, Dan; Pflugfelder, Gert O; Shen, Jie
2013-07-01
The control of organ growth is a fundamental aspect of animal development but remains poorly understood. The morphogen Dpp has long been considered as a general promoter of cell proliferation during Drosophila wing development. It is an ongoing debate whether the Dpp gradient is required for the uniform cell proliferation observed in the wing imaginal disc. Here, we investigated how the Dpp signaling pathway regulates proliferation during wing development. By systematic manipulation of Dpp signaling we observed that it controls proliferation in a region-specific manner: Dpp, via omb, promoted proliferation in the lateral and repressed proliferation in the medial wing disc. Omb controlled the regional proliferation rate by oppositely regulating transcription of the microRNA gene bantam in medial versus lateral wing disc. However, neither the Dpp nor Omb gradient was essential for uniform proliferation along the anteroposterior axis.
NASA Technical Reports Server (NTRS)
Fujiwara, Gustavo; Bragg, Mike; Triphahn, Chris; Wiberg, Brock; Woodard, Brian; Loth, Eric; Malone, Adam; Paul, Bernard; Pitera, David; Wilcox, Pete;
2017-01-01
This report presents the key results from the first two years of a program to develop experimental icing simulation capabilities for full-scale swept wings. This investigation was undertaken as a part of a larger collaborative research effort on ice accretion and aerodynamics for large-scale swept wings. Ice accretion and the resulting aerodynamic effect on large-scale swept wings presents a significant airplane design and certification challenge to air frame manufacturers, certification authorities, and research organizations alike. While the effect of ice accretion on straight wings has been studied in detail for many years, the available data on swept-wing icing are much more limited, especially for larger scales.
Es'kov, E K; Es'kova, M D
2014-01-01
High variability of cells size is used selectively for reproduction of working bees and drones. A decrease in both distance between cells and cells size themselves causes similar effects to body mass and morphometric traits of developing individuals. Adaptation of honey bees to living in shelters has led to their becoming tolerant to hypoxia. Improvement of ethological and physiological mechanisms of thermal regulation is associated with limitation of ecological valence and acquiring of stenothermic features by breed. Optimal thermal conditions for breed are limited by the interval 33-34.5 degrees C. Deviations of temperature by 3-4 degrees C beyond this range have minimum lethal effect at embryonic stage of development and medium effect at the stage of pre-pupa and pupa. Developing at the low bound of the vital range leads to increasing, while developing at the upper bound--to decreasing of body mass, mandibular and hypopharyngeal glands, as well as other organs, which, later, affects the variability of these traits during the adult stage of development. Eliminative and teratogenic efficiency of ecological factors that affect a breed is most often manifested in underdevelopment of wings. However, their size (in case of wing laminas formation). is characterized by relatively low variability and size-dependent asymmetry. Asymmetry variability of wings and other pair organs is expressed through realignment of size excess from right- to left-side one with respect to their increase. Selective elimination by those traits whose emerging probability increases as developmental conditions deviate from the optimal ones promotes restrictions on individual variability. Physiological mechanisms that facilitate adaptability enhancement under conditions of increasing anthropogenic contamination of eivironment and trophic substrates consumed by honey bees, arrear to be toxicants accumulation in rectum and crops' ability to absorb contaminants from nectar in course of its processing to honey.
NASA Astrophysics Data System (ADS)
Kraemer, Kurtis Leigh
Micro air vehicles (MAV) are a class of small uninhabited aircraft with dimensions less than 15 cm (6 in) and mass less than 500g (1.1 lbs). The aim of this research was to develop a fast, accurate, low-cost, and repeatable fabrication process for flapping MAV wings. Through the use of the RepRap Mendel open-source fused-deposition modeling (FDM) rapid prototyping machine ("3-D printer"), various wing prototypes were designed and fabricated using a bio-inspired approach. Testing of the aerodynamic performance of both real locust wings and the 3-D printed wing prototypes was performed through axial spin testing. Bending stiffness measurements were also performed on the 3-D printed wings. Through the use of open-source rapid prototyping technology, a fast and low-cost fabrication process for flapping MAV wings has been developed, out of which further understanding of flapping wing design and fabrication has been gained.
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.
Shape control of an adaptive wing for transonic drag reduction
NASA Astrophysics Data System (ADS)
Austin, Fred; Van Nostrand, William C.
1995-05-01
Theory and experiments to control the static shape of flexible structures by employing internal translational actuators are summarized and plants to extend the work to adaptive wings are presented. Significant reductions in the shock-induced drag are achievable during transonic- cruise by small adaptive modifications to the wing cross-sectional profile. Actuators are employed as truss elements of active ribs to deform the wing cross section. An adaptive-rib model was constructed, and experiments validated the shape-control theory. Plans for future development under an ARPA/AFWAL contract include payoff assessments of the method on an actual aircraft, the development of inchworm TERFENOL-D actuators, and the development of a method to optimize the wing cross-sectional shapes by direct-drag measurements.
Analysis of Mach number 0.8 turboprop slipstream wing/nacelle interactions
NASA Technical Reports Server (NTRS)
Welge, H. R.; Neuhart, D. H.; Dahlin, J. A.
1981-01-01
Data from wind tunnel tests of a powered propeller and nacelle mounted on a supercritical wing are analyzed. Installation of the nacelle significantly affected the wing flow and the flow on the upper surface of the wing is separated near the leading edge under powered conditions. Comparisons of various theories with the data indicated that the Neumann surface panel solution and the Jameson transonic solution gave results adequate for design purposes. A modified wing design was developed (Mod 3) which reduces the wing upper surface pressure coefficients and section lift coefficients at powered conditions to levels below those of the original wing without nacelle or power. A contoured over the wing nacelle that can be installed on the original wing without any appreciable interference to the wing upper surface pressure is described.
Static aeroelastic analysis and tailoring of a single-element racing car wing
NASA Astrophysics Data System (ADS)
Sadd, Christopher James
This thesis presents the research from an Engineering Doctorate research programme in collaboration with Reynard Motorsport Ltd, a manufacturer of racing cars. Racing car wing design has traditionally considered structures to be rigid. However, structures are never perfectly rigid and the interaction between aerodynamic loading and structural flexibility has a direct impact on aerodynamic performance. This interaction is often referred to as static aeroelasticity and the focus of this research has been the development of a computational static aeroelastic analysis method to improve the design of a single-element racing car wing. A static aeroelastic analysis method has been developed by coupling a Reynolds-Averaged Navier-Stokes CFD analysis method with a Finite Element structural analysis method using an iterative scheme. Development of this method has included assessment of CFD and Finite Element analysis methods and development of data transfer and mesh deflection methods. Experimental testing was also completed to further assess the computational analyses. The computational and experimental results show a good correlation and these studies have also shown that a Navier-Stokes static aeroelastic analysis of an isolated wing can be performed at an acceptable computational cost. The static aeroelastic analysis tool was used to assess methods of tailoring the structural flexibility of the wing to increase its aerodynamic performance. These tailoring methods were then used to produce two final wing designs to increase downforce and reduce drag respectively. At the average operating dynamic pressure of the racing car, the computational analysis predicts that the downforce-increasing wing has a downforce of C[1]=-1.377 in comparison to C[1]=-1.265 for the original wing. The computational analysis predicts that the drag-reducing wing has a drag of C[d]=0.115 in comparison to C[d]=0.143 for the original wing.
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.
NASA Technical Reports Server (NTRS)
Soule, V. A.; Badri-Nath, Y.
1973-01-01
The results of a study of the use of composite materials in the wing of a tilt rotor aircraft are presented. An all-metal tilt rotor aircraft was first defined to provide a basis for comparing composite with metal structure. A configuration study was then done in which the wing of the metal aircraft was replaced with composite wings of varying chord and thickness ratio. The results of this study defined the design and performance benefits obtainable with composite materials. Based on these results the aircraft was resized with a composite wing to extend the weight savings to other parts of the aircraft. A wing design was then selected for detailed structural analysis. A development plan including costs and schedules to develop this wing and incorporate it into a proposed flight research tilt rotor vehicle has been devised.
Real-time In-Flight Strain and Deflection Monitoring with Fiber Optic Sensors
NASA Technical Reports Server (NTRS)
Richards, Lance; Parker, Allen R.; Ko, William L.; Piazza, Anthony
2008-01-01
This viewgraph presentation reviews Dryden's efforts to develop in-flight monitoring based on Fiber Optics. One of the motivating factors for this development was the breakup of the Helios aircraft. On Ikhana the use of fiber optics for wing shape sensing is being developed. They are being used to flight validate fiber optic sensor measurements and real-time wing shape sensing predictions on NASA's Ikhana vehicle; validate fiber optic mathematical models and design tools; Assess technical viability and, if applicable, develop methodology and approach to incorporate wing shape measurements within the vehicle flight control system, and develop and flight validate advanced approaches to perform active wing shape control.
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
Static Aeroelastic Scaling and Analysis of a Sub-Scale Flexible Wing Wind Tunnel Model
NASA Technical Reports Server (NTRS)
Ting, Eric; Lebofsky, Sonia; Nguyen, Nhan; Trinh, Khanh
2014-01-01
This paper presents an approach to the development of a scaled wind tunnel model for static aeroelastic similarity with a full-scale wing model. The full-scale aircraft model is based on the NASA Generic Transport Model (GTM) with flexible wing structures referred to as the Elastically Shaped Aircraft Concept (ESAC). The baseline stiffness of the ESAC wing represents a conventionally stiff wing model. Static aeroelastic scaling is conducted on the stiff wing configuration to develop the wind tunnel model, but additional tailoring is also conducted such that the wind tunnel model achieves a 10% wing tip deflection at the wind tunnel test condition. An aeroelastic scaling procedure and analysis is conducted, and a sub-scale flexible wind tunnel model based on the full-scale's undeformed jig-shape is developed. Optimization of the flexible wind tunnel model's undeflected twist along the span, or pre-twist or wash-out, is then conducted for the design test condition. The resulting wind tunnel model is an aeroelastic model designed for the wind tunnel test condition.
Development of the Main Wing Structure of a High Altitude Long Endurance UAV
NASA Astrophysics Data System (ADS)
Park, Sang Wook; Shin, Jeong Woo; Kim, Tae-Uk
2018-04-01
To enhance the flight endurance of a HALE UAV, the main wing of the UAV should have a high aspect ratio and low structural weight. Since a main wing constructed with the thin walled and slender components needed for low structural weight can suffer catastrophic failure during flight, it is important to develop a light-weight airframe without sacrificing structural integrity. In this paper, the design of the main wing of the HALE UAV was conducted using spars which were composed of a carbon-epoxy cylindrical tube and bulkheads to achieve both the weight reduction and structural integrity. The spars were sized using numerical analysis considering non-linear deformation under bending moment. Static strength testing of the wing was conducted under the most critical load condition. Then, the experimental results obtained for the wing were compared to the analytical result from the non-linear finite-element analysis. It was found that the developed main wing reduced its structural weight without any failure under the ultimate load condition of the static strength testing.
Iwata, Masaki; Taira, Wataru; Hiyama, Atsuki; Otaki, Joji M
2015-06-01
The nymphalid groundplan has been proposed to explain diverse butterfly wing color patterns. In this model, each symmetry system is composed of a core element and a pair of paracore elements. The development of this elemental configuration has been explained by the induction model for positional information. However, the diversity of color patterns in other butterfly families in relation to the nymphalid groundplan has not been thoroughly examined. Here, we examined aberrant color pattern phenotypes of a lycaenid butterfly, Zizeeria maha, from mutagenesis and plasticity studies as well as from field surveys. In several mutants, the third and fourth spot arrays were coordinately positioned much closer to the discal spot in comparison to the normal phenotype. In temperature-shock types, the third and fourth array spots were elongated inwardly or outwardly from their normal positions. In field-caught spontaneous mutants, small black spots were located adjacent to normal black spots. Analysis of these aberrant phenotypes indicated that the spots belonging to the third and fourth arrays are synchronously changeable in position and shape around the discal spot. Thus, these arrays constitute paracore elements of the central symmetry system of the lycaenid butterflies, and the discal spot comprises the core element. These aberrant phenotypes can be explained by the black-inducing signals that propagate from the prospective discal spot, as predicted by the induction model. These results suggest the existence of long-range developmental signals that cover a large area of a wing not only in nymphalid butterflies, but also in lycaenid butterflies.
Analytic study of the conditions required for longitudinal stability of dual-wing aircraft
Andrews, Stephen Arthur; Perez, Ruben E.
2017-05-11
Recent studies of new, fuel-efficient transport aircraft have considered designs, which make use of two principal lifting surfaces to provide the required lift as well as trim and static stability. Such designs include open tandem-wings as well as closed joined and box-wings. As a group, these aircraft can be termed dual-wing designs. Our study developed a new analytic model, which takes into account the downwash from the two main wings and is sensitive to three important design variables: the relative areas of each wing, the streamwise separation of the wings, and the center of gravity position. This model was usedmore » to better understand trends in the dual-wing geometry on the stability, maneuverability, and lift-to-drag ratio of the aircraft. Dual-wing aircraft have been shown to have reduced the induced drag compared to the conventional designs. In addition, further drag reductions can be realized as the horizontal tail can be removed if the dual-wings have sufficient streamwise stagger to provide the moments necessary for trim and longitudinal stability. As both wings in a dual-wing system carry a significant fraction of the total lift, trends in such designs that led to longitudinal stability can differ from those of the conventional aircraft and have not been the subject of detailed investigation. Results from the analytic model showed that the longitudinal stability required either a reduction of the fore wing area or shifting the center of gravity forward from the midpoint of both wings' aerodynamic centers. Additionally, for wing configurations of approximately equal fore and aft wing areas, increasing the separation between the two wings decreased the stability of the aircraft. The source of this unusual behavior was the asymmetric distribution of downwash upstream and downstream of the wing. These relationships between dual-wing geometry and stability will provide initial guidance on the conceptual design of dual-wing aircraft and aid in the understanding of the results of more complex studies of such designs, furthering the development of future transport aircraft.« less
Analytic study of the conditions required for longitudinal stability of dual-wing aircraft
DOE Office of Scientific and Technical Information (OSTI.GOV)
Andrews, Stephen Arthur; Perez, Ruben E.
Recent studies of new, fuel-efficient transport aircraft have considered designs, which make use of two principal lifting surfaces to provide the required lift as well as trim and static stability. Such designs include open tandem-wings as well as closed joined and box-wings. As a group, these aircraft can be termed dual-wing designs. Our study developed a new analytic model, which takes into account the downwash from the two main wings and is sensitive to three important design variables: the relative areas of each wing, the streamwise separation of the wings, and the center of gravity position. This model was usedmore » to better understand trends in the dual-wing geometry on the stability, maneuverability, and lift-to-drag ratio of the aircraft. Dual-wing aircraft have been shown to have reduced the induced drag compared to the conventional designs. In addition, further drag reductions can be realized as the horizontal tail can be removed if the dual-wings have sufficient streamwise stagger to provide the moments necessary for trim and longitudinal stability. As both wings in a dual-wing system carry a significant fraction of the total lift, trends in such designs that led to longitudinal stability can differ from those of the conventional aircraft and have not been the subject of detailed investigation. Results from the analytic model showed that the longitudinal stability required either a reduction of the fore wing area or shifting the center of gravity forward from the midpoint of both wings' aerodynamic centers. Additionally, for wing configurations of approximately equal fore and aft wing areas, increasing the separation between the two wings decreased the stability of the aircraft. The source of this unusual behavior was the asymmetric distribution of downwash upstream and downstream of the wing. These relationships between dual-wing geometry and stability will provide initial guidance on the conceptual design of dual-wing aircraft and aid in the understanding of the results of more complex studies of such designs, furthering the development of future transport aircraft.« less
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.
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
Works on theory of flapping wing. [considering boundary layer
NASA Technical Reports Server (NTRS)
Golubev, V. V.
1980-01-01
It is shown mathematically that taking account of the boundary layer is the only way to develop a theory of flapping wings without violating the basic observations and mathematics of hydromechanics. A theory of thrust generation by flapping wings can be developed if the conventional downstream velocity discontinuity surface is replaced with the observed Karman type vortex streets behind a flapping wing. Experiments show that the direction of such vortices is the reverse of that of conventional Karman streets. The streets form by breakdown of the boundary layer. Detailed analysis of the movements of certain birds and insects during flight 'in place' is fully consistent with this theory of the lift, thrust and drag of flapping wings. Further directions for research into flight with flapping wings are indicated.
The fledging of common and thick-billed murres on Middleton Island, Alaska
Hatch, Scott A.
1983-01-01
Three species of alcids, Common and Thick-billed murres (Uria aalge and U. lomvia) and the Razorbill (Alca torda), have post-hatching developmental patterns intermediate to precocial and semi-precocial modes (Sealy 1973). The young leave their cliff nest sites at about one quarter of adult weight and complete their growth at sea. At departure, an event here loosely referred to as "fledging," neither primary nor secondary flight feathers are grown, but well-developed wing coverts enable limited, descending flight.
2010-09-01
Mineral resources include metals, industrial minerals (e.g., aggregate, sand and gravel), oil and gas, and geothermal resources that would be of value...the elderberry shrub ( Sambucus sp.), is not present within the project action area. The proposed project is not likely to adversely affect the bald...impairing essential behavioral patterns, including breeding, feeding, or sheltering. Harass is defined by the Service as an action that creates the
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
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.
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.
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.
Drosophila Lyra mutations are gain-of-function mutations of senseless
NASA Technical Reports Server (NTRS)
Nolo, R.; Abbott, L. A.; Bellen, H. J.
2001-01-01
The Lyra mutation was first described by Jerry Coyne in 1935. Lyra causes recessive pupal lethality and adult heterozygous Lyra mutants exhibit a dominant loss of the anterior and posterior wing margins. Unlike many mutations that cause loss of wing tissue (e.g., scalloped, Beadex, cut, and apterous-Xasta), Lyra wing discs do not exhibit increased necrotic or apoptotic cell death, nor do they show altered BrdU incorporation. However, during wing disc eversion, loss of the anterior and posterior wing margins is apparent. We have previously shown that senseless, a gene that is necessary and sufficient for peripheral nervous system (PNS) development, is allelic to Lyra. Here we show by several genetic criteria that Lyra alleles are neomorphic alleles of senseless that cause ectopic expression of SENSELESS in the wing pouch. Similarly, overexpression of SENSELESS in the wing disc causes loss of wing margin tissue, thereby mimicking the Lyra phenotype. Lyra mutants display aberrant expression of DELTA, VESTIGIAL, WINGLESS, and CUT. As in Lyra mutants, overexpression of SENSELESS in some areas of the wing pouch also leads to loss of WINGLESS and CUT. In summary, our data indicate that overexpression of SENSELESS causes a severe reduction in NOTCH signaling that in turn may lead to decreased transcription of several key genes required for wing development, leading to a failure in cell proliferation and loss of wing margin tissue.
NASA Astrophysics Data System (ADS)
Geisler, T.
2016-12-01
Beetle wings perform a flapping movement, consisting of the rotation relative to the two axes. This paper presents the results of observations and measurements of wings operating parameters in different planes of some beetle species. High speed photos and videos were used. The concept of the mechanism performing a complex wing movement was proposed and developed.
Computer Code Aids Design Of Wings
NASA Technical Reports Server (NTRS)
Carlson, Harry W.; Darden, Christine M.
1993-01-01
AERO2S computer code developed to aid design engineers in selection and evaluation of aerodynamically efficient wing/canard and wing/horizontal-tail configurations that includes simple hinged-flap systems. Code rapidly estimates longitudinal aerodynamic characteristics of conceptual airplane lifting-surface arrangements. Developed in FORTRAN V on CDC 6000 computer system, and ported to MS-DOS environment.
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.
Large capacity oblique all-wing transport aircraft
NASA Technical Reports Server (NTRS)
Galloway, Thomas L.; Phillips, James A.; Kennelly, Robert A., Jr.; Waters, Mark H.
1996-01-01
Dr. R. T. Jones first developed the theory for oblique wing aircraft in 1952, and in subsequent years numerous analytical and experimental projects conducted at NASA Ames and elsewhere have established that the Jones' oblique wing theory is correct. Until the late 1980's all proposed oblique wing configurations were wing/body aircraft with the wing mounted on a pivot. With the emerging requirement for commercial transports with very large payloads, 450-800 passengers, Jones proposed a supersonic oblique flying wing in 1988. For such an aircraft all payload, fuel, and systems are carried within the wing, and the wing is designed with a variable sweep to maintain a fixed subsonic normal Mach number. Engines and vertical tails are mounted on pivots supported from the primary structure of the wing. The oblique flying wing transport has come to be known as the Oblique All-Wing (OAW) transport. This presentation gives the highlights of the OAW project that was to study the total concept of the OAW as a commercial transport.
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.
NASA Technical Reports Server (NTRS)
Nguyen, Nhan; Kaul, Upender; Lebofsky, Sonia; Ting, Eric; Chaparro, Daniel; Urnes, James
2015-01-01
This paper summarizes the recent development of an adaptive aeroelastic wing shaping control technology called variable camber continuous trailing edge flap (VCCTEF). As wing flexibility increases, aeroelastic interactions with aerodynamic forces and moments become an increasingly important consideration in aircraft design and aerodynamic performance. Furthermore, aeroelastic interactions with flight dynamics can result in issues with vehicle stability and control. The initial VCCTEF concept was developed in 2010 by NASA under a NASA Innovation Fund study entitled "Elastically Shaped Future Air Vehicle Concept," which showed that highly flexible wing aerodynamic surfaces can be elastically shaped in-flight by active control of wing twist and bending deflection in order to optimize the spanwise lift distribution for drag reduction. A collaboration between NASA and Boeing Research & Technology was subsequently funded by NASA from 2012 to 2014 to further develop the VCCTEF concept. This paper summarizes some of the key research areas conducted by NASA during the collaboration with Boeing Research and Technology. These research areas include VCCTEF design concepts, aerodynamic analysis of VCCTEF camber shapes, aerodynamic optimization of lift distribution for drag minimization, wind tunnel test results for cruise and high-lift configurations, flutter analysis and suppression control of flexible wing aircraft, and multi-objective flight control for adaptive aeroelastic wing shaping control.
NASA Technical Reports Server (NTRS)
Ehlers, F. E.; Weatherill, W. H.; Yip, E. L.
1984-01-01
A finite difference method to solve the unsteady transonic flow about harmonically oscillating wings was investigated. The procedure is based on separating the velocity potential into steady and unsteady parts and linearizing the resulting unsteady differential equation for small disturbances. The differential equation for the unsteady velocity potential is linear with spatially varying coefficients and with the time variable eliminated by assuming harmonic motion. An alternating direction implicit procedure was investigated, and a pilot program was developed for both two and three dimensional wings. This program provides a relatively efficient relaxation solution without previously encountered solution instability problems. Pressure distributions for two rectangular wings are calculated. Conjugate gradient techniques were developed for the asymmetric, indefinite problem. The conjugate gradient procedure is evaluated for applications to the unsteady transonic problem. Different equations for the alternating direction procedure are derived using a coordinate transformation for swept and tapered wing planforms. Pressure distributions for swept, untaped wings of vanishing thickness are correlated with linear results for sweep angles up to 45 degrees.
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.
Flight Test of the F/A-18 Active Aeroelastic Wing Airplane
NASA Technical Reports Server (NTRS)
Clarke, Robert; Allen, Michael J.; Dibley, Ryan P.; Gera, Joseph; Hodgkinson, John
2005-01-01
Successful flight-testing of the Active Aeroelastic Wing airplane was completed in March 2005. This program, which started in 1996, was a joint activity sponsored by NASA, Air Force Research Laboratory, and industry contractors. The test program contained two flight test phases conducted in early 2003 and early 2005. During the first phase of flight test, aerodynamic models and load models of the wing control surfaces and wing structure were developed. Design teams built new research control laws for the Active Aeroelastic Wing airplane using these flight-validated models; and throughout the final phase of flight test, these new control laws were demonstrated. The control laws were designed to optimize strategies for moving the wing control surfaces to maximize roll rates in the transonic and supersonic flight regimes. Control surface hinge moments and wing loads were constrained to remain within hydraulic and load limits. This paper describes briefly the flight control system architecture as well as the design approach used by Active Aeroelastic Wing project engineers to develop flight control system gains. Additionally, this paper presents flight test techniques and comparison between flight test results and predictions.
Recent progress in the analysis of iced airfoils and wings
NASA Technical Reports Server (NTRS)
Cebeci, Tuncer; Chen, Hsun H.; Kaups, Kalle; Schimke, Sue
1992-01-01
Recent work on the analysis of iced airfoils and wings is described. Ice shapes for multielement airfoils and wings are computed using an extension of the LEWICE code that was developed for single airfoils. The aerodynamic properties of the iced wing are determined with an interactive scheme in which the solutions of the inviscid flow equations are obtained from a panel method and the solutions of the viscous flow equations are obtained from an inverse three-dimensional finite-difference boundary-layer method. A new interaction law is used to couple the inviscid and viscous flow solutions. The newly developed LEWICE multielement code is amplified to a high-lift configuration to calculate the ice shapes on the slat and on the main airfoil and on a four-element airfoil. The application of the LEWICE wing code to the calculation of ice shapes on a MS-317 swept wing shows good agreement with measurements. The interactive boundary-layer method is applied to a tapered iced wing in order to study the effect of icing on the aerodynamic properties of the wing at several angles of attack.
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.
F-8 oblique wing structural feasibility study
NASA Technical Reports Server (NTRS)
Koltko, E.; Katz, A.; Bell, M. A.; Smith, W. D.; Lauridia, R.; Overstreet, C. T.; Klapprott, C.; Orr, T. F.; Jobe, C. L.; Wyatt, F. G.
1975-01-01
The feasibility of fitting a rotating oblique wing on an F-8 aircraft to produce a full scale manned prototype capable of operating in the transonic and supersonic speed range was investigated. The strength, aeroelasticity, and fatigue life of such a prototype are analyzed. Concepts are developed for a new wing, a pivot, a skewing mechanism, control systems that operate through the pivot, and a wing support assembly that attaches in the F-8 wing cavity. The modification of the two-place NTF-8A aircraft to the oblique wing configuration is discussed.
A Discrete-Vortex Method for Studying the Wing Rock of Delta Wings
NASA Technical Reports Server (NTRS)
Gainer, Thomas G.
2002-01-01
A discrete-vortex method is developed to investigate the wing rock problem associated with highly swept wings. The method uses two logarithmic vortices placed above the wing to represent the vortex flow field and uses boundary conditions based on conical flow, vortex rate of change of momentum, and other considerations to position the vortices and determine their strengths. A relationship based on the time analogy and conical-flow assumptions is used to determine the hysteretic positions of the vortices during roll oscillations. Static and dynamic vortex positions and wing rock amplitudes and frequencies calculated by using the method are generally in good agreement with available experimental data. The results verify that wing rock is caused by hysteretic deflections of the vortices and indicate that the stabilizing moments that limit wing rock amplitudes are the result of the one primary vortex moving outboard of the wing where it has little influence on the wing.
Exploring the Role of Habitat on the Wettability of Cicada Wings.
Oh, Junho; Dana, Catherine E; Hong, Sungmin; Román, Jessica K; Jo, Kyoo Dong; Hong, Je Won; Nguyen, Jonah; Cropek, Donald M; Alleyne, Marianne; Miljkovic, Nenad
2017-08-16
Evolutionary pressure has pushed many extant species to develop micro/nanostructures that can significantly affect wettability and enable functionalities such as droplet jumping, self-cleaning, antifogging, antimicrobial, and antireflectivity. In particular, significant effort is underway to understand the insect wing surface structure to establish rational design tools for the development of novel engineered materials. Most studies, however, have focused on superhydrophobic wings obtained from a single insect species, in particular, the Psaltoda claripennis cicada. Here, we investigate the relationship between the spatially dependent wing wettability, topology, and droplet jumping behavior of multiple cicada species and their habitat, lifecycle, and interspecies relatedness. We focus on cicada wings of four different species: Neotibicen pruinosus, N. tibicen, Megatibicen dorsatus, and Magicicada septendecim and take a comparative approach. Using spatially resolved microgoniometry, scanning electron microscopy, atomic force microscopy, and high speed optical microscopy, we show that within cicada species, the wettability of wings is spatially homogeneous across wing cells. All four species were shown to have truncated conical pillars with widely varying length scales ranging from 50 to 400 nm in height. Comparison of the wettability revealed three cicada species with wings that are superhydrophobic (>150°) with low contact angle hysteresis (<5°), resulting in stable droplet jumping behavior. The fourth, more distantly related species (Ma. septendecim) showed only moderate hydrophobic behavior, eliminating some of the beneficial surface functional aspects for this cicada. Correlation between cicada habitat and wing wettability yielded little connection as wetter, swampy environments do not necessarily equate to higher measured wing hydrophobicity. The results, however, do point to species relatedness and reproductive strategy as a closer proxy for predicting wettability and surface structure and resultant enhanced wing surface functionality. This work not only elucidates the differences between inter- and intraspecies cicada wing topology, wettability, and water shedding behavior but also enables the development of rational design tools for the manufacture of artificial surfaces for energy and water applications.
NASA Astrophysics Data System (ADS)
Pearson, Roger A.
Rapidly advancing technology has developed multiple thin filmed devices capable of expanding the abilities of Small Unmanned Aircraft Systems (SUAS). This research develops a viable solution for integrating thin film solar cells into a currently operational SUAS. A wing was designed and produced that was capable of replacing the existing wing while providing additional functionality with embedded solar arrays. The study investigates the challenges of meeting the original requirements of the original equipment manufacturer wing while adapting it to fully protect and support structurally embedded payloads. In total, seven complete wings were produced and tested. Combinations of functional and simulated payloads were fully integrated into two of these wings. The merits of these designs were quantified and validated through both ground testing and flight testing with the SUAS.
Development of a stitched/RFI composite transport wing
NASA Technical Reports Server (NTRS)
Kropp, Yury
1995-01-01
Development of a composite wing primary structure for commercial transport aircraft is being undertaken at McDonnell Douglas under NASA contract. The focus of the program is to design and manufacture a low cost composite wing which can effectively compete with conventional metal wing structures in terms of cost, weight, and ability to withstand damage. These goals are being accomplished by utilizing the stitched/RFI manufacturing process during which the dry fiber preforms consisting of several stacks of warp-knit material are stitched together, impregnated with resin and cured. The stitched/RFI wing skin panels have exceptional damage tolerance and fatigue characteristics, are easily repairable, and can carry higher gross stress than their metal counterparts. This paper gives an overview of the program, describes the key features of the composite wing design and addresses major issues on analysis and manufacturing.
Development of elastomeric flight muscles for flapping wing micro air vehicles
NASA Astrophysics Data System (ADS)
Lau, Gih-Keong; Chin, Yao-Wei; La, Thanh-Giang
2017-04-01
Common drivers of flapping wings are a motorized crank mechanisms, which convert the motor rotation into wing reciprocation. Energetic efficiency of the motorized wing flappers can be quite low due to the lack of elastic storage and high friction. This paper relook into the flapping flight apparatus of natural flyers and draw inspiration to develop flight muscles capable of elastic storage, in addition to the frictionless thoracic compliant mechanisms. We review the recent findings on the use of dielectric elastomer actuators as flight muscles. We also discuss the challenges and the prospects of using dielectric elastomer minimum energy structure to create large and fast bending/unbending, possibly for wing flapping.
Study of Effects of Sweep on the Flutter of Cantilever Wings
NASA Technical Reports Server (NTRS)
Barmby, J G; Cunningham, H J; Garrick, I E
1951-01-01
An experimental and analytical investigation of the flutter of sweptback cantilever wings is reported. The experiments employed groups of wings swept back by rotating and by shearing. The angle of sweep range from 0 degree to 60 degrees and Mach numbers extended to approximately 0.85. A theoretical analysis of the air forces on an oscillating swept wing of high length-chord ratio is developed, and the approximations inherent in the assumptions are discussed. Comparison with experiment indicates that the analysis developed in the present report is satisfactory for giving the main effects of sweep, at least for nearly uniform cantilever wings of high and moderate length-chord ratios.
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.
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
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.
Development of an Automatic Grid Generator for Multi-Element High-Lift Wings
NASA Technical Reports Server (NTRS)
Eberhardt, Scott; Wibowo, Pratomo; Tu, Eugene
1996-01-01
The procedure to generate the grid around a complex wing configuration is presented in this report. The automatic grid generation utilizes the Modified Advancing Front Method as a predictor and an elliptic scheme as a corrector. The scheme will advance the surface grid one cell outward and the newly obtained grid is corrected using the Laplace equation. The predictor-corrector step ensures that the grid produced will be smooth for every configuration. The predictor-corrector scheme is extended for a complex wing configuration. A new technique is developed to deal with the grid generation in the wing-gaps and on the flaps. It will create the grids that fill the gap on the wing surface and the gap created by the flaps. The scheme recognizes these configurations automatically so that minimal user input is required. By utilizing an appropriate sequence in advancing the grid points on a wing surface, the automatic grid generation for complex wing configurations is achieved.
The Aerodynamics of Deforming Wings at Low Reynolds Number
NASA Astrophysics Data System (ADS)
Medina, Albert
Flapping flight has gained much attention in the past decade driven by the desire to understand capabilities observed in nature and the desire to develop agile small-scale aerial vehicles. Advancing our current understanding of unsteady aerodynamics is an essential component in the development of micro-air vehicles (MAV) intended to utilize flight mechanics akin to insect flight. Thus the efforts undertaken that of bio-mimicry. The complexities of insect wing motion are dissected and simplified to more tractable problems to elucidate the fundamentals of unsteady aerodynamics in biologically inspired kinematics. The MAV's fruition would satisfy long established needs in both the military and civilian sectors. Although recent studies have provided great insight into the lift generating mechanisms of flapping wings the deflection response of such wings remains poorly understood. This dissertation numerically and experimentally investigates the aerodynamic performance of passively and actively deflected wings in hover and rotary kinematics. Flexibility is distilled to discrete lines of flexion which acknowledging major flexion lines in insect wings to be the primary avenue for deformation. Of primary concern is the development of the leading-edge vortex (LEV), a high circulation region of low pressure above the wing to which much of the wing's lift generation is attributed. Two-dimensional simulations of wings with chord-wise flexibility in a freestream reveal a lift generating mechanism unavailable to rigid wings with origins in vortical symmetry breaking. The inclusion of flexibility in translating wings accelerated from rest revealed the formation time of the initial LEV was very weakly dependent on the flexible stiffness of the wing, maintaining a universal time scale of four to five chords of travel before shedding. The frequency of oscillatory shedding of the leading and trailing-edge vortices that develops after the initial vortex shedding was shown to be responsive to flexibility satisfying an inverse proportionality to stiffness. In hover, an effective pitch angle can be defined in a flexible wing that accounts for deflection which shifts results toward trend lines of rigid wings. Three-dimensional simulations examining the effects of two distinct deformation modes undergoing prescribed deformation associated with root and tip deflection demonstrated a greater aerodynamic response to tip deflection in hover. Efficiency gains in flexion wings over rigid wing counterpart were shown to be dependent on Reynolds number with efficiency in both modes increasing with increased Reynolds number. Additionally, while the leading-edge vortex axis proved insensitive to deformation, the shape and orientation of the LEV core is modified. Experiments on three-dimensional dynamically-scaled fruit fly wings with passive deformation operating in the bursting limit Reynolds number regime revealed enhanced leading-edge vortex bursting with tip deflection promoting greater LEV core flow deceleration in stroke. Experimental studies on rotary wings highlights a universal formation time of the leading-edge vortex independent of Reynolds number, acceleration profile and aspect ratio. Efforts to replicate LEV bursting phenomena of higher aspect ratio wings in a unity aspect ratio wing such that LEV growth is no limited by span but by the LEV traversing the chord revealed a flow regime of oscillatory lift generation reminiscent of behavior exhibited in translating wings that also maintains magnitude peak to peak.
NASA Technical Reports Server (NTRS)
Krist, Steven E.; Bauer, Steven X. S.
1999-01-01
The design process for developing the natural flow wing design on the HSR arrow wing configuration utilized several design tools and analysis methods. Initial fuselage/wing designs were generated with inviscid analysis and optimization methods in conjunction with the natural flow wing design philosophy. A number of designs were generated, satisfying different system constraints. Of the three natural flow wing designs developed, the NFWAc2 configuration is the design which satisfies the constraints utilized by McDonnell Douglas Aerospace (MDA) in developing a series of optimized configurations; a wind tunnel model of the MDA designed OPT5 configuration was constructed and tested. The present paper is concerned with the viscous analysis and inverse design of the arrow wing configurations, including the effects of the installed diverters/nacelles. Analyses were conducted with OVERFLOW, a Navier-Stokes flow solver for overset grids. Inverse designs were conducted with OVERDISC, which couples OVERFLOW with the CDISC inverse design method. An initial system of overset grids was generated for the OPT5 configuration with installed diverters/nacelles. An automated regridding process was then developed to use the OPT5 component grids to create grids for the natural flow wing designs. The inverse design process was initiated using the NFWAc2 configuration as a starting point, eventually culminating in the NFWAc4 design-for which a wind tunnel model was constructed. Due to the time constraints on the design effort, initial analyses and designs were conducted with a fairly coarse grid; subsequent analyses have been conducted on a refined system of grids. Comparisons of the computational results to experiment are provided at the end of this paper.
Experimental Influence Coefficients and Vibration Modes
NASA Technical Reports Server (NTRS)
Weidman, Deene J.; Kordes, Eldon E.
1959-01-01
Test results are presented for both symmetrical and antisymmetrical static loading of a wing model mounted on a three-point support system. The first six free-free vibration modes were determined experimentally. A comparison is made of the symmetrical nodal patterns and frequencies with the symmetrical nodal patterns and frequencies calculated from the experimental influence coefficients.
2012-01-01
Background To explain eyespot colour-pattern determination in butterfly wings, the induction model has been discussed based on colour-pattern analyses of various butterfly eyespots. However, a detailed structural analysis of eyespots that can serve as a foundation for future studies is still lacking. In this study, fundamental structural rules related to butterfly eyespots are proposed, and the induction model is elaborated in terms of the possible dynamics of morphogenic signals involved in the development of eyespots and parafocal elements (PFEs) based on colour-pattern analysis of the nymphalid butterfly Junonia almana. Results In a well-developed eyespot, the inner black core ring is much wider than the outer black ring; this is termed the inside-wide rule. It appears that signals are wider near the focus of the eyespot and become narrower as they expand. Although fundamental signal dynamics are likely to be based on a reaction-diffusion mechanism, they were described well mathematically as a type of simple uniformly decelerated motion in which signals associated with the outer and inner black rings of eyespots and PFEs are released at different time points, durations, intervals, and initial velocities into a two-dimensional field of fundamentally uniform or graded resistance; this produces eyespots and PFEs that are diverse in size and structure. The inside-wide rule, eyespot distortion, structural differences between small and large eyespots, and structural changes in eyespots and PFEs in response to physiological treatments were explained well using mathematical simulations. Natural colour patterns and previous experimental findings that are not easily explained by the conventional gradient model were also explained reasonably well by the formal mathematical simulations performed in this study. Conclusions In a mode free from speculative molecular interactions, the present study clarifies fundamental structural rules related to butterfly eyespots, delineates a theoretical basis for the induction model, and proposes a mathematically simple mode of long-range signalling that may reflect developmental mechanisms associated with butterfly eyespots. PMID:22409965
Otaki, Joji M
2012-03-13
To explain eyespot colour-pattern determination in butterfly wings, the induction model has been discussed based on colour-pattern analyses of various butterfly eyespots. However, a detailed structural analysis of eyespots that can serve as a foundation for future studies is still lacking. In this study, fundamental structural rules related to butterfly eyespots are proposed, and the induction model is elaborated in terms of the possible dynamics of morphogenic signals involved in the development of eyespots and parafocal elements (PFEs) based on colour-pattern analysis of the nymphalid butterfly Junonia almana. In a well-developed eyespot, the inner black core ring is much wider than the outer black ring; this is termed the inside-wide rule. It appears that signals are wider near the focus of the eyespot and become narrower as they expand. Although fundamental signal dynamics are likely to be based on a reaction-diffusion mechanism, they were described well mathematically as a type of simple uniformly decelerated motion in which signals associated with the outer and inner black rings of eyespots and PFEs are released at different time points, durations, intervals, and initial velocities into a two-dimensional field of fundamentally uniform or graded resistance; this produces eyespots and PFEs that are diverse in size and structure. The inside-wide rule, eyespot distortion, structural differences between small and large eyespots, and structural changes in eyespots and PFEs in response to physiological treatments were explained well using mathematical simulations. Natural colour patterns and previous experimental findings that are not easily explained by the conventional gradient model were also explained reasonably well by the formal mathematical simulations performed in this study. In a mode free from speculative molecular interactions, the present study clarifies fundamental structural rules related to butterfly eyespots, delineates a theoretical basis for the induction model, and proposes a mathematically simple mode of long-range signalling that may reflect developmental mechanisms associated with butterfly eyespots.
Wind tunnel tests of a free-wing/free-trimmer model
NASA Technical Reports Server (NTRS)
Sandlin, D. R.
1982-01-01
The riding qualities of an aircraft with low wing loading can be improved by freeing the wing to rotate about its spanwise axis. A trimming surface also free to rotate about its spanwise axis can be added at the wing tips to permit the use of high lift devices. Wind tunnel tests of the free wing/free trimmer model with the trimmer attached to the wing tips aft of the wing chord were conducted to validate a mathematical model developed to predict the dynamic characteristics of a free wing/free trimmer aircraft. A model consisting of a semispan wing with the trimmer mounted on with the wing on an air bearing and the trimmer on a ball bearing was displaced to various angles of attack and released. The damped oscillations of the wing and trimmer were recorded. Real and imaginary parts of the characteristic equations of motion were determined and compared to values predicted using the mathematical model.
NASA Technical Reports Server (NTRS)
Boelens, Okko J.; Luckring, James M.; Breitsamter, Christian; Hovelmann, Andreas; Knoth, Florian; Malloy, Donald J.; Deck, Sebatien
2015-01-01
A diamond-wing configuration has been developed to isolate and study blunt-leading edge vortex separation with both computations and experiments. The wing has been designed so that the results are relevant to a more complex Uninhabited Combat Air Vehicle concept known as SACCON. The numerical and theoretical development process for this diamond wing is presented, including a view toward planned wind tunnel experiments. This work was conducted under the NATO Science and Technology Organization, Applied Vehicle Technology panel. All information is in the public domain.
Development of a Non-autorotative Airplane Capable of Steep Landing
NASA Technical Reports Server (NTRS)
Schmidt, Wilhelm
1931-01-01
In the following we develop a non-autorotating monoplane wing. The conditions imposed on such a wing, aside from its freedom from autorotation,with respect to its polars and its construction, are taken into account as far as possible. It is indicated that the autorotation characteristics of a wing are dependent upon the speed of air flow as well as on the angle of yaw. This report postulates the knowledge of the behavior of certain conventional wings of different chords and cambers with respect to their air loads at large angles of attack.
Superfast high-resolution absolute 3D recovery of a stabilized flapping flight process.
Li, Beiwen; Zhang, Song
2017-10-30
Scientific research of a stabilized flapping flight process (e.g. hovering) has been of great interest to a variety of fields including biology, aerodynamics, and bio-inspired robotics. Different from the current passive photogrammetry based methods, the digital fringe projection (DFP) technique has the capability of performing dense superfast (e.g. kHz) 3D topological reconstructions with the projection of defocused binary patterns, yet it is still a challenge to measure a flapping flight process with the presence of rapid flapping wings. This paper presents a novel absolute 3D reconstruction method for a stabilized flapping flight process. Essentially, the slow motion parts (e.g. body) and the fast-motion parts (e.g. wings) are segmented and separately reconstructed with phase shifting techniques and the Fourier transform, respectively. The topological relations between the wings and the body are utilized to ensure absolute 3D reconstruction. Experiments demonstrate the success of our computational framework by testing a flapping wing robot at different flapping speeds.
Underwater flight by the planktonic sea butterfly.
Murphy, David W; Adhikari, Deepak; Webster, Donald R; Yen, Jeannette
2016-02-01
In a remarkable example of convergent evolution, we show that the zooplanktonic sea butterfly Limacina helicina 'flies' underwater in the same way that very small insects fly in the air. Both sea butterflies and flying insects stroke their wings in a characteristic figure-of-eight pattern to produce lift, and both generate extra lift by peeling their wings apart at the beginning of the power stroke (the well-known Weis-Fogh 'clap-and-fling' mechanism). It is highly surprising to find a zooplankter 'mimicking' insect flight as almost all zooplankton swim in this intermediate Reynolds number range (Re=10-100) by using their appendages as paddles rather than wings. The sea butterfly is also unique in that it accomplishes its insect-like figure-of-eight wing stroke by extreme rotation of its body (what we call 'hyper-pitching'), a paradigm that has implications for micro aerial vehicle (MAV) design. No other animal, to our knowledge, pitches to this extent under normal locomotion. © 2016. Published by The Company of Biologists Ltd.
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.
Flexible-Wing-Based Micro Air Vehicles
NASA Technical Reports Server (NTRS)
Ifju, Peter G.; Jenkins, David A.; Ettinger, Scott; Lian, Yong-Sheng; Shyy, Wei; Waszak, Martin R.
2002-01-01
This paper documents the development and evaluation of an original flexible-wing-based Micro Air Vehicle (MAV) technology that reduces adverse effects of gusty wind conditions and unsteady aerodynamics, exhibits desirable flight stability, and enhances structural durability. The flexible wing concept has been demonstrated on aircraft with wingspans ranging from 18 inches to 5 inches. Salient features of the flexible-wing-based MAV, including the vehicle concept, flexible wing design, novel fabrication methods, aerodynamic assessment, and flight data analysis are presented.
Numerical and experimental investigations on unsteady aerodynamics of flapping wings
NASA Astrophysics Data System (ADS)
Yu, Meilin
The development of a dynamic unstructured grid high-order accurate spectral difference (SD) method for the three dimensional compressible Navier-Stokes (N-S) equations and its applications in flapping-wing aerodynamics are carried out in this work. Grid deformation is achieved via an algebraic blending strategy to save computational cost. The Geometric Conservation Law (GCL) is imposed to ensure that grid deformation will not contaminate the flow physics. A low Mach number preconditioning procedure is conducted in the developed solver to handle the bio-inspired flow. The capability of the low Mach number preconditioned SD solver is demonstrated by a series of two dimensional (2D) and three dimensional (3D) simulations of the unsteady vortex dominated flow. Several topics in the flapping wing aerodynamics are numerically and experimentally investigated in this work. These topics cover some of the cutting-edge issues in flapping wing aerodynamics, including the wake structure analysis, airfoil thickness and kinematics effects on the aerodynamic performances, vortex structure analysis around 3D flapping wings and the kinematics optimization. Wake structures behind a sinusoidally pitching NACA0012 airfoil are studied with both experimental and numerical approaches. The experiments are carried out with Particle Image Velocimetry (PIV) and two types of wake transition processes, namely the transition from a drag-indicative wake to a thrust-indicative wake and that from the symmetric wake to the asymmetric wake are distinguished. The numerical results from the developed SD solver agree well with the experimental results. It is numerically found that the deflective direction of the asymmetric wake is determined by the initial conditions, e.g. initial phase angle. As most insects use thin wings (i. e., wing thickness is only a few percent of the chord length) in flapping flight, the effects of airfoil thickness on thrust generation are numerically investigated by simulating the flow fields around a series of plunging NACA symmetric airfoils with thickness ratio ranging from 4.0% to 20.0% of the airfoil chord length. The contribution of viscous force to flapping propulsion is accessed and it is found that viscous force becomes thrust producing, instead of drag producing, and plays a non-negligible role in thrust generation for thin airfoils. This is closely related to the variations of the dynamics of the unsteady vortex structures around the plunging airfoils. As nature flyers use complex wing kinematics in flapping flight, kinematics effects on the aerodynamic performance with different airfoil thicknesses are numerically studied by using a series of NACA symmetric airfoils. It is found that the combined plunging and pitching motion can outperform the pure plunging or pitching motion by sophisticatedly adjusting the airfoil gestures during the oscillation stroke. The thin airfoil better manipulates leading edge vortices (LEVs) than the thick airfoil (NACA0030) does in studied cases, and there exists an optimal thickness for large thrust generation with reasonable propulsive efficiency. With the present kinematics and dynamic parameters, relatively low reduced frequency is conducive for thrust production and propulsive efficiency for all tested airfoil thicknesses. In order to obtain the optimal kinematics parameters of flapping flight, a kinematics optimization is then performed. A gradient-based optimization algorithm is coupled with a second-order SD Navier-Stokes solver to search for the optimal kinematics of a certain airfoil undergoing a combined plunging and pitching motion. Then a high-order SD scheme is used to verify the optimization results and reveal the detailed vortex structures associated with the optimal kinematics of the flapping flight. It is found that for the case with maximum propulsive efficiency, there exists no leading edge separation during most of the oscillation cycle. In order to provide constructive suggestions to the design of micro-air-vehicles (MAVs), 3D simulations of the flapping wings are carried out in this work. Both the rectangular and bio-inspired wings with different kinematics are investigated. The formation process of two-jet-like wake patterns behind the finite-span flapping wing is found to be closely related to the interaction between trailing edge vortices and tip vortices. Then the effects of the wing planforms on the aerodynamics performance of the finite-span flapping wings are elucidated in terms of the evolution and dynamic interaction of unsteady vortex structures.
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.
Scaling the Drosophila Wing: TOR-Dependent Target Gene Access by the Hippo Pathway Transducer Yorkie
Parker, Joseph; Struhl, Gary
2015-01-01
Organ growth is controlled by patterning signals that operate locally (e.g., Wingless/Ints [Wnts], Bone Morphogenetic Proteins [BMPs], and Hedgehogs [Hhs]) and scaled by nutrient-dependent signals that act systemically (e.g., Insulin-like peptides [ILPs] transduced by the Target of Rapamycin [TOR] pathway). How cells integrate these distinct inputs to generate organs of the appropriate size and shape is largely unknown. The transcriptional coactivator Yorkie (Yki, a YES-Associated Protein, or YAP) acts downstream of patterning morphogens and other tissue-intrinsic signals to promote organ growth. Yki activity is regulated primarily by the Warts/Hippo (Wts/Hpo) tumour suppressor pathway, which impedes nuclear access of Yki by a cytoplasmic tethering mechanism. Here, we show that the TOR pathway regulates Yki by a separate and novel mechanism in the Drosophila wing. Instead of controlling Yki nuclear access, TOR signaling governs Yki action after it reaches the nucleus by allowing it to gain access to its target genes. When TOR activity is inhibited, Yki accumulates in the nucleus but is sequestered from its normal growth-promoting target genes—a phenomenon we term “nuclear seclusion.” Hence, we posit that in addition to its well-known role in stimulating cellular metabolism in response to nutrients, TOR also promotes wing growth by liberating Yki from nuclear seclusion, a parallel pathway that we propose contributes to the scaling of wing size with nutrient availability. PMID:26474042
NASA Astrophysics Data System (ADS)
Bai, Linge; Widmann, Thomas; Jülicher, Frank; Dahmann, Christian; Breen, David
2013-01-01
Quantifying and visualizing the shape of developing biological tissues provide information about the morphogenetic processes in multicellular organisms. The size and shape of biological tissues depend on the number, size, shape, and arrangement of the constituting cells. To better understand the mechanisms that guide tissues into their final shape, it is important to investigate the cellular arrangement within tissues. Here we present a data processing pipeline to generate 3D volumetric surface models of epithelial tissues, as well as geometric descriptions of the tissues' apical cell cross-sections. The data processing pipeline includes image acquisition, editing, processing and analysis, 2D cell mesh generation, 3D contourbased surface reconstruction, cell mesh projection, followed by geometric calculations and color-based visualization of morphological parameters. In their first utilization we have applied these procedures to construct a 3D volumetric surface model at cellular resolution of the wing imaginal disc of Drosophila melanogaster. The ultimate goal of the reported effort is to produce tools for the creation of detailed 3D geometric models of the individual cells in epithelial tissues. To date, 3D volumetric surface models of the whole wing imaginal disc have been created, and the apicolateral cell boundaries have been identified, allowing for the calculation and visualization of cell parameters, e.g. apical cross-sectional area of cells. The calculation and visualization of morphological parameters show position-dependent patterns of cell shape in the wing imaginal disc. Our procedures should offer a general data processing pipeline for the construction of 3D volumetric surface models of a wide variety of epithelial tissues.
Experimental Optimization of a Free-to-Rotate Wing for Small UAS
NASA Technical Reports Server (NTRS)
Logan, Michael J.; DeLoach, Richard; Copeland, Tiwana; Vo, Steven
2014-01-01
This paper discusses an experimental investigation conducted to optimize a free-to-rotate wing for use on a small unmanned aircraft system (UAS). Although free-to-rotate wings have been used for decades on various small UAS and small manned aircraft, little is known about how to optimize these unusual wings for a specific application. The paper discusses some of the design rationale of the basic wing. In addition, three main parameters were selected for "optimization", wing camber, wing pivot location, and wing center of gravity (c.g.) location. A small apparatus was constructed to enable some simple experimental analysis of these parameters. A design-of-experiment series of tests were first conducted to discern which of the main optimization parameters were most likely to have the greatest impact on the outputs of interest, namely, some measure of "stability", some measure of the lift being generated at the neutral position, and how quickly the wing "recovers" from an upset. A second set of tests were conducted to develop a response-surface numerical representation of these outputs as functions of the three primary inputs. The response surface numerical representations are then used to develop an "optimum" within the trade space investigated. The results of the optimization are then tested experimentally to validate the predictions.
Modeling and Optimization for Morphing Wing Concept Generation
NASA Technical Reports Server (NTRS)
Skillen, Michael D.; Crossley, William A.
2007-01-01
This report consists of two major parts: 1) the approach to develop morphing wing weight equations, and 2) the approach to size morphing aircraft. Combined, these techniques allow the morphing aircraft to be sized with estimates of the morphing wing weight that are more credible than estimates currently available; aircraft sizing results prior to this study incorporated morphing wing weight estimates based on general heuristics for fixed-wing flaps (a comparable "morphing" component) but, in general, these results were unsubstantiated. This report will show that the method of morphing wing weight prediction does, in fact, drive the aircraft sizing code to different results and that accurate morphing wing weight estimates are essential to credible aircraft sizing results.
USDA-ARS?s Scientific Manuscript database
The arrival of the term Eco-Evo-Devo highlights the need to incorporate ecology and development into modern evolutionary research to better understand processes such as adaptation and speciation as well as the effect of environmental changes a species. As basal winged insects (pterygotes), dragonfli...
Federal Register 2010, 2011, 2012, 2013, 2014
2011-02-07
..., an ignition source can develop in the wing tank vapour space during fuel transfer from bag tank CWT..., an ignition source can develop in the wing tank vapour space during fuel transfer from bag tank CWT..., all serial numbers, equipped with a center wing tank (CWT); and Model F.28 [[Page 6543
The NASA supercritical-wing technology
NASA Technical Reports Server (NTRS)
Bartlett, D. W.; Patterson, J. C., Jr.
1978-01-01
A number of high aspect ratio supercritical wings in combination with a representative wide body type fuselage were tested in the Langley 8 foot transonic pressure tunnel. The wing parameters investigated include aspect ratio, sweep, thickness to chord ratio, and camber. Subsequent to these initial series of tests, a particular wing configuration was selected for further study and development. Tests on the selected wing involved the incorporation of a larger inboard trailing edge extension, an inboard leading edge extension, and flow through nacelles. Range factors for the various supercritical wing configurations are compared with those for a reference wide body transport configuration.
NASA Technical Reports Server (NTRS)
Watts, D. J.
1978-01-01
The overall wing study objectives are to study and plan the effort by commercial transport aircraft manufacturers to accomplish the transition from current conventional materials and practices to extensive use of advanced composites in wings of aircraft that will enter service in the 1985-1990 time period. Specific wing study objectives are to define the technology and data needed to support an aircraft manufacturer's commitment to utilize composites primary wing structure in future production aircraft and to develop plans for a composite wing technology program which will provide the needed technology and data.
Aeroelastic tailoring and structural optimization of joined-wing configurations
NASA Astrophysics Data System (ADS)
Lee, Dong-Hwan
2002-08-01
Methodology for integrated aero-structural design was developed using formal optimization. ASTROS (Automated STRuctural Optimization System) was used as an analyzer and an optimizer for performing joined-wing weight optimization with stress, displacement, cantilever or body-freedom flutter constraints. As a pre/post processor, MATLAB was used for generating input file of ASTROS and for displaying the results of the ASTROS. The effects of the aeroelastic constraints on the isotropic and composite joined-wing weight were examined using this developed methodology. The aeroelastic features of a joined-wing aircraft were examined using both the Rayleigh-Ritz method and a finite element based aeroelastic stability and weight optimization procedure. Aircraft rigid-body modes are included to analyze of body-freedom flutter of the joined-wing aircraft. Several parametric studies were performed to determine the most important parameters that affect the aeroelastic behavior of a joined-wing aircraft. The special feature of a joined-wing aircraft is body-freedom flutter involving frequency interaction of the first elastic mode and the aircraft short period mode. In most parametric study cases, the body-freedom flutter speed was less than the cantilever flutter speed that is independent of fuselage inertia. As fuselage pitching moment of inertia was increased, the body-freedom flutter speed increased. When the pitching moment of inertia reaches a critical value, transition from body-freedom flutter to cantilever flutter occurred. The effects of composite laminate orientation on the front and rear wings of a joined-wing configuration were studied. An aircraft pitch divergence mode, which occurred because of forward movement of center of pressure due to wing deformation, was found. Body-freedom flutter and cantilever-like flutter were also found depending on combination of front and rear wing ply orientations. Optimized wing weight behaviors of the planar and non-planar configurations with isotropic and composite materials were investigated. Wing weight optimization of the composite joined-wing result in less weight compared to the metallic wing. Fuselage flexibility affects joined-wing flutter characteristics. Elastic mode shapes of the wing were affected by fuselage deformation and change the flutter speeds compared to the rigid fuselage. Body-freedom flutter speeds decrease as fuselage flexibility increases. Optimum wing weights increase as fuselage flexibility increases. Flutter analysis of a box wing configuration investigated the effects of center of gravity location and pitch moment of inertia on flutter speed.
Extended Range Aerial Delivery Using an Unpowered Autonomous Tailless UAV
NASA Astrophysics Data System (ADS)
Kraft, Tyler E.
An alternative approach for precision aerial delivery utilizing a flying wing for controllable forward glide is presented. Although effective, current delivery methods either display a lack of control, or require close standoff distances, potentially endangering aircraft personnel as well as bystanders. Hardware-in-the-loop simulations provide an efficient method for evaluating various wing designs and actuation configurations. Four control surface configurations are presented and evaluated, encompassing traditional aircraft and ram-air parafoil control approaches. Fixed-wing and multirotor unmanned aircraft-based flight tests were conducted to evaluate the controllability and handling performance of the various configurations of both a fixed wing model and a model with collapsing wings. A manufacturing process was developed to allow repeatable results in the field using cheap, mostly disposable materials. A powered flying wing model was used to maximize data collection in later stages of software development. Data collected during flight tests was used to create a model of the system and develop a Nonlinear Dynamic Inversion controller for autonomous flight. The NDI controller was able to provide stable flight in pitch, but will need more development to control yaw, instead an intentional bias was built in to show proof of concept for direct yaw control. The results demonstrate the feasibility of the flying wing-based aerial delivery; however, significant challenges remain regarding the stability and scalability of the system.
NASA Technical Reports Server (NTRS)
Ahmadi, A. R.
1981-01-01
A low frequency unsteady lifting-line theory is developed for a harmonically oscillating wing of large aspect ratio. The wing is assumed to be chordwise rigid but completely flexible in the span direction. The theory is developed by use of the method of matched asymptotic expansions which reduces the problem from a singular integral equation to quadrature. The wing displacements are prescribed and the pressure field, airloads, and unsteady induced downwash are obtained in closed form. The influence of reduced frequency, aspect ratio, planform shape, and mode of oscillation on wing aerodynamics is demonstrated through numerical examples. Compared with lifting-surface theory, computation time is reduced significantly. Using the present theory, the energetic quantities associated with the propulsive performance of a finite wing oscillating in combined pitch and heave are obtained in closed form. Numerical examples are presented for an elliptic wing.
Control of forward swept wing configurations dominated by flight-dynamic/aeroelastic interactions
NASA Technical Reports Server (NTRS)
Rimer, M.; Chipman, R.; Muniz, B.
1984-01-01
An active control system concept for an aeroelastic wind-tunnel model of a statically unstable FSW configuration with wing-mounted stores is developed to provide acceptable longitudinal flying qualities while maintaining adequate flutter speed margin. On FSW configurations, the inherent aeroelastic wing divergence tendency causes strong flight-dynamic/aeroelastic interactions that in certain cases can produce a dynamic instability known as body-freedom flutter (BFF). The carriage of wing-mounted stores is shown to severely aggravate this problem. The control system developed combines a canard-based SAS with an Active Divergence/Flutter Suppression (ADFS) system which relies on wing-mounted sensors and a trailing-edge device (flaperon). Synergism between these two systems is exploited to obtain the flying qualities and flutter speed objectives.
NASA Innovation Fund 2010 Project Elastically Shaped Future Air Vehicle Concept
NASA Technical Reports Server (NTRS)
Nguyen, Nhan
2010-01-01
This report describes a study conducted in 2010 under the NASA Innovation Fund Award to develop innovative future air vehicle concepts. Aerodynamic optimization was performed to produce three different aircraft configuration concepts for low drag, namely drooped wing, inflected wing, and squashed fuselage. A novel wing shaping control concept is introduced. This concept describes a new capability of actively controlling wing shape in-flight to minimize drag. In addition, a novel flight control effector concept is developed to enable wing shaping control. This concept is called a variable camber continuous trailing edge flap that can reduce drag by as much as 50% over a conventional flap. In totality, the potential benefits of fuel savings offered by these concepts can be significant.
Structural tests and development of a laminar flow control wing surface composite chordwise joint
NASA Technical Reports Server (NTRS)
Lineberger, L. B.
1984-01-01
The dramatic increases in fuel costs and the potential for periods of limited fuel availability provided the impetus to explore technologies to reduce transport aircraft fuel consumption. NASA sponsored the Aircraft Energy Efficiency (ACEE) program beginning in 1976 to develop technologies to improve fuel efficiency. The Lockheed-Georgia Company accomplished under NAS1-16235 Laminar-Flow-Control (LFC) Wing Panel Structural Design and Development (WSSD); design, manufacturing, and testing activities. An in-depth preliminary design of the baseline 1993 LFC wing was accomplished. A surface panel using the Lockheed graphite/epoxy integrated LFC wing box structural concept was designed. The concept was shown by analysis to be structurally efficient and cost effective. Critical details of the surface and surface joint was demonstrated by fabricating and testing complex, concept selection specimens. The Lockheed-Georgia Company accomplishments, Development of LFC Wind Surface Composite Structures (WSCS), are documented. Tests were conducted on two CV2 panels to verify the static tension and fatigue strength of LFC wing surface chordwise joints.
Application of winglets and/or wing tip extensions with active load control on the Boeing 747
NASA Technical Reports Server (NTRS)
Allison, R. L.; Perkin, B. R.; Schoenman, R. L.
1978-01-01
The application of wing tip modifications and active control technology to the Boeing 747 airplane for the purpose of improving fuel efficiency is considered. Wing tip extensions, wing tip winglets, and the use of the outboard ailerons for active wing load alleviation are described. Modest performance improvements are indicated. A costs versus benefits approach is taken to decide which, if any, of the concepts warrant further development and flight test leading to possible incorporation into production airplanes.
NASA Technical Reports Server (NTRS)
Lovejoy, Andrew E.
2015-01-01
A structural concept called pultruded rod stitched efficient unitized structure (PRSEUS) was developed by the Boeing Company to address the complex structural design aspects associated with a pressurized hybrid wing body (HWB) aircraft configuration. While PRSEUS was an enabling technology for the pressurized HWB structure, limited investigation of PRSEUS for other aircraft structures, such as circular fuselages and wings, has been done. Therefore, a study was undertaken to investigate the potential weight savings afforded by using the PRSEUS concept for a commercial transport wing. The study applied PRSEUS to the Advanced Subsonic Technology (AST) Program composite semi-span test article, which was sized using three load cases. The initial PRSEUS design was developed by matching cross-sectional stiffnesses for each stringer/skin combination within the wing covers, then the design was modified to ensure that the PRSEUS design satisfied the design criteria. It was found that the PRSEUS wing design exhibited weight savings over the blade-stiffened composite AST Program wing of nearly 9%, and a weight savings of 49% and 29% for the lower and upper covers, respectively, compared to an equivalent metallic wing.
Development and Testing of Control Laws for the Active Aeroelastic Wing Program
NASA Technical Reports Server (NTRS)
Dibley, Ryan P.; Allen, Michael J.; Clarke, Robert; Gera, Joseph; Hodgkinson, John
2005-01-01
The Active Aeroelastic Wing research program was a joint program between the U.S. Air Force Research Laboratory and NASA established to investigate the characteristics of an aeroelastic wing and the technique of using wing twist for roll control. The flight test program employed the use of an F/A-18 aircraft modified by reducing the wing torsional stiffness and adding a custom research flight control system. The research flight control system was optimized to maximize roll rate using only wing surfaces to twist the wing while simultaneously maintaining design load limits, stability margins, and handling qualities. NASA Dryden Flight Research Center developed control laws using the software design tool called CONDUIT, which employs a multi-objective function optimization to tune selected control system design parameters. Modifications were made to the Active Aeroelastic Wing implementation in this new software design tool to incorporate the NASA Dryden Flight Research Center nonlinear F/A-18 simulation for time history analysis. This paper describes the design process, including how the control law requirements were incorporated into constraints for the optimization of this specific software design tool. Predicted performance is also compared to results from flight.
Su, Shiyu; Lim, Matthew; Kunte, Krushnamegh
2015-11-01
Predation exerts strong selection on mimetic butterfly wing color patterns, which also serve other functions such as sexual selection. Therefore, specific selection pressures may affect the sexes and signal components differentially. We tested three predictions about the evolution of mimetic resemblance by comparing wing coloration of aposematic butterflies and their Batesian mimics: (a) females gain greater mimetic advantage than males and therefore are better mimics, (b) due to intersexual genetic correlations, sexually monomorphic mimics are better mimics than female-limited mimics, and (c) mimetic resemblance is better on the dorsal wing surface that is visible to predators in flight. Using a physiological model of avian color vision, we quantified mimetic resemblance from predators' perspective, which showed that female butterflies were better mimics than males. Mimetic resemblance in female-limited mimics was comparable to that in sexually monomorphic mimics, suggesting that intersexual genetic correlations did not constrain adaptive response to selection for female-limited mimicry. Mimetic resemblance on the ventral wing surface was better than that on the dorsal wing surface, implying stronger natural and sexual selection on ventral and dorsal surfaces, respectively. These results suggest that mimetic resemblance in butterfly mimicry rings has evolved under various selective pressures acting in a sex- and wing surface-specific manner. © 2015 The Author(s). Evolution © 2015 The Society for the Study of Evolution.
Development of tailless and all-wing gliders and airplanes
NASA Technical Reports Server (NTRS)
Lademann, Robert W E
1932-01-01
Tailless airplanes are characterized by having all their control surfaces, especially the elevator, incorporated in the wings. This paper provides a discussion of the history of their development and current state of development.
Multidisciplinary Shape Optimization of a Composite Blended Wing Body Aircraft
NASA Astrophysics Data System (ADS)
Boozer, Charles Maxwell
A multidisciplinary shape optimization tool coupling aerodynamics, structure, and performance was developed for battery powered aircraft. Utilizing high-fidelity computational fluid dynamics analysis tools and a structural wing weight tool, coupled based on the multidisciplinary feasible optimization architecture; aircraft geometry is modified in the optimization of the aircraft's range or endurance. The developed tool is applied to three geometries: a hybrid blended wing body, delta wing UAS, the ONERA M6 wing, and a modified ONERA M6 wing. First, the optimization problem is presented with the objective function, constraints, and design vector. Next, the tool's architecture and the analysis tools that are utilized are described. Finally, various optimizations are described and their results analyzed for all test subjects. Results show that less computationally expensive inviscid optimizations yield positive performance improvements using planform, airfoil, and three-dimensional degrees of freedom. From the results obtained through a series of optimizations, it is concluded that the newly developed tool is both effective at improving performance and serves as a platform ready to receive additional performance modules, further improving its computational design support potential.
The Application of the NFW Design Philosophy to the HSR Arrow Wing Configuration
NASA Technical Reports Server (NTRS)
Bauer, Steven X. S.; Krist, Steven E.
1999-01-01
The Natural Flow Wing design philosophy was developed for improving performance characteristics of highly-swept fighter aircraft at cruise and maneuvering conditions across the Mach number range (from Subsonic through Supersonic). The basic philosophy recognizes the flow characteristics that develop on highly swept wings and contours the surface to take advantage of those flow characteristics (e.g., forward facing surfaces in low pressure regions and aft facing surfaces in higher pressure regions for low drag). Because the wing leading edge and trailing edge have multiple sweep angles and because of shocks generated on nacelles and diverters, a viscous code was required to accurately define the surface pressure distributions on the wing. A method of generating the surface geometry to take advantage of those surface pressures (as well as not violating any structural constraints) was developed and the resulting geometries were analyzed and compared to a baseline configuration. This paper will include discussions of the basic Natural Flow Wing design philosophy, the application of the philosophy to an HSCT vehicle, and preliminary wind-tunnel assessment of the NFW HSCT vehicle.
VIEW OF 77710A REACTOR WING, LOOKING NORTH, SHOWING DOOR TO ...
VIEW OF 777-10A REACTOR WING, LOOKING NORTH, SHOWING DOOR TO PROCESS DEVELOPMENT PILE ROOM AND LABORATORY WING ON RIGHT IN BACKGROUND - Physics Assembly Laboratory, Area A/M, Savannah River Site, Aiken, Aiken County, SC
Characteristics of the flow around tandem flapping wings
NASA Astrophysics Data System (ADS)
Muscutt, Luke; Ganapathisubramani, Bharathram; Weymouth, Gabriel; The University of Southampton Team
2014-11-01
Vortex recapture is a fundamental fluid mechanics phenomenon which is important to many fields. Any large scale vorticity contained within a freestream flow may affect the aerodynamic properties of a downstream body. In the case of tandem flapping wings, the front wing generates strong large scale vorticity which impinges on the hind wing. The characteristics of this interaction are greatly affected by the spacing, and the phase of flapping between the front and rear wings. The interaction of the vorticity of the rear wing with the shed vorticity of the front wing may be constructive or destructive, increasing thrust or efficiency of the hind wing when compared to a wing operating in isolation. Knowledge of the parameter space where the maximum increases in these are obtained is important for the development of tandem wing unmanned air and underwater vehicles, commercial aerospace and renewable energy applications. This question is addressed with a combined computational and experimental approach, and a discussion of these is presented.
Separation behavior of boundary layers on three-dimensional wings
NASA Technical Reports Server (NTRS)
Stock, H. W.
1981-01-01
An inverse boundary layer procedure for calculating separated, turbulent boundary layers at infinitely long, crabbing wing was developed. The procedure was developed for calculating three dimensional, incompressible turbulent boundary layers was expanded to adiabatic, compressible flows. Example calculations with transsonic wings were made including viscose effects. In this case an approximated calculation method described for areas of separated, turbulent boundary layers, permitting calculation of this displacement thickness. The laminar boundary layer development was calculated with inclined ellipsoids.
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
Preliminary development of a wing in ground effect vehicle
NASA Astrophysics Data System (ADS)
Abidin, Razali; Ahamat, Mohamad Asmidzam; Ahmad, Tarmizi; Saad, Mohd Rasdan; Hafizi, Ezzat
2018-02-01
Wing in ground vehicle is one of the mode of transportation that allows high speed movement over water by travelling few meters above the water level. Through this manouver strategy, a cushion of compressed air exists between the wing in ground vehicle wings and water. This significantly increase the lift force, thus reducing the necessity in having a long wing span. Our project deals with the development of wing in ground vehicle with the capability of transporting four people. The total weight of this wing in ground vehicle was estimated at 5.4 kN to enable the prediction on required wing area, minimum takeoff velocity, drag force and engine power requirement. The required takeoff velocity is decreases as the lift coefficient increases, and our current mathematical model shows the takeoff velocity at 50 m/s avoid the significant increase in lift coefficient for the wing area of 5 m2. At the velocity of 50 m/s, the drag force created by this wing in ground vehicle is well below 1 kN, which required a 100-120 kW of engine power if the propeller has the efficiency of 0.7. Assessment on the stresses and deflection of the hull structural indicate the capability of plywood to withstand the expected load. However, excessive deflection was expected in the rear section which requires a minor structural modification. In the near future, we expect that the wind tunnel tests of this wing in ground vehicle model would enable more definite prediction on the important parameters related to its performance.
Quasi-steady aerodynamic model of clap-and-fling flapping MAV and validation using free-flight data.
Armanini, S F; Caetano, J V; Croon, G C H E de; Visser, C C de; Mulder, M
2016-06-30
Flapping-wing aerodynamic models that are accurate, computationally efficient and physically meaningful, are challenging to obtain. Such models are essential to design flapping-wing micro air vehicles and to develop advanced controllers enhancing the autonomy of such vehicles. In this work, a phenomenological model is developed for the time-resolved aerodynamic forces on clap-and-fling ornithopters. The model is based on quasi-steady theory and accounts for inertial, circulatory, added mass and viscous forces. It extends existing quasi-steady approaches by: including a fling circulation factor to account for unsteady wing-wing interaction, considering real platform-specific wing kinematics and different flight regimes. The model parameters are estimated from wind tunnel measurements conducted on a real test platform. Comparison to wind tunnel data shows that the model predicts the lift forces on the test platform accurately, and accounts for wing-wing interaction effectively. Additionally, validation tests with real free-flight data show that lift forces can be predicted with considerable accuracy in different flight regimes. The complete parameter-varying model represents a wide range of flight conditions, is computationally simple, physically meaningful and requires few measurements. It is therefore potentially useful for both control design and preliminary conceptual studies for developing new platforms.
Blue Pattern Flower in Common Bean Expressed by Interaction of Prpi-2 with a New Gene tbp
USDA-ARS?s Scientific Manuscript database
The inheritance of blue pattern flower (BPF) expression was investigated in common bean (Phaseolus vulgaris L.). The BPF trait was derived from accession line G07262, and the flowers express blue banner petal and white wings with blue veins. Crosses between a BPF stock and three other parents - t ...
2013-03-01
Dominated Research 21 2.2.3 Trend 3 : Biomimicked Wing Design 23 2.2.4 Knowledge Gap: Beneficial Flexibility 25 2.2.5 Controversy: The Aeroelastic...Aerovironment Inc. ........................................................................................ 3 Figure 2: The “Hummingbird” developed...5 Figure 3 The number of UAS hours flown by the DOD from 1996
NASA Technical Reports Server (NTRS)
Srokowski, A. J.
1978-01-01
The problem of obtaining accurate estimates of suction requirements on swept laminar flow control wings was discussed. A fast accurate computer code developed to predict suction requirements by integrating disturbance amplification rates was described. Assumptions and approximations used in the present computer code are examined in light of flow conditions on the swept wing which may limit their validity.
NASA Technical Reports Server (NTRS)
Riley, D. R.
1985-01-01
A six-degree-of-freedom nonlinear simulation was developed for a two-place, single-engine, low-wing general aviation airplane for the stall and initial departure regions of flight. Two configurations, one with and one without an outboard wing-leading-edge modification, were modeled. The math models developed are presented simulation predictions and flight-test data for validation purposes and simulation results for the two configurations for various maneuvers and power settings are compared to show the beneficial influence of adding the wing-leading-edge modification.
Design development of graphite primary structures enables SSTO success
NASA Astrophysics Data System (ADS)
Biagiotti, V. A.; Yahiro, J. S.; Suh, Daniel E.; Hodges, Eric R.; Prior, Donald J.
1997-01-01
This paper describes the development of a graphite composite wing and a graphite composite intertank primary structure for application toward Single-Stage to Orbit space vehicles such as those under development in NASA's X-33/Reusable Launch Vehicle (RLV) Program. The trade study and designs are based on a Rockwell vertical take-off and horizontal landing (VTHL) wing-body RLV vehicle. Northrop Grumman's approach using a building block development technique is described. Composite Graphite/Bismaleimide (Gr/BMI) material characterization test results are presented. Unique intertank and wing composite subcomponent test article designs are described and test results to date are presented. Wing and intertank Full Scale Section Test Article (FSTA) objectives and designs are outlined. Trade studies, supporting building block testing, and FSTA demonstrations combine to develop graphite primary structure composite technology that enables developing X-33/RLV design programs to meet critical SSTO structural weight and operations performance criteria.
How the embryo makes a limb: determination, polarity and identity
Tickle, Cheryll
2015-01-01
The vertebrate limb with its complex anatomy develops from a small bud of undifferentiated mesoderm cells encased in ectoderm. The bud has its own intrinsic polarity and can develop autonomously into a limb without reference to the rest of the embryo. In this review, recent advances are integrated with classical embryology, carried out mainly in chick embryos, to present an overview of how the embryo makes a limb bud. We will focus on how mesoderm cells in precise locations in the embryo become determined to form a limb and express the key transcription factors Tbx4 (leg/hindlimb) or Tbx5 (wing/forelimb). These Tbx transcription factors have equivalent functions in the control of bud formation by initiating a signalling cascade involving Wnts and fibroblast growth factors (FGFs) and by regulating recruitment of mesenchymal cells from the coelomic epithelium into the bud. The mesoderm that will form limb buds and the polarity of the buds is determined with respect to both antero-posterior and dorso-ventral axes of the body. The position in which a bud develops along the antero-posterior axis of the body will also determine its identity – wing/forelimb or leg/hindlimb. Hox gene activity, under the influence of retinoic acid signalling, is directly linked with the initiation of Tbx5 gene expression in the region along the antero-posterior axis of the body that will form wings/forelimbs and determines antero-posterior polarity of the buds. In contrast, Tbx4 expression in the regions that will form legs/hindlimbs is regulated by the homeoprotein Pitx1 and there is no evidence that Hox genes determine antero-posterior polarity of the buds. Bone morphogenetic protein (BMP) signalling determines the region along the dorso-ventral axis of the body in which both wings/forelimbs and legs/hindlimbs develop and dorso-ventral polarity of the buds. The polarity of the buds leads to the establishment of signalling regions – the dorsal and ventral ectoderm, producing Wnts and BMPs, respectively, the apical ectodermal ridge producing fibroblast growth factors and the polarizing region, Sonic hedgehog (Shh). These signals are the same in both wings/forelimbs and legs/hindlimbs and control growth and pattern formation by providing the mesoderm cells of the limb bud as it develops with positional information. The precise anatomy of the limb depends on the mesoderm cells in the developing bud interpreting positional information according to their identity – determined by Pitx1 in hindlimbs – and genotype. The competence to form a limb extends along the entire antero-posterior axis of the trunk – with Hox gene activity inhibiting the formation of forelimbs in the interlimb region – and also along the dorso-ventral axis. PMID:26249743
How the embryo makes a limb: determination, polarity and identity.
Tickle, Cheryll
2015-10-01
The vertebrate limb with its complex anatomy develops from a small bud of undifferentiated mesoderm cells encased in ectoderm. The bud has its own intrinsic polarity and can develop autonomously into a limb without reference to the rest of the embryo. In this review, recent advances are integrated with classical embryology, carried out mainly in chick embryos, to present an overview of how the embryo makes a limb bud. We will focus on how mesoderm cells in precise locations in the embryo become determined to form a limb and express the key transcription factors Tbx4 (leg/hindlimb) or Tbx5 (wing/forelimb). These Tbx transcription factors have equivalent functions in the control of bud formation by initiating a signalling cascade involving Wnts and fibroblast growth factors (FGFs) and by regulating recruitment of mesenchymal cells from the coelomic epithelium into the bud. The mesoderm that will form limb buds and the polarity of the buds is determined with respect to both antero-posterior and dorso-ventral axes of the body. The position in which a bud develops along the antero-posterior axis of the body will also determine its identity - wing/forelimb or leg/hindlimb. Hox gene activity, under the influence of retinoic acid signalling, is directly linked with the initiation of Tbx5 gene expression in the region along the antero-posterior axis of the body that will form wings/forelimbs and determines antero-posterior polarity of the buds. In contrast, Tbx4 expression in the regions that will form legs/hindlimbs is regulated by the homeoprotein Pitx1 and there is no evidence that Hox genes determine antero-posterior polarity of the buds. Bone morphogenetic protein (BMP) signalling determines the region along the dorso-ventral axis of the body in which both wings/forelimbs and legs/hindlimbs develop and dorso-ventral polarity of the buds. The polarity of the buds leads to the establishment of signalling regions - the dorsal and ventral ectoderm, producing Wnts and BMPs, respectively, the apical ectodermal ridge producing fibroblast growth factors and the polarizing region, Sonic hedgehog (Shh). These signals are the same in both wings/forelimbs and legs/hindlimbs and control growth and pattern formation by providing the mesoderm cells of the limb bud as it develops with positional information. The precise anatomy of the limb depends on the mesoderm cells in the developing bud interpreting positional information according to their identity - determined by Pitx1 in hindlimbs - and genotype. The competence to form a limb extends along the entire antero-posterior axis of the trunk - with Hox gene activity inhibiting the formation of forelimbs in the interlimb region - and also along the dorso-ventral axis. © 2015 Anatomical Society.
Garvon, Jason M; Fedynich, Alan M; Peterson, Markus J; Pence, Danny B
2011-01-01
The influence of spatially distinct host subpopulations on helminth community structure and pattern was examined in a migratory avian host species. Forty helminth species represented by 24,082 individuals were collected from 184 blue-winged teal (Anas discors; BWT) from 2 primary migratory corridors in Florida (eastern migratory corridor; EMC) and Louisiana and Texas (western migratory corridor; WMC). Mean species richness was greater in BWT from the WMC (x̅±SE = 10.2 ± 0.3 species) than the EMC (8.6 ± 0.2). The helminth community from the WMC had higher abundances of 6 common/intermediate species. Corridor helminth communities were similar in species composition but less similar when incorporating abundances of those species. Overlapping distributions of phylogenetically related host species that share generalist helminth species across ecologically similar habitats seem to mitigate the isolating mechanisms that are necessary for the distinct coevolutionary pathways to develop between adjacent corridors.
Nacelle Aerodynamic and Inertial Loads (NAIL) project
NASA Technical Reports Server (NTRS)
1982-01-01
A flight test survey of pressures measured on wing, pylon, and nacelle surfaces and of the operating loads on Boeing 747/Pratt & Whitney JT9D-7A nacelles was made to provide information on airflow patterns surrounding the propulsion system installations and to clarify processes responsible for inservice deterioration of fuel economy. Airloads at takeoff rotation were found to be larger than at any other normal service condition because of the combined effects of high angle of attack and high engine airflow. Inertial loads were smaller than previous estimates indicated. A procedure is given for estimating inlet airloads at low speeds and high angles of attack for any underwing high bypass ratio turbofan installation approximately resembling the one tested. Flight procedure modifications are suggested that may result in better fuel economy retention in service. Pressures were recorded on the core cowls and pylons of both engine installations and on adjacent wing surfaces for use in development of computer codes for analysis of installed propulsion system aerodynamic drag interference effects.
Growth of mallards fed phosphamidon for 13-day periods during three different developmental stages
Haseltine, S.; Hensler, G.L.
1981-01-01
Mallard ducklings (Anas platyrhynchos) were exposed to a 13-day dietary treatment of O, 0.5, or 5.0 ppm phosphamidon at one of three successive age intervals (5-17 days, 18-30 days, or 31-43 days) during a 10-week growth period. Weekly measurements of body weight, wing length, primary feather length, and bill length revealed slower development of primary feathers in those birds treated from 5 to 17 days; treatment effects on body weight and wing length from 6 to 8 weeks of age were observed among those birds treated from 18 to 30 days of age. Some differences in growth patterns among birds treated with the same phosphamidon level, but at different growth stages, were attributed to the varying size of the group with which a duckling was housed at different times in the growth process. No brain cholinesterase depression was observed in any group either 24 h after phosphamidon treatment was terminated or at 10 weeks of age.
High-Reynolds Number Circulation Control Testing in the National Transonic Facility
NASA Technical Reports Server (NTRS)
Milholen, William E., II; Jones, Gregory S.; Chan, David T.; Goodliff, Scott L.
2012-01-01
A new capability to test active flow control concepts and propulsion simulations at high Reynolds numbers in the National Transonic Facility at the NASA Langley Research Center is being developed. The first active flow control experiment was completed using the new FAST-MAC semi-span model to study Reynolds number scaling effects for several circulation control concepts. Testing was conducted over a wide range of Mach numbers, up to chord Reynolds numbers of 30 million. The model was equipped with four onboard flow control valves allowing independent control of the circulation control plenums, which were directed over a 15% chord simple-hinged flap. Preliminary analysis of the uncorrected lift data showed that the circulation control increased the low-speed maximum lift coefficient by 33%. At transonic speeds, the circulation control was capable of positively altering the shockwave pattern on the upper wing surface and reducing flow separation. Furthermore, application of the technique to only the outboard portion of the wing demonstrated the feasibility of a pneumatic based roll control capability.
SMOC can act as both an antagonist and an expander of BMP signaling.
Thomas, J Terrig; Eric Dollins, D; Andrykovich, Kristin R; Chu, Tehyen; Stultz, Brian G; Hursh, Deborah A; Moos, Malcolm
2017-03-21
The matricellular protein SMOC (Secreted Modular Calcium binding protein) is conserved phylogenetically from vertebrates to arthropods. We showed previously that SMOC inhibits bone morphogenetic protein (BMP) signaling downstream of its receptor via activation of mitogen-activated protein kinase (MAPK) signaling. In contrast, the most prominent effect of the Drosophila orthologue, pentagone ( pent ), is expanding the range of BMP signaling during wing patterning. Using SMOC deletion constructs we found that SMOC-∆EC, lacking the extracellular calcium binding (EC) domain, inhibited BMP2 signaling, whereas SMOC-EC (EC domain only) enhanced BMP2 signaling. The SMOC-EC domain bound HSPGs with a similar affinity to BMP2 and could expand the range of BMP signaling in an in vitro assay by competition for HSPG-binding. Together with data from studies in vivo we propose a model to explain how these two activities contribute to the function of Pent in Drosophila wing development and SMOC in mammalian joint formation.
Biologically Inspired, Anisoptropic Flexible Wing for Optimal Flapping Flight
2013-01-31
Anisotropic Flexible Wing for Optimal Flapping Flight FA9550-07-1-0547 Sb. GRANT NUMBER Sc. PROGRAM ELEMENT NUMBER 6. AUTHOR(S) Sd. PROJECT NUMBER...anisotropic structural flexibility ; c) Conducted coordinated experimental and computational modeling to determine the roles of aerodynamic loading, wing inertia...and structural flexibility and elasticity; and d) Developed surrogate tools for flapping wing MA V design and optimization. Detailed research
Federal Register 2010, 2011, 2012, 2013, 2014
2010-11-19
... source can develop in the wing tank vapour space during fuel transfer from bag tank CWT [center wing tank... vapour space during fuel transfer from bag tank CWT [center wing tank], if the electrical power for... with a center wing tank (CWT); and Model F28 Mark 0100 airplanes, serial numbers 11244 through 11441...
Smith, N Adam; Clarke, Julia A
2014-02-01
Although studies of osteological morphology, gross myology, myological histology, neuroanatomy, and wing-scaling have all documented anatomical modifications associated with wing-propelled diving, the osteohistological study of this highly derived method of locomotion has been limited to penguins. Herein we present the first osteohistological study of the derived forelimbs and hind limbs of wing-propelled diving Pan-Alcidae (Aves, Charadriiformes). In addition to detailing differences between wing-propelled diving charadriiforms and nondiving charadriiforms, microstructural modifications to the humeri, ulnae and femora of extinct flightless pan-alcids are contrasted with those of volant alcids. Histological thin-sections of four species of pan-alcids (Alca torda, †Alca grandis, †Pinguinus impennis, †Mancalla cedrosensis) and one outgroup charadriiform (Stercorarius longicaudus) were compared. The forelimb bones of wing-propelled diving charadriiforms were found to have significantly thicker (∼22%) cortical bone walls. Additionally, as in penguins, the forelimbs of flightless pan-alcids are found to be osteosclerotic. However, unlike the pattern documented in penguins that display thickened cortices in both forelimbs and hind limbs, the forelimb and hind limb elements of pan-alcids display contrasting microstructural morphologies with thickened forelimb cortices and relatively thinner femoral cortices. Additionally, the identification of medullary bone in the sampled †Pinguinus impennis specimen suggests that further osteohistological investigation could provide an answer to longstanding questions regarding sexual dimorphism of Great Auks. Finally, these results suggest that it is possible to discern volant from flightless wing-propelled divers from fragmentary fossil remains. Copyright © 2013 Wiley Periodicals, Inc.
Static Structural Analysis of a Variable Span Morphing Wing for Unmanned Aerial Vehicle
NASA Astrophysics Data System (ADS)
Bashir, M.; Rajendran, P.
2018-05-01
While the primary reason to develop an adaptive wing is the aerodynamic benefits, the primary hindrance is the structural and vibrational considerations due to the unsteady nature of the airflow during the flight. Hence this study forms an important part of the morphable wing technology. In this paper, the design of a moderate aspect ratio variable span wing will be performed. The morphing wing is modeled structurally to observe the effect of spanwise load distribution on the wing structure. For the structural design and analysis of the unmanned aerial vehicle (UAV) under this study, commercial software Solidworks and Ansys/Static Structural/Modal are used. The static structural analyses of the wing are performed under different load conditions. The results of these analyses show that the designed structure is safe within the flight envelope. It is observed that the wing-root bending moment increases drastically due to an increase in the wingspan. Thus, the bending moment along the wingspan of the morphing wing is much larger than that of the conventional wing which results in an increase in the deflection of the free-end. The maximum stress for the un-extended wing configuration increases for the extended wing configuration.
The angiotensin-converting enzyme (ACE) gene family of Bombyx mori.
Yan, Hai-Yan; Mita, Kazuei; Zhao, Xia; Tanaka, Yoshikazu; Moriyama, Minoru; Wang, Huabin; Iwanaga, Masashi; Kawasaki, Hideki
2017-04-15
We previously reported regarding an ecdysone-inducible angiotensin-converting enzyme (ACE) gene. We found another four ACE genes in the Bombyx genome. The present study was undertaken to clarify the evolutionally changed function of the ACE of Bombyx mori. Core regions of deduced amino acid sequences of ACE genes were compared with those of other insect ACE genes. Five Bombyx genes have the conserved Zn 2+ -binding-site motif (HEXXH); however, BmAcer4 has only one and BmAcer3 has no catalytic ligand. BmAcer1 and BmAcer2 were expressed in several organs. BmAcer3 was expressed in testes, and BmAcer4 and BmAcer5 were expressed in compound eyes; however, the transcription levels of these three genes were very low. Quantitative RT-PCR and Western analysis were conducted to determine the tissue distribution and developmental expression of BmAcer1and BmAcer2. Transcripts of BmAcer1 and BmAcer2 were found in the reproductive organs during the larval and pupal stages. BmAcer1 was dominant in fat bodies during the feeding stage and showed high expression in the epidermis, wing discs, and pupal wing tissues after the wandering stage. Its expression patterns in epidermis, wing discs, and wing tissues resembled the hemolymph ecdysteroid titer in the larval and pupal stages. Acer1 was observed in the hemolymph at all stages, appearing to be the source of it are fat bodies, wings, and epidermis, and functioning after being secreted into the hemolymph. BmAcer2 was abundant in the midgut during the feeding stage and after the wandering stage and in silk glands after the pupal stage. We conclude that the evolution of BmAcer occurred through duplication, and, thereafter, functional diversification developed. Copyright © 2017 Elsevier B.V. All rights reserved.
Impact of extended defects on optical properties of (1-101)GaN grown on patterned Si
NASA Astrophysics Data System (ADS)
Okur, S.; Izyumskaya, N.; Zhang, F.; Avrutin, V.; Metzner, S.; Karbaum, C.; Bertram, F.; Christen, J.; Morkoç, H.; Özgür, Ü.
2014-03-01
The optical quality of semipolar (1 101)GaN layers was explored by time- and polarization-resolved photoluminescence spectroscopy. High intensity bandedge emission was observed in +c-wing regions of the stripes as a result of better structural quality, while -c-wing regions were found to be of poorer optical quality due to basal plane and prismatic stacking faults (BSFs and PSFs) in addition to a high density of TDs. The high optical quality region formed on the +cwings was evidenced also from the much slower biexponential PL decays (0.22 ns and 1.70 ns) and an order of magnitude smaller amplitude ratio of the fast decay (nonradiative origin) to the slow decay component (radiative origin) compared to the -c-wing regions. In regard to defect-related emission, decay times for the BSF and PSF emission lines at 25 K (~ 0.80 ns and ~ 3.5 ns, respectively) were independent of the excitation density within the range employed (5 - 420 W/cm2), and much longer than that for the donor bound excitons (0.13 ns at 5 W/cm2 and 0.22 ns at 420 W/cm2). It was also found that the emission from BSFs had lower polarization degree (0.22) than that from donor bound excitons (0.35). The diminution of the polarization degree when photogenerated carriers recombine within the BSFs is another indication of the negative effects of stacking faults on the optical quality of the semipolar (1101)GaN. In addition, spatial distribution of defects in semipolar (1101)-oriented InGaN active region layers grown on stripe patterned Si substrates was investigated using near-field scanning optical microscopy. The optical quality of -c- wing regions was found to be worse compared to +c-wing regions due to the presence of higher density of stacking faults and threading dislocations. The emission from the +c-wings was very bright and relatively uniform across the sample, which is indicative of a homogeneous In distribution.
Aerodynamic and structural studies of joined-wing aircraft
NASA Technical Reports Server (NTRS)
Kroo, Ilan; Smith, Stephen; Gallman, John
1991-01-01
A method for rapidly evaluating the structural and aerodynamic characteristics of joined-wing aircraft was developed and used to study the fundamental advantages attributed to this concept. The technique involves a rapid turnaround aerodynamic analysis method for computing minimum trimmed drag combined with a simple structural optimization. A variety of joined-wing designs are compared on the basis of trimmed drag, structural weight, and, finally, trimmed drag with fixed structural weight. The range of joined-wing design parameters resulting in best cruise performance is identified. Structural weight savings and net drag reductions are predicted for certain joined-wing configurations compared with conventional cantilever-wing configurations.
ACEE composite structures technology
NASA Technical Reports Server (NTRS)
James, A. M.
1984-01-01
Topics addressed include: strength and hygrothermal response of L-1011 fin components; wing fuel containment and damage tolerance development; impact dynamics; acoustic transmission; fuselage structure; composite transport wing technology development; spar/assembly concepts.
Swept-Wing Ice Accretion Characterization and Aerodynamics
NASA Technical Reports Server (NTRS)
Broeren, Andy P.; Potapczuk, Mark G.; Riley, James T.; Villedieu, Philippe; Moens, Frederic; Bragg, Michael B.
2013-01-01
NASA, FAA, ONERA, the University of Illinois and Boeing have embarked on a significant, collaborative research effort to address the technical challenges associated with icing on large-scale, three-dimensional swept wings. The overall goal is to improve the fidelity of experimental and computational simulation methods for swept-wing ice accretion formation and resulting aerodynamic effect. A seven-phase research effort has been designed that incorporates ice-accretion and aerodynamic experiments and computational simulations. As the baseline, full-scale, swept-wing-reference geometry, this research will utilize the 65% scale Common Research Model configuration. Ice-accretion testing will be conducted in the NASA Icing Research Tunnel for three hybrid swept-wing models representing the 20%, 64% and 83% semispan stations of the baseline-reference wing. Three-dimensional measurement techniques are being developed and validated to document the experimental ice-accretion geometries. Artificial ice shapes of varying geometric fidelity will be developed for aerodynamic testing over a large Reynolds number range in the ONERA F1 pressurized wind tunnel and in a smaller-scale atmospheric wind tunnel. Concurrent research will be conducted to explore and further develop the use of computational simulation tools for ice accretion and aerodynamics on swept wings. The combined results of this research effort will result in an improved understanding of the ice formation and aerodynamic effects on swept wings. The purpose of this paper is to describe this research effort in more detail and report on the current results and status to date. 1
Swept-Wing Ice Accretion Characterization and Aerodynamics
NASA Technical Reports Server (NTRS)
Broeren, Andy P.; Potapczuk, Mark G.; Riley, James T.; Villedieu, Philippe; Moens, Frederic; Bragg, Michael B.
2013-01-01
NASA, FAA, ONERA, the University of Illinois and Boeing have embarked on a significant, collaborative research effort to address the technical challenges associated with icing on large-scale, three-dimensional swept wings. The overall goal is to improve the fidelity of experimental and computational simulation methods for swept-wing ice accretion formation and resulting aerodynamic effect. A seven-phase research effort has been designed that incorporates ice-accretion and aerodynamic experiments and computational simulations. As the baseline, full-scale, swept-wing-reference geometry, this research will utilize the 65 percent scale Common Research Model configuration. Ice-accretion testing will be conducted in the NASA Icing Research Tunnel for three hybrid swept-wing models representing the 20, 64 and 83 percent semispan stations of the baseline-reference wing. Threedimensional measurement techniques are being developed and validated to document the experimental ice-accretion geometries. Artificial ice shapes of varying geometric fidelity will be developed for aerodynamic testing over a large Reynolds number range in the ONERA F1 pressurized wind tunnel and in a smaller-scale atmospheric wind tunnel. Concurrent research will be conducted to explore and further develop the use of computational simulation tools for ice accretion and aerodynamics on swept wings. The combined results of this research effort will result in an improved understanding of the ice formation and aerodynamic effects on swept wings. The purpose of this paper is to describe this research effort in more detail and report on the current results and status to date.
Development of Micro Air Vehicle Technology With In-Flight Adaptive-Wing Structure
NASA Technical Reports Server (NTRS)
Waszak, Martin R. (Technical Monitor); Shkarayev, Sergey; Null, William; Wagner, Matthew
2004-01-01
This is a final report on the research studies, "Development of Micro Air Vehicle Technology with In-Flight Adaptrive-Wing Structure". This project involved the development of variable-camber technology to achieve efficient design of micro air vehicles. Specifically, it focused on the following topics: 1) Low Reynolds number wind tunnel testing of cambered-plate wings. 2) Theoretical performance analysis of micro air vehicles. 3) Design of a variable-camber MAV actuated by micro servos. 4) Test flights of a variable-camber MAV.
Direct Harmonic Linear Navier-Stokes Methods for Efficient Simulation of Wave Packets
NASA Technical Reports Server (NTRS)
Streett, C. L.
1998-01-01
Wave packets produced by localized disturbances play an important role in transition in three-dimensional boundary layers, such as that on a swept wing. Starting with the receptivity process, we show the effects of wave-space energy distribution on the development of packets and other three-dimensional disturbance patterns. Nonlinearity in the receptivity process is specifically addressed, including demonstration of an effect which can enhance receptivity of traveling crossflow disturbances. An efficient spatial numerical simulation method is allowing most of the simulations presented to be carried out on a workstation.
Oil-flow study of a Space Shuttle orbiter tip-fin controller
NASA Technical Reports Server (NTRS)
Helms, V. T., III
1983-01-01
Possible use of tip-fin controllers instead of a vertical tail on advanced winged entry vehicles was examined. Elimination of the vertical tail and using tip-fins offers the advantages of positive yaw control at high angles of attack and a potential weight savings. Oil-flow technique was used to obtain surface flow patterns on a tip-fin installed on a 0.01-scale Space Shuttle orbiter model for the purpose of assessing the extent of flow interference effects on the wing and tip-fin which might lead to serious heating problems. Tests were conducted in air at Mach 10 for a free-stream Reynolds numbers of .000113 at 20, 30, and 40 degree angle of attack and sideslip angles of 0 and 2 degree. Elevon deflections of -10, 0, and 10 degree and tip-fin control-surface deflections of 0, 20, and 40 degree were employed. Test results were also used to aid in the interpretation of heating data obtained on a Shuttle orbiter tip-fin on another model in a different facility. A limited comparison of oil-flow patterns and heat-transfer data is included. It was determined that elevon deflection angles from -10 to 10 degree and sideslip angles up to 2 degree have very little effect on tip-fin surface flow patterns. Also, there is a minimum of interference between the tip-fin and the wing. The most significant flow interactions occur on the tip-fin onboard surface as a result of its control-surface deflections.
Cha, Dong H; Hesler, Stephen P; Linn, Charles E; Zhang, Aijun; Teal, Peter E A; Knight, Alan L; Roelofs, Wendell L; Loeb, Gregory M
2013-02-01
Oil-coated clear panel traps baited with a host plant-based kairomone lure have successfully been used for monitoring female grape berry moth, Paralobesia viteana (Clemens) (Lepidoptera: Tortricidae), but low capture rates as well as difficulty in servicing these traps makes them unsuitable for commercial use. We compared the performance of different trap designs in a flight tunnel and in a vineyard by using a 7-component synthetic kairomone blend, with a focus on trap visual cues. In flight tunnel experiments, a clear delta trap performed better than other traps. When we tested clear delta, green delta, or clear wing traps baited with a cut grape shoot, >50% of female grape berry moths made complete upwind flights. However, the clear delta trap was the only design that resulted in female moths entering the trap. Similar results were observed when females were tested with different traps (clear delta, green delta, white delta, clear wing, or green wing traps) baited with the kairomone lure. Adding a visual pattern that mimicked grape shoots to the outside surface of the clear delta trap resulted in 66% of the females that made upwind flights entering the trap. However, the positive effect of adding a visual pattern to the trap was not observed in a vineyard setting, where clear delta traps with or without a visual pattern caught similar numbers of females. Still, the number of male and female grape berry moths captured in clear delta traps with or without a visual pattern was not significantly different from the number of male and female grape berry moths captured in panel traps, suggesting that the use of these delta traps could be a less cumbersome alternative to oil-coated panel traps for monitoring female grape berry moth.
Abbasi, R; Marcus, J M
2015-11-01
Ocelli are serially repeated colour patterns on the wings of many butterflies. Eyespots are elaborate ocelli that function in predator avoidance and deterrence as well as in mate choice. A phylogenetic approach was used to study ocelli and eyespot evolution in Vanessa butterflies, a genus exhibiting diverse phenotypes among these serial homologs. Forty-four morphological characters based on eyespot number, arrangement, shape and the number of elements in each eyespot were defined and scored. Ocelli from eight wing cells on the dorsal and ventral surfaces of the forewing and hindwing were evaluated. The evolution of these characters was traced over a phylogeny of Vanessa based on 7750 DNA base pairs from 10 genes. Our reconstruction predicts that the ancestral Vanessa had 5 serially arranged ocelli on all four wing surfaces. The ancestral state on the dorsal forewing and ventral hindwing was ocelli arranged in two heterogeneous groups. On the dorsal hindwing, the ancestral state was either homogenous or ocelli arranged in two heterogeneous groups. On the ventral forewing, we determined that the ancestral state was organized into three heterogeneous groups. In Vanessa, almost all ocelli are individuated and capable of independent evolution relative to other colour patterns except for the ocelli in cells -1 and 0 on the dorsal and ventral forewings, which appear to be constrained to evolve in parallel. The genus Vanessa is a good model system for the study of serial homology and the interaction of selective forces with developmental architecture to produce diversity in butterfly colour patterns. © 2015 European Society For Evolutionary Biology. Journal of Evolutionary Biology © 2015 European Society For Evolutionary Biology.
NASA Astrophysics Data System (ADS)
Diasinos, S.; Gatto, A.
2008-09-01
This paper details a quantitative 3D investigation using LDA into the interaction aerodynamics on a sub-scale open wheel race car inverted front wing and wheel. Of primary importance to this study was the influence of changing wing angle of attack and span on the resulting near-field and far-field flow characteristics. Results obtained showed that both variables do have a significant influence on the resultant flow-field, particularly on wing vortex and wheel wake development and propagation.
Function of Lipid Storage Droplet 1 (Lsd1) in Wing Development of Drosophila melanogaster.
Men, Tran Thanh; Binh, Tran Duy; Yamaguchi, Masamitsu; Huy, Nguyen Tien; Kamei, Kaeko
2016-04-29
Perilipins are evolutionarily conserved from Drosophila to humans, the lipid storage droplet 1 (Lsd1) is a Drosophila homolog of human perilipin 1. The function of Lsd1 as a regulator of lipolysis in Drosophila has been demonstrated, as the Lsd1 mutant causes an increase of lipid droplet size. However, the functions of this gene during development are still under investigation. In order to determine the function of Lsd1 during development, Lsd1 was knocked down in Drosophila using the GAL4-UAS system. Selective knockdown of Lsd1 in the dorsal wing disc caused an atrophied wing phenotype. The generation of reactive oxygen species in the wing pouch compartment of the Lsd1-knockdown flies was significantly higher than in the control. Immunostaining with caspase-3 antibody revealed a greater number of apoptotic cells in Lsd1-knockdown wing discs than in the control. Cell death by autophagy was also induced in the knockdown flies. Moreover, cells deprived of Lsd1 showed mitochondrial expansion and decreased ATP levels. These results strongly suggest that knockdown of Lsd1 induces mitochondrial stress and the production of reactive oxygen species that result in cell death, via apoptosis and the autophagy pathway. These results highlight the roles of Drosophila Lsd1 during wing development.
How differential deflection of the inboard and outboard leading-edge flaps affected the handling qua
NASA Technical Reports Server (NTRS)
2002-01-01
How differential deflection of the inboard and outboard leading-edge flaps affected the handling qualities of this modified F/A-18A was evaluated during the first check flight in the Active Aeroelastic Wing program at NASA's Dryden Flight Research Center. The Active Aeroelastic Wing program at NASA's Dryden Flight Research Center seeks to determine the advantages of twisting flexible wings for primary maneuvering roll control at transonic and supersonic speeds, with traditional control surfaces such as ailerons and leading-edge flaps used to aerodynamically induce the twist. From flight test and simulation data, the program intends to develop structural modeling techniques and tools to help design lighter, more flexible high aspect-ratio wings for future high-performance aircraft, which could translate to more economical operation or greater payload capability. AAW flight tests began in November, 2002 with checkout and parameter-identification flights. Based on data obtained during the first flight series, new flight control software will be developed and a second series of research flights will then evaluate the AAW concept in a real-world environment. The program uses wings that were modified to the flexibility of the original pre-production F-18 wing. Other modifications include a new actuator to operate the outboard leading edge flap over a greater range and rate, and a research flight control system to host the aeroelastic wing control laws. The Active Aeroelastic Wing Program is jointly funded and managed by the Air Force Research Laboratory and NASA Dryden Flight Research Center, with Boeing's Phantom Works as prime contractor for wing modifications and flight control software development. The F/A-18A aircraft was provided by the Naval Aviation Systems Test Team and modified for its research role by NASA Dryden technicians.
NASA Astrophysics Data System (ADS)
Tsushima, Natsuki
The purpose of this dissertation is to develop an analytical framework to analyze highly flexible multifunctional wings with integral active and passive control and energy harvesting using piezoelectric transduction. Such multifunctional wings can be designed to enhance aircraft flight performance, especially to support long-endurance flights and to be adaptive to various flight conditions. This work also demonstrates the feasibility of the concept of piezoelectric multifunctional wings for the concurrent active control and energy harvesting to improve the aeroelastic performance of high-altitude long-endurance unmanned air vehicles. Functions of flutter suppression, gust alleviation, energy generation, and energy storage are realized for the performance improvement. The multifunctional wings utilize active and passive piezoelectric effects for the efficient adaptive control and energy harvesting. An energy storage with thin-film lithium-ion battery cells is designed for harvested energy accumulation. Piezoelectric effects are included in a strain-based geometrically nonlinear beam formulation for the numerical studies. The resulting structural dynamic equations are coupled with a finite-state unsteady aerodynamic formulation, allowing for piezoelectric energy harvesting and active actuation with the nonlinear aeroelastic system. This development helps to provide an integral electro-aeroelastic solution of concurrent active piezoelectric control and energy harvesting for wing vibrations, with the consideration of the geometrical nonlinear effects of slender multifunctional wings. A multifunctional structure for active actuation is designed by introducing anisotropic piezoelectric laminates. Linear quadratic regulator and linear quadratic Gaussian controllers are implemented for the active control of wing vibrations including post-flutter limit-cycle oscillations and gust perturbation. An adaptive control algorithm for gust perturbation is then developed. In this research, the active piezoelectric actuation is applied as the primary approach for flutter suppression, with energy harvesting, as a secondary passive approach, concurrently working to provide an additional damping effect on the wing vibration. The multifunctional wing also generates extra energy from residual wing vibration. This research presents a comprehensive approach for an effective flutter suppression and gust alleviation of highly flexible piezoelectric wings, while allowing to harvest the residual vibration energy. Numerical results with the multifunctional wing concept show the potential to improve the aircraft performance from both aeroelastic stability and energy consumption aspects.
A new genus of long-legged flies displaying remarkable wing directional asymmetry
Justin B. Runyon; Richard L. Hurley
2004-01-01
A previously unknown group of flies is described whose males exhibit directional asymmetry, in that the left wing is larger than, and of a different shape from, the right wing. To our knowledge, wing asymmetry of this degree has not previously been reported in an animal capable of flight. Such consistent asymmetry must result from a leftÃÂright axis during development...
Semi-automated quantitative Drosophila wings measurements.
Loh, Sheng Yang Michael; Ogawa, Yoshitaka; Kawana, Sara; Tamura, Koichiro; Lee, Hwee Kuan
2017-06-28
Drosophila melanogaster is an important organism used in many fields of biological research such as genetics and developmental biology. Drosophila wings have been widely used to study the genetics of development, morphometrics and evolution. Therefore there is much interest in quantifying wing structures of Drosophila. Advancement in technology has increased the ease in which images of Drosophila can be acquired. However such studies have been limited by the slow and tedious process of acquiring phenotypic data. We have developed a system that automatically detects and measures key points and vein segments on a Drosophila wing. Key points are detected by performing image transformations and template matching on Drosophila wing images while vein segments are detected using an Active Contour algorithm. The accuracy of our key point detection was compared against key point annotations of users. We also performed key point detection using different training data sets of Drosophila wing images. We compared our software with an existing automated image analysis system for Drosophila wings and showed that our system performs better than the state of the art. Vein segments were manually measured and compared against the measurements obtained from our system. Our system was able to detect specific key points and vein segments from Drosophila wing images with high accuracy.
Scaling law and enhancement of lift generation of an insect-size hovering flexible wing
Kang, Chang-kwon; Shyy, Wei
2013-01-01
We report a comprehensive scaling law and novel lift generation mechanisms relevant to the aerodynamic functions of structural flexibility in insect flight. Using a Navier–Stokes equation solver, fully coupled to a structural dynamics solver, we consider the hovering motion of a wing of insect size, in which the dynamics of fluid–structure interaction leads to passive wing rotation. Lift generated on the flexible wing scales with the relative shape deformation parameter, whereas the optimal lift is obtained when the wing deformation synchronizes with the imposed translation, consistent with previously reported observations for fruit flies and honeybees. Systematic comparisons with rigid wings illustrate that the nonlinear response in wing motion results in a greater peak angle compared with a simple harmonic motion, yielding higher lift. Moreover, the compliant wing streamlines its shape via camber deformation to mitigate the nonlinear lift-degrading wing–wake interaction to further enhance lift. These bioinspired aeroelastic mechanisms can be used in the development of flapping wing micro-robots. PMID:23760300
NASA Technical Reports Server (NTRS)
Johnston, J. F.
1979-01-01
Active wing load alleviation to extend the wing span by 5.8 percent, giving a 3 percent reduction in cruise drag is covered. The active wing load alleviation used symmetric motions of the outboard ailerons for maneuver load control (MLC) and elastic mode suppression (EMS), and stabilizer motions for gust load alleviation (GLA). Slow maneuvers verified the MLC, and open and closed-loop flight frequency response tests verified the aircraft dynamic response to symmetric aileron and stabilizer drives as well as the active system performance. Flight tests in turbulence verified the effectiveness of the active controls in reducing gust-induced wing loads. It is concluded that active wing load alleviation/extended span is proven in the L-1011 and is ready for application to airline service; it is a very practical way to obtain the increased efficiency of a higher aspect ratio wing with minimum structural impact.
A Conceptual Wing Flutter Analysis Tool for Systems Analysis and Parametric Design Study
NASA Technical Reports Server (NTRS)
Mukhopadhyay, Vivek
2003-01-01
An interactive computer program was developed for wing flutter analysis in the conceptual design stage. The objective was to estimate flutt er instability boundaries of a typical wing, when detailed structural and aerodynamic data are not available. Effects of change in key flu tter parameters can also be estimated in order to guide the conceptual design. This userfriendly software was developed using MathCad and M atlab codes. The analysis method was based on non-dimensional paramet ric plots of two primary flutter parameters, namely Regier number and Flutter number, with normalization factors based on wing torsion stiffness, sweep, mass ratio, taper ratio, aspect ratio, center of gravit y location and pitch-inertia radius of gyration. These parametric plo ts were compiled in a Chance-Vought Corporation report from database of past experiments and wind tunnel test results. An example was prese nted for conceptual flutter analysis of outer-wing of a Blended-Wing- Body aircraft.
Transonic empirical configuration design process
NASA Technical Reports Server (NTRS)
Whitcomb, R. T.
1983-01-01
This lecture describes some of the experimental research pertaining to transonic configuration development conducted by the Transonic Aerodynamics Branch of the NASA Langley Research Center. Discussions are presented of the following: use of florescent oil films for the study of surface boundary layer flows; the severe effect of wind tunnel wall interference on the measured configuration drag rise near the speed of sound as determined by a comparison between wind tunnel and free air results; the development of a near sonic transport configuration incorporating a supercritical wing and an indented fuselage, designed on the basis of the area rule with a modification to account for the presence of local supersonic flow above the wing; a device for improving the transonic pitch up of swept wings with very little added drag at the cruise condition; a means for reducing the large transonic aerodynamic interference between the wing, fuselage, nacelle and pylon for a for a fuselage mounted nacelle having the inlet above the wing; and methods for reducing the transonic interference between flows over a winglet and the wing.
Rotor/Wing Interactions in Hover
NASA Technical Reports Server (NTRS)
Young, Larry A.; Derby, Michael R.
2002-01-01
Hover predictions of tiltrotor aircraft are hampered by the lack of accurate and computationally efficient models for rotor/wing interactional aerodynamics. This paper summarizes the development of an approximate, potential flow solution for the rotor-on-rotor and wing-on-rotor interactions. This analysis is based on actuator disk and vortex theory and the method of images. The analysis is applicable for out-of-ground-effect predictions. The analysis is particularly suited for aircraft preliminary design studies. Flow field predictions from this simple analytical model are validated against experimental data from previous studies. The paper concludes with an analytical assessment of the influence of rotor-on-rotor and wing-on-rotor interactions. This assessment examines the effect of rotor-to-wing offset distance, wing sweep, wing span, and flaperon incidence angle on tiltrotor inflow and performance.
Beyond magic traits: Multimodal mating cues in Heliconius butterflies.
Mérot, Claire; Frérot, Brigitte; Leppik, Ene; Joron, Mathieu
2015-11-01
Species coexistence involves the evolution of reproductive barriers opposing gene flow. Heliconius butterflies display colorful patterns affecting mate choice and survival through warning signaling and mimicry. These patterns are called "magic traits" for speciation because divergent natural selection may promote mimicry shifts in pattern whose role as mating cue facilitates reproductive isolation. By contrast, between comimetic species, natural selection promotes pattern convergence. We addressed whether visual convergence interferes with reproductive isolation by testing for sexual isolation between two closely related species with similar patterns, H. timareta thelxinoe and H. melpomene amaryllis. Experiments with models confirmed visual attraction based on wing phenotype, leading to indiscriminate approach. Nevertheless, mate choice experiments showed assortative mating. Monitoring male behavior toward live females revealed asymmetry in male preference, H. melpomene males courting both species equally while H. timareta males strongly preferred conspecifics. Experiments with hybrid males suggested an important genetic component for such asymmetry. Behavioral observations support a key role for short-distance cues in determining male choice in H. timareta. Scents extracts from wings and genitalia revealed interspecific divergence in chemical signatures, and hybrid female scent composition was significantly associated with courtship intensity by H. timareta males, providing candidate chemical mating cues involved in sexual isolation. © 2015 The Author(s). Evolution © 2015 The Society for the Study of Evolution.
NASA Astrophysics Data System (ADS)
Reed, John L., Jr.; Hemmelgarn, Christopher D.; Pelley, Bryan M.; Havens, Ernie
2005-05-01
Cornerstone Research Group, Inc. (CRG) is developing a unique adaptive wing structure intended to enhance the capability of loitering Unmanned Air Vehicles (UAVs). In order to tailor the wing design to a specific application, CRG has developed a wing structure capable of morphing in chord and increasing planform area by 80 percent. With these features, aircraft will be capable of optimizing their flight efficiency throughout the entire mission profile. The key benefit from this morphing design is increased maneuverability, resulting in improved effectiveness over the current design. During the development process CRG has overcome several challenges in the design of such a structure while incorporating advanced materials capable of maintaining aerodynamic shape and transferring aerodynamic loads while enabling crucial changes in planform shape. To overcome some of these challenges, CRG is working on integration of their shape memory polymer materials into the wing skin to enable seamless morphing. This paper will address the challenges associated with the development of a morphing aerospace structure capable of such large shape change, the materials necessary for enabling morphing capabilities, and the current status of the morphing program within CRG.
A Study of the Zero-Lift Drag-Rise Characteristics of Wing-Body Combinations Near the Speed of Sound
NASA Technical Reports Server (NTRS)
Whitcomb, Richard T
1956-01-01
Comparisons have been made of the shock phenomena and drag-rise increments for representative wing and central-body combinations with those for bodies of revolution having the same axial developments of cross-sectional areas normal to the airstream. On the basis of these comparisons, it is concluded that near the speed of sound the zero-lift drag rise of a low-aspect-ratio thin-wing and body combination is primarily dependent on the axial development of the cross-sectional areas normal to the airstream. It follows that the drag rise for any such configuration is approximately the same as that for any other with the same development of cross-sectional areas. Investigations have also been made of representative wing-body combinations with the body so indented that the axial developments of cross-sectional areas for the combinations were the same as that for the original body alone. Such indentations greatly reduced or eliminated the zero-lift drag-rise increments associated with the wings near the speed of sound.
A study on the utilization of advanced composites in commercial aircraft wing structure
NASA Technical Reports Server (NTRS)
Watts, D. J.
1978-01-01
A study was conducted to define the technology and data needed to support the introduction of advanced composite materials in the wing structure of future production aircraft. The study accomplished the following: (1) definition of acceptance factors, (2) identification of technology issues, (3) evaluation of six candidate wing structures, (4) evaluation of five program options, (5) definition of a composite wing technology development plan, (6) identification of full-scale tests, (7) estimation of program costs for the total development plan, (8) forecast of future utilization of composites in commercial transport aircraft and (9) identification of critical technologies for timely program planning.
The plane problem of the flapping wing
NASA Technical Reports Server (NTRS)
Birnbaum, Walter
1954-01-01
In connection with an earlier report on the lifting vortex sheet which forms the basis of the following investigations this will show how the methods developed there are also suitable for dealing with the air forces for a wing with a circulation variable with time. The theory of a propulsive wing flapping up and down periodically in the manner of a bird's wing is developed. This study shows how the lift and its moment result as a function of the flapping motion, what thrust is attainable, and how high is the degree of efficiency of this flapping propulsion unit if the air friction is disregarded.
Flying wings / flying fuselages
NASA Technical Reports Server (NTRS)
Wood, Richard M.; Bauer, Steven X. S.
2001-01-01
The present paper has documented the historical relationships between various classes of all lifting vehicles, which includes the flying wing, all wing, tailless, lifting body, and lifting fuselage. The diversity in vehicle focus was to ensure that all vehicle types that map have contributed to or been influenced by the development of the classical flying wing concept was investigated. The paper has provided context and perspective for present and future aircraft design studies that may employ the all lifting vehicle concept. The paper also demonstrated the benefit of developing an understanding of the past in order to obtain the required knowledge to create future concepts with significantly improved aerodynamic performance.
Design, testing, and damage tolerance study of bonded stiffened composite wing cover panels
NASA Technical Reports Server (NTRS)
Madan, Ram C.; Sutton, Jason O.
1988-01-01
Results are presented from the application of damage tolerance criteria for composite panels to multistringer composite wing cover panels developed under NASA's Composite Transport Wing Technology Development contract. This conceptual wing design integrated aeroelastic stiffness constraints with an enhanced damage tolerance material system, in order to yield optimized producibility and structural performance. Damage tolerance was demonstrated in a test program using full-sized cover panel subcomponents; panel skins were impacted at midbay between stiffeners, directly over a stiffener, and over the stiffener flange edge. None of the impacts produced visible damage. NASTRAN analyses were performed to simulate NDI-detected invisible damage.
NASA Technical Reports Server (NTRS)
1976-01-01
An experimental and analytical aerodynamic program to develop predesign guides for irregular planform wings is reported. The benefits are linearization of subsonic lift curve slope to high angles of attack and avoidance of subsonic pitch instabilities at high lift by proper tailoring of the planform fillet wing combination while providing the desired hypersonic trim angle and stability. The two prime areas of concern are to optimize shuttle orbiter landing and entry characteristics. Basic longitudinal aerodynamic characteristics at high supersonic speeds are developed.
Stability and Control Properties of an Aeroelastic Fixed Wing Micro Aerial Vehicle
NASA Technical Reports Server (NTRS)
Waszak, Martin R.; Jenkins, Luther N.; Ifju, Peter
2001-01-01
Micro aerial vehicles have been the subject of considerable interest and development over the last several years. The majority of current vehicle concepts rely on rigid fixed wings or rotors. An alternate design based on an aeroelastic membrane wing concept has also been developed that has exhibited desired characteristics in flight test demonstrations and competition. This paper presents results from a wind tunnel investigation that sought to quantify stability and control properties for a family of vehicles using the aeroelastic design. The results indicate that the membrane wing does exhibit potential benefits that could be exploited to enhance the design of future flight vehicles.
Utilizing adaptive wing technology in the control of a micro air vehicle
NASA Astrophysics Data System (ADS)
Null, William R.; Wagner, Matthew G.; Shkarayev, Sergey V.; Jouse, Wayne C.; Brock, Keith M.
2002-07-01
Evolution of the design of micro air vehicles (MAVs) towards miniaturization has been severely constrained by the size and mass of the electronic components needed to control the vehicles. Recent research, experimentation, and development in the area of smart materials have led to the possibility of embedding control actuators, fabricated from smart materials, in the wing of the vehicle, reducing both the size and mass of these components. Further advantages can be realized by developing adaptive wing structures. Small size and mass, and low airspeeds, can lead to considerable buffeting during flight, and may result in a loss of flight control. In order to counter these effects, we are developing a thin, variable-cambered airfoil design with actuators embedded within the wing. In addition to reducing the mass and size of the vehicle or, conversely, increasing its available payload, an important benefit from the adaptive wing concept is the possibility of in-flight modification of the flight envelope. Reduced airspeeds, which are crucial during loiter, can be realized by an in-flight increase in wing camber. Conversely, decreases in camber provide for an airframe best suited for rapid ingress/egress and extension of the mission range.
Teaching Basic Algebra Courses at the College Level
ERIC Educational Resources Information Center
Mallenby, Michel L.; Mallenby, Douglas W.
2004-01-01
Three dysfunctional behaviors of basic algebra students are described: Silence as Camouflage, Wing and a Prayer, and Ignorance is OK. These behavior patterns are explained, and beneficial teaching methods that address the weaknesses are presented.
Truong, Vi Khanh; Geeganagamage, Nipuni Mahanamanam; Baulin, Vladimir A; Vongsvivut, Jitraporn; Tobin, Mark J; Luque, Pere; Crawford, Russell J; Ivanova, Elena P
2017-06-01
Nanostructured insect wing surfaces have been reported to possess the ability to resist bacterial colonization through the mechanical rupture of bacterial cells coming into contact with the surface. In this work, the susceptibility of physiologically young, mature and old Staphylococcus aureus CIP 65.8 and Pseudomonas aeruginosa ATCC 9721 bacterial cells, to the action of the bactericidal nano-pattern of damselfly Calopteryx haemorrhoidalis wing surfaces, was investigated. The results were obtained using several surface characterization techniques including optical profilometry, scanning electron microscopy, synchrotron-sourced Fourier transform infrared microspectroscopy, water contact angle measurements and antibacterial assays. The data indicated that the attachment propensity of physiologically young S. aureus CIP 65.8 T and mature P. aeruginosa ATCC 9721 bacterial cells was greater than that of the cells at other stages of growth. Both the S. aureus CIP 65.8 T and P. aeruginosa ATCC 9721 cells, grown at the early (1 h) and late stationary phase (24 h), were found to be most susceptible to the action of the wings, with up to 89.7 and 61.3% as well as 97.9 and 97.1% dead cells resulting from contact with the wing surface, respectively.
NASA Astrophysics Data System (ADS)
Johansson, L. Christoffer; Håkansson, Jonas; Jakobsen, Lasse; Hedenström, Anders
2016-04-01
Large ears enhance perception of echolocation and prey generated sounds in bats. However, external ears likely impair aerodynamic performance of bats compared to birds. But large ears may generate lift on their own, mitigating the negative effects. We studied flying brown long-eared bats, using high resolution, time resolved particle image velocimetry, to determine the aerodynamics of flying with large ears. We show that the ears and body generate lift at medium to cruising speeds (3-5 m/s), but at the cost of an interaction with the wing root vortices, likely reducing inner wing performance. We also propose that the bats use a novel wing pitch mechanism at the end of the upstroke generating thrust at low speeds, which should provide effective pitch and yaw control. In addition, the wing tip vortices show a distinct spiraling pattern. The tip vortex of the previous wingbeat remains into the next wingbeat and rotates together with a newly formed tip vortex. Several smaller vortices, related to changes in circulation around the wing also spiral the tip vortex. Our results thus show a new level of complexity in bat wakes and suggest large eared bats are less aerodynamically limited than previous wake studies have suggested.
Drury, J. P.; Grether, G. F.
2014-01-01
Traits that mediate intraspecific social interactions may overlap in closely related sympatric species, resulting in costly between-species interactions. Such interactions have principally interested investigators studying the evolution of reproductive isolation via reproductive character displacement (RCD) or reinforcement, yet in addition to reproductive interference, interspecific trait overlap can lead to costly between-species aggression. Previous research on rubyspot damselflies (Hetaerina spp.) demonstrated that sympatric shifts in male wing colour patterns and competitor recognition reduce interspecific aggression, supporting the hypothesis that agonistic character displacement (ACD) drove trait shifts. However, a recent theoretical model shows that RCD overshadows ACD if the same male trait is used for both female mate recognition and male competitor recognition. To determine whether female mate recognition is based on male wing coloration in Hetaerina, we conducted a phenotype manipulation experiment. Compared to control males, male H. americana with wings manipulated to resemble a sympatric congener (H. titia) suffered no reduction in mating success. Thus, female mate recognition is not based on species differences in male wing coloration. Experimental males did, however, experience higher interspecific fighting rates and reduced survival compared to controls. These results greatly strengthen the case for ACD and highlight the mechanistic distinction between ACD and RCD. PMID:25339724
Roff, Derek A; Fairbairn, Daphne J
2007-01-01
Predicting evolutionary change is the central goal of evolutionary biology because it is the primary means by which we can test evolutionary hypotheses. In this article, we analyze the pattern of evolutionary change in a laboratory population of the wing-dimorphic sand cricket Gryllus firmus resulting from relaxation of selection favoring the migratory (long-winged) morph. Based on a well-characterized trade-off between fecundity and flight capability, we predict that evolution in the laboratory environment should result in a reduction in the proportion of long-winged morphs. We also predict increased fecundity and reduced functionality and weight of the major flight muscles in long-winged females but little change in short-winged (flightless) females. Based on quantitative genetic theory, we predict that the regression equation describing the trade-off between ovary weight and weight of the major flight muscles will show a change in its intercept but not in its slope. Comparisons across generations verify all of these predictions. Further, using values of genetic parameters estimated from previous studies, we show that a quantitative genetic simulation model can account for not only the qualitative changes but also the evolutionary trajectory. These results demonstrate the power of combining quantitative genetic and physiological approaches for understanding the evolution of complex traits.
NASA Technical Reports Server (NTRS)
Mann, M. J.; Huffman, J. K.; Fox, C. H., Jr.; Campbell, R. L.
1983-01-01
Wind tunnel tests were conducted to examine the use of wing leading-edge devices for improved subsonic and transonic maneuver performance. These devices were tested on a fighter configuration which utilized supercritical-wing technology. The configuration had a leading-edge sweep of 45 deg and an aspect ratio of 3.28. The tests were conducted at Mach numbers of 0.60 and 0.85 with angles of attack from -0.5 deg to 22 deg. At both Mach numbers, sharp leading-edge flaps produced vortices which greatly altered the flow pattern on the wing and resulted in substantial reductions in drag at high lift. Underwing or pylon-type vortex generators also reduced drag at high lift. The vortex generators worked better at a Mach number of 0.60. The vortex generators gave the best overall results with zero toe-in angle and when mounted on either the outboard part of the wing or at both an outboard location and halfway out the semispan. Both the flaps and the vortex generators had a minor effect on the pitching moment. Fluorescent minitufts were found to be useful for flow visualization at transonic maneuver conditions.
Study of lee-side flows over conically cambered Delta wings at supersonic speeds, part 2
NASA Technical Reports Server (NTRS)
Wood, Richard M.; Watson, Carolyn B.
1987-01-01
An experimental investigation was performed in which surface pressure data, flow visualization data, and force and moment data were obtained on four conical delta wing models which differed in leading edge camber only. Wing leading edge camber was achieved through a deflection of the outboard 30% of the local wing semispan of a reference 75 deg swept flat delta wing. The four wing models have leading edge deflection angles delta sub F of 0, 5, 10, and 15 deg measured streamwise. Data for the wings with delta sub F = 10 and 15 deg showed that hinge line separation dominated the lee-side wing loading and prohibited the development of leading edge separation on the deflected portion of wing leading edge. However, data for the wing with delta sub F = 5 deg showed that at an angle of attack of 5 deg, a vortex was positioned on the deflected leading edge with reattachment at the hinge line. Flow visualization results were presented which detail the influence of Mach number, angle of attack, and camber on the lee-side flow characteristics of conically cambered delta wings. Analysis of photographic data identified the existence of 12 distinctive lee-side flow types.
Quantifying the dynamic wing morphing of hovering hummingbird
Nakata, Toshiyuki; Kitamura, Ikuo; Tanaka, Hiroto
2017-01-01
Animal wings are lightweight and flexible; hence, during flapping flight their shapes change. It has been known that such dynamic wing morphing reduces aerodynamic cost in insects, but the consequences in vertebrate flyers, particularly birds, are not well understood. We have developed a method to reconstruct a three-dimensional wing model of a bird from the wing outline and the feather shafts (rachides). The morphological and kinematic parameters can be obtained using the wing model, and the numerical or mechanical simulations may also be carried out. To test the effectiveness of the method, we recorded the hovering flight of a hummingbird (Amazilia amazilia) using high-speed cameras and reconstructed the right wing. The wing shape varied substantially within a stroke cycle. Specifically, the maximum and minimum wing areas differed by 18%, presumably due to feather sliding; the wing was bent near the wrist joint, towards the upward direction and opposite to the stroke direction; positive upward camber and the ‘washout’ twist (monotonic decrease in the angle of incidence from the proximal to distal wing) were observed during both half-strokes; the spanwise distribution of the twist was uniform during downstroke, but an abrupt increase near the wrist joint was found during upstroke. PMID:28989736
NASA Astrophysics Data System (ADS)
Mieloszyk, M.; Krawczuk, M.; Zak, A.; Ostachowicz, W.
2010-08-01
In this paper a concept of an adaptive wing for small-aircraft applications with an array of fibre Bragg grating (FBG) sensors has been presented and discussed. In this concept the shape of the wing can be controlled and altered thanks to the wing design and the use of integrated shape memory alloy actuators. The concept has been tested numerically by the use of the finite element method. For numerical calculations the commercial finite element package ABAQUS® has been employed. A finite element model of the wing has been prepared in order to estimate the values of the wing twisting angles and distributions of the twist for various activation scenarios. Based on the results of numerical analysis the locations and numbers of the FBG sensors have also been determined. The results of numerical calculations obtained by the authors confirmed the usefulness of the assumed wing control strategy. Based on them and the concept developed of the adaptive wing, a wing demonstration stand has been designed and built. The stand has been used to verify experimentally the performance of the adaptive wing and the usefulness of the FBG sensors for evaluation of the wing condition.
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.
NASA Technical Reports Server (NTRS)
Pao, J. L.; Mehrotra, S. C.; Lan, C. E.
1982-01-01
A computer code base on an improved vortex filament/vortex core method for predicting aerodynamic characteristics of slender wings with edge vortex separations is developed. The code is applicable to camber wings, straked wings or wings with leading edge vortex flaps at subsonic speeds. The prediction of lifting pressure distribution and the computer time are improved by using a pair of concentrated vortex cores above the wing surface. The main features of this computer program are: (1) arbitrary camber shape may be defined and an option for exactly defining leading edge flap geometry is also provided; (2) the side edge vortex system is incorporated.
Roughness Effects on the Formation of a Leading Edge Vortex
NASA Astrophysics Data System (ADS)
Elliott, Cassidy; Lang, Amy; Wahidi, Redha; Wilroy, Jacob
2017-11-01
Microscopic scales cover the wings of Monarch butterflies, creating a patterned surface that acts as a natural energy capture mechanism. This patterning is thought to delay the growth of the leading edge vortex (LEV) produced by the flapping motion of a wing. Increased skin friction caused by the scales leads to a weaker LEV being shed into the butterfly's wake, lessening drag and increasing flight efficiency. To test how this roughness effects LEV formation, a plate of random roughness was designed in SolidWorks and printed on the Objet 30 Pro 3D printer. A 2x3x5 cubic foot tow tank was used to test the rough plate at Reynold's numbers of 1500, 3000, and 6000 (velocities of 8, 16, and 32 mm/s) at an angle of attack of 45 degrees. Images were captured of the LEV generated when the plate was towed upwards through the particle-seeded flow. These images were used to determine the XY velocity of the particles using a technique called Digital Particle Image Velocimetry (DPIV). Codes written in MATLAB were used to track and measure the strength of the LEV. Circulation values for the randomly-rough plate were then compared to the same values generated in a previous experiment that used a smooth plate and a grooved plate to determine the effect of the patterning on vortex development. Funding for this research project was provided by the National Science Foundation under the Research Experience for Undergraduates (REU) program (REU Supplement CBET 1628600 under CBET 1335848).
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.
NASA Technical Reports Server (NTRS)
Anusonti-Inthra, Phuriwat
2010-01-01
A novel Computational Fluid Dynamics (CFD) coupling framework using a conventional Reynolds-Averaged Navier-Stokes (BANS) solver to resolve the near-body flow field and a Particle-based Vorticity Transport Method (PVTM) to predict the evolution of the far field wake is developed, refined, and evaluated for fixed and rotary wing cases. For the rotary wing case, the RANS/PVTM modules are loosely coupled to a Computational Structural Dynamics (CSD) module that provides blade motion and vehicle trim information. The PVTM module is refined by the addition of vortex diffusion, stretching, and reorientation models as well as an efficient memory model. Results from the coupled framework are compared with several experimental data sets (a fixed-wing wind tunnel test and a rotary-wing hover test).
NASA Technical Reports Server (NTRS)
Larson, R. R.
1986-01-01
The wing on the NASA F-111 transonic aircraft technology airplane was modified to provide flexible leading and trailing edge flaps. This wing is known as the mission adaptive wing (MAW) because aerodynamic efficiency can be maintained at all speeds. Unlike a conventional wing, the MAW has no spoilers, external flap hinges, or fairings to break the smooth contour. The leading edge flaps and three-segment trailing edge flaps are controlled by a redundant fly-by-wire control system that features a dual digital primary system architecture providing roll and symmetric commands to the MAW control surfaces. A segregated analog backup system is provided in the event of a primary system failure. This paper discusses the design, development, testing, qualification, and flight test experience of the MAW primary and backup flight control systems.
Feeding ecology of northern pintails and green-winged teal wintering in California
Euliss, Ned H.; Harris, Stanley W.
1987-01-01
The feeding ecology of northern pintails (Anas acuta) and green-winged teal (A. crecca) was examined from October through February 1979-81 in 4 major seasonal marsh types in the Central Valley, California. The esophagi of 262 pintails contained 72.3% plant seeds and 27.7% animal matter. The esophagi of 173 green-winged teal contained 62.3% plant seeds and 37.6% animal matter. Swamp timothy (Heleochloa schoenoides) caryopses, chironomid midge larvae, and common barnyardgrass (Echinochloa crusgalli) caryopses formed >50% of the diet of both species. Both species were highly opportunistic and generally shifted their food habits seasonally to the most available foods. Animal matter increased seasonally in the diets of both and formed about 60% of the foods eaten during January and February compared to only about 8% in October and 17% in December. Both species used open water marsh habitats almost exclusively in daytime but they used densely vegetated marshes almost exclusively at night. Management recommendations based on the food habits and habitat use patterns of pintails and green-winged teal are offered.
NASA Astrophysics Data System (ADS)
Ehl, Stefan; Ebertshäuser, Marlene; Gros, Patrick; Schmitt, Thomas
2017-11-01
High mountain ecosystems are extreme habitats, and adaptation strategies to this ecosystem are still poorly understood in most groups. To unravel such strategies, we performed a MRR study in the Hohe Tauern National Park (Salzburg, Austria) with two nymphalid butterfly species, Boloria pales and B. napaea. We analysed their population structure over one flight period by studying the development of population size and wing wear. B. pales had more individuals and a higher survival probability than B. napaea; the sensitivity to extreme weather conditions or other external influences was higher in B. napaea. We only observed proterandry in B. pales. Imagines of both species survived under snow for at least some days. Additionally, we observed a kind of risk-spreading, in that individuals of both species, and especially B. pales, have regularly emerged throughout the flight period. This emergence pattern divided the population's age structure into three phases: an initial phase with decreasing wing quality (emergence > mortality), followed by an equilibrium phase with mostly constant average wing condition (emergence = mortality) and a final ageing phase with strongly deteriorating wing condition (mortality » emergence). Consequently, neither species would likely become extinct because of particularly unsuitable weather conditions during a single flight period. The observed differences between the two species suggest a better regional adaptation of B. pales, which is restricted to high mountain systems of Europe. In contrast, the arctic-alpine B. napaea might be best adapted to conditions in the Arctic and not the more southern high mountain systems. However, this needs to be examined during future research in the Arctic.
Supersonic aerodynamics of delta wings
NASA Technical Reports Server (NTRS)
Wood, Richard M.
1988-01-01
Through the empirical correlation of experimental data and theoretical analysis, a set of graphs has been developed which summarize the inviscid aerodynamics of delta wings at supersonic speeds. The various graphs which detail the aerodynamic performance of delta wings at both zero-lift and lifting conditions were then employed to define a preliminary wing design approach in which both the low-lift and high-lift design criteria were combined to define a feasible design space.
NASA Technical Reports Server (NTRS)
Lamar, John E.; Brandon, Jay; Stacy, Kathryn; Johnson, Thomas D., Jr.; Severance, Kurt; Childers, Brooks A.
1993-01-01
A flight research program to study the flow structure and separated-flow origins over an F-106B aircraft wing is described. The flight parameters presented include Mach numbers from 0.26 to 0.81, angles of attack from 8.5 deg to 22.5 deg, Reynolds numbers from 22.6 x 10(exp 6) to 57.3 x 10(exp 6) and load factors from 0.9 to 3.9 times the acceleration due to gravity. Techniques for vapor screens, image enhancement, photogrammetry, and computer graphics are integrated to analyze vortex-flow systems. Emphasis is placed on the development and application of the techniques. The spatial location of vortex cores and their tracks over the wing are derived from the analysis. Multiple vortices are observed and are likely attributed to small surface distortions in the wing leading-edge region. A major thrust is to correlate locations of reattachment lines obtained from the off-surface (vapor-screen) observations with those obtained from on-surface oil-flow patterns and pressure-port data. Applying vapor-screen image data to approximate reattachment lines is experimental, but depending on the angle of attack, the agreement with oil-flow results is generally good. Although surface pressure-port data are limited, the vapor-screen data indicate reattachment point occurrences consistent with the available data.
Accomplishments of the Abrupt Wing Stall (AWS) Program and Future Research Requirements
NASA Technical Reports Server (NTRS)
Hall, Robert M.; Woodson, Shawn H.; Chambers, Joseph R.
2003-01-01
The Abrupt Wing Stall (AWS) Program has addressed the problem of uncommanded lateral motions, such as wing drop and wing rock, at transonic speeds. The genesis of this Program was the experience of the F/A-1 8E/F Program in the late 1990's, when wing drop was discovered in the heart of the maneuver envelope for the pre-production aircraft. While the F/A-1 8E/F problem was subsequently corrected by a leading-edge flap scheduling change and the addition of a porous door to the wing fold fairing, the AWS Program was initiated as a national response to the lack of technology readiness available at the time of the F/A-18E/F Development Program. The AWS Program objectives were to define causal factors for the F/A-18E/F experience, to gain insights into the flow physics associated with wing drop, and to develop methods and analytical tools so that future programs could identify this type of problem before going to flight test. The paper reviews, for the major goals of the AWS Program, the status of the technology before the
Zhang, Sichao; Chen, Yifang
2015-01-01
The bright and iridescent blue color from Morpho butterfly wings has attracted worldwide attentions to explore its mysterious nature for long time. Although the physics of structural color by the nanophotonic structures built on the wing scales has been well established, replications of the wing structure by standard top-down lithography still remains a challenge. This paper reports a technical breakthrough to mimic the blue color of Morpho butterfly wings, by developing a novel nanofabrication process, based on electron beam lithography combined with alternate PMMA/LOR development/dissolution, for photonic structures with aligned lamellae multilayers in colorless polymers. The relationship between the coloration and geometric dimensions as well as shapes is systematically analyzed by solving Maxwell’s Equations with a finite domain time difference simulator. Careful characterization of the mimicked blue by spectral measurements under both normal and oblique angles are carried out. Structural color in blue reflected by the fabricated wing scales, is demonstrated and further extended to green as an application exercise of the new technique. The effects of the regularity in the replicas on coloration are analyzed. In principle, this approach establishes a starting point for mimicking structural colors beyond the blue in Morpho butterfly wings. PMID:26577813
Features of owl wings that promote silent flight
Weger, Matthias; Klaas, Michael; Schröder, Wolfgang
2017-01-01
Owls are an order of birds of prey that are known for the development of a silent flight. We review here the morphological adaptations of owls leading to silent flight and discuss also aerodynamic properties of owl wings. We start with early observations (until 2005), and then turn to recent advances. The large wings of these birds, resulting in low wing loading and a low aspect ratio, contribute to noise reduction by allowing slow flight. The serrations on the leading edge of the wing and the velvet-like surface have an effect on noise reduction and also lead to an improvement of aerodynamic performance. The fringes at the inner feather vanes reduce noise by gliding into the grooves at the lower wing surface that are formed by barb shafts. The fringed trailing edge of the wing has been shown to reduce trailing edge noise. These adaptations to silent flight have been an inspiration for biologists and engineers for the development of devices with reduced noise production. Today several biomimetic applications such as a serrated pantograph or a fringed ventilator are available. Finally, we discuss unresolved questions and possible future directions. PMID:28163870
Zhang, Sichao; Chen, Yifang
2015-11-18
The bright and iridescent blue color from Morpho butterfly wings has attracted worldwide attentions to explore its mysterious nature for long time. Although the physics of structural color by the nanophotonic structures built on the wing scales has been well established, replications of the wing structure by standard top-down lithography still remains a challenge. This paper reports a technical breakthrough to mimic the blue color of Morpho butterfly wings, by developing a novel nanofabrication process, based on electron beam lithography combined with alternate PMMA/LOR development/dissolution, for photonic structures with aligned lamellae multilayers in colorless polymers. The relationship between the coloration and geometric dimensions as well as shapes is systematically analyzed by solving Maxwell's Equations with a finite domain time difference simulator. Careful characterization of the mimicked blue by spectral measurements under both normal and oblique angles are carried out. Structural color in blue reflected by the fabricated wing scales, is demonstrated and further extended to green as an application exercise of the new technique. The effects of the regularity in the replicas on coloration are analyzed. In principle, this approach establishes a starting point for mimicking structural colors beyond the blue in Morpho butterfly wings.
NASA Astrophysics Data System (ADS)
Zhang, Sichao; Chen, Yifang
2015-11-01
The bright and iridescent blue color from Morpho butterfly wings has attracted worldwide attentions to explore its mysterious nature for long time. Although the physics of structural color by the nanophotonic structures built on the wing scales has been well established, replications of the wing structure by standard top-down lithography still remains a challenge. This paper reports a technical breakthrough to mimic the blue color of Morpho butterfly wings, by developing a novel nanofabrication process, based on electron beam lithography combined with alternate PMMA/LOR development/dissolution, for photonic structures with aligned lamellae multilayers in colorless polymers. The relationship between the coloration and geometric dimensions as well as shapes is systematically analyzed by solving Maxwell’s Equations with a finite domain time difference simulator. Careful characterization of the mimicked blue by spectral measurements under both normal and oblique angles are carried out. Structural color in blue reflected by the fabricated wing scales, is demonstrated and further extended to green as an application exercise of the new technique. The effects of the regularity in the replicas on coloration are analyzed. In principle, this approach establishes a starting point for mimicking structural colors beyond the blue in Morpho butterfly wings.
Aerodynamic Characteristics of Tube-Launched Tandem Wing Unmanned Aerial Vehicle
NASA Astrophysics Data System (ADS)
Rosid, Nurhayyan H.; Irsyad Lukman, E.; Fadlillah, M. Ahmad; Agoes Moelyadi, M.
2018-04-01
Tube Launched UAV with expandable tandem-wing configuration becomes one of the most interesting topic to be investigated. Folding wing mechanism is used due to the requirements that the UAV should be folded into tubular launcher. This paper focuses on investigating the aerodynamics characteristics because of the effects of folding wing mechanism, tandem wing configuration, and rapid deploying process from tube launcher. The aerodynamic characteristics investigation is conducted using computational fluid dynamics (CFD) at low Reynolds numbers (Re < 200000). The results of the simulation are used for the development of ITB Tube-launched UAV prototype and for future studies.
Three-dimensional aerodynamic shape optimization of supersonic delta wings
NASA Technical Reports Server (NTRS)
Burgreen, Greg W.; Baysal, Oktay
1994-01-01
A recently developed three-dimensional aerodynamic shape optimization procedure AeSOP(sub 3D) is described. This procedure incorporates some of the most promising concepts from the area of computational aerodynamic analysis and design, specifically, discrete sensitivity analysis, a fully implicit 3D Computational Fluid Dynamics (CFD) methodology, and 3D Bezier-Bernstein surface parameterizations. The new procedure is demonstrated in the preliminary design of supersonic delta wings. Starting from a symmetric clipped delta wing geometry, a Mach 1.62 asymmetric delta wing and two Mach 1. 5 cranked delta wings were designed subject to various aerodynamic and geometric constraints.
Properties of oscillating refractive optical wings with one reflective surface
NASA Astrophysics Data System (ADS)
Artusio-Glimpse, Alexandra B.; Swartzlander, Grover A.
2013-09-01
A new modality for optical micromanipulation is under investigation. Optical wings are shaped refractive objects that experience a force and torque owing to the reflection and transmission of uniform light at the object surface. We present wing designs that provide a restoring torque that returns the wing to a source facing orientation while preserving efficient thrust from radiation pressure. The torsional stiffness and orbital period of a set of optical wing cross-sectional shapes are determined from numerical ray-tracing analyses. These results demonstrate the potential to develop an efficient optomechanical device for applications in microbiology and space flight systems.
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.
Aeroelastic loads prediction for an arrow wing. Task 1: Evaluation of R. P. White's method
NASA Technical Reports Server (NTRS)
Borland, C. J.; Manro, M. E.
1983-01-01
The separated flow method is evaluated. This method was developed for moderately swept wings with multiple, constant strength vortex systems. The flow on the highly swept wing used in this evaluation is characterized by a single vortex system of continuously varying strength.
NASA Astrophysics Data System (ADS)
DeLuca, Anthony M.
Considerable research and investigation has been conducted on the aerodynamic performance, and the predominate flow physics of the Manduca Sexta size of biomimetically designed and fabricated wings as part of the AFIT FWMAV design project. Despite a burgeoning interest and research into the diverse field of flapping wing flight and biomimicry, the aerodynamics of flapping wing flight remains a nebulous field of science with considerable variance into the theoretical abstractions surrounding aerodynamic mechanisms responsible for aerial performance. Traditional FWMAV flight models assume a form of a quasi-steady approximation of wing aerodynamics based on an infinite wing blade element model (BEM). An accurate estimation of the lift, drag, and side force coefficients is a critical component of autonomous stability and control models. This research focused on two separate experimental avenues into the aerodynamics of AFIT's engineered hawkmoth wings|forces and flow visualization. 1. Six degree of freedom force balance testing, and high speed video analysis was conducted on 30°, 45°, and 60° angle stop wings. A novel, non-intrusive optical tracking algorithm was developed utilizing a combination of a Gaussian Mixture Model (GMM) and ComputerVision (OpenCV) tools to track the wing in motion from multiple cameras. A complete mapping of the wing's kinematic angles as a function of driving amplitude was performed. The stroke angle, elevation angle, and angle of attack were tabulated for all three wings at driving amplitudes ranging from A=0.3 to A=0.6. The wing kinematics together with the force balance data was used to develop several aerodynamic force coefficient models. A combined translational and rotational aerodynamic model predicted lift forces within 10%, and vertical forces within 6%. The total power consumption was calculated for each of the three wings, and a Figure of Merit was calculated for each wing as a general expression of the overall efficiency of the wing. Th 60° angle stop wing achieved the largest total stroke angle and generated the most lift for the lowest power consumption of the wings tested. 2. Phase averaged stereo Particle Image Velocimetry (PIV) data was collected at eight phases through the flap cycle on the 30°, 45°, and 60° angle stop wings. Wings were mounted transverse and parallel to the interrogating laser sheet, and planar velocity intersections at the wing mid-span, one chord below the wing, were compared to one another to verify data fidelity. A Rankine-Froude actuator disk model was adapted to calculate the approximate vertical thrust generated from the total momentum flux through the flapping semi-disk using the velocity field measurements. Three component stereo u, v, and w-velocity contour measurements confirmed the presence of extensive vortical structures in the vicinity of the wing. The leading edge vortex was successfully tracked through the stroke cycle appearing at approximately 25% span, increasing in circulatory strength and translational velocity down the span toward the tip, and dissipating just after 75% span. Thrust calculations showed the vertically mounted wing more accurately represented the vertical forces when compared to its corresponding force balance measurement than the horizontally mounted wing. The mid-span showed the highest vertical velocity profile below the wing; and hence, was the location responsible for the majority of lift production along the span.