The development of wing shape in Lepidoptera: mitotic density, not orientation, is the primary determinant of shape.

PubMed

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.

Convergent evolution in the genetic basis of Müllerian mimicry in heliconius butterflies.

PubMed

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.

Convergent, modular expression of ebony and tan in the mimetic wing patterns of Heliconius butterflies.

PubMed

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

Wing shape variation associated with mimicry in butterflies.

PubMed

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.

Wnt signaling underlies evolution and development of the butterfly wing pattern symmetry systems.

PubMed

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.

An introgressed wing pattern acts as a mating cue.

PubMed

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.

Transcriptome analysis of the painted lady butterfly, Vanessa cardui during wing color pattern development.

PubMed

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.

Wing patterning genes and coevolution of Müllerian mimicry in Heliconius butterflies: Support from phylogeography, cophylogeny, and divergence times.

PubMed

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.

RNAi screening of developmental toolkit genes: a search for novel wing genes in the red flour beetle, Tribolium castaneum.

PubMed

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.

Single master regulatory gene coordinates the evolution and development of butterfly color and iridescence

PubMed Central

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

Single master regulatory gene coordinates the evolution and development of butterfly color and iridescence.

PubMed

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.

Inhibition of Shh signalling in the chick wing gives insights into digit patterning and evolution.

PubMed

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.

Characterisation and expression of microRNAs in developing wings of the neotropical butterfly Heliconius melpomene

PubMed Central

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

Characterisation and expression of microRNAs in developing wings of the neotropical butterfly Heliconius melpomene.

PubMed

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.

Morphogenesis in bat wings: linking development, evolution and ecology.

PubMed

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

Effects of ornamentation and phylogeny on the evolution of wing shape in stalk-eyed flies (Diopsidae).

PubMed

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.

Comparative developmental analysis of Drosophila and Tribolium reveals conserved and diverged roles of abrupt in insect wing evolution.

PubMed

Ravisankar, Padmapriyadarshini; Lai, Yi-Ting; Sambrani, Nagraj; Tomoyasu, Yoshinori

2016-01-15

Morphological innovation is a fundamental process in evolution, yet its molecular basis is still elusive. Acquisition of elytra, highly modified beetle forewings, is an important innovation that has driven the successful radiation of beetles. Our RNAi screening for candidate genes has identified abrupt (ab) as a potential key player in elytron evolution. In this study, we performed a series of RNA interference (RNAi) experiments in both Tribolium and Drosophila to understand the contributions of ab to the evolution of beetle elytra. We found that (i) ab is essential for proper wing vein patterning both in Tribolium and Drosophila, (ii) ab has gained a novel function in determining the unique elytron shape in the beetle lineage, (iii) unlike Hippo and Insulin, other shape determining pathways, the shape determining function of ab is specific to the elytron and not required in the hindwing, (iv) ab has a previously undescribed role in the Notch signal-associated wing formation processes, which appears to be conserved between beetles and flies. These data suggest that ab has gained a new function during elytron evolution in beetles without compromising the conserved wing-related functions. Gaining a new function without losing evolutionarily conserved functions may be a key theme in the evolution of morphologically novel structures. Copyright © 2015 Elsevier Inc. All rights reserved.

Colour pattern homology and evolution in Vanessa butterflies (Nymphalidae: Nymphalini): eyespot characters.

PubMed

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.

Evolution of female-specific wingless forms in bagworm moths.

PubMed

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.

Cruising the rain forest floor: butterfly wing shape evolution and gliding in ground effect.

PubMed

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.

Prey from the eyes of predators: Color discriminability of aposematic and mimetic butterflies from an avian visual perspective.

PubMed

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.

Beyond magic traits: Multimodal mating cues in Heliconius butterflies.

PubMed

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.

Laboratory evolution of the migratory polymorphism in the sand cricket: combining physiology with quantitative genetics.

PubMed

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.

Crossflow-Vortex Breakdown on Swept Wings: Correlation of Nonlinear Physics

NASA Technical Reports Server (NTRS)

Joslin, R. D.; Streett, C. L.

1994-01-01

The spatial evolution of cross flow-vortex packets in a laminar boundary layer on a swept wing are computed by the direct numerical simulation of the incompressible Navier- Stokes equations. A wall-normal velocity distribution of steady suction and blowing at the wing surface is used to generate a strip of equally spaced and periodic disturbances along the span. Three simulations are conducted to study the effect of initial amplitude on the disturbance evolution, to determine the role of traveling cross ow modes in transition, and to devise a correlation function to guide theories of transition prediction. In each simulation, the vortex packets first enter a chordwise region of linear independent growth, then, the individual packets coalesce downstream and interact with adjacent packets, and, finally, the vortex packets nonlinearly interact to generate inflectional velocity profiles. As the initial amplitude of the disturbance is increased, the length of the evolution to breakdown decreases. For this pressure gradient, stationary modes dominate the disturbance evolution. A two-coeffcient function was devised to correlate the simulation results. The coefficients, combined with a single simulation result, provide sufficient information to generate the evolution pattern for disturbances of any initial amplitude.

Tipping the scales: Evolution of the allometric slope independent of average trait size.

PubMed

Stillwell, R Craig; Shingleton, Alexander W; Dworkin, Ian; Frankino, W Anthony

2016-02-01

The scaling of body parts is central to the expression of morphology across body sizes and to the generation of morphological diversity within and among species. Although patterns of scaling-relationship evolution have been well documented for over one hundred years, little is known regarding how selection acts to generate these patterns. In part, this is because it is unclear the extent to which the elements of log-linear scaling relationships-the intercept or mean trait size and the slope-can evolve independently. Here, using the wing-body size scaling relationship in Drosophila melanogaster as an empirical model, we use artificial selection to demonstrate that the slope of a morphological scaling relationship between an organ (the wing) and body size can evolve independently of mean organ or body size. We discuss our findings in the context of how selection likely operates on morphological scaling relationships in nature, the developmental basis for evolved changes in scaling, and the general approach of using individual-based selection experiments to study the expression and evolution of morphological scaling. © 2016 The Author(s). Evolution © 2016 The Society for the Study of Evolution.

Qualitative skeletal correlates of wing shape in extant birds (Aves: Neoaves).

PubMed

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.

Genome of Drosophila suzukii, the Spotted Wing Drosophila

PubMed Central

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

Numerical analyses of evolution of unsteady flow structures in the wake of flapping starling wing model

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.

The Functional Basis of Wing Patterning in Heliconius Butterflies: The Molecules Behind Mimicry

PubMed Central

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

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

PubMed Central

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

Slender wing theory including regions of embedded total pressure loss

NASA Technical Reports Server (NTRS)

Mccune, James E.; Tavares, T. Sean; Lee, Norman K. W.; Weissbein, David

1988-01-01

An aerodynamic theory of the flow about slender delta wings is described. The theory includes a treatment of the self-consistent development of the vortex wake patterns above the wing necessary to maintain smooth flow at the wing edges. The paper focuses especially on the formation within the wake of vortex 'cores' as embedded regions of total pressure loss, fed and maintained by umbilical vortex sheets emanating from the wing edges. Criteria are developed for determining the growing size and location of these cores, as well as the distribution and strength of the vorticity within them. In this paper, however, the possibility of vortex breakup is omitted. The aerodynamic consequences of the presence and evolution of the cores and the associated wake structure are illustrated and discussed. It is noted that wake history effects can have substantial influence on the distribution of normal force on the wing as well as on its magnitude.

The evolution of host specificity in dove body lice.

PubMed

Johnson, Kevin P; Weckstein, Jason D; Bush, Sarah E; Clayton, Dale H

2011-11-01

Conventional wisdom suggests that parasites evolve increased host specialization over time. Host specificity, which describes the number of host species parasitized, is one aspect of host specialization. Recent studies of vertebrate parasites indicate that highly host-specific parasite lineages are not, in fact, evolutionary dead ends; host generalists can evolve from host specialists. Using phylogenetic reconstruction methods, we evaluate these patterns in the body lice (Insecta: Phthiraptera) of pigeons and doves, which are permanent ectoparasites that complete their entire life cycle on the body of the host. We find that species of body lice that parasitize more than one species of host (generalists) are invariably derived from lice parasitizing only one species of host (specialists). A previous study of the wing lice of pigeons and doves also found that generalists were derived from specialists, and that these changes were correlated with the presence of a potentially competing species of wing louse on the same host. For body lice we did not find such a correlation with competition. Instead, the evolution of host generalists in body lice was correlated with host ecology. When we compared body lice that parasitize terrestrial versus arboreal hosts, we found that the evolution of host generalists was associated with terrestrial hosts. In contrast, wing lice showed no correlation between the evolution of generalists and host ecology. The correlation in body lice suggests that dispersal between host species may occur via the ground. This, in turn, suggests that body lice may fall to the ground more often than wing lice. To test this hypothesis, we conducted an experiment to compare the rate at which body and wing lice are dislodged from the bodies of preening pigeons. Interestingly, our results showed that body lice are dislodged four times more often than wing lice. Therefore, species of terrestrial doves are far more likely to encounter body lice than wing lice on the ground.

Complex dynamics underlie the evolution of imperfect wing pattern convergence in butterflies.

PubMed

Finkbeiner, Susan D; Briscoe, Adriana D; Mullen, Sean P

2017-04-01

Adaptive radiation is characterized by rapid diversification that is strongly associated with ecological specialization. However, understanding the evolutionary mechanisms fueling adaptive diversification requires a detailed knowledge of how natural selection acts at multiple life-history stages. Butterflies within the genus Adelpha represent one of the largest and most diverse butterfly lineages in the Neotropics. Although Adelpha species feed on an extraordinary diversity of larval hosts, convergent evolution is widespread in this group, suggesting that selection for mimicry may contribute to adaptive divergence among species. To investigate this hypothesis, we conducted predation studies in Costa Rica using artificial butterfly facsimiles. Specifically, we predicted that nontoxic, palatable Adelpha species that do not feed on host plants in the family Rubiaceae would benefit from sharing a locally convergent wing pattern with the presumably toxic Rubiaceae-feeding species via reduced predation. Contrary to expectations, we found that the presumed mimic was attacked significantly more than its locally convergent model at a frequency paralleling attack rates on both novel and palatable prey. Although these data reveal the first evidence for protection from avian predators by the supposed toxic, Rubiaceae-feeding Adelpha species, we conclude that imprecise mimetic patterns have high costs for Batesian mimics in the tropics. © 2017 The Author(s). Evolution © 2017 The Society for the Study of Evolution.

Varying and unchanging whiteness on the wings of dusk-active and shade-inhabiting Carystoides escalantei butterflies.

PubMed

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.

Wing serial homologs and the origin and evolution of the insect wing.

PubMed

Ohde, Takahiro; Yaginuma, Toshinobu; Niimi, Teruyuki

2014-04-01

The origin and evolution of insect wings has been the subject of extensive debate. The issue has remained controversial largely because of the absence of definitive fossil evidence or direct developmental evidence of homology between wings and a putative wing origin. Recent identification of wing serial homologs (WSHs) has provided researchers with a potential strategy for identifying WSHs in other species. Future comparative developmental analyses between wings and WSHs may clarify the important steps underlying the evolution of insect wings. Copyright © 2013 The Authors. Published by Elsevier GmbH.. All rights reserved.

Mutation predicts 40 million years of fly wing evolution.

PubMed

Houle, David; Bolstad, Geir H; van der Linde, Kim; Hansen, Thomas F

2017-08-24

Mutation enables evolution, but the idea that adaptation is also shaped by mutational variation is controversial. Simple evolutionary hypotheses predict such a relationship if the supply of mutations constrains evolution, but it is not clear that constraints exist, and, even if they do, they may be overcome by long-term natural selection. Quantification of the relationship between mutation and phenotypic divergence among species will help to resolve these issues. Here we use precise data on over 50,000 Drosophilid fly wings to demonstrate unexpectedly strong positive relationships between variation produced by mutation, standing genetic variation, and the rate of evolution over the last 40 million years. Our results are inconsistent with simple constraint hypotheses because the rate of evolution is very low relative to what both mutational and standing variation could allow. In principle, the constraint hypothesis could be rescued if the vast majority of mutations are so deleterious that they cannot contribute to evolution, but this also requires the implausible assumption that deleterious mutations have the same pattern of effects as potentially advantageous ones. Our evidence for a strong relationship between mutation and divergence in a slowly evolving structure challenges the existing models of mutation in evolution.

Aerodynamic forces and flow structures of the leading edge vortex on a flapping wing considering ground effect.

PubMed

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.

Evolution of insect wings and development - new details from Palaeozoic nymphs.

PubMed

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.

A Burst of miRNA Innovation in the Early Evolution of Butterflies and Moths

PubMed Central

Quah, Shan; Hui, Jerome H.L.; Holland, Peter W.H.

2015-01-01

MicroRNAs (miRNAs) are involved in posttranscriptional regulation of gene expression. Because several miRNAs are known to affect the stability or translation of developmental regulatory genes, the origin of novel miRNAs may have contributed to the evolution of developmental processes and morphology. Lepidoptera (butterflies and moths) is a species-rich clade with a well-established phylogeny and abundant genomic resources, thereby representing an ideal system in which to study miRNA evolution. We sequenced small RNA libraries from developmental stages of two divergent lepidopterans, Cameraria ohridella (Horse chestnut Leafminer) and Pararge aegeria (Speckled Wood butterfly), discovering 90 and 81 conserved miRNAs, respectively, and many species-specific miRNA sequences. Mapping miRNAs onto the lepidopteran phylogeny reveals rapid miRNA turnover and an episode of miRNA fixation early in lepidopteran evolution, implying that miRNA acquisition accompanied the early radiation of the Lepidoptera. One lepidopteran-specific miRNA gene, miR-2768, is located within an intron of the homeobox gene invected, involved in insect segmental and wing patterning. We identified cubitus interruptus (ci) as a likely direct target of miR-2768, and validated this suppression using a luciferase assay system. We propose a model by which miR-2768 modulates expression of ci in the segmentation pathway and in patterning of lepidopteran wing primordia. PMID:25576364

Gradual and contingent evolutionary emergence of leaf mimicry in butterfly wing patterns.

PubMed

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.

The homology of wing base sclerites and flight muscles in Ephemeroptera and Neoptera and the morphology of the pterothorax of Habroleptoides confusa (Insecta: Ephemeroptera: Leptophlebiidae).

PubMed

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.

New developmental evidence supports a homeotic frameshift of digit identity in the evolution of the bird wing

PubMed Central

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

New developmental evidence supports a homeotic frameshift of digit identity in the evolution of the bird wing.

PubMed

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

PubMed Central

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

Wings versus legs in the avian bauplan: development and evolution of alternative locomotor strategies.

PubMed

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.

Making quantitative morphological variation from basic developmental processes: where are we? The case of the Drosophila wing

PubMed Central

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

Underwater flight by the planktonic sea butterfly.

PubMed

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.

The generation and diversification of butterfly eyespot color patterns.

PubMed

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.

Mutualistic mimicry and filtering by altitude shape the structure of Andean butterfly communities.

PubMed

Chazot, Nicolas; Willmott, Keith R; Santacruz Endara, Paola G; Toporov, Alexandre; Hill, Ryan I; Jiggins, Chris D; Elias, Marianne

2014-01-01

Both the abiotic environment and abiotic interactions among species contribute to shaping species assemblages. While the roles of habitat filtering and competitive interactions are clearly established, less is known about how positive interactions, whereby species benefit from the presence of one another, affect community structure. Here we assess the importance of positive interactions by studying Andean communities of butterflies that interact mutualistically via Müllerian mimicry. We show that communities at similar altitudes have a similar phylogenetic composition, confirming that filtering by altitude is an important process. We also provide evidence that species that interact mutualistically (i.e., species that share the same mimicry wing pattern) coexist at large scales more often than expected by chance. Furthermore, we detect an association between mimicry structure and altitude that is stronger than expected even when phylogeny is corrected for, indicating adaptive convergence for wing pattern and/or altitudinal range driven by mutualistic interactions. Positive interactions extend far beyond Müllerian mimicry, with many examples in plants and animals, and their role in the evolution and assembly of communities may be more pervasive than is currently appreciated. Our findings have strong implications for the evolution and resilience of community structure in a changing world.

Differential expression of Meis2, Mab21l2 and Tbx3 during limb development associated with diversification of limb morphology in mammals.

PubMed

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.

Differential Expression of Meis2, Mab21l2 and Tbx3 during Limb Development Associated with Diversification of Limb Morphology in Mammals

PubMed Central

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

Dynamic Pattern Formation for Wings of Pterygota in an Eclosion ---Pattern Analysis for Wings with the Imago---

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.

Vertical distribution, flight behaviour and evolution of wing morphology in Morpho butterflies.

PubMed

Devries, P J; Penz, Carla M; Hill, Ryan I

2010-09-01

1. Flight is a key innovation in the evolution of insects that is crucial to their dispersal, migration, territoriality, courtship and predator avoidance. Male butterflies have characteristic territoriality and courtship flight behaviours, and females use a characteristic flight behaviour when searching for host plants. This implies that selection acts on wing morphology to maximize flight performance for conducting important behaviours among sexes. 2. Butterflies in the genus Morpho are obvious components of neotropical forests, and many observations indicate that they show two broad categories of flight behaviour and flight height. Although species can be categorized as using gliding or flapping flight, and flying at either canopy or understorey height, the association of flight behaviour and flight height with wing shape evolution has never been explored. 3. Two clades within Morpho differ in flight behaviour and height. Males and females of one clade inhabit the forest understorey and use flapping flight, whereas in the other clade, males use gliding flight at canopy level and females use flapping flight in both canopy and understorey. 4. We used independent contrasts to answer whether wing shape is associated with flight behaviour and height. Given a single switch to canopy habitation and gliding flight, we compared contrasts for the node at which the switch to canopy flight occurred with the distribution of values in the two focal clades. We found significant changes in wing shape at the transition to canopy flight only in males, and no change in size for either sex. A second node within the canopy clade suggests that other factors may also be involved in wing shape evolution. Our results reinforce the hypothesis that natural selection acts differently on male and female butterfly wing shape and indicate that the transition to canopy flight cannot explain all wing shape diversity in Morpho. 5. This study provides a starting point for characterizing evolution of wing morphology in forest butterflies in the contexts of habitat selection and flight behaviour. Further, these observations suggest that exploring wing shape evolution for canopy and understorey species in other insects may help understand the effects of habitat destruction on biological diversity.

Comparative genomics of the mimicry switch in Papilio dardanus.

PubMed

Timmermans, Martijn J T N; Baxter, Simon W; Clark, Rebecca; Heckel, David G; Vogel, Heiko; Collins, Steve; Papanicolaou, Alexie; Fukova, Iva; Joron, Mathieu; Thompson, Martin J; Jiggins, Chris D; ffrench-Constant, Richard H; Vogler, Alfried P

2014-07-22

The African Mocker Swallowtail, Papilio dardanus, is a textbook example in evolutionary genetics. Classical breeding experiments have shown that wing pattern variation in this polymorphic Batesian mimic is determined by the polyallelic H locus that controls a set of distinct mimetic phenotypes. Using bacterial artificial chromosome (BAC) sequencing, recombination analyses and comparative genomics, we show that H co-segregates with an interval of less than 500 kb that is collinear with two other Lepidoptera genomes and contains 24 genes, including the transcription factor genes engrailed (en) and invected (inv). H is located in a region of conserved gene order, which argues against any role for genomic translocations in the evolution of a hypothesized multi-gene mimicry locus. Natural populations of P. dardanus show significant associations of specific morphs with single nucleotide polymorphisms (SNPs), centred on en. In addition, SNP variation in the H region reveals evidence of non-neutral molecular evolution in the en gene alone. We find evidence for a duplication potentially driving physical constraints on recombination in the lamborni morph. Absence of perfect linkage disequilibrium between different genes in the other morphs suggests that H is limited to nucleotide positions in the regulatory and coding regions of en. Our results therefore support the hypothesis that a single gene underlies wing pattern variation in P. dardanus.

Evolution of limited seed dispersal ability on gypsum islands.

PubMed

Schenk, John J

2013-09-01

Dispersal is a major feature of plant evolution that has many advantages but is not always favored. Wide dispersal, for example, leads to greater seed loss in oceanic-island endemics, and evolution has favored morphologies that limit dispersal. I tested the hypothesis that selection favored limited dispersal on gypsum islands in western North America, where edaphic communities are sparsely vegetated except for a specialized flora that competes poorly with the surrounding flora. • I applied a series of comparative phylogenetic approaches to gypsophilic species of Mentzelia section Bartonia (Loasaceae) to investigate the evolution of limited dispersal function in seed wings, which increase primary dispersal by wind. Through these tests, I determined whether narrowed wings were selected for in gypsophilic species. • Gypsophily was derived four to seven times. Seed area was not significantly correlated with gypsophily or wing area. Wing area was significantly smaller in the derived gypsum endemics, supporting the hypothesis in favor of limited dispersal function. A model-fitting approach identified two trait optima in wing area, with gypsum endemics having a lower optimum. • Evolution into novel ecologies influences morphological evolution. Morphological characters have been selected for limited dispersal following evolution onto gypsum islands. Selection for limited dispersal ability has occurred across animals and plants, both in oceanic and terrestrial systems, which suggests that reduced dispersal ability may be a general process: selection favors limited dispersal if the difference in survival between the habitat of the parent and the surrounding area is great enough.

The aerodynamic benefit of wing-wing interaction depends on stroke trajectory in flapping insect wings.

PubMed

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.

Longwing (Heliconius) butterflies combine a restricted set of pigmentary and structural coloration mechanisms.

PubMed

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.

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.

Interspecific aggression, not interspecific mating, drives character displacement in the wing coloration of male rubyspot damselflies (Hetaerina)

PubMed Central

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

Influence of wing tip morphology on vortex dynamics of flapping flight

NASA Astrophysics Data System (ADS)

Krishna, Swathi; Mulleners, Karen

2013-11-01

The mechanism of flapping wing flight provides insects with extraordinary flight capabilities. The uniquely shaped wing tips give insects an edge in flight performance and the interaction between the leading edge vortices and wing tip vortices enhance their propelling efficiencies and manoeuvrability. These are qualities that are sought after in current-day Micro Air Vehicles. A detailed understanding of the vortex dynamics of flapping flight and the influence of the wing tip planform is imperative for technical application. An experimental study is conducted to investigate the effects of different wing tip planforms on the formation, evolution and interaction of vortical structures. We thereby focus on the interaction between the coherent structures evolving from the leading edge and the wing tip during pitching and flapping motions.The spatial and temporal evolution of the three-dimensional flow structures are determined using Scanning (Stereo) Particle Image Velocimetry and an in-depth coherent structure analysis. By comparing the vortex dynamics, the aerodynamic performance of various wing tip planforms are evaluated.

Insights into bird wing evolution and digit specification from polarizing region fate maps.

PubMed

Towers, Matthew; Signolet, Jason; Sherman, Adrian; Sang, Helen; Tickle, Cheryll

2011-08-09

The proposal that birds descended from theropod dinosaurs with digits 2, 3 and 4 was recently given support by short-term fate maps, suggesting that the chick wing polarizing region-a group that Sonic hedgehog-expressing cells-gives rise to digit 4. Here we show using long-term fate maps that Green fluorescent protein-expressing chick wing polarizing region grafts contribute only to soft tissues along the posterior margin of digit 4, supporting fossil data that birds descended from theropods that had digits 1, 2 and 3. In contrast, digit IV of the chick leg with four digits (I-IV) arises from the polarizing region. To determine how digit identity is specified over time, we inhibited Sonic hedgehog signalling. Fate maps show that polarizing region and adjacent cells are specified in parallel through a series of anterior to posterior digit fates-a process of digit specification that we suggest is involved in patterning all vertebrate limbs with more than three digits.

Traits and evolution of wing venation pattern in paraneopteran insects.

PubMed

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.

Expression of Genes Involved in Drosophila Wing Morphogenesis and Vein Patterning Are Altered by Spaceflight

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.

Growth and patterning are evolutionarily dissociated in the vestigial wing discs of workers of the red imported fire ant, Solenopsis invicta.

PubMed

Bowsher, Julia H; Wray, Gregory A; Abouheif, Ehab

2007-12-15

Over the last decade, it has become clear that organismal form is largely determined by developmental and evolutionary changes in the growth and pattern formation of tissues. Yet, there is little known about how these two integrated processes respond to environmental cues or how they evolve relative to one another. Here, we present the discovery of vestigial wing imaginal discs in worker larvae of the red imported fire ant, Solenopsis invicta. These vestigial wing discs are present in all worker larvae, which is uncommon for a species with a large worker size distribution. Furthermore, the growth trajectory of these vestigial discs is distinct from all of the ant species examined to date because they grow at a rate slower than the leg discs. We predicted that the growth trajectory of the vestigial wing discs would be mirrored by evolutionary changes in their patterning. We tested this prediction by examining the expression of three patterning genes, extradenticle, ultrabithorax, and engrailed, known to underlie the wing polyphenism in ants. Surprisingly, the expression patterns of these three genes in the vestigial wing discs was the same as those found in ant species with different worker size distributions and wing disc growth than fire ants. We conclude that growth and patterning are evolutionarily dissociated in the vestigial wing discs of S. invicta because patterning in these discs is conserved, whereas their growth trajectories are not. The evolutionary dissociation of growth and patterning may be an important feature of gene networks that underlie polyphenic traits. 2007 Wiley-Liss, Inc

Integration of wings and their eyespots in the speckled wood butterfly Pararge aegeria.

PubMed

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.

Forelimb posture in dinosaurs and the evolution of the avian flapping flight-stroke.

PubMed

Nudds, Robert L; Dyke, Gareth J

2009-04-01

Ontogenetic and behavioral studies using birds currently do not document the early evolution of flight because birds (including juveniles) used in such studies employ forelimb oscillation frequencies over 10 Hz, forelimb stroke-angles in excess of 130 degrees , and possess uniquely avian flight musculatures. Living birds are an advanced morphological stage in the development of flapping flight. To gain insight into the early stages of flight evolution (i.e., prebird), in the absence of a living analogue, a new approach using Strouhal number was used. Strouhal number is a nondimensional number that describes the relationship between wing-stroke amplitude (A), wing-beat frequency (f), and flight speed (U). Calculations indicated that even moderate wing movements are enough to generate rudimentary thrust and that a propulsive flapping flight-stroke could have evolved via gradual incremental changes in wing movement and wing morphology. More fundamental to the origin of the avian flapping flight-stroke is the question of how a symmetrical forelimb posture-required for gliding and flapping flight-evolved from an alternating forelimb motion, evident in all extant bipeds when running except birds.

Distal-less induces elemental color patterns in Junonia butterfly wings.

PubMed

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.

Phylogenetic Analysis of Pelecaniformes (Aves) Based on Osteological Data: Implications for Waterbird Phylogeny and Fossil Calibration Studies

PubMed Central

Smith, Nathan D.

2010-01-01

Background Debate regarding the monophyly and relationships of the avian order Pelecaniformes represents a classic example of discord between morphological and molecular estimates of phylogeny. This lack of consensus hampers interpretation of the group's fossil record, which has major implications for understanding patterns of character evolution (e.g., the evolution of wing-propelled diving) and temporal diversification (e.g., the origins of modern families). Relationships of the Pelecaniformes were inferred through parsimony analyses of an osteological dataset encompassing 59 taxa and 464 characters. The relationships of the Plotopteridae, an extinct family of wing-propelled divers, and several other fossil pelecaniforms (Limnofregata, Prophaethon, Lithoptila, ?Borvocarbo stoeffelensis) were also assessed. The antiquity of these taxa and their purported status as stem members of extant families makes them valuable for studies of higher-level avian diversification. Methodology/Principal Findings Pelecaniform monophyly is not recovered, with Phaethontidae recovered as distantly related to all other pelecaniforms, which are supported as a monophyletic Steganopodes. Some anatomical partitions of the dataset possess different phylogenetic signals, and partitioned analyses reveal that these discrepancies are localized outside of Steganopodes, and primarily due to a few labile taxa. The Plotopteridae are recovered as the sister taxon to Phalacrocoracoidea, and the relationships of other fossil pelecaniforms representing key calibration points are well supported, including Limnofregata (sister taxon to Fregatidae), Prophaethon and Lithoptila (successive sister taxa to Phaethontidae), and ?Borvocarbo stoeffelensis (sister taxon to Phalacrocoracidae). These relationships are invariant when ‘backbone’ constraints based on recent avian phylogenies are imposed. Conclusions/Significance Relationships of extant pelecaniforms inferred from morphology are more congruent with molecular phylogenies than previously assumed, though notable conflicts remain. The phylogenetic position of the Plotopteridae implies that wing-propelled diving evolved independently in plotopterids and penguins, representing a remarkable case of convergent evolution. Despite robust support for the placement of fossil taxa representing key calibration points, the successive outgroup relationships of several “stem fossil + crown family” clades are variable and poorly supported across recent studies of avian phylogeny. Thus, the impact these fossils have on inferred patterns of temporal diversification depends heavily on the resolution of deep nodes in avian phylogeny. PMID:20976229

The angiotensin-converting enzyme (ACE) gene family of Bombyx mori.

PubMed

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.

Convergent evolution of sexual shape dimorphism in Diptera.

PubMed

Bonduriansky, Russell

2006-05-01

Several patterns of sexual shape dimorphism, such as male body elongation, eye stalks, or extensions of the exoskeleton, have evolved repeatedly in the true flies (Diptera). Although these dimorphisms may have evolved in response to sexual selection on male body shape, conserved genetic factors may have contributed to this convergent evolution, resulting in stronger phenotypic convergence than might be expected from functional requirements alone. I compared phenotypic variation in body shape in two distantly related species exhibiting sexually dimorphic body elongation: Prochyliza xanthostoma (Piophilidae) and Telostylinus angusticollis (Neriidae). Although sexual selection appears to act differently on male body shape in these species, they exhibited strikingly similar patterns of sexual dimorphism. Likewise, patterns of within-sex shape variation were similar in the two species, particularly in males: relative elongation of the male head capsule, antenna, and legs was associated with reduced head capsule width and wing length, but was nearly independent of variation in thorax length. However, the two species presented contrasting patterns of static allometry: male sexual traits exhibited elevated allometric slopes in T. angusticollis, but not in P. xanthostoma. These results suggest that a shared pattern of covariation among traits may have channeled the evolution of sexually dimorphic body elongation in these species. Nonetheless, static allometries may have been shaped by species-specific selection pressures or genetic architectures. Copyright 2006 Wiley-Liss, Inc.

Transport Mechanisms Governing initial Leading-Edge Vortex Development on a Pitching Wing

NASA Astrophysics Data System (ADS)

Wabick, Kevin; Berdon, Randall; Buchholz, James; Johnson, Kyle; Thurow, Brian

2017-11-01

The formation and evolution of Leading Edge Vortices (LEVs) are ubiquitous in natural fliers and maneuvering wings, and have a profound impact on aerodynamic loads. The formation of an LEV is experimentally investigated on a pitching flat-plate wing of aspect-ratio 2, and dimensionless pitch rates of k = Ωc / 2 U of 0.1, 0.2, and 0.5, at a Reynolds number of 104. The sources and sinks of vorticity that contribute to the growth and evolution of the LEV are investigated at spanwise regions of interest, and their relative balance is compared to other wing kinematics, and the case of a two-dimensional pitching wing. This work is supported by the Air Force Office of Scientific Research (Grant Number FA9550-16-1-0107, Dr. Douglas Smith, program manager).

Asymmetric hindwing foldings in rove beetles.

PubMed

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.

Climate variability slows evolutionary responses of Colias butterflies to recent climate change.

PubMed

Kingsolver, Joel G; Buckley, Lauren B

2015-03-07

How does recent climate warming and climate variability alter fitness, phenotypic selection and evolution in natural populations? We combine biophysical, demographic and evolutionary models with recent climate data to address this question for the subalpine and alpine butterfly, Colias meadii, in the southern Rocky Mountains. We focus on predicting patterns of selection and evolution for a key thermoregulatory trait, melanin (solar absorptivity) on the posterior ventral hindwings, which affects patterns of body temperature, flight activity, adult and egg survival, and reproductive success in Colias. Both mean annual summer temperatures and thermal variability within summers have increased during the past 60 years at subalpine and alpine sites. At the subalpine site, predicted directional selection on wing absorptivity has shifted from generally positive (favouring increased wing melanin) to generally negative during the past 60 years, but there is substantial variation among years in the predicted magnitude and direction of selection and the optimal absorptivity. The predicted magnitude of directional selection at the alpine site declined during the past 60 years and varies substantially among years, but selection has generally been positive at this site. Predicted evolutionary responses to mean climate warming at the subalpine site since 1980 is small, because of the variability in selection and asymmetry of the fitness function. At both sites, the predicted effects of adaptive evolution on mean population fitness are much smaller than the fluctuations in mean fitness due to climate variability among years. Our analyses suggest that variation in climate within and among years may strongly limit evolutionary responses of ectotherms to mean climate warming in these habitats. © 2015 The Author(s) Published by the Royal Society. All rights reserved.

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.

Structural or pigmentary? Origin of the distinctive white stripe on the blue wing of a Morpho butterfly.

PubMed

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.

Genomic Sequence around Butterfly Wing Development Genes: Annotation and Comparative Analysis

PubMed Central

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

Character displacement in the fighting colours of Hetaerina damselflies.

PubMed

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.

Did Adult Diurnal Activity Influence the Evolution of Wing Morphology in Opoptera Butterflies?

PubMed

Penz, C M; Heine, K B

2016-02-01

The butterfly genus Opoptera includes eight species, three of which have diurnal habits while the others are crepuscular (the usual activity period for members of the tribe Brassolini). Although never measured in the field, it is presumed that diurnal Opoptera species potentially spend more time flying than their crepuscular relatives. If a shift to diurnal habits potentially leads to a higher level of activity and energy expenditure during flight, then selection should operate on increased aerodynamic and energetic efficiency, leading to changes in wing shape. Accordingly, we ask whether diurnal habits have influenced the evolution of wing morphology in Opoptera. Using phylogenetically independent contrasts and Wilcoxon rank sum tests, we confirmed our expectation that the wings of diurnal species have higher aspect ratios (ARs) and lower wing centroids (WCs) than crepuscular congeners. These wing shape characteristics are known to promote energy efficiency during flight. Three Opoptera wing morphotypes established a priori significantly differed in AR and WC values. The crepuscular, cloud forest dweller Opoptera staudingeri (Godman & Salvin) was exceptional in having an extended forewing tip and the highest AR and lowest WC within Opoptera, possibly to facilitate flight in a cooler environment. Our study is the first to investigate how butterfly wing morphology might evolve as a response to a behavioral shift in adult time of activity.

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.

Adaptive introgression across species boundaries in Heliconius butterflies.

PubMed

Pardo-Diaz, Carolina; Salazar, Camilo; Baxter, Simon W; Merot, Claire; Figueiredo-Ready, Wilsea; Joron, Mathieu; McMillan, W Owen; Jiggins, Chris D

2012-01-01

It is widely documented that hybridisation occurs between many closely related species, but the importance of introgression in adaptive evolution remains unclear, especially in animals. Here, we have examined the role of introgressive hybridisation in transferring adaptations between mimetic Heliconius butterflies, taking advantage of the recent identification of a gene regulating red wing patterns in this genus. By sequencing regions both linked and unlinked to the red colour locus, we found a region that displays an almost perfect genotype by phenotype association across four species, H. melpomene, H. cydno, H. timareta, and H. heurippa. This particular segment is located 70 kb downstream of the red colour specification gene optix, and coalescent analysis indicates repeated introgression of adaptive alleles from H. melpomene into the H. cydno species clade. Our analytical methods complement recent genome scale data for the same region and suggest adaptive introgression has a crucial role in generating adaptive wing colour diversity in this group of butterflies.

Independently Controlled Wing Stroke Patterns in the Fruit Fly Drosophila melanogaster

PubMed Central

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.

PubMed

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.

A wing expressed sequence tag resource for Bicyclus anynana butterflies, an evo-devo model

PubMed Central

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

Color pattern specific proteins in black scales in developing wings ofPrecis coenia Hübner (Nymphalidae, Lepidoptera).

PubMed

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.

Wing Defects in Drosophila xenicid Mutant Clones Are Caused by C-Terminal Deletion of Additional Sex Combs (Asx)

PubMed Central

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

Kinematics and Flow Evolution of a Flexible Wing in Stall Flutter

NASA Astrophysics Data System (ADS)

Farnsworth, John; Akkala, James; Buchholz, James; McLaughlin, Thomas

2014-11-01

Large amplitude stall flutter limit cycle oscillations were observed on an aspect ratio six finite span NACA0018 flexible wing model at a free stream velocity of 23 m/s and an initial angle of attack of six degrees. The wing motion was characterized by periodic oscillations of predominately a torsional mode at a reduced frequency of k = 0.1. The kinematics were quantified via stereoscopic tracking of the wing surface with high speed camera imaging and direct linear transformation. Simultaneously acquired accelerometer measurements were used to track the wing motion and trigger the collection of two-dimensional particle image velocimetry field measurements to the phase angle of the periodic motion. Aerodynamically, the flutter motion is driven by the development and shedding of a dynamic stall vortex system, the evolution of which is characterized and discussed. This work was supported by the AFOSR Flow Interactions and Control Portfolio monitored by Dr. Douglas Smith and the AFOSR/ASEE Summer Faculty Fellowship Program (JA and JB).

Convergent evolution of mechanically optimal locomotion in aquatic invertebrates and vertebrates.

PubMed

Bale, Rahul; Neveln, Izaak D; Bhalla, Amneet Pal Singh; MacIver, Malcolm A; Patankar, Neelesh A

2015-04-01

Examples of animals evolving similar traits despite the absence of that trait in the last common ancestor, such as the wing and camera-type lens eye in vertebrates and invertebrates, are called cases of convergent evolution. Instances of convergent evolution of locomotory patterns that quantitatively agree with the mechanically optimal solution are very rare. Here, we show that, with respect to a very diverse group of aquatic animals, a mechanically optimal method of swimming with elongated fins has evolved independently at least eight times in both vertebrate and invertebrate swimmers across three different phyla. Specifically, if we take the length of an undulation along an animal's fin during swimming and divide it by the mean amplitude of undulations along the fin length, the result is consistently around twenty. We call this value the optimal specific wavelength (OSW). We show that the OSW maximizes the force generated by the body, which also maximizes swimming speed. We hypothesize a mechanical basis for this optimality and suggest reasons for its repeated emergence through evolution.

Comparative population genetics of a mimicry locus among hybridizing Heliconius butterfly species.

PubMed

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.

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.

Exploring Flight Research with Experimental Gliders

NASA Technical Reports Server (NTRS)

1999-01-01

A look at the research aircraft flown by NASA and its predecessor, the National Advisory Committee for Aeronautics (NACA), since the 1940's reveals an evolution of wing designs. In fact, each of the first series of NACA experimental research aircraft ("X-planes") used different wing and tail configurations to tackle the problems of supersonic flight, These early jet aircraft had straight wings (X-1), wings that angled (swept) toward the tail (X-2), triangular (delta) wings (XF-92), and wings that could be moved in flight to change the angle of backward sweep (X-5). Each design added to our knowledge of high-speed flight.

Rapid convergent evolution in wild crickets.

PubMed

Pascoal, Sonia; Cezard, Timothee; Eik-Nes, Aasta; Gharbi, Karim; Majewska, Jagoda; Payne, Elizabeth; Ritchie, Michael G; Zuk, Marlene; Bailey, Nathan W

2014-06-16

The earliest stages of convergent evolution are difficult to observe in the wild, limiting our understanding of the incipient genomic architecture underlying convergent phenotypes. To address this, we capitalized on a novel trait, flatwing, that arose and proliferated at the start of the 21st century in a population of field crickets (Teleogryllus oceanicus) on the Hawaiian island of Kauai. Flatwing erases sound-producing structures on male forewings. Mutant males cannot sing to attract females, but they are protected from fatal attack by an acoustically orienting parasitoid fly (Ormia ochracea). Two years later, the silent morph appeared on the neighboring island of Oahu. We tested two hypotheses for the evolutionary origin of flatwings in Hawaii: (1) that the silent morph originated on Kauai and subsequently introgressed into Oahu and (2) that flatwing originated independently on each island. Morphometric analysis of male wings revealed that Kauai flatwings almost completely lack typical derived structures, whereas Oahu flatwings retain noticeably more wild-type wing venation. Using standard genetic crosses, we confirmed that the mutation segregates as a single-locus, sex-linked Mendelian trait on both islands. However, genome-wide scans using RAD-seq recovered almost completely distinct markers linked with flatwing on each island. The patterns of allelic association with flatwing on either island reveal different genomic architectures consistent with the timing of two mutational events on the X chromosome. Divergent wing morphologies linked to different loci thus cause identical behavioral outcomes--silence--illustrating the power of selection to rapidly shape convergent adaptations from distinct genomic starting points. Copyright © 2014 Elsevier Ltd. All rights reserved.

Genomic Signatures of Speciation in Sympatric and Allopatric Hawaiian Picture-Winged Drosophila.

PubMed

Kang, Lin; Settlage, Robert; McMahon, Wyatt; Michalak, Katarzyna; Tae, Hongseok; Garner, Harold R; Stacy, Elizabeth A; Price, Donald K; Michalak, Pawel

2016-05-30

The Hawaiian archipelago provides a natural arena for understanding adaptive radiation and speciation. The Hawaiian Drosophila are one of the most diverse endemic groups in Hawaiì with up to 1,000 species. We sequenced and analyzed entire genomes of recently diverged species of Hawaiian picture-winged Drosophila, Drosophila silvestris and Drosophila heteroneura from Hawaiì Island, in comparison with Drosophila planitibia, their sister species from Maui, a neighboring island where a common ancestor of all three had likely occurred. Genome-wide single nucleotide polymorphism patterns suggest the more recent origin of D. silvestris and D. heteroneura, as well as a pervasive influence of positive selection on divergence of the three species, with the signatures of positive selection more prominent in sympatry than allopatry. Positively selected genes were significantly enriched for functional terms related to sensory detection and mating, suggesting that sexual selection played an important role in speciation of these species. In particular, sequence variation in Olfactory receptor and Gustatory receptor genes seems to play a major role in adaptive radiation in Hawaiian pictured-winged Drosophila. © The Author 2016. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution.

Local pharmacological effects of tungstate on the color-pattern determination of butterfly wings: a possible relationship between the eyespot and parafocal element.

PubMed

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.

Tungstate-induced color-pattern modifications of butterfly wings are independent of stress response and ecdysteroid effect.

PubMed

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.

Ancient homology underlies adaptive mimetic diversity across butterflies

PubMed Central

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

Mixed signals? Morphological and molecular evidence suggest a color polymorphism in some neotropical polythore damselflies.

PubMed

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.

Mixed Signals? Morphological and Molecular Evidence Suggest a Color Polymorphism in Some Neotropical Polythore Damselflies

PubMed Central

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

Transcriptome profiling with focus on potential key genes for wing development and evolution in Megaloprepus caerulatus, the damselfly species with the world´s largest wings

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...

Artificial evolution of the morphology and kinematics in a flapping-wing mini-UAV.

PubMed

de Margerie, E; Mouret, J B; Doncieux, S; Meyer, J-A

2007-12-01

Birds demonstrate that flapping-wing flight (FWF) is a versatile flight mode, compatible with hovering, forward flight and gliding to save energy. This extended flight domain would be especially useful on mini-UAVs. However, design is challenging because aerodynamic efficiency is conditioned by complex movements of the wings, and because many interactions exist between morphological (wing area, aspect ratio) and kinematic parameters (flapping frequency, stroke amplitude, wing unfolding). Here we used artificial evolution to optimize these morpho-kinematic features on a simulated 1 kg UAV, equipped with wings articulated at the shoulder and wrist. Flight tests were conducted in a dedicated steady aerodynamics simulator. Parameters generating horizontal flight for minimal mechanical power were retained. Results showed that flight at medium speed (10-12 m s(-1)) can be obtained for reasonable mechanical power (20 W kg(-1)), while flight at higher speed (16-20 m s(-1)) implied increased power (30-50 W kg(-1)). Flight at low speed (6-8 m s(-1)) necessitated unrealistic power levels (70-500 W kg(-1)), probably because our simulator neglected unsteady aerodynamics. The underlying adaptation of morphology and kinematics to varying flight speed were compared to available biological data on the flight of birds.

The influence of aspect ratio and stroke pattern on force generation of a bat-inspired membrane wing.

PubMed

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.

PubMed

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

PubMed Central

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

The wings before the bird: an evaluation of flapping-based locomotory hypotheses in bird antecedents.

PubMed

Dececchi, T Alexander; Larsson, Hans C E; Habib, Michael B

2016-01-01

Powered flight is implicated as a major driver for the success of birds. Here we examine the effectiveness of three hypothesized pathways for the evolution of the flight stroke, the forelimb motion that powers aerial locomotion, in a terrestrial setting across a range of stem and basal avians: flap running, Wing Assisted Incline Running (WAIR), and wing-assisted leaping. Using biomechanical mathematical models based on known aerodynamic principals and in vivo experiments and ground truthed using extant avians we seek to test if an incipient flight stroke may have contributed sufficient force to permit flap running, WAIR, or leaping takeoff along the phylogenetic lineage from Coelurosauria to birds. None of these behaviours were found to meet the biomechanical threshold requirements before Paraves. Neither was there a continuous trend of refinement for any of these biomechanical performances across phylogeny nor a signal of universal applicability near the origin of birds. None of these flap-based locomotory models appear to have been a major influence on pre-flight character acquisition such as pennaceous feathers, suggesting non-locomotory behaviours, and less stringent locomotory behaviours such as balancing and braking, played a role in the evolution of the maniraptoran wing and nascent flight stroke. We find no support for widespread prevalence of WAIR in non-avian theropods, but can't reject its presence in large winged, small-bodied taxa like Microraptor and Archaeopteryx. Using our first principles approach we find that "near flight" locomotor behaviors are most sensitive to wing area, and that non-locomotory related selection regimes likely expanded wing area well before WAIR and other such behaviors were possible in derived avians. These results suggest that investigations of the drivers for wing expansion and feather elongation in theropods need not be intrinsically linked to locomotory adaptations, and this separation is critical for our understanding of the origin of powered flight and avian evolution.

The wings before the bird: an evaluation of flapping-based locomotory hypotheses in bird antecedents

PubMed Central

Larsson, Hans C.E.; Habib, Michael B.

2016-01-01

Background: Powered flight is implicated as a major driver for the success of birds. Here we examine the effectiveness of three hypothesized pathways for the evolution of the flight stroke, the forelimb motion that powers aerial locomotion, in a terrestrial setting across a range of stem and basal avians: flap running, Wing Assisted Incline Running (WAIR), and wing-assisted leaping. Methods: Using biomechanical mathematical models based on known aerodynamic principals and in vivo experiments and ground truthed using extant avians we seek to test if an incipient flight stroke may have contributed sufficient force to permit flap running, WAIR, or leaping takeoff along the phylogenetic lineage from Coelurosauria to birds. Results: None of these behaviours were found to meet the biomechanical threshold requirements before Paraves. Neither was there a continuous trend of refinement for any of these biomechanical performances across phylogeny nor a signal of universal applicability near the origin of birds. None of these flap-based locomotory models appear to have been a major influence on pre-flight character acquisition such as pennaceous feathers, suggesting non-locomotory behaviours, and less stringent locomotory behaviours such as balancing and braking, played a role in the evolution of the maniraptoran wing and nascent flight stroke. We find no support for widespread prevalence of WAIR in non-avian theropods, but can’t reject its presence in large winged, small-bodied taxa like Microraptor and Archaeopteryx. Discussion: Using our first principles approach we find that “near flight” locomotor behaviors are most sensitive to wing area, and that non-locomotory related selection regimes likely expanded wing area well before WAIR and other such behaviors were possible in derived avians. These results suggest that investigations of the drivers for wing expansion and feather elongation in theropods need not be intrinsically linked to locomotory adaptations, and this separation is critical for our understanding of the origin of powered flight and avian evolution. PMID:27441115

Spontaneous long-range calcium waves in developing butterfly wings.

PubMed

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.

New specimen of Archaeopteryx provides insights into the evolution of pennaceous feathers.

PubMed

Foth, Christian; Tischlinger, Helmut; Rauhut, Oliver W M

2014-07-03

Discoveries of bird-like theropod dinosaurs and basal avialans in recent decades have helped to put the iconic 'Urvogel' Archaeopteryx into context and have yielded important new data on the origin and early evolution of feathers. However, the biological context under which pennaceous feathers evolved is still debated. Here we describe a new specimen of Archaeopteryx with extensive feather preservation, not only on the wings and tail, but also on the body and legs. The new specimen shows that the entire body was covered in pennaceous feathers, and that the hindlimbs had long, symmetrical feathers along the tibiotarsus but short feathers on the tarsometatarsus. Furthermore, the wing plumage demonstrates that several recent interpretations are problematic. An analysis of the phylogenetic distribution of pennaceous feathers on the tail, hindlimb and arms of advanced maniraptorans and basal avialans strongly indicates that these structures evolved in a functional context other than flight, most probably in relation to display, as suggested by some previous studies. Pennaceous feathers thus represented an exaptation and were later, in several lineages and following different patterns, recruited for aerodynamic functions. This indicates that the origin of flight in avialans was more complex than previously thought and might have involved several convergent achievements of aerial abilities.

Neighboring genes shaping a single adaptive mimetic trait.

PubMed

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.

Development of a cyber physical apparatus for investigating fluid structure interaction on leading edge vortex evolution

NASA Astrophysics Data System (ADS)

Raghu Gowda, Belagumba Venkatachalaiah

This dissertation examines how simple structural compliance impacts a specific transient vortex phenomenon that occurs on high angle of attack lifting surfaces termed dynamic stall. In many Fluid structure interaction (FSI) research efforts, a purely physical or purely computational approach is taken. In this work a low cost cyber-physical (CPFD) system is designed and developed for representing the FSI in the leading edge vortex (LEV) development problem. The leading edge compliance appears to be favorable in a specific spring constant range for a given wing. When the leading edge compliance prescribed via CPFD system is too low compared with the moment due to dynamic pressure or fluid unsteady effect, the LEV behavior is similar to that of a rigid wing system. When the leading edge compliance is too high, excessive compliance is introduced into the wing system and the leading edge vortex evolution is affected by the large change in wing angle. At moderate leading edge compliance, a balance appears to be achieved in which the leading edge vorticity shedding rate supports the long term evolution of the leading edge vortex. Further investigation is required to determine specific parameters governing these leading edge compliance ranges.

Baculovirus-mediated gene transfer in butterfly wings in vivo: an efficient expression system with an anti-gp64 antibody.

PubMed

Dhungel, Bidur; Ohno, Yoshikazu; Matayoshi, Rie; Otaki, Joji M

2013-03-25

Candidate genes for color pattern formation in butterfly wings have been known based on gene expression patterns since the 1990s, but their functions remain elusive due to a lack of a functional assay. Several methods of transferring and expressing a foreign gene in butterfly wings have been reported, but they have suffered from low success rates or low expression levels. Here, we developed a simple, practical method to efficiently deliver and express a foreign gene using baculovirus-mediated gene transfer in butterfly wings in vivo. A recombinant baculovirus containing a gene for green fluorescent protein (GFP) was injected into pupae of the blue pansy butterfly Junonia orithya (Nymphalidae). GFP fluorescence was detected in the pupal wings and other body parts of the injected individuals three to five days post-injection at various degrees of fluorescence. We obtained a high GFP expression rate at relatively high virus titers, but it was associated with pupal death before color pattern formation in wings. To reduce the high mortality rate caused by the baculovirus treatment, we administered an anti-gp64 antibody, which was raised against baculovirus coat protein gp64, to infected pupae after the baculovirus injection. This treatment greatly reduced the mortality rate of the infected pupae. GFP fluorescence was observed in pupal and adult wings and other body parts of the antibody-treated individuals at various degrees of fluorescence. Importantly, we obtained completely developed wings with a normal color pattern, in which fluorescent signals originated directly from scales or the basal membrane after the removal of scales. GFP fluorescence in wing tissues spatially coincided with anti-GFP antibody staining, confirming that the fluorescent signals originated from the expressed GFP molecules. Our baculovirus-mediated gene transfer system with an anti-gp64 antibody is reasonably efficient, and it can be an invaluable tool to transfer, express, and functionally examine foreign genes in butterfly wings and also in other non-model insect systems.

A conserved supergene locus controls colour pattern diversity in Heliconius butterflies.

PubMed

Joron, Mathieu; Papa, Riccardo; Beltrán, Margarita; Chamberlain, Nicola; Mavárez, Jesús; Baxter, Simon; Abanto, Moisés; Bermingham, Eldredge; Humphray, Sean J; Rogers, Jane; Beasley, Helen; Barlow, Karen; ffrench-Constant, Richard H; Mallet, James; McMillan, W Owen; Jiggins, Chris D

2006-10-01

We studied whether similar developmental genetic mechanisms are involved in both convergent and divergent evolution. Mimetic insects are known for their diversity of patterns as well as their remarkable evolutionary convergence, and they have played an important role in controversies over the respective roles of selection and constraints in adaptive evolution. Here we contrast three butterfly species, all classic examples of Müllerian mimicry. We used a genetic linkage map to show that a locus, Yb, which controls the presence of a yellow band in geographic races of Heliconius melpomene, maps precisely to the same location as the locus Cr, which has very similar phenotypic effects in its co-mimic H. erato. Furthermore, the same genomic location acts as a "supergene", determining multiple sympatric morphs in a third species, H. numata. H. numata is a species with a very different phenotypic appearance, whose many forms mimic different unrelated ithomiine butterflies in the genus Melinaea. Other unlinked colour pattern loci map to a homologous linkage group in the co-mimics H. melpomene and H. erato, but they are not involved in mimetic polymorphism in H. numata. Hence, a single region from the multilocus colour pattern architecture of H. melpomene and H. erato appears to have gained control of the entire wing-pattern variability in H. numata, presumably as a result of selection for mimetic "supergene" polymorphism without intermediates. Although we cannot at this stage confirm the homology of the loci segregating in the three species, our results imply that a conserved yet relatively unconstrained mechanism underlying pattern switching can affect mimicry in radically different ways. We also show that adaptive evolution, both convergent and diversifying, can occur by the repeated involvement of the same genomic regions.

Forelimb kinematics and motor patterns of swimming loggerhead sea turtles (Caretta caretta): are motor patterns conserved in the evolution of new locomotor strategies?

PubMed

Rivera, Angela R V; Wyneken, Jeanette; Blob, Richard W

2011-10-01

Novel functions in animals may evolve through changes in morphology, muscle activity or a combination of both. The idea that new functions or behavior can arise solely through changes in structure, without concurrent changes in the patterns of muscle activity that control movement of those structures, has been formalized as the neuromotor conservation hypothesis. In vertebrate locomotor systems, evidence for neuromotor conservation is found across evolutionary transitions in the behavior of terrestrial species, and in evolutionary transitions from terrestrial species to flying species. However, evolutionary transitions in the locomotion of aquatic species have received little comparable study to determine whether changes in morphology and muscle function were coordinated through the evolution of new locomotor behavior. To evaluate the potential for neuromotor conservation in an ancient aquatic system, we quantified forelimb kinematics and muscle activity during swimming in the loggerhead sea turtle, Caretta caretta. Loggerhead forelimbs are hypertrophied into wing-like flippers that produce thrust via dorsoventral forelimb flapping. We compared kinematic and motor patterns from loggerheads with previous data from the red-eared slider, Trachemys scripta, a generalized freshwater species exhibiting unspecialized forelimb morphology and anteroposterior rowing motions during swimming. For some forelimb muscles, comparisons between C. caretta and T. scripta support neuromotor conservation; for example, the coracobrachialis and the latissimus dorsi show similar activation patterns. However, other muscles (deltoideus, pectoralis and triceps) do not show neuromotor conservation; for example, the deltoideus changes dramatically from a limb protractor/elevator in sliders to a joint stabilizer in loggerheads. Thus, during the evolution of flapping in sea turtles, drastic restructuring of the forelimb was accompanied by both conservation and evolutionary novelty in limb motor patterns.

Three-dimensional flow structures and evolution of the leading-edge vortices on a flapping wing.

PubMed

Lu, Yuan; Shen, Gong Xin

2008-04-01

Following the identification and confirmation of the substructures of the leading-edge vortex (LEV) system on flapping wings, it is apparent that the actual LEV structures could be more complex than had been estimated in previous investigations. In this experimental study, we reveal for the first time the detailed three-dimensional (3-D) flow structures and evolution of the LEVs on a flapping wing in the hovering condition at high Reynolds number (Re=1624). This was accomplished by utilizing an electromechanical model dragonfly wing flapping in a water tank (mid-stroke angle of attack=60 degrees) and applying phase-lock based multi-slice digital stereoscopic particle image velocimetry (DSPIV) to measure the target flow fields at three typical stroke phases: at 0.125 T (T=stroke period), when the wing was accelerating; at 0.25 T, when the wing had maximum speed; and at 0.375 T, when the wing was decelerating. The result shows that the LEV system is a collection of four vortical elements: one primary vortex and three minor vortices, instead of a single conical or tube-like vortex as reported or hypothesized in previous studies. These vortical elements are highly time-dependent in structure and show distinct ;stay properties' at different spanwise sections. The spanwise flows are also time-dependent, not only in the velocity magnitude but also in direction.

Digit loss in archosaur evolution and the interplay between selection and constraints.

PubMed

de Bakker, Merijn A G; Fowler, Donald A; den Oude, Kelly; Dondorp, Esther M; Navas, M Carmen Garrido; Horbanczuk, Jaroslaw O; Sire, Jean-Yves; Szczerbińska, Danuta; Richardson, Michael K

2013-08-22

Evolution involves interplay between natural selection and developmental constraints. This is seen, for example, when digits are lost from the limbs during evolution. Extant archosaurs (crocodiles and birds) show several instances of digit loss under different selective regimes, and show limbs with one, two, three, four or the ancestral number of five digits. The 'lost' digits sometimes persist for millions of years as developmental vestiges. Here we examine digit loss in the Nile crocodile and five birds, using markers of three successive stages of digit development. In two independent lineages under different selection, wing digit I and all its markers disappear. In contrast, hindlimb digit V persists in all species sampled, both as cartilage, and as Sox9- expressing precartilage domains, 250 million years after the adult digit disappeared. There is therefore a mismatch between evolution of the embryonic and adult phenotypes. All limbs, regardless of digit number, showed similar expression of sonic hedgehog (Shh). Even in the one-fingered emu wing, expression of posterior genes Hoxd11 and Hoxd12 was conserved, whereas expression of anterior genes Gli3 and Alx4 was not. We suggest that the persistence of digit V in the embryo may reflect constraints, particularly the conserved posterior gene networks associated with the zone of polarizing activity (ZPA). The more rapid and complete disappearance of digit I may reflect its ZPA-independent specification, and hence, weaker developmental constraints. Interacting with these constraints are selection pressures for limb functions such as flying and perching. This model may help to explain the diverse patterns of digit loss in tetrapods. Our study may also help to understand how selection on adults leads to changes in development.

EB Ford revisited: assessing the long-term stability of wing-spot patterns and population genetic structure of the meadow brown butterfly on the Isles of Scilly

PubMed Central

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

The digits of the wing of birds are 1, 2, and 3. A review.

PubMed

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

Color pattern analysis of nymphalid butterfly wings: revision of the nymphalid groundplan.

PubMed

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.

PubMed

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).

Adaptive Introgression across Species Boundaries in Heliconius Butterflies

PubMed Central

Pardo-Diaz, Carolina; Salazar, Camilo; Baxter, Simon W.; Merot, Claire; Figueiredo-Ready, Wilsea; Joron, Mathieu; McMillan, W. Owen; Jiggins, Chris D.

2012-01-01

It is widely documented that hybridisation occurs between many closely related species, but the importance of introgression in adaptive evolution remains unclear, especially in animals. Here, we have examined the role of introgressive hybridisation in transferring adaptations between mimetic Heliconius butterflies, taking advantage of the recent identification of a gene regulating red wing patterns in this genus. By sequencing regions both linked and unlinked to the red colour locus, we found a region that displays an almost perfect genotype by phenotype association across four species, H. melpomene, H. cydno, H. timareta, and H. heurippa. This particular segment is located 70 kb downstream of the red colour specification gene optix, and coalescent analysis indicates repeated introgression of adaptive alleles from H. melpomene into the H. cydno species clade. Our analytical methods complement recent genome scale data for the same region and suggest adaptive introgression has a crucial role in generating adaptive wing colour diversity in this group of butterflies. PMID:22737081

Interplay of cell dynamics and epithelial tension during morphogenesis of the Drosophila pupal wing

PubMed Central

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

Patterning of a compound eye on an extinct dipteran wing.

PubMed

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 ...

Convergent Evolution of Mechanically Optimal Locomotion in Aquatic Invertebrates and Vertebrates

PubMed Central

Bale, Rahul; Neveln, Izaak D.; Bhalla, Amneet Pal Singh

2015-01-01

Examples of animals evolving similar traits despite the absence of that trait in the last common ancestor, such as the wing and camera-type lens eye in vertebrates and invertebrates, are called cases of convergent evolution. Instances of convergent evolution of locomotory patterns that quantitatively agree with the mechanically optimal solution are very rare. Here, we show that, with respect to a very diverse group of aquatic animals, a mechanically optimal method of swimming with elongated fins has evolved independently at least eight times in both vertebrate and invertebrate swimmers across three different phyla. Specifically, if we take the length of an undulation along an animal’s fin during swimming and divide it by the mean amplitude of undulations along the fin length, the result is consistently around twenty. We call this value the optimal specific wavelength (OSW). We show that the OSW maximizes the force generated by the body, which also maximizes swimming speed. We hypothesize a mechanical basis for this optimality and suggest reasons for its repeated emergence through evolution. PMID:25919026

Characterization of the Decision Network for Wing Expansion in Drosophila Using Targeted Expression of the TRPM8 Channel

PubMed Central

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

Keeping the band together: evidence for false boundary disruptive coloration in a butterfly.

PubMed

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.

PubMed

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.

Sexual Dimorphism and Retinal Mosaic Diversification following the Evolution of a Violet Receptor in Butterflies.

PubMed

McCulloch, Kyle J; Yuan, Furong; Zhen, Ying; Aardema, Matthew L; Smith, Gilbert; Llorente-Bousquets, Jorge; Andolfatto, Peter; Briscoe, Adriana D

2017-09-01

Numerous animal lineages have expanded and diversified the opsin-based photoreceptors in their eyes underlying color vision behavior. However, the selective pressures giving rise to new photoreceptors and their spectral tuning remain mostly obscure. Previously, we identified a violet receptor (UV2) that is the result of a UV opsin gene duplication specific to Heliconius butterflies. At the same time the violet receptor evolved, Heliconius evolved UV-yellow coloration on their wings, due to the pigment 3-hydroxykynurenine (3-OHK) and the nanostructure architecture of the scale cells. In order to better understand the selective pressures giving rise to the violet receptor, we characterized opsin expression patterns using immunostaining (14 species) and RNA-Seq (18 species), and reconstructed evolutionary histories of visual traits in five major lineages within Heliconius and one species from the genus Eueides. Opsin expression patterns are hyperdiverse within Heliconius. We identified six unique retinal mosaics and three distinct forms of sexual dimorphism based on ommatidial types within the genus Heliconius. Additionally, phylogenetic analysis revealed independent losses of opsin expression, pseudogenization events, and relaxation of selection on UVRh2 in one lineage. Despite this diversity, the newly evolved violet receptor is retained across most species and sexes surveyed. Discriminability modeling of behaviorally preferred 3-OHK yellow wing coloration suggests that the violet receptor may facilitate Heliconius color vision in the context of conspecific recognition. Our observations give insights into the selective pressures underlying the origins of new visual receptors. © The Author 2017. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.

PARALLEL RACE FORMATION AND THE EVOLUTION OF MIMICRY IN HELICONIUS BUTTERFLIES: A PHYLOGENETIC HYPOTHESIS FROM MITOCHONDRIAL DNA SEQUENCES.

PubMed

Brower, Andrew V Z

1996-02-01

Mimicry has been a fundamental focus of research since the birth of evolutionary biology yet rarely has been studied from a phylogenetic perspective beyond the simple recognition that mimics are not similar due to common descent. The difficulty of finding characters to discern relationships among closely related and convergent taxa has challenged systematists for more than a century. The phenotypic diversity of wing pattens among mimetic Heliconius adds an additional twist to the problem, because single species contain more than a dozen radically different-looking geographical races even though the mimetic advantage is theoretically highest when all individuals within and between species appear the same. Mitochondrial DNA (mtDNA) offers an independent way to address these issues. In this study, Cytochrome Oxidase I and II sequences from multiple, parallel races of Heliconius erato and Heliconius melpomene are examined, to estimate intraspecific phylogeny and gauge sequence divergence and ages of clades among races within each species. Although phenotypes of sympatric races exhibit remarkable concordance between the two species, the mitochondrial cladograms show that the species have not shared a common evolutionary history. H. erato exhibits a basal split between trans- and cis-Andean groups of races, whereas H. melpomene originates in the Guiana Shield. Diverse races in either species appear to have evolved within the last 200,000 yr, and convergent phenotypes have evolved independently within as well as between species. These results contradict prior theories of the evolution of mimicry based on analysis of wing-pattern genetics. © 1996 The Society for the Study of Evolution.

Cellular basis for singing motor pattern generation in the field cricket (Gryllus bimaculatus DeGeer)

PubMed Central

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

Baculovirus-mediated gene transfer in butterfly wings in vivo: an efficient expression system with an anti-gp64 antibody

PubMed Central

2013-01-01

Background Candidate genes for color pattern formation in butterfly wings have been known based on gene expression patterns since the 1990s, but their functions remain elusive due to a lack of a functional assay. Several methods of transferring and expressing a foreign gene in butterfly wings have been reported, but they have suffered from low success rates or low expression levels. Here, we developed a simple, practical method to efficiently deliver and express a foreign gene using baculovirus-mediated gene transfer in butterfly wings in vivo. Results A recombinant baculovirus containing a gene for green fluorescent protein (GFP) was injected into pupae of the blue pansy butterfly Junonia orithya (Nymphalidae). GFP fluorescence was detected in the pupal wings and other body parts of the injected individuals three to five days post-injection at various degrees of fluorescence. We obtained a high GFP expression rate at relatively high virus titers, but it was associated with pupal death before color pattern formation in wings. To reduce the high mortality rate caused by the baculovirus treatment, we administered an anti-gp64 antibody, which was raised against baculovirus coat protein gp64, to infected pupae after the baculovirus injection. This treatment greatly reduced the mortality rate of the infected pupae. GFP fluorescence was observed in pupal and adult wings and other body parts of the antibody-treated individuals at various degrees of fluorescence. Importantly, we obtained completely developed wings with a normal color pattern, in which fluorescent signals originated directly from scales or the basal membrane after the removal of scales. GFP fluorescence in wing tissues spatially coincided with anti-GFP antibody staining, confirming that the fluorescent signals originated from the expressed GFP molecules. Conclusions Our baculovirus-mediated gene transfer system with an anti-gp64 antibody is reasonably efficient, and it can be an invaluable tool to transfer, express, and functionally examine foreign genes in butterfly wings and also in other non-model insect systems. PMID:23522444

Interaction of a trailing vortex with an oscillating wing

NASA Astrophysics Data System (ADS)

McKenna, C.; Fishman, G.; Rockwell, D.

2018-01-01

A technique of particle image velocimetry is employed to characterize the flow structure of a trailing vortex incident upon the tip region of an oscillating wing (plate). The amplitude and velocity of the wing are nearly two orders of magnitude smaller than the wing chord and free stream velocity, respectively. Depending upon the outboard displacement of the incident vortex relative to the wing tip, distinctive patterns of upwash, downwash, and shed vorticity are observed. These patterns are a strong function of the phase of the wing motion during its oscillation cycle. At a given phase, the wing oscillation induces upwash that is reinforced by the upwash of the incident vortex, giving a maximum net upwash. Conversely, when these two origins of upwash counteract, rather than reinforce, one another during the oscillation cycle, the net upwash attains minimum value. Analogous interpretations hold for regions of maximum and minimum net downwash located outboard of the regions of upwash. The magnitude and scale of the vorticity shed from the tip of the wing are directly correlated with the net upwash, which takes different forms related to the outboard displacement of the incident vortex. As the location of the incident vortex is displaced towards the wing tip, both the maximum upwash and the maximum vorticity of the tip vortex initially increase and then decrease. For the limiting case where the incident vortex impinges directly upon the tip of the wing, there is no tip vortex or induced region of upwash. Furthermore, at small values of vortex displacement from the wing tip, the position of the incident vortex varies significantly from its nominal position during the oscillation cycle. All of the foregoing features are interpreted in conjunction with the flow topology in the form of streamlines and critical points, superposed on patterns of vorticity. It is shown that despite the small amplitude of the wing motion, the flow topology is fundamentally different at maximum positive and negative values of the velocity of the wing tip, that is, they are not symmetric.

6. DETAIL OF MASONRY ON SOUTHWEST WING WALL. MASONRY ON ...

Library of Congress Historic Buildings Survey, Historic Engineering Record, Historic Landscapes Survey

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

Comparative cophylogenetics of Australian phabine pigeons and doves (Aves: Columbidae) and their feather lice (Insecta: Phthiraptera)

USGS Publications Warehouse

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.

Comparative cophylogenetics of Australian phabine pigeons and doves (Aves: Columbidae) and their feather lice (Insecta: Phthiraptera).

PubMed

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.

Airplane wing deformation and flight flutter detection method by using three-dimensional speckle image correlation technology.

PubMed

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.

Physiologically induced color-pattern changes in butterfly wings: mechanistic and evolutionary implications.

PubMed

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.

What serial homologs can tell us about the origin of insect wings

PubMed Central

2017-01-01

Although the insect wing is a textbook example of morphological novelty, the origin of insect wings remains a mystery and is regarded as a chief conundrum in biology. Centuries of debates have culminated into two prominent hypotheses: the tergal origin hypothesis and the pleural origin hypothesis. However, between these two hypotheses, there is little consensus in regard to the origin tissue of the wing as well as the evolutionary route from the origin tissue to the functional flight device. Recent evolutionary developmental (evo-devo) studies have shed new light on the origin of insect wings. A key concept in these studies is “serial homology”. In this review, we discuss how the wing serial homologs identified in recent evo-devo studies have provided a new angle through which this century-old conundrum can be explored. We also review what we have learned so far from wing serial homologs and discuss what we can do to go beyond simply identifying wing serial homologs and delve further into the developmental and genetic mechanisms that have facilitated the evolution of insect wings. PMID:28357056

Correlated evolution of host and parasite body size: tests of Harrison's rule using birds and lice.

PubMed

Johnson, Kevin P; Bush, Sarah E; Clayton, Dale H

2005-08-01

Large-bodied species of hosts often harbor large-bodied parasites, a pattern known as Harrison's rule. Harrison's rule has been documented for a variety of animal parasites and herbivorous insects, yet the adaptive basis of the body-size correlation is poorly understood. We used phylogenetically independent methods to test for Harrison's rule across a large assemblage of bird lice (Insecta: Phthiraptera). The analysis revealed a significant relationship between louse and host size, despite considerable variation among taxa. We explored factors underlying this variation by testing Harrison's rule within two groups of feather-specialist lice that share hosts (pigeons and doves). The two groups, wing lice (Columbicola spp.) and body lice (Physconelloidinae spp.), have similar life histories, despite spending much of their time on different feather tracts. Wing lice showed strong support for Harrison's rule, whereas body lice showed no significant correlation with host size. Wing louse size was correlated with wing feather size, which was in turn correlated with overall host size. In contrast, body louse size showed no correlation with body feather size, which also was not correlated with overall host size. The reason why body lice did not fit Harrison's rule may be related to the fact that different species of body lice use different microhabitats within body feathers. More detailed measurements of body feathers may be needed to explore the precise relationship of body louse size to relevant components of feather size. Whatever the reason, Harrison's rule does not hold in body lice, possibly because selection on body size is mediated by community-level interactions between body lice.

The marginal band system in nymphalid butterfly wings.

PubMed

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.

The Aerodynamic Performance of the Houck Configuration Flow Guides

DTIC Science & Technology

2007-06-01

Vortices.............................................................................................13 2.5 Winglets ...associated with the Houck configuration. This includes winglets , biplanes, and joined-wing aircraft. After that the chapter will discuss the evolution...efficiency factor (e = 1 for elliptical wing). 2.5 Winglets A winglet is best described by Jean Chattot’s quote: “ Winglets are aerodynamic components

Leg regeneration stunts wing growth and hinders flight performance in a stick insect (Sipyloidea sipylus).

PubMed

Maginnis, Tara L

2006-07-22

Major morphological structures are sometimes produced not once, but twice. For example, stick insects routinely shed legs to escape a predator or tangled moult, and these legs are subsequently re-grown. Here, I show that in Sipyloidea sipylus, re-growth of a leg during development causes adults to have disproportionately smaller wings and increases wing loading. These morphological consequences of leg regeneration led to significant reductions in several biologically relevant measures of individual flight performance. This previously unrecognized tradeoff between legs and wings reveals the integrated nature of phasmid phenotypes, and I propose how this tradeoff may have shaped phasmid evolution.

The effect of small angle of attack on the laminar-turbulent transition in boundary layer on swept wing at Mach number M=2

NASA Astrophysics Data System (ADS)

Semionov, N. V.; Yermolaev, Yu. G.; Kosinov, A. D.; Semenov, A. N.; Smorodsky, B. V.; Yatskikh, A. A.

2017-10-01

The paper is devoted to an experimental and theoretical study of effect of small angle of attack on disturbances evolution and laminar-turbulent transition in a supersonic boundary layer on swept wing at Mach number M=2. The experiments are conducted at the low nose supersonic wind tunnel T-325 of ITAM. Model is a symmetrical wing with a 45° sweep angle, a 3 percent-thick circular-arc airfoil. The transition location is determined using a hot-wire anemometer. Confirmed monotonous growth of the transition Reynolds numbers with increasing of angle of attack from -2° to 2.5°. The experimental data on the influence of the angle of attack on the disturbances evolution in the supersonic boundary layer on the swept wing model are obtained. Calculations on the effect of small angles of attack on the development of perturbations are made in the framework of the linear theory of stability. A good qualitative correspondence of theoretical and experimental data are obtained.

Genome-wide SNPs reveal fine-scale differentiation among wingless alpine stonefly populations and introgression between winged and wingless forms.

PubMed

Dussex, Nicolas; Chuah, Aaron; Waters, Jonathan M

2016-01-01

Insect flight loss is a repeated phenomenon in alpine habitats, where wing reduction is thought to enhance local recruitment and increase fecundity. One predicted consequence of flight loss is reduced dispersal ability, which should lead to population genetic differentiation and perhaps ultimately to speciation. Using a dataset of 15,123 SNP loci, we present comparative analyses of fine-scale population structure in codistributed Zelandoperla stonefly species, across three parallel altitudinal transects in New Zealand's Rock and Pillar mountain range. We find that winged populations (altitude 200-500 m; Zelandoperla decorata) show no genetic structuring within or among streams, suggesting substantial dispersal mediated by flight. By contrast, wingless populations (Zelandoperla fenestrata; altitude 200-1100 m) exhibit distinct genetic clusters associated with each stream, and additional evidence of isolation by distance within streams. Our data support the hypothesis that wing-loss can initiate diversification in alpine insect populations over small spatial scales. The often deep phylogenetic placement of lowland Z. fenestrata within their stream-specific clades suggests the possibility of independent alpine colonization events for each stream. Additionally, the detection of winged, interspecific hybrid individuals raises the intriguing possibility that a previously flightless lineage could reacquire flight via introgression. © 2015 The Author(s). Evolution © 2015 The Society for the Study of Evolution.

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?

Wing-wake interaction reduces power consumption in insect tandem wings

NASA Astrophysics Data System (ADS)

Lehmann, Fritz-Olaf

Insects are capable of a remarkable diversity of flight techniques. Dragonflies, in particular, are notable for their powerful aerial manoeuvres and endurance during prey catching or territory flights. While most insects such as flies, bees and wasps either reduced their hinds wings or mechanically coupled fore and hind wings, dragonflies have maintained two independent-controlled pairs of wings throughout their evolution. An extraordinary feature of dragonfly wing kinematics is wing phasing, the shift in flapping phase between the fore and hind wing periods. Wing phasing has previously been associated with an increase in thrust production, readiness for manoeuvrability and hunting performance. Recent studies have shown that wing phasing in tandem wings produces a twofold modulation in hind wing lift, but slightly reduces the maximum combined lift of fore and hind wings, compared to two wings flapping in isolation. Despite this disadvantage, however, wing phasing is effective in improving aerodynamic efficiency during flight by the removal of kinetic energy from the wake. Computational analyses demonstrate that this increase in flight efficiency may save up to 22% aerodynamic power expenditure compared to insects flapping only two wings. In terms of engineering, energetic benefits in four-wing flapping are of substantial interest in the field of biomimetic aircraft design, because the performance of man-made air vehicles is often limited by high-power expenditure rather than by lift production. This manuscript provides a summary on power expenditures and aerodynamic efficiency in flapping tandem wings by investigating wing phasing in a dynamically scaled robotic model of a hovering dragonfly.

Wing-wake interaction reduces power consumption in insect tandem wings

NASA Astrophysics Data System (ADS)

Lehmann, Fritz-Olaf

2009-05-01

Insects are capable of a remarkable diversity of flight techniques. Dragonflies, in particular, are notable for their powerful aerial manoeuvres and endurance during prey catching or territory flights. While most insects such as flies, bees and wasps either reduced their hinds wings or mechanically coupled fore and hind wings, dragonflies have maintained two independent-controlled pairs of wings throughout their evolution. An extraordinary feature of dragonfly wing kinematics is wing phasing, the shift in flapping phase between the fore and hind wing periods. Wing phasing has previously been associated with an increase in thrust production, readiness for manoeuvrability and hunting performance. Recent studies have shown that wing phasing in tandem wings produces a twofold modulation in hind wing lift, but slightly reduces the maximum combined lift of fore and hind wings, compared to two wings flapping in isolation. Despite this disadvantage, however, wing phasing is effective in improving aerodynamic efficiency during flight by the removal of kinetic energy from the wake. Computational analyses demonstrate that this increase in flight efficiency may save up to 22% aerodynamic power expenditure compared to insects flapping only two wings. In terms of engineering, energetic benefits in four-wing flapping are of substantial interest in the field of biomimetic aircraft design, because the performance of man-made air vehicles is often limited by high-power expenditure rather than by lift production. This manuscript provides a summary on power expenditures and aerodynamic efficiency in flapping tandem wings by investigating wing phasing in a dynamically scaled robotic model of a hovering dragonfly.

Cellular basis of morphological variation and temperature-related plasticity in Drosophila melanogaster strains with divergent wing shapes.

PubMed

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.

Three-dimensional vortex wake structure of flapping wings in hovering flight.

PubMed

Cheng, Bo; Roll, Jesse; Liu, Yun; Troolin, Daniel R; Deng, Xinyan

2014-02-06

Flapping wings continuously create and send vortices into their wake, while imparting downward momentum into the surrounding fluid. However, experimental studies concerning the details of the three-dimensional vorticity distribution and evolution in the far wake are limited. In this study, the three-dimensional vortex wake structure in both the near and far field of a dynamically scaled flapping wing was investigated experimentally, using volumetric three-component velocimetry. A single wing, with shape and kinematics similar to those of a fruitfly, was examined. The overall result of the wing action is to create an integrated vortex structure consisting of a tip vortex (TV), trailing-edge shear layer (TESL) and leading-edge vortex. The TESL rolls up into a root vortex (RV) as it is shed from the wing, and together with the TV, contracts radially and stretches tangentially in the downstream wake. The downwash is distributed in an arc-shaped region enclosed by the stretched tangential vorticity of the TVs and the RVs. A closed vortex ring structure is not observed in the current study owing to the lack of well-established starting and stopping vortex structures that smoothly connect the TV and RV. An evaluation of the vorticity transport equation shows that both the TV and the RV undergo vortex stretching while convecting downwards: a three-dimensional phenomenon in rotating flows. It also confirms that convection and secondary tilting and stretching effects dominate the evolution of vorticity.

Live Cell Imaging of Butterfly Pupal and Larval Wings In Vivo

PubMed Central

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.

PubMed

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.

Ontogeny of aerial righting and wing flapping in juvenile birds.

PubMed

Evangelista, Dennis; Cam, Sharlene; Huynh, Tony; Krivitskiy, Igor; Dudley, Robert

2014-08-01

Mechanisms of aerial righting in juvenile chukar partridge (Alectoris chukar) were studied from hatching to 14 days-post-hatching (dph). Asymmetric movements of the wings were used from 1 to 8 dph to effect progressively more successful righting behaviour via body roll. Following 8 dph, wing motions transitioned to bilaterally symmetric flapping that yielded aerial righting via nose-down pitch, along with substantial increases in vertical force production during descent. Ontogenetically, the use of such wing motions to effect aerial righting precedes both symmetric flapping and a previously documented behaviour in chukar (i.e. wing-assisted incline running) hypothesized to be relevant to incipient flight evolution in birds. These findings highlight the importance of asymmetric wing activation and controlled aerial manoeuvres during bird development and are potentially relevant to understanding the origins of avian flight. © 2014 The Author(s) Published by the Royal Society. All rights reserved.

Evidence for seasonal patterns in the relative abundance of avian influenza virus subtypes in blue-winged teal (Anas discors)

USGS Publications Warehouse

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.

Parametric Dependence of Initial LEV Behavior on Maneuvering Wings

NASA Astrophysics Data System (ADS)

Berdon, Randall; Wabick, Kevin; Buchholz, James; Johnson, Kyle; Thurow, Brian; University of Iowa Team; Auburn University Team

2017-11-01

A maneuvering rectangular wing of aspect ratio 2 is examined experimentally using dye visualization and PIV to characterize the initial development of the leading-edge vortex (LEV) during a rolling maneuver in a uniform free stream. Understanding the underlying physics during the early evolution of the vortex is important for developing strategies to manipulate vortex evolution. Varying the dimensionless radius of gyration of the wing (Rg/c, where Rg is the radius of gyration and c is the chord) and the advance ratio (J=U/ ΩRg, where U is the free-stream velocity and Ω is the roll rate) affects the structure of the vortex and its propensity to remain attached. The influence of these parameters will be discussed, toward identification of similarity parameters governing vortex development. This work is supported by the Air Force Office of Scientific Research (Grant Number FA9550-16-1-0107, Dr. Douglas Smith, program manager).

Pupal development and pigmentation process of a polka-dotted fruit fly, Drosophila guttifera (Insecta, Diptera).

PubMed

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.

Features of flow around the flying wing model at various attack and slip angle

NASA Astrophysics Data System (ADS)

Pavlenko, A. M.; Zanin, B. Yu.; Katasonov, M. M.

2017-10-01

Experimental study of flow features around aircraft model having "flying wing" form and belonging to the category of small-unmanned aerial vehicleswas carried out. Hot-wire anemometry and flow visualization techniques were used in the investigation to get quantitative data and streamlines pictures ofthe flow near the model surface. Evolution of vortex structures depending on the attack and slip angle was demonstrated. The possibility of flow control and reduction of flow separation zones on the wing surface by means of ledges in the form of cones was also investigated. It was shown, that the laminar-turbulent transition scenario on the flying wing model is identical to the one on a straight wing and occurs through the development of a package of unstable oscillations in the boundary layer separation.

Strong geographical variation in wing aspect ratio of a damselfly, Calopteryx maculata (Odonata: Zygoptera)

PubMed Central

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

Does Skipping a Meal Matter to a Butterfly's Appearance? Effects of Larval Food Stress on Wing Morphology and Color in Monarch Butterflies

PubMed Central

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

Does skipping a meal matter to a butterfly's appearance? Effects of larval food stress on wing morphology and color in monarch butterflies.

PubMed

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.

Coevolution between flight morphology, vertical stratification and sexual dimorphism: what can we learn from tropical butterflies?

PubMed

Graça, M B; Pequeno, P A C L; Franklin, E; Morais, J W

2017-10-01

Occurrence patterns are partly shaped by the affinity of species with habitat conditions. For winged organisms, flight-related attributes are vital for ecological performance. However, due to the different reproductive roles of each sex, we expect divergence in flight energy budget, and consequently different selection responses between sexes. We used tropical frugivorous butterflies as models to investigate coevolution between flight morphology, sex dimorphism and vertical stratification. We studied 94 species of Amazonian fruit-feeding butterflies sampled in seven sites across 3341 ha. We used wing-thorax ratio as a proxy for flight capacity and hierarchical Bayesian modelling to estimate stratum preference. We detected a strong phylogenetic signal in wing-thorax ratio in both sexes. Stouter fast-flying species preferred the canopy, whereas more slender slow-flying species preferred the understorey. However, this relationship was stronger in females than in males, suggesting that female phenotype associates more intimately with habitat conditions. Within species, males were stouter than females and sexual dimorphism was sharper in understorey species. Because trait-habitat relationships were independent from phylogeny, the matching between flight morphology and stratum preference is more likely to reflect adaptive radiation than shared ancestry. This study sheds light on the impact of flight and sexual dimorphism on the evolution and ecological adaptation of flying organisms. © 2017 European Society For Evolutionary Biology. Journal of Evolutionary Biology © 2017 European Society For Evolutionary Biology.

Spectrally tuned structural and pigmentary coloration of birdwing butterfly wing scales.

PubMed

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

PubMed Central

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

Experimental Characterization of Wings for a Hawkmoth-Sized Micro Air Vehicle

DTIC Science & Technology

2014-03-27

131 viii List of Figures Figure Page 2.1 Mechanization of Hawkmoth Thorax . . . . . . . . . . . . . . . . . . . . . . . 7 2.2 Different Insect ...Wing Created by O’Hara . . . . . . . . . . . . . . . . 21 2.15 Evolution of FEA Elements . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 3.1...biological counterparts, birds and insects . Ellington [17] illustrates the differences between these two mechanisms. Insects generally fly under laminar flow

Germline transformation of the butterfly Bicyclus anynana.

PubMed

Marcus, Jeffrey M; Ramos, Diane M; Monteiro, Antónia

2004-08-07

Ecological and evolutionary theory has frequently been inspired by the diversity of colour patterns on the wings of butterflies. More recently, these varied patterns have also become model systems for studying the evolution of developmental mechanisms. A technique that will facilitate our understanding of butterfly colour-pattern development is germline transformation. Germline transformation permits functional tests of candidate gene products and of cis-regulatory regions, and provides a means of generating new colour-pattern mutants by insertional mutagenesis. We report the successful transformation of the African satyrid butterfly Bicyclus anynana with two different transposable element vectors, Hermes and piggyBac, each carrying EGFP coding sequences driven by the 3XP3 synthetic enhancer that drives gene expression in the eyes. Candidate lines identified by screening for EGFP in adult eyes were later confirmed by PCR amplification of a fragment of the EGFP coding sequence from genomic DNA. Flanking DNA surrounding the insertions was amplified by inverse PCR and sequenced. Transformation rates were 5% for piggyBac and 10.2% for Hermes. Ultimately, the new data generated by these techniques may permit an integrated understanding of the developmental genetics of colour-pattern formation and of the ecological and evolutionary processes in which these patterns play a role.

Spatial reflection patterns of iridescent wings of male pierid butterflies: curved scales reflect at a wider angle than flat scales.

PubMed

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.

System-dependent regulations of colour-pattern development: a mutagenesis study of the pale grass blue butterfly

PubMed Central

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

Gliding Swifts Attain Laminar Flow over Rough Wings

PubMed Central

Lentink, David; de Kat, Roeland

2014-01-01

Swifts are among the most aerodynamically refined gliding birds. However, the overlapping vanes and protruding shafts of their primary feathers make swift wings remarkably rough for their size. Wing roughness height is 1–2% of chord length on the upper surface—10,000 times rougher than sailplane wings. Sailplanes depend on extreme wing smoothness to increase the area of laminar flow on the wing surface and minimize drag for extended glides. To understand why the swift does not rely on smooth wings, we used a stethoscope to map laminar flow over preserved wings in a low-turbulence wind tunnel. By combining laminar area, lift, and drag measurements, we show that average area of laminar flow on swift wings is 69% (n = 3; std 13%) of their total area during glides that maximize flight distance and duration—similar to high-performance sailplanes. Our aerodynamic analysis indicates that swifts attain laminar flow over their rough wings because their wing size is comparable to the distance the air travels (after a roughness-induced perturbation) before it transitions from laminar to turbulent. To interpret the function of swift wing roughness, we simulated its effect on smooth model wings using physical models. This manipulation shows that laminar flow is reduced and drag increased at high speeds. At the speeds at which swifts cruise, however, swift-like roughness prolongs laminar flow and reduces drag. This feature gives small birds with rudimentary wings an edge during the evolution of glide performance. PMID:24964089

Cryptically Patterned Moths Perceive Bark Structure When Choosing Body Orientations That Match Wing Color Pattern to the Bark Pattern

PubMed Central

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

Cryptically patterned moths perceive bark structure when choosing body orientations that match wing color pattern to the bark pattern.

PubMed

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.

Waterproof and translucent wings at the same time: problems and solutions in butterflies.

PubMed

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

PubMed Central

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.

PubMed

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.

Nonecholocating fruit bats produce biosonar clicks with their wings.

PubMed

Boonman, Arjan; Bumrungsri, Sara; Yovel, Yossi

2014-12-15

Because evolution mostly acts over millions of years, the intermediate steps leading to a functional sensory system remain enigmatic. Accordingly, there is an ongoing debate regarding the evolution of bat echolocation. In search of the origin of bat echolocation, we studied how Old World fruit bats, which have always been classified as nonecholocating, orient in complete darkness. We found that two of these nonecholocating species used click-like sounds to detect and discriminate objects in complete darkness. However, we discovered that this click-based echo sensing is rudimentary and does not allow these bats to estimate distance accurately as all other echolocating bats can. Moreover, unlike all other echolocating bats, which generate pulses using the larynx or the tongue, these bats generated clicks with their wings. We provide evidence suggesting that all Old World fruit bats can click with their wings. Although this click-based echo sensing used by Old World fruit bats may not represent the ancestral form of current (laryngeal) bat echolocation, we argue that clicking fruit bats could be considered behavioral fossils, opening a window to study the evolution of echolocation. Copyright © 2014 Elsevier Ltd. All rights reserved.

Developments and Validations of Fully Coupled CFD and Practical Vortex Transport Method for High-Fidelity Wake Modeling in Fixed and Rotary Wing Applications

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).

Subtractive Structural Modification of Morpho Butterfly Wings.

PubMed

Shen, Qingchen; He, Jiaqing; Ni, Mengtian; Song, Chengyi; Zhou, Lingye; Hu, Hang; Zhang, Ruoxi; Luo, Zhen; Wang, Ge; Tao, Peng; Deng, Tao; Shang, Wen

2015-11-11

Different from studies of butterfly wings through additive modification, this work for the first time studies the property change of butterfly wings through subtractive modification using oxygen plasma etching. The controlled modification of butterfly wings through such subtractive process results in gradual change of the optical properties, and helps the further understanding of structural optimization through natural evolution. The brilliant color of Morpho butterfly wings is originated from the hierarchical nanostructure on the wing scales. Such nanoarchitecture has attracted a lot of research effort, including the study of its optical properties, its potential use in sensing and infrared imaging, and also the use of such structure as template for the fabrication of high-performance photocatalytic materials. The controlled subtractive processes provide a new path to modify such nanoarchitecture and its optical property. Distinct from previous studies on the optical property of the Morpho wing structure, this study provides additional experimental evidence for the origination of the optical property of the natural butterfly wing scales. The study also offers a facile approach to generate new 3D nanostructures using butterfly wings as the templates and may lead to simpler structure models for large-scale man-made structures than those offered by original butterfly wings. © 2015 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim.

Hovering hummingbird wing aerodynamics during the annual cycle. I. Complete wing.

PubMed

Achache, Yonathan; Sapir, Nir; Elimelech, Yossef

2017-08-01

The diverse hummingbird family (Trochilidae) has unique adaptations for nectarivory, among which is the ability to sustain hover-feeding. As hummingbirds mainly feed while hovering, it is crucial to maintain this ability throughout the annual cycle-especially during flight-feather moult, in which wing area is reduced. To quantify the aerodynamic characteristics and flow mechanisms of a hummingbird wing throughout the annual cycle, time-accurate aerodynamic loads and flow field measurements were correlated over a dynamically scaled wing model of Anna's hummingbird ( Calypte anna ). We present measurements recorded over a model of a complete wing to evaluate the baseline aerodynamic characteristics and flow mechanisms. We found that the vorticity concentration that had developed from the wing's leading-edge differs from the attached vorticity structure that was typically found over insects' wings; firstly, it is more elongated along the wing chord, and secondly, it encounters high levels of fluctuations rather than a steady vortex. Lift characteristics resemble those of insects; however, a 20% increase in the lift-to-torque ratio was obtained for the hummingbird wing model. Time-accurate aerodynamic loads were also used to evaluate the time-evolution of the specific power required from the flight muscles, and the overall wingbeat power requirements nicely matched previous studies.

Unsteady fluid dynamics around a hovering wing

NASA Astrophysics Data System (ADS)

Krishna, Swathi; Green, Melissa; Mulleners, Karen

2017-11-01

The unsteady flow around a hovering flat plate wing has been investigated experimentally using particle image velocimetry and direct force measurements. The measurements are conducted on a wing that rotates symmetrically about the stroke reversal at a reduced frequency of k = 0.32 and Reynolds number of Re = 220 . The Lagrangian finite-time Lyapunov exponent method is used to analyse the unsteady flow fields by identifying dynamically relevant flow features such as the primary leading edge vortex (LEV), secondary vortices, and topological saddles, and their evolution within a flapping cycle. The flow evolution is divided into four stages that are characterised by the LEV (a)emergence, (b)growth, (c)lift-off, and (d)breakdown and decay. Tracking saddle points is shown to be helpful in defining the LEV lift-off which occurs at the maximum stroke velocity. The flow fields are correlated with the aerodynamic forces revealing that the maximum lift and drag are observed just before LEV lift-off. The end of wing rotation in the beginning of the stroke stimulates a change in the direction of the LEV growth and the start of rotation at the end of the stroke triggers the breakdown of the LEV.

Decapentaplegic and growth control in the developing Drosophila wing.

PubMed

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.

Wing Rock Motion and its Flow Mechanism over a Chined-Body Configuration

NASA Astrophysics Data System (ADS)

Wang, Yankui; Li, Qian; Shi, Wei

2015-11-01

Wing rock motion is one kind of uncommanded oscillation around the body axis over the most of the aircraft at enough high angle of attack and has a strong threat to the flight safety. The purpose of this paper is to investigate the wing rock motion over a typical body-wing configuration with a chined fuselage at fixed angle of attack firstly and four kinds of wing rock motion are revealed based on the flow phenomena, namely non-oscillation, lateral deflection, limit-cycle oscillation and irregular oscillation. Simultaneously, some special relationship between the wing rock motion and the flow over the chined body configuration are discussed. In addition, the evolution of wing rock motion and its corresponding flows when the model undergoes pitching up are also given out. All the experiments have been conducted in a low-speed wind tunnel at a Reynolds number of 1.87*10E5 and angle of attack from 0deg to 65deg. National Natural Science Foundation of China(11472028) and Open fund from State Key Laboratory of Aerodynamics.

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.

76 FR 49541 - Endangered and Threatened Wildlife and Plants; Emergency Listing of the Miami Blue Butterfly as...

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...

77 FR 20947 - Endangered and Threatened Wildlife and Plants; Listing of the Miami Blue Butterfly as Endangered...

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...

Beneficial aerodynamic effect of wing scales on the climbing flight of butterflies.

PubMed

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.

Recent Progress on the Stretched Lens Array (SLA)

NASA Technical Reports Server (NTRS)

O'Neill, Markl; McDanal, A. J.; Piszczor, Michael; George, Patrick; Eskenazi, Michael; Botke, Matthew; Edwards, David; Hoppe, David; Brandhorst, Henry

2005-01-01

At the last Space Photovoltaic Research and Technology Conference, SPRAT XVII, held during the fateful week of 9/11/01, our team presented a paper on the early developments related to the new Stretched Lens Array (SLA), including its evolution from the successful SCARLET array on the NASA/JPL Deep Space 1 spacecraft. Within the past two years, the SLA team has made significant progress in the SLA technology, including the successful fabrication and testing of a complete four-panel prototype solar array wing (Fig. 1). The prototype wing verified the mechanical and structural design of the rigid-panel SLA approach, including multiple successful demonstrations of automatic wing deployment. One panel in the prototype wing included four fully functional photovoltaic receivers, employing triple-junction solar cells.

Computation of spanwise distribution of circulation and lift coefficient for flapped wings of arbitrary planform

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.

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.

Hovering hummingbird wing aerodynamics during the annual cycle. II. Implications of wing feather moult

PubMed Central

Sapir, Nir; Elimelech, Yossef

2018-01-01

Birds usually moult their feathers in a particular sequence which may incur aerodynamic, physiological and behavioural implications. Among birds, hummingbirds are unique species in their sustained hovering flight. Because hummingbirds frequently hover-feed, they must maintain sufficiently high flight capacities even when moulting their flight feathers. A hummingbird wing consists of 10 primary flight feathers whose absence during moult may strongly affect wing performance. Using dynamic similarity rules, we compared time-accurate aerodynamic loads and flow field measurements over several wing geometries that follow the natural feather moult sequence of Calypte anna, a common hummingbird species in western North America. Our results suggest a drop of more than 20% in lift production during the early stages of the moult sequence in which mid-wing flight feathers are moulted. We also found that the wing's ability to generate lift strongly depended on the morphological integrity of the outer primaries and leading-edge. These findings may explain the evolution of wing morphology and moult attributes. Specifically, the high overlap between adjacent wing feathers, especially at the wing tip, and the slow sequential replacement of the wing feathers result in a relatively small reduction in wing surface area during moult with limited aerodynamic implications. We present power and efficiency analyses for hover flight during moult under several plausible scenarios, suggesting that body mass reduction could be a compensatory mechanism that preserves the energetic costs of hover flight. PMID:29515884

Hovering hummingbird wing aerodynamics during the annual cycle. II. Implications of wing feather moult.

PubMed

Achache, Yonathan; Sapir, Nir; Elimelech, Yossef

2018-02-01

Birds usually moult their feathers in a particular sequence which may incur aerodynamic, physiological and behavioural implications. Among birds, hummingbirds are unique species in their sustained hovering flight. Because hummingbirds frequently hover-feed, they must maintain sufficiently high flight capacities even when moulting their flight feathers. A hummingbird wing consists of 10 primary flight feathers whose absence during moult may strongly affect wing performance. Using dynamic similarity rules, we compared time-accurate aerodynamic loads and flow field measurements over several wing geometries that follow the natural feather moult sequence of Calypte anna , a common hummingbird species in western North America. Our results suggest a drop of more than 20% in lift production during the early stages of the moult sequence in which mid-wing flight feathers are moulted. We also found that the wing's ability to generate lift strongly depended on the morphological integrity of the outer primaries and leading-edge. These findings may explain the evolution of wing morphology and moult attributes. Specifically, the high overlap between adjacent wing feathers, especially at the wing tip, and the slow sequential replacement of the wing feathers result in a relatively small reduction in wing surface area during moult with limited aerodynamic implications. We present power and efficiency analyses for hover flight during moult under several plausible scenarios, suggesting that body mass reduction could be a compensatory mechanism that preserves the energetic costs of hover flight.

Community interactions govern host-switching with implications for host–parasite coevolutionary history

PubMed Central

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

Technology Innovation of Power Transmission Gearing in Aviation

NASA Technical Reports Server (NTRS)

Handschuh, Robert F.

2009-01-01

An overview of rotary wing evolution and innovations over the last 20 years was presented. This overview is provided from a drive system perspective. Examples of technology innovations that have changed and advanced drive systems of rotary wing vehicles will be provided. These innovations include full 6-axis CNC gear manufacture, face gear development to aerospace standards, health and usage monitoring, and gear geometry and bearing improvements. Also, an overview of current state-of-the-art activities being conducted at NASA Glenn is presented with a short look to fixed and rotary wing aircraft and systems needed for the future.

Chromospheres and mass loss in metal-deficient giant stars

NASA Technical Reports Server (NTRS)

Dupree, A. K.; Hartmann, L.; Avrett, E. H.

1984-01-01

Semiempirical atmospheric models indicate that the characteristic emission in the wings of the H-alpha line observed in Population II giant stars can arise naturally within static chromospheres. Radial expansion gives an asymmetric, blueshifted H-alpha core accompanied by greater emission in the red line wing than in the blue wing. Wind models with extended atmospheres suggest mass loss rates much smaller than 2 x 10 to the -9th solar mass per yr. Thus H-alpha provides no evidence that steady mass loss can significantly affect the evolution of stars on the red giant branch of globular clusters.

Spatial patterns of correlated scale size and scale color in relation to color pattern elements in butterfly wings.

PubMed

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.

Geographic variation in ultraviolet reflectance of the wings of the female cabbage butterfly, Pieris rapae.

PubMed

Obara, Yoshiaki; Ozawa, Gaku; Fukano, Yuya

2008-11-01

The British and Japanese subspecies of the cabbage butterfly, Pieris rapae , differ in terms of the UV reflectance of their wings ( Obara and Majerus, 2000 ). We studied the biogeographical distribution of the female cabbage butterfly having wings with UV reflectance around the Eurasian continent, and between Britain and Japan. For the study, we collected specimens from various locations. A gradient in the UV reflectance of the wings appears to exist along the west-east axis; reflectance was higher toward the east and reached a peak in butterflies in Japan. The UV-reflecting Japanese subspecies Pieris rapae crucivora was found exclusively along the east coast of the Eurasian continent. This suggests that the Japanese subspecies has evolved from a continental ancestor, with females having UV-absorbing wings. We discuss the results of our study with regard to the evolution and adaptive significance of UV coloration in the Japanese subspecies.

Hovering hummingbird wing aerodynamics during the annual cycle. I. Complete wing

PubMed Central

Sapir, Nir; Elimelech, Yossef

2017-01-01

The diverse hummingbird family (Trochilidae) has unique adaptations for nectarivory, among which is the ability to sustain hover-feeding. As hummingbirds mainly feed while hovering, it is crucial to maintain this ability throughout the annual cycle—especially during flight-feather moult, in which wing area is reduced. To quantify the aerodynamic characteristics and flow mechanisms of a hummingbird wing throughout the annual cycle, time-accurate aerodynamic loads and flow field measurements were correlated over a dynamically scaled wing model of Anna’s hummingbird (Calypte anna). We present measurements recorded over a model of a complete wing to evaluate the baseline aerodynamic characteristics and flow mechanisms. We found that the vorticity concentration that had developed from the wing’s leading-edge differs from the attached vorticity structure that was typically found over insects’ wings; firstly, it is more elongated along the wing chord, and secondly, it encounters high levels of fluctuations rather than a steady vortex. Lift characteristics resemble those of insects; however, a 20% increase in the lift-to-torque ratio was obtained for the hummingbird wing model. Time-accurate aerodynamic loads were also used to evaluate the time-evolution of the specific power required from the flight muscles, and the overall wingbeat power requirements nicely matched previous studies. PMID:28878971

Morphological response of songbirds to 100 years of landscape change in North America.

PubMed

Desrochers, A

2010-06-01

Major landscape changes caused by humans may create strong selection pressures and induce rapid evolution in natural populations. In the last 100 years, eastern North America has experienced extensive clear-cutting in boreal areas, while afforestation has occurred in most temperate areas. Based on museum specimens, I show that wings of several boreal forest songbirds and temperate songbirds of non-forest habitats have become more pointed over the last 100 years. In contrast, wings of most temperate forest and early-successional boreal forests species have become less pointed over the same period. In contrast to wing shape, the bill length of most species did not change significantly through time. These results are consistent with the "habitat isolation hypothesis", i.e., songbirds evolved in response to recent changes in the amount of available habitat and associated implications for mobility. Rapid morphological evolution may mitigate, without necessarily preventing, negative consequences of habitat loss caused by humans through direct exploitation or climate change.

Homologization of the flight musculature of zygoptera (insecta: odonata) and neoptera (insecta).

PubMed

Büsse, Sebastian; Genet, Cécile; Hörnschemeyer, Thomas

2013-01-01

Among the winged insects (Pterygota) the Dragonflies and Damselflies (Odonata) are unique for several reasons. Behaviourally they are aerial predators that hunt and catch their prey in flight, only. Morphologically the flight apparatus of Odonata is significantly different from what is found in the remaining Pterygota. However, to understand the phylogenetic relationships of winged insects and the origin and evolution of insect flight in general, it is essential to know how the elements of the odonatan flight apparatus relate to those of the other Pterygota. Here we present a comprehensive, comparative morphological investigation of the thoracic flight musculature of damselflies (Zygoptera). Based on our new data we propose a homologization scheme for the thoracic musculature throughout Pterygota. The new homology hypotheses will allow for future comparative work and especially for phylogenetic analyses using characters of the thoracic musculature throughout all winged insects. This will contribute to understand the early evolution of pterygote insects and their basal phylogenetic relationship.

Homologization of the Flight Musculature of Zygoptera (Insecta: Odonata) and Neoptera (Insecta)

PubMed Central

Büsse, Sebastian; Genet, Cécile; Hörnschemeyer, Thomas

2013-01-01

Among the winged insects (Pterygota) the Dragonflies and Damselflies (Odonata) are unique for several reasons. Behaviourally they are aerial predators that hunt and catch their prey in flight, only. Morphologically the flight apparatus of Odonata is significantly different from what is found in the remaining Pterygota. However, to understand the phylogenetic relationships of winged insects and the origin and evolution of insect flight in general, it is essential to know how the elements of the odonatan flight apparatus relate to those of the other Pterygota. Here we present a comprehensive, comparative morphological investigation of the thoracic flight musculature of damselflies (Zygoptera). Based on our new data we propose a homologization scheme for the thoracic musculature throughout Pterygota. The new homology hypotheses will allow for future comparative work and especially for phylogenetic analyses using characters of the thoracic musculature throughout all winged insects. This will contribute to understand the early evolution of pterygote insects and their basal phylogenetic relationship. PMID:23457479

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

PubMed Central

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.

PubMed

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.

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.

Gene expression underlying adaptive variation in Heliconius wing patterns: non-modular regulation of overlapping cinnabar and vermilion prepatterns.

PubMed

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.

Positional dependence of scale size and shape in butterfly wings: wing-wide phenotypic coordination of color-pattern elements and background.

PubMed

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.

Force generation and wing deformation characteristics of a flapping-wing micro air vehicle 'DelFly II' in hovering flight.

PubMed

Percin, M; van Oudheusden, B W; de Croon, G C H E; Remes, B

2016-05-19

The study investigates the aerodynamic performance and the relation between wing deformation and unsteady force generation of a flapping-wing micro air vehicle in hovering flight configuration. Different experiments were performed where fluid forces were acquired with a force sensor, while the three-dimensional wing deformation was measured with a stereo-vision system. In these measurements, time-resolved power consumption and flapping-wing kinematics were also obtained under both in-air and in-vacuum conditions. Comparison of the results for different flapping frequencies reveals different wing kinematics and deformation characteristics. The high flapping frequency case produces higher forces throughout the complete flapping cycle. Moreover, a phase difference occurs in the variation of the forces, such that the low flapping frequency case precedes the high frequency case. A similar phase lag is observed in the temporal evolution of the wing deformation characteristics, suggesting that there is a direct link between the two phenomena. A considerable camber formation occurs during stroke reversals, which is mainly determined by the stiffener orientation. The wing with the thinner surface membrane displays very similar characteristics to the baseline wing, which implies the dominance of the stiffeners in terms of providing rigidity to the wing. Wing span has a significant effect on the aerodynamic efficiency such that increasing the span length by 4 cm results in a 6% enhancement in the cycle-averaged X-force to power consumption ratio compared to the standard DelFly II wings with a span length of 28 cm.

Biomechanical basis of wing and haltere coordination in flies

PubMed Central

Deora, Tanvi; Singh, Amit Kumar; Sane, Sanjay P.

2015-01-01

The spectacular success and diversification of insects rests critically on two major evolutionary adaptations. First, the evolution of flight, which enhanced the ability of insects to colonize novel ecological habitats, evade predators, or hunt prey; and second, the miniaturization of their body size, which profoundly influenced all aspects of their biology from development to behavior. However, miniaturization imposes steep demands on the flight system because smaller insects must flap their wings at higher frequencies to generate sufficient aerodynamic forces to stay aloft; it also poses challenges to the sensorimotor system because precise control of wing kinematics and body trajectories requires fast sensory feedback. These tradeoffs are best studied in Dipteran flies in which rapid mechanosensory feedback to wing motor system is provided by halteres, reduced hind wings that evolved into gyroscopic sensors. Halteres oscillate at the same frequency as and precisely antiphase to the wings; they detect body rotations during flight, thus providing feedback that is essential for controlling wing motion during aerial maneuvers. Although tight phase synchrony between halteres and wings is essential for providing proper timing cues, the mechanisms underlying this coordination are not well understood. Here, we identify specific mechanical linkages within the thorax that passively mediate both wing–wing and wing–haltere phase synchronization. We demonstrate that the wing hinge must possess a clutch system that enables flies to independently engage or disengage each wing from the mechanically linked thorax. In concert with a previously described gearbox located within the wing hinge, the clutch system enables independent control of each wing. These biomechanical features are essential for flight control in flies. PMID:25605915

Insights into insect wing origin provided by functional analysis of vestigial in the red flour beetle, Tribolium castaneum.

PubMed

Clark-Hachtel, Courtney M; Linz, David M; Tomoyasu, Yoshinori

2013-10-15

Despite accumulating efforts to unveil the origin of insect wings, it remains one of the principal mysteries in evolution. Currently, there are two prominent models regarding insect wing origin: one connecting the origin to the paranotal lobe and the other to the proximodorsal leg branch (exite). However, neither hypothesis has been able to surpass the other. To approach this conundrum, we focused our analysis on vestigial (vg), a critical wing gene initially identified in Drosophila. Our investigation in Tribolium (Coleoptera) has revealed that, despite the well-accepted view of vg as an essential wing gene, there are two groups of vg-dependent tissues in the "wingless" first thoracic segment (T1). We show that one of these tissues, the carinated margin, also depends on other factors essential for wing development (such as Wingless signal and apterous), and has nubbin enhancer activity. In addition, our homeotic mutant analysis shows that wing transformation in T1 originates from both the carinated margin and the other vg-dependent tissue, the pleural structures (trochantin and epimeron). Intriguingly, these two tissues may actually be homologous to the two proposed wing origins (paranotal lobes and exite bearing proximal leg segments). Therefore, our findings suggest that the vg-dependent tissues in T1 could be wing serial homologs present in a more ancestral state, thus providing compelling functional evidence for the dual origin of insect wings.

A Single-Wing Removal Method to Assess Correspondence Between Gene Expression and Phenotype in Butterflies: The Case of Distal-less.

PubMed

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.

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).

Three-dimensional, high-resolution skeletal kinematics of the avian wing and shoulder during ascending flapping flight and uphill flap-running.

PubMed

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).

Subsonic Glideback Rocket Demonstrator Flight Testing

NASA Technical Reports Server (NTRS)

DeTurris, Dianne J.; Foster, Trevor J.; Barthel, Paul E.; Macy, Daniel J.; Droney, Christopher K.; Talay, Theodore A. (Technical Monitor)

2001-01-01

For the past two years, Cal Poly's rocket program has been aggressively exploring the concept of remotely controlled, fixed wing, flyable rocket boosters. This program, embodied by a group of student engineers known as Cal Poly Space Systems, has successfully demonstrated the idea of a rocket design that incorporates a vertical launch pattern followed by a horizontal return flight and landing. Though the design is meant for supersonic flight, CPSS demonstrators are deployed at a subsonic speed. Many steps have been taken by the club that allowed the evolution of the StarBooster prototype to reach its current size: a ten-foot tall, one-foot diameter, composite material rocket. Progress is currently being made that involves multiple boosters along with a second stage, third rocket.

Statistical Properties of Line Centroid Velocity Increments in the rho Ophiuchi Cloud

NASA Technical Reports Server (NTRS)

Lis, D. C.; Keene, Jocelyn; Li, Y.; Phillips, T. G.; Pety, J.

1998-01-01

We present a comparison of histograms of CO (2-1) line centroid velocity increments in the rho Ophiuchi molecular cloud with those computed for spectra synthesized from a three-dimensional, compressible, but non-starforming and non-gravitating hydrodynamic simulation. Histograms of centroid velocity increments in the rho Ophiuchi cloud show clearly non-Gaussian wings, similar to those found in histograms of velocity increments and derivatives in experimental studies of laboratory and atmospheric flows, as well as numerical simulations of turbulence. The magnitude of these wings increases monotonically with decreasing separation, down to the angular resolution of the data. This behavior is consistent with that found in the phase of the simulation which has most of the properties of incompressible turbulence. The time evolution of the magnitude of the non-Gaussian wings in the histograms of centroid velocity increments in the simulation is consistent with the evolution of the vorticity in the flow. However, we cannot exclude the possibility that the wings are associated with the shock interaction regions. Moreover, in an active starforming region like the rho Ophiuchi cloud, the effects of shocks may be more important than in the simulation. However, being able to identify shock interaction regions in the interstellar medium is also important, since numerical simulations show that vorticity is generated in shock interactions.

Complex coevolution of wing, tail, and vocal sounds of courting male bee hummingbirds.

PubMed

Clark, Christopher J; McGuire, Jimmy A; Bonaccorso, Elisa; Berv, Jacob S; Prum, Richard O

2018-03-01

Phenotypic characters with a complex physical basis may have a correspondingly complex evolutionary history. Males in the "bee" hummingbird clade court females with sound from tail-feathers, which flutter during display dives. On a phylogeny of 35 species, flutter sound frequency evolves as a gradual, continuous character on most branches. But on at least six internal branches fall two types of major, saltational changes: mode of flutter changes, or the feather that is the sound source changes, causing frequency to jump from one discrete value to another. In addition to their tail "instruments," males also court females with sound from their syrinx and wing feathers, and may transfer or switch instruments over evolutionary time. In support of this, we found a negative phylogenetic correlation between presence of wing trills and singing. We hypothesize this transference occurs because wing trills and vocal songs serve similar functions and are thus redundant. There are also three independent origins of self-convergence of multiple signals, in which the same species produces both a vocal (sung) frequency sweep, and a highly similar nonvocal sound. Moreover, production of vocal, learned song has been lost repeatedly. Male bee hummingbirds court females with a diverse, coevolving array of acoustic traits. © 2018 The Author(s). Evolution © 2018 The Society for the Study of Evolution.

A wing-assisted running robot and implications for avian flight evolution.

PubMed

Peterson, K; Birkmeyer, P; Dudley, R; Fearing, R S

2011-12-01

DASH+Wings is a small hexapedal winged robot that uses flapping wings to increase its locomotion capabilities. To examine the effects of flapping wings, multiple experimental controls for the same locomotor platform are provided by wing removal, by the use of inertially similar lateral spars, and by passive rather than actively flapping wings. We used accelerometers and high-speed cameras to measure the performance of this hybrid robot in both horizontal running and while ascending inclines. To examine consequences of wing flapping for aerial performance, we measured lift and drag forces on the robot at constant airspeeds and body orientations in a wind tunnel; we also determined equilibrium glide performance in free flight. The addition of flapping wings increased the maximum horizontal running speed from 0.68 to 1.29 m s⁻¹, and also increased the maximum incline angle of ascent from 5.6° to 16.9°. Free flight measurements show a decrease of 10.3° in equilibrium glide slope between the flapping and gliding robot. In air, flapping improved the mean lift:drag ratio of the robot compared to gliding at all measured body orientations and airspeeds. Low-amplitude wing flapping thus provides advantages in both cursorial and aerial locomotion. We note that current support for the diverse theories of avian flight origins derive from limited fossil evidence, the adult behavior of extant flying birds, and developmental stages of already volant taxa. By contrast, addition of wings to a cursorial robot allows direct evaluation of the consequences of wing flapping for locomotor performance in both running and flying.

Habitat variation and wing coloration affect wing shape evolution in dragonflies.

PubMed

Outomuro, D; Dijkstra, K-D B; Johansson, F

2013-09-01

Habitats are spatially and temporally variable, and organisms must be able to track these changes. One potential mechanism for this is dispersal by flight. Therefore, we would expect flying animals to show adaptations in wing shape related to habitat variation. In this work, we explored variation in wing shape in relation to preferred water body (flowing water or standing water with tolerance for temporary conditions) and landscape (forested to open) using 32 species of dragonflies of the genus Trithemis (80% of the known species). We included a potential source of variation linked to sexual selection: the extent of wing coloration on hindwings. We used geometric morphometric methods for studying wing shape. We also explored the phenotypic correlation of wing shape between the sexes. We found that wing shape showed a phylogenetic structure and therefore also ran phylogenetic independent contrasts. After correcting for the phylogenetic effects, we found (i) no significant effect of water body on wing shape; (ii) male forewings and female hindwings differed with regard to landscape, being progressively broader from forested to open habitats; (iii) hindwings showed a wider base in wings with more coloration, especially in males; and (iv) evidence for phenotypic correlation of wing shape between the sexes across species. Hence, our results suggest that natural and sexual selection are acting partially independently on fore- and hindwings and with differences between the sexes, despite evidence for phenotypic correlation of wing shape between males and females. © 2013 The Authors. Journal of Evolutionary Biology © 2013 European Society For Evolutionary Biology.

A Simulation Study of Mutations in the Genetic Regulatory Hierarchy for Butterfly Eyespot Focus Determination

PubMed Central

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

Dpp Signaling Activity Requires Pentagone to Scale with Tissue Size in the Growing Drosophila Wing Imaginal Disc

PubMed Central

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.

Geometric morphometrics analysis of the hind wing of leaf beetles: proximal and distal parts are separate modules.

PubMed

Ren, Jing; Bai, Ming; Yang, Xing-Ke; Zhang, Run-Zhi; Ge, Si-Qin

2017-01-01

The success of beetles is mainly attributed to the possibility to hide the hindwings under the sclerotised elytra. The acquisition of the transverse folding function of the hind wing is an important event in the evolutionary history of beetles. In this study, the morphological and functional variances in the hind wings of 94 leaf beetle species (Coleoptera: Chrysomelinae) is explored using geometric morphometrics based on 36 landmarks. Principal component analysis and Canonical variate analysis indicate that changes of apical area, anal area, and middle area are three useful phylogenetic features at a subtribe level of leaf beetles. Variances of the apical area are the most obvious, which strongly influence the entire venation variance. Partial least squares analysis indicates that the proximal and distal parts of hind wings are weakly associated. Modularity tests confirm that the proximal and distal compartments of hind wings are separate modules. It is deduced that for leaf beetles, or even other beetles, the hind wing possibly exhibits significant functional divergences that occurred during the evolution of transverse folding that resulted in the proximal and distal compartments of hind wings evolving into separate functional modules.

Semi-automated quantitative Drosophila wings measurements.

PubMed

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.

Unique wing scale photonics of male Rajah Brooke's birdwing butterflies.

PubMed

Wilts, Bodo D; Giraldo, Marco A; Stavenga, Doekele G

2016-01-01

Ultrastructures in butterfly wing scales can take many shapes, resulting in the often striking coloration of many butterflies due to interference of light. The plethora of coloration mechanisms is dazzling, but often only single mechanisms are described for specific animals. We have here investigated the male Rajah Brooke's birdwing, Trogonoptera brookiana, a large butterfly from Malaysia, which is marked by striking, colorful wing patterns. The dorsal side is decorated with large, iridescent green patterning, while the ventral side of the wings is primarily brown-black with small white, blue and green patches on the hindwings. Dense arrays of red hairs, creating a distinct collar as well as contrasting areas ventrally around the thorax, enhance the butterfly's beauty. The remarkable coloration is realized by a diverse number of intricate and complicated nanostructures in the hairs as well as the wing scales. The red collar hairs contain a broad-band absorbing pigment as well as UV-reflecting multilayers resembling the photonic structures of Morpho butterflies; the white wing patches consist of scales with prominent thin film reflectors; the blue patches have scales with ridge multilayers and these scales also have centrally concentrated melanin. The green wing areas consist of strongly curved scales, which possess a uniquely arranged photonic structure consisting of multilayers and melanin baffles that produces highly directional reflections. Rajah Brooke's birdwing employs a variety of structural and pigmentary coloration mechanisms to achieve its stunning optical appearance. The intriguing usage of order and disorder in related photonic structures in the butterfly wing scales may inspire novel optical materials as well as investigations into the development of these nanostructures in vivo.

Differential involvement of Hedgehog signaling in butterfly wing and eyespot development.

PubMed

Tong, Xiaoling; Lindemann, Anna; Monteiro, Antónia

2012-01-01

Butterfly eyespots may have evolved from the recruitment of pre-existent gene circuits or regulatory networks into novel locations on the wing. Gene expression data suggests one such circuit, the Hedgehog (Hh) signaling pathway and its target gene engrailed (en), was recruited from a role in patterning the anterior-posterior insect wing axis to a role patterning butterfly eyespots. However, while Junonia coenia expresses hh and en both in the posterior compartment of the wing and in eyespot centers, Bicyclus anynana lacks hh eyespot-specific expression. This suggests that Hh signaling may not be functioning in eyespot development in either species or that it functions in J. coenia but not in B. anynana. In order to test these hypotheses, we performed functional tests of Hh signaling in these species. We investigated the effects of Hh protein sequestration during the larval stage on en expression levels, and on wing size and eyespot size in adults. Hh sequestration led to significantly reduced en expression and to significantly smaller wings and eyespots in both species. But while eyespot size in B. anynana was reduced proportionately to wing size, in J. coenia, eyespots were reduced disproportionately, indicating an independent role of Hh signaling in eyespot development in J. coenia. We conclude that while Hh signaling retains a conserved role in promoting wing growth across nymphalid butterflies, it plays an additional role in eyespot development in some, but not all, lineages of nymphalid butterflies. We discuss our findings in the context of alternative evolutionary scenarios that led to the differential expression of hh and other Hh pathway signaling members across nymphalid species.

Artificial insect wings of diverse morphology for flapping-wing micro air vehicles.

PubMed

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.

The effective compliance of spatially evolving planar wing-cracks

NASA Astrophysics Data System (ADS)

Ayyagari, R. S.; Daphalapurkar, N. P.; Ramesh, K. T.

2018-02-01

We present an analytic closed form solution for anisotropic change in compliance due to the spatial evolution of planar wing-cracks in a material subjected to largely compressive loading. A fully three-dimensional anisotropic compliance tensor is defined and evaluated considering the wing-crack mechanism, using a mixed-approach based on kinematic and energetic arguments to derive the coefficients in incremental compliance. Material, kinematic and kinetic parametric influences on the increments in compliance are studied in order to understand their physical implications on material failure. Model verification is carried out through comparisons to experimental uniaxial compression results to showcase the predictive capabilities of the current study.

Aerodynamic Characteristics of a Feathered Dinosaur Measured Using Physical Models. Effects of Form on Static Stability and Control Effectiveness

PubMed Central

Evangelista, Dennis; Cardona, Griselda; Guenther-Gleason, Eric; Huynh, Tony; Kwong, Austin; Marks, Dylan; Ray, Neil; Tisbe, Adrian; Tse, Kyle; Koehl, Mimi

2014-01-01

We report the effects of posture and morphology on the static aerodynamic stability and control effectiveness of physical models based on the feathered dinosaur, Microraptor gui, from the Cretaceous of China. Postures had similar lift and drag coefficients and were broadly similar when simplified metrics of gliding were considered, but they exhibited different stability characteristics depending on the position of the legs and the presence of feathers on the legs and the tail. Both stability and the function of appendages in generating maneuvering forces and torques changed as the glide angle or angle of attack were changed. These are significant because they represent an aerial environment that may have shifted during the evolution of directed aerial descent and other aerial behaviors. Certain movements were particularly effective (symmetric movements of the wings and tail in pitch, asymmetric wing movements, some tail movements). Other appendages altered their function from creating yaws at high angle of attack to rolls at low angle of attack, or reversed their function entirely. While M. gui lived after Archaeopteryx and likely represents a side experiment with feathered morphology, the general patterns of stability and control effectiveness suggested from the manipulations of forelimb, hindlimb and tail morphology here may help understand the evolution of flight control aerodynamics in vertebrates. Though these results rest on a single specimen, as further fossils with different morphologies are tested, the findings here could be applied in a phylogenetic context to reveal biomechanical constraints on extinct flyers arising from the need to maneuver. PMID:24454820

Helical vortices generated by flapping wings of bumblebees

NASA Astrophysics Data System (ADS)

Farge, Marie; Engels, Thomas; Kolomenskiy, Dmitry; Schneider, Kai; Lehmann, Fritz; Sesterhenn, Jörn

2016-11-01

We analyze high resolution numerical simulation data of a bumblebee with fixed body and prescribed wing motion, flying in a numerical wind tunnel, presented in. The inflow condition of the tunnel varies from unperturbed laminar to strongly turbulent. The flow generated by the flapping wings indicates the important role of the leading edge vortex (LEV), responsible for elevated lift production and which is not significantly altered by the inflow turbulence. The LEV has a conical structure due to the three-dimensional motion of the wings. This flow configuration produces strong vorticity on the sharp leading edge and the outwards velocity (from the root to the tip of the wing) in the spanwise direction. Flow visualizations show that the generated vortical structures are characterized by a strong helicity. We study the evolution of the mean helicity for each wing and analyze the impact of turbulent inflow. We thankfully acknowledge financial support from the French-German AIFIT project funded by DFG and ANR (Grant 15-CE40-0019). DK gratefully acknowledges financial support from the JSPS postdoctoral fellowship.

Polymorphism at a mimicry supergene maintained by opposing frequency-dependent selection pressures.

PubMed

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

PubMed Central

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

Insights into insect wing origin provided by functional analysis of vestigial in the red flour beetle, Tribolium castaneum

PubMed Central

Clark-Hachtel, Courtney M.; Linz, David M.; Tomoyasu, Yoshinori

2013-01-01

Despite accumulating efforts to unveil the origin of insect wings, it remains one of the principal mysteries in evolution. Currently, there are two prominent models regarding insect wing origin: one connecting the origin to the paranotal lobe and the other to the proximodorsal leg branch (exite). However, neither hypothesis has been able to surpass the other. To approach this conundrum, we focused our analysis on vestigial (vg), a critical wing gene initially identified in Drosophila. Our investigation in Tribolium (Coleoptera) has revealed that, despite the well-accepted view of vg as an essential wing gene, there are two groups of vg-dependent tissues in the “wingless” first thoracic segment (T1). We show that one of these tissues, the carinated margin, also depends on other factors essential for wing development (such as Wingless signal and apterous), and has nubbin enhancer activity. In addition, our homeotic mutant analysis shows that wing transformation in T1 originates from both the carinated margin and the other vg-dependent tissue, the pleural structures (trochantin and epimeron). Intriguingly, these two tissues may actually be homologous to the two proposed wing origins (paranotal lobes and exite bearing proximal leg segments). Therefore, our findings suggest that the vg-dependent tissues in T1 could be wing serial homologs present in a more ancestral state, thus providing compelling functional evidence for the dual origin of insect wings. PMID:24085843

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.

Differences in the selection response of serially repeated color pattern characters: standing variation, development, and evolution.

PubMed

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.

Real-Time In Vivo Imaging of Butterfly Wing Development: Revealing the Cellular Dynamics of the Pupal Wing Tissue

PubMed Central

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

Real-time in vivo imaging of butterfly wing development: revealing the cellular dynamics of the pupal wing tissue.

PubMed

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.

Life-history evolution and the microevolution of intermediary metabolism: activities of lipid-metabolizing enzymes in life-history morphs of a wing-dimorphic cricket.

PubMed

Zera, Anthony J; Zhao, Zhangwu

2003-03-01

Although a considerable amount of information is available on the ecology, genetics, and physiology of life-history traits, much more limited data are available on the biochemical and genetic correlates of life-history variation within species. Specific activities of five enzymes of lipid biosynthesis and two enzymes of amino acid catabolism were compared among lines selected for flight-capable (LW[f]) versus flightless (SW) morphs of the cricket Gryllus firmus. These morphs, which exist in natural populations, differ genetically in ovarian growth (100-400% higher in SW) and aspects of flight capability including the size of wings and flight muscles, and the concentration of triglyceride flight fuel (40% greater in LW[f]). Consistently higher activity of each enzyme in LW(f) versus SW-selected lines, and strong co-segregation between morph and enzyme activity, demonstrated genetically based co-variance between wing morph and enzyme activity. Developmental profiles of enzyme activities strongly paralleled profiles of triglyceride accumulation during adulthood and previous measures of in vivo lipid biosynthesis. These data strongly imply that genetically based elevation in activities of lipogenic enzymes, and enzymes controlling the conversion of amino acids into lipids, is an important cause underlying the elevated accumulation of triglyceride in the LW(f) morph, a key biochemical component of the trade-off between elevated early fecundity and flight capability. Global changes in lipid and amino-acid metabolism appear to have resulted from microevolutionary alteration of regulators of metabolism. Finally, strong genotype x environment (diet) interactions were observed for most enzyme activities. Future progress in understanding the functional causes of life-history evolution requires a more detailed synthesis of the fields of life-history evolution and metabolic biochemistry. Wing polymorphism is a powerful experimental model in such integrative studies.

Food load manipulation ability shapes flight morphology in females of central-place foraging Hymenoptera

PubMed Central

2013-01-01

Background Ecological constraints related to foraging are expected to affect the evolution of morphological traits relevant to food capture, manipulation and transport. Females of central-place foraging Hymenoptera vary in their food load manipulation ability. Bees and social wasps modulate the amount of food taken per foraging trip (in terms of e.g. number of pollen grains or parts of prey), while solitary wasps carry exclusively entire prey items. We hypothesized that the foraging constraints acting on females of the latter species, imposed by the upper limit to the load size they are able to transport in flight, should promote the evolution of a greater load-lifting capacity and manoeuvrability, specifically in terms of greater flight muscle to body mass ratio and lower wing loading. Results Our comparative study of 28 species confirms that, accounting for shared ancestry, female flight muscle ratio was significantly higher and wing loading lower in species taking entire prey compared to those that are able to modulate load size. Body mass had no effect on flight muscle ratio, though it strongly and negatively co-varied with wing loading. Across species, flight muscle ratio and wing loading were negatively correlated, suggesting coevolution of these traits. Conclusions Natural selection has led to the coevolution of resource load manipulation ability and morphological traits affecting flying ability with additional loads in females of central-place foraging Hymenoptera. Release from load-carrying constraints related to foraging, which took place with the evolution of food load manipulation ability, has selected against the maintenance of a powerful flight apparatus. This could be the case since investment in flight muscles may have to be traded against other life-history traits, such as reproductive investment. PMID:23805850

The effect of wing flexibility on sound generation of flapping wings.

PubMed

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.

The wing and the eye: a parsimonious theory for scaling and growth control?

PubMed

Romanova-Michaelides, Maria; Aguilar-Hidalgo, Daniel; Jülicher, Frank; Gonzalez-Gaitan, Marcos

2015-01-01

How a developing organ grows and patterns to its final shape is an important question in developmental biology. Studies of growth and patterning in the Drosophila wing imaginal disc have identified a key player, the morphogen Decapentaplegic (Dpp). These studies provided insights into our understanding of growth control and scaling: expansion of the Dpp gradient correlated with the growth of the tissue. A recent report on growth of a Drosophila organ other than the wing, the eye imaginal disc, prompts a reconsideration of our models of growth control. Despite striking differences between the two, the Dpp gradient scales with the target tissues of both organs and the growth of both the wing and the eye is controlled by Dpp. The goal of this review is to discuss whether a parsimonious model of scaling and growth control can explain the relationship between the Dpp gradient and growth in these two different developmental systems. © 2015 Wiley Periodicals, Inc.

Water Tunnel Flow Visualization Study Through Poststall of 12 Novel Planform Shapes

NASA Technical Reports Server (NTRS)

Gatlin, Gregory M.; Neuhart, Dan H.

1996-01-01

To determine the flow field characteristics of 12 planform geometries, a flow visualization investigation was conducted in the Langley 16- by 24-Inch Water Tunnel. Concepts studied included flat plate representations of diamond wings, twin bodies, double wings, cutout wing configurations, and serrated forebodies. The off-surface flow patterns were identified by injecting colored dyes from the model surface into the free-stream flow. These dyes generally were injected so that the localized vortical flow patterns were visualized. Photographs were obtained for angles of attack ranging from 10' to 50', and all investigations were conducted at a test section speed of 0.25 ft per sec. Results from the investigation indicate that the formation of strong vortices on highly swept forebodies can improve poststall lift characteristics; however, the asymmetric bursting of these vortices could produce substantial control problems. A wing cutout was found to significantly alter the position of the forebody vortex on the wing by shifting the vortex inboard. Serrated forebodies were found to effectively generate multiple vortices over the configuration. Vortices from 65' swept forebody serrations tended to roll together, while vortices from 40' swept serrations were more effective in generating additional lift caused by their more independent nature.

A Model for Selection of Eyespots on Butterfly Wings.

PubMed

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.

Dynamics of F-actin prefigure the structure of butterfly wing scales.

PubMed

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.

A simulation study of mutations in the genetic regulatory hierarchy for butterfly eyespot focus determination.

PubMed

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.

Solar powered aircraft

NASA Technical Reports Server (NTRS)

Phillips, W. H. (Inventor)

1983-01-01

A cruciform wing structure for a solar powered aircraft is disclosed. Solar cells are mounted on horizontal wing surfaces. Wing surfaces with spanwise axis perpendicular to surfaces maintain these surfaces normal to the Sun's rays by allowing aircraft to be flown in a controlled pattern at a large bank angle. The solar airplane may be of conventional design with respect to fuselage, propeller and tail, or may be constructed around a core and driven by propeller mechanisms attached near the tips of the airfoils.

Simulation of Cell Patterning Triggered by Cell Death and Differential Adhesion in Drosophila Wing.

PubMed

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.

DOE Office of Scientific and Technical Information (OSTI.GOV)

Huang, J. Y.; E, J. C.; Huang, J. W.

Impact fracture of single-crystal Si is critical to long-term reliability of electronic devices and solar cells for its wide use as components or substrates in semiconductor industry. Single-crystal Si is loaded along two different crystallographic directions with a split Hopkinson pressure bar integrated with an in situ x-ray imaging and diffraction system. Bulk stress histories are measured, simultaneously with x-ray phase contrast imaging (XPCI) and Laue diffraction. Damage evolution is quantified with grayscale maps from XPCI. Single-crystal Si exhibits pronounced anisotropy in fracture modes, and thus fracture strengths and damage evolution. For loading along [11¯ 0] and viewing along [001],more » (1¯1¯0)[11¯ 0] cleavage is activated and induces horizontal primary cracks followed by perpendicular wing cracks. However, for loading along [011¯] and viewing along [111], random nucleation and growth of shear and tensile-splitting crack networks lead to catastrophic failure of materials with no cleavage. The primary-wing crack mode leads to a lower characteristic fracture strength due to predamage, but a more concentrated strength distribution, i.e., a higher Weibull modulus, compared to the second loading case. Furthermore, the sequential primary cracking, wing cracking and wing-crack coalescence processes result in a gradual increase of damage with time, deviating from theoretical predictions. Particle size and aspect ratios of fragments are discussed with postmortem fragment analysis, which verifies fracture modes observed in XPCI.« less

Paleozoic Nymphal Wing Pads Support Dual Model of Insect Wing Origins.

PubMed

Prokop, Jakub; Pecharová, Martina; Nel, André; Hörnschemeyer, Thomas; Krzemińska, Ewa; Krzemiński, Wiesław; Engel, Michael S

2017-01-23

The appearance of wings in insects, early in their evolution [1], has been one of the more critical innovations contributing to their extraordinary diversity. Despite the conspicuousness and importance of wings, the origin of these structures has been difficult to resolve and represented one of the "abominable mysteries" in evolutionary biology [2]. More than a century of debate has boiled the matter down to two competing alternatives-one of wings representing an extension of the thoracic notum, the other stating that they are appendicular derivations from the lateral body wall. Recently, a dual model has been supported by genomic and developmental data [3-6], representing an amalgamation of elements from both the notal and pleural hypotheses. Here, we reveal crucial information from the wing pad joints of Carboniferous palaeodictyopteran insect nymphs using classical and high-tech techniques. These nymphs had three pairs of wing pads that were medially articulated to the thorax but also broadly contiguous with the notum anteriorly and posteriorly (details unobservable in modern insects), supporting their overall origin from the thoracic notum as well as the expected medial, pleural series of axillary sclerites. Our study provides support for the formation of the insect wing from the thoracic notum as well as the already known pleural elements of the arthropodan leg. These results support the unique, dual model for insect wing origins and the convergent reduction of notal fusion in more derived clades, presumably due to wing rotation during development, and they help to bring resolution to this long-standing debate. Copyright © 2017 Elsevier Ltd. All rights reserved.

Rather than resonance, flapping wing flyers may play on aerodynamics to improve performance.

PubMed

Ramananarivo, Sophie; Godoy-Diana, Ramiro; Thiria, Benjamin

2011-04-12

Saving energy and enhancing performance are secular preoccupations shared by both nature and human beings. In animal locomotion, flapping flyers or swimmers rely on the flexibility of their wings or body to passively increase their efficiency using an appropriate cycle of storing and releasing elastic energy. Despite the convergence of many observations pointing out this feature, the underlying mechanisms explaining how the elastic nature of the wings is related to propulsive efficiency remain unclear. Here we use an experiment with a self-propelled simplified insect model allowing to show how wing compliance governs the performance of flapping flyers. Reducing the description of the flapping wing to a forced oscillator model, we pinpoint different nonlinear effects that can account for the observed behavior--in particular a set of cubic nonlinearities coming from the clamped-free beam equation used to model the wing and a quadratic damping term representing the fluid drag associated to the fast flapping motion. In contrast to what has been repeatedly suggested in the literature, we show that flapping flyers optimize their performance not by especially looking for resonance to achieve larger flapping amplitudes with less effort, but by tuning the temporal evolution of the wing shape (i.e., the phase dynamics in the oscillator model) to optimize the aerodynamics.

Rather than resonance, flapping wing flyers may play on aerodynamics to improve performance

PubMed Central

Ramananarivo, Sophie; Godoy-Diana, Ramiro; Thiria, Benjamin

2011-01-01

Saving energy and enhancing performance are secular preoccupations shared by both nature and human beings. In animal locomotion, flapping flyers or swimmers rely on the flexibility of their wings or body to passively increase their efficiency using an appropriate cycle of storing and releasing elastic energy. Despite the convergence of many observations pointing out this feature, the underlying mechanisms explaining how the elastic nature of the wings is related to propulsive efficiency remain unclear. Here we use an experiment with a self-propelled simplified insect model allowing to show how wing compliance governs the performance of flapping flyers. Reducing the description of the flapping wing to a forced oscillator model, we pinpoint different nonlinear effects that can account for the observed behavior—in particular a set of cubic nonlinearities coming from the clamped-free beam equation used to model the wing and a quadratic damping term representing the fluid drag associated to the fast flapping motion. In contrast to what has been repeatedly suggested in the literature, we show that flapping flyers optimize their performance not by especially looking for resonance to achieve larger flapping amplitudes with less effort, but by tuning the temporal evolution of the wing shape (i.e., the phase dynamics in the oscillator model) to optimize the aerodynamics. PMID:21444774

Inspiration for wing design: how forelimb specialization enables active flight in modern vertebrates

PubMed Central

2017-01-01

Harnessing flight strategies refined by millions of years of evolution can help expedite the design of more efficient, manoeuvrable and robust flying robots. This review synthesizes recent advances and highlights remaining gaps in our understanding of how bird and bat wing adaptations enable effective flight. Included in this discussion is an evaluation of how current robotic analogues measure up to their biological sources of inspiration. Studies of vertebrate wings have revealed skeletal systems well suited for enduring the loads required during flight, but the mechanisms that drive coordinated motions between bones and connected integuments remain ill-described. Similarly, vertebrate flight muscles have adapted to sustain increased wing loading, but a lack of in vivo studies limits our understanding of specific muscular functions. Forelimb adaptations diverge at the integument level, but both bird feathers and bat membranes yield aerodynamic surfaces with a level of robustness unparalleled by engineered wings. These morphological adaptations enable a diverse range of kinematics tuned for different flight speeds and manoeuvres. By integrating vertebrate flight specializations—particularly those that enable greater robustness and adaptability—into the design and control of robotic wings, engineers can begin narrowing the wide margin that currently exists between flying robots and vertebrates. In turn, these robotic wings can help biologists create experiments that would be impossible in vivo. PMID:28592663

Transitional fossil earwigs--a missing link in Dermaptera evolution.

PubMed

Zhao, Jingxia; Zhao, Yunyun; Shih, Chungkun; Ren, Dong; Wang, Yongjie

2010-11-10

The Dermaptera belongs to a group of winged insects of uncertain relationship within Polyneoptera, which has expanded anal region and adds numerous anal veins in the hind wing. Evolutional history and origin of Dermaptera have been in contention. In this paper, we report two new fossil earwigs in a new family of Bellodermatidae fam. nov. The fossils were collected from the Jiulongshan Formation (Middle Jurassic) in Inner Mongolia, northeast China. This new family, characterized by an unexpected combination of primitive and derived characters, is bridging the missing link between suborders of Archidermaptera and Eodermaptera. Phylogenetic analyses support the new family to be a new clade at the base of previously defined Eodermaptera and to be a stem group of (Eodermaptera+Neodermaptera). Evolutional history and origin of Dermaptera have been in contention, with dramatically different viewpoints by contemporary authors. It is suggested that the oldest Dermaptera might possibly be traced back to the Late Triassic-Early Jurassic and they had divided into Archidermaptera and (Eodermaptera+Neodermaptera) in the Middle Jurassic.

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.

Cicada-Wing-Inspired Self-Cleaning Antireflection Coatings on Polymer Substrates.

PubMed

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)

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.

Free flight odor tracking in Drosophila: Effect of wing chemosensors, sex and pheromonal gene regulation

PubMed Central

Houot, Benjamin; Gigot, Vincent; Robichon, Alain; Ferveur, Jean-François

2017-01-01

The evolution of powered flight in insects had major consequences for global biodiversity and involved the acquisition of adaptive processes allowing individuals to disperse to new ecological niches. Flies use both vision and olfactory input from their antennae to guide their flight; chemosensors on fly wings have been described, but their function remains mysterious. We studied Drosophila flight in a wind tunnel. By genetically manipulating wing chemosensors, we show that these structures play an essential role in flight performance with a sex-specific effect. Pheromonal systems are also involved in Drosophila flight guidance: transgenic expression of the pheromone production and detection gene, desat1, produced low, rapid flight that was absent in control flies. Our study suggests that the sex-specific modulation of free-flight odor tracking depends on gene expression in various fly tissues including wings and pheromonal-related tissues. PMID:28067325

Forward-swept wing configuration designed for high maneuverability by use of a transonic computational method

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.

Recent Loads Calibration Experience With a Delta Wing Airplane

NASA Technical Reports Server (NTRS)

Jenkins, Jerald M.; Kuhl, Albert E.

1977-01-01

Aircraft which are designed for supersonic and hypersonic flight are evolving with delta wing configurations. An integral part of the evolution of all new aircraft is the flight test phase. Included in the flight test phase is an effort to identify and evaluate the loads environment of the aircraft. The most effective way of examining the loads environment is to utilize calibrated strain gages to provide load magnitudes. Using strain gage data to accomplish this has turned out to be anything but a straightforward task. The delta wing configuration has turned out to be a very difficult type of wing structure to calibrate. Elevated structural temperatures result in thermal effects which contaminate strain gage data being used to deduce flight loads. The concept of thermally calibrating a strain gage system is an approach to solving this problem. This paper will address how these problems were approached on a program directed toward measuring loads on the wing of a large, flexible supersonic aircraft. Structural configurations typical of high-speed delta wing aircraft will be examined. The temperature environment will be examined to see how it induces thermal stresses which subsequently cause errors in loads equations used to deduce the flight loads.

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.

The Morphological Characterization of the Forewing of the Manduca sexta Species for the Application of Biomimetic Flapping Wing Micro Air Vehicles

DTIC Science & Technology

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

How the pterosaur got its wings.

PubMed

Tokita, Masayoshi

2015-11-01

Throughout the evolutionary history of life, only three vertebrate lineages took to the air by acquiring a body plan suitable for powered flight: birds, bats, and pterosaurs. Because pterosaurs were the earliest vertebrate lineage capable of powered flight and included the largest volant animal in the history of the earth, understanding how they evolved their flight apparatus, the wing, is an important issue in evolutionary biology. Herein, I speculate on the potential basis of pterosaur wing evolution using recent advances in the developmental biology of flying and non-flying vertebrates. The most significant morphological features of pterosaur wings are: (i) a disproportionately elongated fourth finger, and (ii) a wing membrane called the brachiopatagium, which stretches from the posterior surface of the arm and elongated fourth finger to the anterior surface of the leg. At limb-forming stages of pterosaur embryos, the zone of polarizing activity (ZPA) cells, from which the fourth finger eventually differentiates, could up-regulate, restrict, and prolong expression of 5'-located Homeobox D (Hoxd) genes (e.g. Hoxd11, Hoxd12, and Hoxd13) around the ZPA through pterosaur-specific exploitation of sonic hedgehog (SHH) signalling. 5'Hoxd genes could then influence downstream bone morphogenetic protein (BMP) signalling to facilitate chondrocyte proliferation in long bones. Potential expression of Fgf10 and Tbx3 in the primordium of the brachiopatagium formed posterior to the forelimb bud might also facilitate elongation of the phalanges of the fourth finger. To establish the flight-adapted musculoskeletal morphology shared by all volant vertebrates, pterosaurs probably underwent regulatory changes in the expression of genes controlling forelimb and pectoral girdle musculoskeletal development (e.g. Tbx5), as well as certain changes in the mode of cell-cell interactions between muscular and connective tissues in the early phase of their evolution. Developmental data now accumulating for extant vertebrate taxa could be helpful in understanding the cellular and molecular mechanisms of body-plan evolution in extinct vertebrates as well as extant vertebrates with unique morphology whose embryonic materials are hard to obtain. © 2014 The Author. Biological Reviews © 2014 Cambridge Philosophical Society.

Galactic cold cores. VII. Filament formation and evolution: Methods and observational constraints

NASA Astrophysics Data System (ADS)

Rivera-Ingraham, A.; Ristorcelli, I.; Juvela, M.; Montillaud, J.; Men'shchikov, A.; Malinen, J.; Pelkonen, V.-M.; Marston, A.; Martin, P. G.; Pagani, L.; Paladini, R.; Paradis, D.; Ysard, N.; Ward-Thompson, D.; Bernard, J.-P.; Marshall, D. J.; Montier, L.; Tóth, L. V.

2016-06-01

Context. The association of filaments with protostellar objects has made these structures a priority target in star formation studies. However, little is known about the link between filament properties and their local environment. Aims: The datasets from the Herschel Galactic Cold cores key programme allow for a statistical study of filaments with a wide range of intrinsic and environmental characteristics. Characterisation of this sample can therefore be used to identify key physical parameters and quantify the role of the environment in the formation of supercritical filaments. These results are necessary to constrain theoretical models of filament formation and evolution. Methods: Filaments were extracted from fields at distance D< 500 pc with the getfilaments algorithm and characterised according to their column density profiles and intrinsic properties. Each profile was fitted with a beam-convolved Plummer-like function, and the filament structure was quantified based on the relative contributions from the filament "core", represented by a Gaussian, and "wing" component, dominated by the power-law behaviour of the Plummer-like function. These filament parameters were examined for populations associated with different background levels. Results: Filaments increase their core (Mline,core) and wing (Mline,wing) contributions while increasing their total linear mass density (Mline,tot). Both components appear to be linked to the local environment, with filaments in higher backgrounds having systematically more massive Mline,core and Mline,wing. This dependence on the environment supports an accretion-based model of filament evolution in the local neighbourhood (D ≤ 500 pc). Structures located in the highest backgrounds develop the highest central AV, Mline,core, and Mline,wing as Mline,tot increases with time, favoured by the local availability of material and the enhanced gravitational potential. Our results indicate that filaments acquiring a significantly massive central region with Mline,core≳Mcrit/2 may become supercritical and form stars. This translates into a need for filaments to become at least moderately self-gravitating to undergo localised star formation or become star-forming filaments. Herschel is an ESA space observatory with science instruments provided by European-led Principal Investigator consortia and with important participation from NASA.

Experimental and numerical studies of beetle-inspired flapping wing in hovering flight.

PubMed

Van Truong, Tien; Le, Tuyen Quang; Park, Hoon Cheol; Byun, Doyoung

2017-05-17

In this paper, we measure unsteady forces and visualize 3D vortices around a beetle-like flapping wing model in hovering flight by experiment and numerical simulation. The measurement of unsteady forces and flow patterns around the wing were conducted using a dynamically scaled wing model in the mineral-oil tank. The wing kinematics were directly derived from the experiment of a real beetle. The 3D flow structures of the flapping wing were captured by using air bubble visualization while forces were measured by a sensor attached at the wing base. In comparison, the size and topology of spiral leading edge vortex, trailing edge vortex and tip vortex are well matched from experimental and numerical studies. In addition, the time history of forces calculated from numerical simulation is also similar to that from theforce measurement. A difference of average force is in order of 10 percent. The results indicate that the leading edge vortex due to rotational acceleration at the end of the stroke during flapping wing causes significant reduction of lift. The present study provides useful information on hover flight to develop a beetle-like flapping wing Micro Air Vehicle.

Global-Local Analysis and Optimization of a Composite Civil Tilt-Rotor Wing

NASA Technical Reports Server (NTRS)

Rais-Rohani, Masound

1999-01-01

This report gives highlights of an investigation on the design and optimization of a thin composite wing box structure for a civil tilt-rotor aircraft. Two different concepts are considered for the cantilever wing: (a) a thin monolithic skin design, and (b) a thick sandwich skin design. Each concept is examined with three different skin ply patterns based on various combinations of 0, +/-45, and 90 degree plies. The global-local technique is used in the analysis and optimization of the six design models. The global analysis is based on a finite element model of the wing-pylon configuration while the local analysis uses a uniformly supported plate representing a wing panel. Design allowables include those on vibration frequencies, panel buckling, and material strength. The design optimization problem is formulated as one of minimizing the structural weight subject to strength, stiffness, and d,vnamic constraints. Six different loading conditions based on three different flight modes are considered in the design optimization. The results of this investigation reveal that of all the loading conditions the one corresponding to the rolling pull-out in the airplane mode is the most stringent. Also the frequency constraints are found to drive the skin thickness limits, rendering the buckling constraints inactive. The optimum skin ply pattern for the monolithic skin concept is found to be (((0/+/-45/90/(0/90)(sub 2))(sub s))(sub s), while for the sandwich skin concept the optimal ply pattern is found to be ((0/+/-45/90)(sub 2s))(sub s).

Genome sequence of a novel H14N7 subtype influenza A virus isolated from a blue-winged teal (Anas discors) harvested in Texas, USA

USGS Publications Warehouse

Ramey, Andy M.; Reeves, Andrew; Poulson, Rebecca L.; Carter, Deborah L.; Davis-Fields, Nicholas; Stallknecht, David E.

2016-01-01

We report here the complete genome sequence of a novel H14N7 subtype influenza A virus (IAV) isolated from a blue-winged teal (Anas discors) harvested in Texas, USA. The genomic characteristics of this IAV strain with a previously undetected subtype combination suggest recent viral evolution within the New World wild-bird IAV reservoir.                   

Sex chromosome linked genetic variance and the evolution of sexual dimorphism of quantitative traits.

PubMed

Husby, Arild; Schielzeth, Holger; Forstmeier, Wolfgang; Gustafsson, Lars; Qvarnström, Anna

2013-03-01

Theory predicts that sex chromsome linkage should reduce intersexual genetic correlations thereby allowing the evolution of sexual dimorphism. Empirical evidence for sex linkage has come largely from crosses and few studies have examined how sexual dimorphism and sex linkage are related within outbred populations. Here, we use data on an array of different traits measured on over 10,000 individuals from two pedigreed populations of birds (collared flycatcher and zebra finch) to estimate the amount of sex-linked genetic variance (h(2)z ). Of 17 traits examined, eight showed a nonzero h(2)Z estimate but only four were significantly different from zero (wing patch size and tarsus length in collared flycatchers, wing length and beak color in zebra finches). We further tested how sexual dimorphism and the mode of selection operating on the trait relate to the proportion of sex-linked genetic variance. Sexually selected traits did not show higher h(2)Z than morphological traits and there was only a weak positive relationship between h(2)Z and sexual dimorphism. However, given the relative scarcity of empirical studies, it is premature to make conclusions about the role of sex chromosome linkage in the evolution of sexual dimorphism. © 2012 The Author(s). Evolution© 2012 The Society for the Study of Evolution.

SN2012ab: a peculiar Type IIn supernova with aspherical circumstellar material

NASA Astrophysics Data System (ADS)

Bilinski, Christopher; Smith, Nathan; Williams, G. Grant; Smith, Paul; Zheng, WeiKang; Graham, Melissa L.; Mauerhan, Jon C.; Andrews, Jennifer E.; Filippenko, Alexei V.; Akerlof, Carl; Chatzopoulos, E.; Hoffman, Jennifer L.; Huk, Leah; Leonard, Douglas C.; Marion, G. H.; Milne, Peter; Quimby, Robert M.; Silverman, Jeffrey M.; Vinkó, Jozsef; Wheeler, J. Craig; Yuan, Fang

2018-03-01

We present photometry, spectra, and spectropolarimetry of supernova (SN) 2012ab, mostly obtained over the course of ˜300 d after discovery. SN 2012ab was a Type IIn (SN IIn) event discovered near the nucleus of spiral galaxy 2MASXJ12224762+0536247. While its light curve resembles that of SN 1998S, its spectral evolution does not. We see indications of CSM interaction in the strong intermediate-width emission features, the high luminosity (peak at absolute magnitude M = -19.5), and the lack of broad absorption features in the spectrum. The Hα emission undergoes a peculiar transition. At early times it shows a broad blue emission wing out to -14 000 km s-1 and a truncated red wing. Then at late times (>100 d) it shows a truncated blue wing and a very broad red emission wing out to roughly +20 000 km s-1. This late-time broad red wing probably arises in the reverse shock. Spectra also show an asymmetric intermediate-width Hα component with stronger emission on the red side at late times. The evolution of the asymmetric profiles requires a density structure in the distant CSM that is highly aspherical. Our spectropolarimetric data also suggest asphericity with a strong continuum polarization of ˜1-3 per cent and depolarization in the Hα line, indicating asphericity in the CSM at a level comparable to that in other SNe IIn. We estimate a mass-loss rate of \\dot{M} = 0.050 M_{⊙} yr^{-1} for vpre = 100 km s-1 extending back at least 75 yr prior to the SN. The strong departure from axisymmetry in the CSM of SN 2012ab may suggest that the progenitor was an eccentric binary system undergoing eruptive mass loss.

Structural dynamics and aerodynamics measurements of biologically inspired flexible flapping wings.

PubMed

Wu, P; Stanford, B K; Sällström, E; Ukeiley, L; Ifju, P G

2011-03-01

Flapping wing flight as seen in hummingbirds and insects poses an interesting unsteady aerodynamic problem: coupling of wing kinematics, structural dynamics and aerodynamics. There have been numerous studies on the kinematics and aerodynamics in both experimental and computational cases with both natural and artificial wings. These studies tend to ignore wing flexibility; however, observation in nature affirms that passive wing deformation is predominant and may be crucial to the aerodynamic performance. This paper presents a multidisciplinary experimental endeavor in correlating a flapping micro air vehicle wing's aeroelasticity and thrust production, by quantifying and comparing overall thrust, structural deformation and airflow of six pairs of hummingbird-shaped membrane wings of different properties. The results show that for a specific spatial distribution of flexibility, there is an effective frequency range in thrust production. The wing deformation at the thrust-productive frequencies indicates the importance of flexibility: both bending and twisting motion can interact with aerodynamic loads to enhance wing performance under certain conditions, such as the deformation phase and amplitude. By measuring structural deformations under the same aerodynamic conditions, beneficial effects of passive wing deformation can be observed from the visualized airflow and averaged thrust. The measurements and their presentation enable observation and understanding of the required structural properties for a thrust effective flapping wing. The intended passive responses of the different wings follow a particular pattern in correlation to their aerodynamic performance. Consequently, both the experimental technique and data analysis method can lead to further studies to determine the design principles for micro air vehicle flapping wings.

Localised JAK/STAT Pathway Activation Is Required for Drosophila Wing Hinge Development

PubMed Central

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

Leading-edge flow reattachment and the lateral static stability of low-aspect-ratio rectangular wings

NASA Astrophysics Data System (ADS)

Linehan, Thomas; Mohseni, Kamran

2017-11-01

The relationship between lateral static stability derivative, Clβ,lift coefficient, CL, and angle of attack was investigated for rectangular wings of aspect ratio A R =0.75 ,1 ,1.5 , and 3 using Stereo-Digital Particle Image Velocimetry (S-DPIV) and direct force and moment measurements. When the product Cl βA R is plotted with respect to CL, the lateral stability curves of each wing collapse to a single line for CL<0.7 . For CL>0.7 , the linearity and scaling of Clβwith respect to CL is lost. S-DPIV is used to elucidate the flow physics in this nonlinear regime. At α =10∘ , the leading-edge separation region emerges on the leeward portion of the sideslipped wing by means of vortex shedding. For the A R ≤1.5 wings at α >15∘ , the tip vortex downwash is sufficient to restrict the shedding of leading-edge vorticity thereby sustaining the lift of the leading-edge separation region at high angles of attack. Concurrently, the windward tip vortex grows in size and strength with increasing angle of attack, displacing the leading-edge separation region further toward the leeward wing. This reorganization of lift-generating vorticity results in the initial nonlinearities between Cl β and CL at angles of attack for which CL is still increasing. At angles of attack near that of maximum lift for the A R ≤1 wings, the windward tip vortex lifts off the wing, decreasing the lateral static stability of the wing prior to lift stall. For the A R =3 wing at α >10∘ , nonlinear trends in Cl β versus CL occur due to the spanwise evolution of stalled flow.

The evolution of avian wing shape and previously unrecognized trends in covert feathering

PubMed Central

Wang, Xia; Clarke, Julia A.

2015-01-01

Avian wing shape has been related to flight performance, migration, foraging behaviour and display. Historically, linear measurements of the feathered aerofoil and skeletal proportions have been used to describe this shape. While the distribution of covert feathers, layered over the anterior wing, has long been assumed to contribute to aerofoil properties, to our knowledge no previous studies of trends in avian wing shape assessed their variation. Here, these trends are explored using a geometric–morphometric approach with landmarks describing the wing outline as well as the extent of dorsal and ventral covert feathers for 105 avian species. We find that most of the observed variation is explained by phylogeny and ecology but shows only a weak relationship with previously described flight style categories, wing loading and an investigated set of aerodynamic variables. Most of the recovered variation is in greater primary covert feather extent, followed by secondary feather length and the shape of the wing tip. Although often considered a plastic character strongly linked to flight style, the estimated ancestral wing morphology is found to be generally conservative among basal parts of most major avian lineages. The radiation of birds is characterized by successive diversification into largely distinct areas of morphospace. However, aquatic taxa show convergence in feathering despite differences in flight style, and songbirds move into a region of morphospace also occupied by basal taxa but at markedly different body sizes. These results have implications for the proposed inference of flight style in extinct taxa. PMID:26446812

An experimental study of the unsteady vortex structures in the wake of a root-fixed flapping wing

NASA Astrophysics Data System (ADS)

Hu, Hui; Clemons, Lucas; Igarashi, Hirofumi

2011-08-01

An experimental study was conducted to characterize the evolution of the unsteady vortex structures in the wake of a root-fixed flapping wing with the wing size, stroke amplitude, and flapping frequency within the range of insect characteristics for the development of novel insect-sized nano-air-vehicles (NAVs). The experiments were conducted in a low-speed wing tunnel with a miniaturized piezoelectric wing (i.e., chord length, C = 12.7 mm) flapping at a frequency of 60 Hz (i.e., f = 60 Hz). The non-dimensional parameters of the flapping wing are chord Reynolds number of Re = 1,200, reduced frequency of k = 3.5, and non-dimensional flapping amplitude at wingtip h = A/C = 1.35. The corresponding Strouhal number (Str) is 0.33 , which is well within the optimal range of 0.2 < Str < 0.4 used by flying insects and birds and swimming fishes for locomotion. A digital particle image velocimetry (PIV) system was used to achieve phased-locked and time-averaged flow field measurements to quantify the transient behavior of the wake vortices in relation to the positions of the flapping wing during the upstroke and down stroke flapping cycles. The characteristics of the wake vortex structures in the chordwise cross planes at different wingspan locations were compared quantitatively to elucidate underlying physics for a better understanding of the unsteady aerodynamics of flapping flight and to explore/optimize design paradigms for the development of novel insect-sized, flapping-wing-based NAVs.

Wings of the butterfly: Sunspot groups for 1826-2015

NASA Astrophysics Data System (ADS)

Leussu, R.; Usoskin, I. G.; Senthamizh Pavai, V.; Diercke, A.; Arlt, R.; Denker, C.; Mursula, K.

2017-03-01

The spatio-temporal evolution of sunspot activity, the so-called Maunder butterfly diagram, has been continously available since 1874 using data from the Royal Greenwich Observatory, extended by SOON network data after 1976. Here we present a new extended butterfly diagram of sunspot group occurrence since 1826, using the recently digitized data from Schwabe (1826-1867) and Spörer (1866-1880). The wings of the diagram are separated using a recently developed method based on an analysis of long gaps in sunspot group occurrence in different latitude bands. We define characteristic latitudes, corresponding to the start, end, and the largest extent of the wings (the F, L, and H latitudes). The H latitudes (30°-45°) are highly significantly correlated with the strength of the wings (quantified by the total sum of the monthly numbers of sunspot groups). The F latitudes (20°-30°) depict a weak tendency, especially in the southern hemisphere, to follow the wing strength. The L latitudes (2°-10°) show no clear relation to the wing strength. Overall, stronger cycle wings tend to start at higher latitudes and have a greater wing extent. A strong (5-6)-cycle periodic oscillation is found in the start and end times of the wings and in the overlap and gaps between successive wings of one hemisphere. While the average wing overlap is zero in the southern hemisphere, it is two to three months in the north. A marginally significant oscillation of about ten solar cycles is found in the asymmetry of the L latitudes. The new long database of butterfly wings provides new observational constraints to solar dynamo models that discuss the spatio-temporal distribution of sunspot occurrence over the solar cycle and longer. Digital data for Fig. 1 are available at the CDS via anonymous ftp to http://cdsarc.u-strasbg.fr (http://130.79.128.5) or via http://cdsarc.u-strasbg.fr/viz-bin/qcat?J/A+A/599/A131

Weights of wild mallard Anas platyrhynchos, gadwall A. strepera, and blue-winged teal A. discors during the breeding season

USGS Publications Warehouse

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.

Host-feeding sources and habitats jointly affect wing developmental stability depending on sex in the major Chagas disease vector Triatoma infestans.

PubMed

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.

Inspiration for wing design: how forelimb specialization enables active flight in modern vertebrates.

PubMed

Chin, Diana D; Matloff, Laura Y; Stowers, Amanda Kay; Tucci, Emily R; Lentink, David

2017-06-01

Harnessing flight strategies refined by millions of years of evolution can help expedite the design of more efficient, manoeuvrable and robust flying robots. This review synthesizes recent advances and highlights remaining gaps in our understanding of how bird and bat wing adaptations enable effective flight. Included in this discussion is an evaluation of how current robotic analogues measure up to their biological sources of inspiration. Studies of vertebrate wings have revealed skeletal systems well suited for enduring the loads required during flight, but the mechanisms that drive coordinated motions between bones and connected integuments remain ill-described. Similarly, vertebrate flight muscles have adapted to sustain increased wing loading, but a lack of in vivo studies limits our understanding of specific muscular functions. Forelimb adaptations diverge at the integument level, but both bird feathers and bat membranes yield aerodynamic surfaces with a level of robustness unparalleled by engineered wings. These morphological adaptations enable a diverse range of kinematics tuned for different flight speeds and manoeuvres. By integrating vertebrate flight specializations-particularly those that enable greater robustness and adaptability-into the design and control of robotic wings, engineers can begin narrowing the wide margin that currently exists between flying robots and vertebrates. In turn, these robotic wings can help biologists create experiments that would be impossible in vivo . © 2017 The Author(s).

Dynamic deformation and fracture of single crystal silicon: Fracture modes, damage laws, and anisotropy

DOE PAGES

Huang, J. Y.; E, J. C.; Huang, J. W.; ...

2016-05-25

Impact fracture of single-crystal Si is critical to long-term reliability of electronic devices and solar cells for its wide use as components or substrates in semiconductor industry. Single-crystal Si is loaded along two different crystallographic directions with a split Hopkinson pressure bar integrated with an in situ x-ray imaging and diffraction system. Bulk stress histories are measured, simultaneously with x-ray phase contrast imaging (XPCI) and Laue diffraction. Damage evolution is quantified with grayscale maps from XPCI. Single-crystal Si exhibits pronounced anisotropy in fracture modes, and thus fracture strengths and damage evolution. For loading along [11¯ 0] and viewing along [001],more » (1¯1¯0)[11¯ 0] cleavage is activated and induces horizontal primary cracks followed by perpendicular wing cracks. However, for loading along [011¯] and viewing along [111], random nucleation and growth of shear and tensile-splitting crack networks lead to catastrophic failure of materials with no cleavage. The primary-wing crack mode leads to a lower characteristic fracture strength due to predamage, but a more concentrated strength distribution, i.e., a higher Weibull modulus, compared to the second loading case. Furthermore, the sequential primary cracking, wing cracking and wing-crack coalescence processes result in a gradual increase of damage with time, deviating from theoretical predictions. Particle size and aspect ratios of fragments are discussed with postmortem fragment analysis, which verifies fracture modes observed in XPCI.« less

Insulin receptor-mediated signaling via phospholipase C-γ regulates growth and differentiation in Drosophila.

PubMed

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.

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.

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.

PubMed

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 ).

An experimental study of mushroom shaped stall cells. [on finite wings with separated flow

NASA Technical Reports Server (NTRS)

Winkelmann, A. E.

1982-01-01

Surface patterns characterized by a pair of counter-rotating swirls have been observed in connection with the conduction of surface flow visualization experiments involving test geometries with separated flows. An example of this phenomenon occurring on a finite wing with trailing edge stall has been referred to by Winkelmann and Barlow (1980) as 'mushroom shaped'. A description is presented of a collection of experimental results which show or suggest the occurrence of mushroom shaped stall cells on a variety of test geometries. Investigations conducted with finite wings, airfoil models, and flat plates are considered, and attention is given to studies involving the use of bluff models, investigations of shock induced boundary layer separation, and mushroom shaped patterns observed in a number of miscellaneous cases. It is concluded that the mushroom shaped stall cell appears commonly in separated flow regions.

Wingless is a positive regulator of eyespot color patterns in Bicyclus anynana butterflies.

PubMed

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.

U.S. Department of Defense Official Website

Science.gov Websites

during the evolution, the members of Marine Aerial Refueler Transport Squadron 352, Marine Aircraft Group 11, 3rd Marine Aircraft Wing, returned from their relief mission with a sense of accomplishment

Wind tunnel investigation of the interaction and breakdown characteristics of slender wing vortices at subsonic, transonic, and supersonic speeds

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.

Molecular pathways to parallel evolution: I. Gene nexuses and their morphological correlates.

PubMed

Zuckerkandl, E

1994-12-01

Aspects of the regulatory interactions among genes are probably as old as most genes are themselves. Correspondingly, similar predispositions to changes in such interactions must have existed for long evolutionary periods. Features of the structure and the evolution of the system of gene regulation furnish the background necessary for a molecular understanding of parallel evolution. Patently "unrelated" organs, such as the fat body of a fly and the liver of a mammal, can exhibit fractional homology, a fraction expected to become subject to quantitation. This also seems to hold for different organs in the same organism, such as wings and legs of a fly. In informational macromolecules, on the other hand, homology is indeed all or none. In the quite different case of organs, analogy is expected usually to represent attenuated homology. Many instances of putative convergence are likely to turn out to be predominantly parallel evolution, presumably including the case of the vertebrate and cephalopod eyes. Homology in morphological features reflects a similarity in networks of active genes. Similar nexuses of active genes can be established in cells of different embryological origins. Thus, parallel development can be considered a counterpart to parallel evolution. Specific macromolecular interactions leading to the regulation of the c-fos gene are given as an example of a "controller node" defined as a regulatory unit. Quantitative changes in gene control are distinguished from relational changes, and frequent parallelism in quantitative changes is noted in Drosophila enzymes. Evolutionary reversions in quantitative gene expression are also expected. The evolution of relational patterns is attributed to several distinct mechanisms, notably the shuffling of protein domains. The growth of such patterns may in part be brought about by a particular process of compensation for "controller gene diseases," a process that would spontaneously tend to lead to increased regulatory and organismal complexity. Despite the inferred increase in gene interaction complexity, whose course over evolutionary time is unknown, the number of homology groups for the functional and structural protein units designated as domains has probably remained rather constant, even as, in some of its branches, evolution moved toward "higher" organisms. In connection with this process, the question is raised of parallel evolution within the purview of activating and repressing master switches and in regard to the number of levels into which the hierarchies of genic master switches will eventually be resolved.

The potential influence of morphology on the evolutionary divergence of an acoustic signal

PubMed Central

Pitchers, W. R.; Klingenberg, C.P.; Tregenza, Tom; Hunt, J.; Dworkin, I.

2014-01-01

The evolution of acoustic behaviour and that of the morphological traits mediating its production are often coupled. Lack of variation in the underlying morphology of signalling traits has the potential to constrain signal evolution. This relationship is particularly likely in field crickets, where males produce acoustic advertisement signals to attract females by stridulating with specialized structures on their forewings. In this study, we characterise the size and geometric shape of the forewings of males from six allopatric populations of the black field cricket (Teleogryllus commodus) known to have divergent advertisement calls. We sample from each of these populations using both wild-caught and common-garden reared cohorts, allowing us to test for multivariate relationships between wing morphology and call structure. We show that the allometry of shape has diverged across populations. However, there was a surprisingly small amount of covariation between wing shape and call structure within populations. Given the importance of male size for sexual selection in crickets, the divergence we observe among populations has the potential to influence the evolution of advertisement calls in this species. PMID:25223712

WINGS-SPE Spectroscopy in the WIde-field Nearby Galaxy-cluster Survey

NASA Astrophysics Data System (ADS)

Cava, A.; Bettoni, D.; Poggianti, B. M.; Couch, W. J.; Moles, M.; Varela, J.; Biviano, A.; D'Onofrio, M.; Dressler, A.; Fasano, G.; Fritz, J.; Kjærgaard, P.; Ramella, M.; Valentinuzzi, T.

2009-03-01

Aims: We present the results from a comprehensive spectroscopic survey of the WINGS (WIde-field Nearby Galaxy-cluster Survey) clusters, a program called WINGS-SPE. The WINGS-SPE sample consists of 48 clusters, 22 of which are in the southern sky and 26 in the north. The main goals of this spectroscopic survey are: (1) to study the dynamics and kinematics of the WINGS clusters and their constituent galaxies, (2) to explore the link between the spectral properties and the morphological evolution in different density environments and across a wide range of cluster X-ray luminosities and optical properties. Methods: Using multi-object fiber-fed spectrographs, we observed our sample of WINGS cluster galaxies at an intermediate resolution of 6-9 Å and, using a cross-correlation technique, we measured redshifts with a mean accuracy of ~45 km s-1. Results: We present redshift measurements for 6137 galaxies and their first analyses. Details of the spectroscopic observations are reported. The WINGS-SPE has ~30% overlap with previously published data sets, allowing us both to perform a complete comparison with the literature and to extend the catalogs. Conclusions: Using our redshifts, we calculate the velocity dispersion for all the clusters in the WINGS-SPE sample. We almost triple the number of member galaxies known in each cluster with respect to previous works. We also investigate the X-ray luminosity vs. velocity dispersion relation for our WINGS-SPE clusters, and find it to be consistent with the form Lx ∝ σ_v^4. Table 4, containing the complete redshift catalog, is only available in electronic form at the CDS via anonymous ftp to cdsarc.u-strasbg.fr (130.79.128.5) or via http://cdsweb.u-strasbg.fr/cgi-bin/qcat?J/A+A/495/707

In situ protocol for butterfly pupal wings using riboprobes.

PubMed

Ramos, Diane; Monteiro, Antonia

2007-01-01

Here we present, in video format, a protocol for in situ hybridizations in pupal wings of the butterfly Bicyclus anynana using riboprobes. In situ hybridizations, a mainstay of developmental biology, are useful to study the spatial and temporal patterns of gene expression in developing tissues at the level of transcription. If antibodies that target the protein products of gene transcription have not yet been developed, and/or there are multiple gene copies of a particular protein in the genome that cannot be differentiated using available antibodies, in situs can be used instead. While an in situ technique for larval wing discs has been available to the butterfly community for several years, the current protocol has been optimized for the larger and more fragile pupal wings.

Boundary Dpp promotes growth of medial and lateral regions of the Drosophila wing.

PubMed

Barrio, Lara; Milán, Marco

2017-07-04

The gradient of Decapentaplegic (Dpp) in the Drosophila wing has served as a paradigm to characterize the role of morphogens in regulating patterning. However, the role of this gradient in regulating tissue size is a topic of intense debate as proliferative growth is homogenous. Here, we combined the Gal4/UAS system and a temperature-sensitive Gal80 molecule to induce RNAi-mediated depletion of dpp and characterise the spatial and temporal requirement of Dpp in promoting growth. We show that Dpp emanating from the AP compartment boundary is required throughout development to promote growth by regulating cell proliferation and tissue size. Dpp regulates growth and proliferation rates equally in central and lateral regions of the developing wing appendage and reduced levels of Dpp affects similarly the width and length of the resulting wing. We also present evidence supporting the proposal that graded activity of Dpp is not an absolute requirement for wing growth.

An exploratory study of apex fence flaps on a 74 deg delta wing

NASA Technical Reports Server (NTRS)

Wahls, R. A.; Vess, R. J.

1985-01-01

An exploratory wind tunnel investigation was performed to observe the flow field effects produced by vertically deployed apex fences on a planar 74 degree delta wing. The delta shaped fences, each comprising approximately 3.375 percent of the wing area, were affixed along the first 25 percent of the wing leading edge in symmetric as well as asymmetric (i.e., fence on one side only) arrangements. The vortex flow field was visualized at angles of attack from 0 to 20 degrees using helium bubble and oil flow techniques; upper surface pressures were also measured along spanwise rows. The results were used to construct a preliminary description of the vortex patterns and induced pressures associated with vertical apex fence deployment. The objective was to obtain an initial evaluation of the potential of apex fences as vortex devices for subsonic lift modulation as well as lateral directional control of delta wing aircraft.

A major gene controls mimicry and crypsis in butterflies and moths

PubMed Central

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.

PubMed

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.

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.

The evolution of avian wing shape and previously unrecognized trends in covert feathering.

PubMed

Wang, Xia; Clarke, Julia A

2015-10-07

Avian wing shape has been related to flight performance, migration, foraging behaviour and display. Historically, linear measurements of the feathered aerofoil and skeletal proportions have been used to describe this shape. While the distribution of covert feathers, layered over the anterior wing, has long been assumed to contribute to aerofoil properties, to our knowledge no previous studies of trends in avian wing shape assessed their variation. Here, these trends are explored using a geometric-morphometric approach with landmarks describing the wing outline as well as the extent of dorsal and ventral covert feathers for 105 avian species. We find that most of the observed variation is explained by phylogeny and ecology but shows only a weak relationship with previously described flight style categories, wing loading and an investigated set of aerodynamic variables. Most of the recovered variation is in greater primary covert feather extent, followed by secondary feather length and the shape of the wing tip. Although often considered a plastic character strongly linked to flight style, the estimated ancestral wing morphology is found to be generally conservative among basal parts of most major avian lineages. The radiation of birds is characterized by successive diversification into largely distinct areas of morphospace. However, aquatic taxa show convergence in feathering despite differences in flight style, and songbirds move into a region of morphospace also occupied by basal taxa but at markedly different body sizes. These results have implications for the proposed inference of flight style in extinct taxa. © 2015 The Author(s).

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.

Effects of the essential oil constituent thymol and other neuroactive chemicals on flight motor activity and wing beat frequency in the blowfly Phaenicia sericata.

PubMed

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.

Control of vortex on a non-slender delta wing by a nanosecond pulse surface dielectric barrier discharge

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.

Hormonal control of GTP cyclohydrolase I gene expression and enzyme activity during color pattern development in wings of Precis coenia.

PubMed

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.

Heat transfer and oil flow studies on a single-stage-to-orbit control-configured winged entry vehicle

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.

The Hydrodynamics of Plesiosaurs

NASA Astrophysics Data System (ADS)

Muscutt, Luke; Ganapathisubramani, Bharathram; Dyke, Gareth; Weymouth, Gabriel

2015-11-01

Plesiosaurs are extinct marine reptiles that existed at the same time as the dinosaurs, and are the only known animals to swim by actively flapping their four wing-like flippers. This can be viewed as a tandem flapping wing problem, where the hind wing is operating in the wake of the fore wing. Experiments using full-scale robotic plesiosaur flippers in a large flume tank have been used to investigate the kinematics and interaction of the flippers. The flippers are actuated in heave and pitch, and a combination of force measurements and flow visualization are used to analyze the characteristics of the vortex interaction between the flippers. Previous two-dimensional numerical simulations have shown that certain kinematics give an increase in thrust of the hind flipper of up to 50%. The current experiments determine if such thrust augmentation is present for a three-dimensional flowfield and determine the kinematics that give the highest possible thrust. This will help to answer paleo-biological questions about the function and evolution of the plesiosaur flippers, along with helping to determine if tandem flapping wings could be a viable propulsion system for autonomous underwater vehicles. Support from EPSRC and Ginko Investments Ltd.

Molecular Mechanisms for High Hydrostatic Pressure-Induced Wing Mutagenesis in Drosophila melanogaster.

PubMed

Wang, Hua; Wang, Kai; Xiao, Guanjun; Ma, Junfeng; Wang, Bingying; Shen, Sile; Fu, Xueqi; Zou, Guangtian; Zou, Bo

2015-10-08

Although High hydrostatic pressure (HHP) as an important physical and chemical tool has been increasingly applied to research of organism, the response mechanisms of organism to HHP have not been elucidated clearly thus far. To identify mutagenic mechanisms of HHP on organisms, here, we treated Drosophila melanogaster (D. melanogaster) eggs with HHP. Approximately 75% of the surviving flies showed significant morphological abnormalities from the egg to the adult stages compared with control flies (p < 0.05). Some eggs displayed abnormal chorionic appendages, some larvae were large and red, and some adult flies showed wing abnormalities. Abnormal wing phenotypes of D. melanogaster induced by HHP were used to investigate the mutagenic mechanisms of HHP on organism. Thus 285 differentially expressed genes associated with wing mutations were identified using Affymetrix Drosophila Genome Array 2.0 and verified with RT-PCR. We also compared wing development-related central genes in the mutant flies with control flies using DNA sequencing to show two point mutations in the vestigial (vg) gene. This study revealed the mutagenic mechanisms of HHP-induced mutagenesis in D. melanogaster and provided a new model for the study of evolution on organisms.

Three-dimensional flow visualization and vorticity dynamics in revolving wings

NASA Astrophysics Data System (ADS)

Cheng, Bo; Sane, Sanjay P.; Barbera, Giovanni; Troolin, Daniel R.; Strand, Tyson; Deng, Xinyan

2013-01-01

We investigated the three-dimensional vorticity dynamics of the flows generated by revolving wings using a volumetric 3-component velocimetry system. The three-dimensional velocity and vorticity fields were represented with respect to the base axes of rotating Cartesian reference frames, and the second invariant of the velocity gradient was evaluated and used as a criterion to identify two core vortex structures. The first structure was a composite of leading, trailing, and tip-edge vortices attached to the wing edges, whereas the second structure was a strong tip vortex tilted from leading-edge vortices and shed into the wake together with the vorticity generated at the tip edge. Using the fundamental vorticity equation, we evaluated the convection, stretching, and tilting of vorticity in the rotating wing frame to understand the generation and evolution of vorticity. Based on these data, we propose that the vorticity generated at the leading edge is carried away by strong tangential flow into the wake and travels downwards with the induced downwash. The convection by spanwise flow is comparatively negligible. The three-dimensional flow in the wake also exhibits considerable vortex tilting and stretching. Together these data underscore the complex and interconnected vortical structures and dynamics generated by revolving wings.

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

Developmental pathways of Glossotermes oculatus (Isoptera, Serritermitidae): at the cross-roads of worker caste evolution in termites.

PubMed

Bourguignon, Thomas; Sobotník, Jan; Hanus, Robert; Roisin, Yves

2009-01-01

The onset of a specialized ("true") worker caste is a crucial step in the evolution of termite societies. Such workers, permanently excluded from wing development, repeatedly evolved from totipotent immatures, called "false" workers or pseudergates. In the family Rhinotermitidae, the presence of true workers and the level of specialization of this caste are highly variable, and key taxa illustrate transitional situations providing clues about worker evolution. Here we focused on the status of working immatures of Glossotermes oculatus, from the family Serritermitidae, now thought to represent either the sister-group of the Rhinotermitidae or a basal lineage nested within them. Contrary to previous assumptions, we show that the apterous immatures performing worker tasks in G. oculatus are the source of the single wing-budded nymphal instar preceding the alate. Consequently, they qualify as pseudergates rather than true workers. However, the sex ratio is strongly male biased in pseudergates and soldiers, which is a trait usually restricted to termites with true workers. We therefore argue that pseudergates of G. oculatus are close to a point where the species could easily shift toward the differentiation of a true worker caste, and that G. oculatus pinpoints a new possible route for the evolution of true workers from pseudergates.

Aerodynamic Interaction between Delta Wing and Hemisphere-Cylinder in Supersonic Flow

NASA Astrophysics Data System (ADS)

Nishino, Atsuhiro; Ishikawa, Takahumi; Nakamura, Yoshiaki

As future space vehicles, Reusable Launch Vehicle (RLV) needs to be developed, where there are two kinds of RLV: Single Stage To Orbit (SSTO) and Two Stage To Orbit (TSTO). In the latter case, the shock/shock interaction and shock/boundary layer interaction play a key role. In the present study, we focus on the supersonic flow field with aerodynamic interaction between a delta wing and a hemisphere-cylinder, which imitate a TSTO, where the clearance, h, between the delta wing and hemisphere-cylinder is a key parameter. As a result, complicated flow patterns were made clear, including separation bubbles.

Propeller swirl effect on single-engine general-aviation aircraft stall-spin tendencies

NASA Technical Reports Server (NTRS)

Katz, Joseph; Feistel, Terry W.

1987-01-01

An investigation is conducted of the effect of a single engine, untapered low wing general aviation aircraft propeller's swirl on the craft's stall pattern. The asymmetrical character of the propeller's swirl can trigger an early stall of one of the wings, aggravating the spin-entry condition. It is shown that the combination of this propeller-induced effect with adverse sideslip can result in large and abrupt changes in the rolling moment, in such conditions as uncoordinated low speed turning maneuvers where the pilot yaws the aircraft with wings level, rather than rolling it.

A Middle Triassic thoracopterid from China highlights the evolutionary origin of overwater gliding in early ray-finned fishes.

PubMed

Xu, Guang-Hui; Zhao, Li-Jun; Shen, Chen-Chen

2015-01-01

Gliding adaptations in thoracopterid flying fishes represent a remarkable case of convergent evolution of overwater gliding strategy with modern exocoetid flying fishes, but the evolutionary origin of this strategy was poorly known in the thoracopterids because of lack of transitional forms. Until recently, all thoracopterids, from the Late Triassic of Austria and Italy and the Middle Triassic of South China, were highly specialized 'four-winged' gliders in having wing-like paired fins and an asymmetrical caudal fin with the lower caudal lobe notably larger than the upper lobe. Here, we show that the new genus Wushaichthys and the previously alleged 'peltopleurid' Peripeltopleurus, from the Middle Triassic (Ladinian, 235-242 Ma) of South China and near the Ladinian/Anisian boundary of southern Switzerland and northern Italy, respectively, represent the most primitive and oldest known thoracopterids. Wushaichthys, the most basal thoracopterid, shows certain derived features of this group in the skull. Peripeltopleurus shows a condition intermediate between Wushaichthys and Thoracopterus in having a slightly asymmetrical caudal fin but still lacking wing-like paired fins. Phylogenetic studies suggest that the evolution of overwater gliding of thoracopterids was gradual in nature; a four-stage adaption following the 'cranial specialization-asymmetrical caudal fin-enlarged paired fins-scale reduction' sequence has been recognized in thoracopterid evolution. Moreover, Wushaichthys and Peripeltopleurus bear hooklets on the anal fin of supposed males, resembling those of modern viviparious teleosts. Early thoracopterids probably had evolved a live-bearing reproductive strategy. © 2015 The Author(s) Published by the Royal Society. All rights reserved.

Low-Frequency Flow Oscillations on Stalled Wings Exhibiting Cellular Separation Topology

NASA Astrophysics Data System (ADS)

Disotell, Kevin James

One of the most pervasive threats to aircraft controllability is wing stall, a condition associated with loss of lift due to separation of air flow from the wing surface at high angles of attack. A recognized need for improved upset recovery training in extended-envelope flight simulators is a physical understanding of the post-stall aerodynamic environment, particularly key flow phenomena which influence the vehicle trajectory. Large-scale flow structures known as stall cells, which scale with the wing chord and are spatially-periodic along the span, have been previously observed on post-stall airfoils with trailing-edge separation present. Despite extensive documentation of stall cells in the literature, the physical mechanisms behind their formation and evolution have proven to be elusive. The undertaken study has sought to characterize the inherently turbulent separated flow existing above the wing surface with cell formation present. In particular, the question of how the unsteady separated flow may interact with the wing to produce time-averaged cellular surface patterns is considered. Time-resolved, two-component particle image velocimetry measurements were acquired at the plane of symmetry of a single stall cell formed on an extruded NACA 0015 airfoil model at chord Reynolds number of 560,000 to obtain insight into the time-dependent flow structure. The evolution of flow unsteadiness was analyzed over a static angle-of-attack range covering the narrow post-stall regime in which stall cells have been observed. Spectral analysis of velocity fields acquired near the stall angle confirmed a low-frequency flow oscillation previously detected in pointwise surface measurements by Yon and Katz (1998), corresponding to a Strouhal number of 0.042 based on frontal projected chord height. Probability density functions of the streamwise velocity component were used to estimate the convective speed of this mode at approximately half the free-stream velocity, in agreement with Yon and Katz. Large-amplitude streamwise Reynolds stresses in the separated shear layer were found to be manifested by the low-frequency oscillation through inspection of the spectral energy distribution. Using the method of Proper Orthogonal Decomposition to construct reduced-order models of the acquired time sequences, the low-frequency unsteadiness appeared to be linked to an interaction between the separated and trailing-edge shear layers, in contrast to a bubble-bursting mechanism which has been observed for different stall behaviors. As the static angle of attack was increased further, the separated flow structure was seen to transition to a faster eddy motion expected for bluff-body wakes. A novel scaling study was conducted to evaluate the potential role of low-frequency unsteadiness in producing the spanwise wavelengths associated with cell formation, which was found to be in favorable agreement with scaling trends in the literature. Finally, instantaneous pressure-sensitive paint measurements were demonstrated on a DU 97-W-300 wind turbine airfoil at chord Reynolds number of 225,000 with leading-edge trip applied, in which the development of spiral node structures associated with cell formation were captured in the trailing-edge separation. The contributed work suggests that further study into the influence of large-scale unsteadiness on the three-dimensional organization of stall cells is merited.

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.

Ecomorphology of the external flight apparatus of blackcaps (Sylvia atricapilla) with different migration behavior.

PubMed

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.

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.

Temperature-Related Divergence in Experimental Populations of DROSOPHILA MELANOGASTER. I. Genetic and Developmental Basis of Wing Size and Shape Variation

PubMed Central

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°).

Spatial Variation in Body Size and Wing Dimorphism Correlates With Environmental Conditions in the Grasshopper Dichroplus vittatus (Orthoptera: Acrididae).

PubMed

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.

A new genus and species of tettigarctid cicada from the early Miocene of New Zealand: Paratettigarcta zealandica (Hemiptera, Auchenorrhyncha, Tettigarctidae)

PubMed Central

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

A new genus and species of tettigarctid cicada from the early Miocene of New Zealand: Paratettigarctazealandica (Hemiptera, Auchenorrhyncha, Tettigarctidae).

PubMed

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

USGS Publications Warehouse

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.

Folding Wings like a Cockroach: A Review of Transverse Wing Folding Ensign Wasps (Hymenoptera: Evaniidae: Afrevania and Trissevania)

PubMed Central

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

Basal Complex and Basal Venation of Odonata Wings: Structural Diversity and Potential Role in the Wing Deformation

PubMed Central

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

Butterfly Wings Are Three-Dimensional: Pupal Cuticle Focal Spots and Their Associated Structures in Junonia Butterflies.

PubMed

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.

Butterfly Wings Are Three-Dimensional: Pupal Cuticle Focal Spots and Their Associated Structures in Junonia Butterflies

PubMed Central

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

Genome differentiation of Drosophila melanogaster from a microclimate contrast in Evolution Canyon, Israel

PubMed Central

Hübner, Sariel; Rashkovetsky, Eugenia; Kim, Young Bun; Oh, Jung Hun; Michalak, Katarzyna; Weiner, Dmitry; Korol, Abraham B.; Nevo, Eviatar; Michalak, Pawel

2013-01-01

The opposite slopes of “Evolution Canyon” in Israel have served as a natural model system of adaptation to a microclimate contrast. Long-term studies of Drosophila melanogaster populations inhabiting the canyon have exhibited significant interslope divergence in thermal and drought stress resistance, candidate genes, mobile elements, habitat choice, mating discrimination, and wing-shape variation, all despite close physical proximity of the contrasting habitats, as well as substantial interslope migration. To examine patterns of genetic differentiation at the genome-wide level, we used high coverage sequencing of the flies’ genomes. A total of 572 genes were significantly different in allele frequency between the slopes, 106 out of which were associated with 74 significantly overrepresented gene ontology (GO) terms, particularly so with response to stimulus and developmental and reproductive processes, thus corroborating previous observations of interslope divergence in stress response, life history, and mating functions. There were at least 37 chromosomal “islands” of interslope divergence and low sequence polymorphism, plausible signatures of selective sweeps, more abundant in flies derived from one (north-facing) of the slopes. Positive correlation between local recombination rate and the level of nucleotide polymorphism was also found. PMID:24324170

The subtropical mesospheric jet observed by the Nimbus 7 Limb Infrared Monitor of the Stratosphere

NASA Technical Reports Server (NTRS)

Dunkerton, T. J.; Delisi, D. P.

1985-01-01

Nimbus 7 Limb Infrared Monitor of the Stratosphere observations of wave-mean flow interactions in the winter 1978-1979 middle atmosphere are surveyed, extending up to 0.05 mbar. These observations describe the evolution of the subtropical mesospheric jet and its polar mixed layer. Quasi-steady mean wind patterns are disrupted by three transitions in this winter: one primarily affecting the mesosphere (December 15, 1978), a minor warming affecting both regions (January 26-February 8, 1979), and a major warming largely confined to the stratosphere (February 22, 1979). The zonally averaged flow is barotropically unstable in the wings of the subtropical mesospheric jet. All the major decelerations of the mean flow are correlated with D(F), the body force per unit mass directly attributable to planetary Rossby waves, indicating that these waves make a significant contribution to the momentum budget in the lower half of the mesosphere.

[Key morphofunctional transformations in the evolution of chiropterans (Bats, Chiroptera)].

PubMed

Kovaleva, I M

2014-01-01

Study on the morphology and morphogenesis of wing membranes in Bats has revealed some peculiarities in their structure and development. Understanding the embryogenesis of these animals, as well as attraction of data obtained on their molecular genetics and paleontology, allows one to single out some factors that could have initiated evolutionary modifications in development programs. A scenario of the key morphofunctional transformations in the forelimbs during the evolution of chiropterans is given.

Divergent mating patterns and a unique mode of external sperm transfer in Zoraptera: an enigmatic group of pterygote insects

NASA Astrophysics Data System (ADS)

Dallai, R.; Gottardo, M.; Mercati, D.; Machida, R.; Mashimo, Y.; Matsumura, Y.; Beutel, R. G.

2013-06-01

A remarkable external sperm transfer is described for the first time in a species of a group of winged insects (Pterygota), the enigmatic Zoraptera. Mating and sperm transfer of two species of the order were examined in detail, documented, and compared with each other and with patterns described for other species belonging to the order. The behavior differs strikingly in Zorotypus impolitus and Zorotypus magnicaudelli. A copula is performed by males and females of the latter, as it is also the case in other zorapteran species and generally in pterygote insects. In striking contrast to this, males of Z. impolitus do not copulate but deposit small (100 μm in diameter) spermatophores externally on the abdomen of the female. Each spermatophore contains only one giant spermatozoon (3 mm long and 3 μm wide), a unique feature in the entire Hexapoda. External sperm transfer in Pterygota is a highly unusual case of evolutionary reversal. The very small relict group Zoraptera displays a uniform general morphology but exhibits very different reproductive structures and patterns of mating behavior. This may be an extreme form of a more general situation in insects, with a specific form of selection resulting in an accelerated rate of evolution in the reproductive system.

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.

Winged scapula incidence and upper limb morbidity after surgery for breast cancer with axillary dissection.

PubMed

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.

Ecological Trade-offs between Migration and Reproduction Are Mediated by the Nutrition-Sensitive Insulin-Signaling Pathway.

PubMed

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.

Boundary Dpp promotes growth of medial and lateral regions of the Drosophila wing

PubMed Central

Barrio, Lara; Milán, Marco

2017-01-01

The gradient of Decapentaplegic (Dpp) in the Drosophila wing has served as a paradigm to characterize the role of morphogens in regulating patterning. However, the role of this gradient in regulating tissue size is a topic of intense debate as proliferative growth is homogenous. Here, we combined the Gal4/UAS system and a temperature-sensitive Gal80 molecule to induce RNAi-mediated depletion of dpp and characterise the spatial and temporal requirement of Dpp in promoting growth. We show that Dpp emanating from the AP compartment boundary is required throughout development to promote growth by regulating cell proliferation and tissue size. Dpp regulates growth and proliferation rates equally in central and lateral regions of the developing wing appendage and reduced levels of Dpp affects similarly the width and length of the resulting wing. We also present evidence supporting the proposal that graded activity of Dpp is not an absolute requirement for wing growth. DOI: http://dx.doi.org/10.7554/eLife.22013.001 PMID:28675372

An experimental study of separated flow on a finite wing

NASA Technical Reports Server (NTRS)

Winkelmann, A. E.

1981-01-01

The flow field associated with the formation of a mushroom shaped trailing edge stall cell on a low-aspect-ratio (AR = 4.0) wing was investigated in a series of low speed wind tunnel tests (Reynolds number based on 15.2 cm chord = 480,000). Flow field surveys of the separation bubble and wake of a partially stalled and fully stalled wing were completed using a hot-wire probe, a split-film probe, and a directional sensitive pressure probe. A new color video display technique was developed to display the flow field survey data. Photographs were obtained of surface oil flow patterns and smoke flow visualization

Exploring bird aerodynamics using radio-controlled models.

PubMed

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.

Iridescence and spectral filtering of the gyroid-type photonic crystals in Parides sesostris wing scales

PubMed Central

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

Investigation of hindwing folding in ladybird beetles by artificial elytron transplantation and microcomputed tomography.

PubMed

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.

Investigation of hindwing folding in ladybird beetles by artificial elytron transplantation and microcomputed tomography

PubMed Central

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

De novo analysis of the Nilaparvata lugens (Stål) antenna transcriptome and expression patterns of olfactory genes.

PubMed

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

PubMed Central

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.

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.

Plant terpenoids: acute toxicities and effects on flight motor activity and wing beat frequency in the blow fly Phaenicia sericata.

PubMed

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.

Extensive transcriptional response associated with seasonal plasticity of butterfly wing patterns.

PubMed

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.

Command and Compensation in a Neuromodulatory Decision Network

PubMed Central

Luan, Haojiang; Diao, Fengqiu; Peabody, Nathan C.; White, Benjamin H.

2012-01-01

The neural circuits that mediate behavioral choices must not only weigh internal demands and environmental circumstances, but also select and implement specific actions, including associated visceral or neuroendocrine functions. Coordinating these multiple processes suggests considerable complexity. As a consequence, even circuits that support simple behavioral decisions remain poorly understood. Here we show that the environmentally-sensitive wing expansion decision of adult fruit flies is coordinated by a single pair of neuromodulatory neurons with command-like function. Targeted suppression of these neurons using the Split Gal4 system abrogates the fly's ability to expand its wings in the face of environmental challenges, while stimulating them forces expansion by coordinately activating both motor and neuroendocrine outputs. The arbitration and implementation of the wing expansion decision by this neuronal pair may illustrate a general strategy by which neuromodulatory neurons orchestrate behavior. Interestingly, the decision network shows a behavioral plasticity that is unmasked under conducive environmental conditions in flies lacking the function of the command-like neuromodulatory neurons. Such flies can often expand their wings using a motor program distinct from that of wildtype animals and controls. This compensatory program may be the vestige of an ancestral, environmentally-insensitive program used for wing expansion that existed prior to the evolution of the environmentally-adaptive program currently used by Drosophila and other cyclorrhaphan flies. PMID:22262886

New insights on basivenal sclerites using 3D tools and homology of wing veins in Odonatoptera (Insecta).

PubMed

Jacquelin, Lauriane; Desutter-Grandcolas, Laure; Chintauan-Marquier, Ioana; Boistel, Renaud; Zheng, Daran; Prokop, Jakub; Nel, André

2018-01-10

Being implied in flight, mimetism, communication, and protection, the insect wings were crucial organs for the mega diversification of this clade. Despite several attempts, the problem of wing evolution remains unresolved because the basal parts of the veins essential for vein identification are hidden in the basivenal sclerites. The homologies between wing characters thus cannot be accurately verified, while they are of primary importance to solve long-standing problems, such as the monophyly of the Palaeoptera, viz. Odonatoptera, Panephemeroptera, and Palaeozoic Palaeodictyopterida mainly known by their wings. Hitherto the tools to homologize venation were suffering several cases of exceptions, rendering them unreliable. Here we reconstruct the odonatopteran venation using fossils and a new 3D imaging tool, resulting congruent with the concept of Riek and Kukalová-Peck, with important novelties, viz. median anterior vein fused to radius and radius posterior nearly as convex as radius anterior (putative synapomorphies of Odonatoptera); subcostal anterior (ScA) fused to costal vein and most basal primary antenodal crossvein being a modified posterior branch of ScA (putative synapomorphies of Palaeoptera). These findings may reveal critical for future analyses of the relationships between fossil and extant Palaeoptera, helping to solve the evolutionary history of the insects as a whole.

Analytical and experimental investigation of microstructural alterations in bearing steel in rolling contact fatigue

NASA Astrophysics Data System (ADS)

Mobasher Moghaddam, Sina

Rolling Contact Fatigue (RCF) is one the most common failure modes in bearings. RCF is usually associated with particular microstructural alterations. Such alterations (i.e. white etching cracks, butterflies, etc.) which lead to RCF failure are known to be among the most concerning matters to bearing industry. In the current work, an analytical as well as experimental approaches are used to investigate "butterfly wing" formation, crack initiation and propagation from inclusions. A new damage evolution equation coupled with a FE model is employed to account for the effect of mean stresses and alternating stresses simultaneously to investigate butterfly formation. The proposed damage evolution law matches experimentally observed butterfly orientation, shape, and size successfully. The model is used to obtain S-N results for butterfly formation at different Hertzian load levels. The results corroborate well with the experimental data available in the open literature. The model is used to predict debonding at the inclusion/matrix interface and the most vulnerable regions for crack initiation on butterfly/matrix interface. A new variable called butterfly formation index (BFI) is introduced to manifest the dependence of wing formation on depth. The value of critical damage inside the butterfly wings was obtained experimentally and was then used to simulate damage evolution. Voronoi tessellation was used to develop the FEM domains to capture the effect of microstructural randomness on butterfly wing formation, crack initiation and propagation. Then, the effects of different inclusion characteristics such as size, depth, and stiffness on RCF life are studied. The results show that stiffness of an inclusion and its location has a significant effect on the RCF life: stiffer inclusions and inclusions located at the depth of maximum shear stress reversal are more detrimental to the RCF life. Stress concentrations are not significantly affected by inclusion size for the cases investigated; however, a stereology study showed that larger inclusions have a higher chance to be located at the critical depth and cause failure. Crack maps were recorded and compared to spall geometries observed experimentally. The results show that crack initiation locations and final spall shapes are similar to what has been observed in failed bearings.

Rapid evolution and gene expression: a rapidly evolving Mendelian trait that silences field crickets has widespread effects on mRNA and protein expression.

PubMed

Pascoal, S; Liu, X; Ly, T; Fang, Y; Rockliffe, N; Paterson, S; Shirran, S L; Botting, C H; Bailey, N W

2016-06-01

A major advance in modern evolutionary biology is the ability to start linking phenotypic evolution in the wild with genomic changes that underlie that evolution. We capitalized on a rapidly evolving Hawaiian population of crickets (Teleogryllus oceanicus) to test hypotheses about the genomic consequences of a recent Mendelian mutation of large effect which disrupts the development of sound-producing structures on male forewings. The resulting silent phenotype, flatwing, persists because of natural selection imposed by an acoustically orienting parasitoid, but it interferes with mate attraction. We examined gene expression differences in developing wing buds of wild-type and flatwing male crickets using RNA-seq and quantitative proteomics. Most differentially expressed (DE) transcripts were down-regulated in flatwing males (625 up vs. 1716 down), whereas up- and down-regulated proteins were equally represented (30 up and 34 down). Differences between morphs were clearly not restricted to a single pathway, and we recovered annotations associated with a broad array of functions that would not be predicted a priori. Using a candidate gene detection test based on homology, we identified 30% of putative Drosophila wing development genes in the cricket transcriptome, but only 10% were DE. In addition to wing-related annotations, endocrine pathways and several biological processes such as reproduction, immunity and locomotion were DE in the mutant crickets at both biological levels. Our results illuminate the breadth of genetic pathways that are potentially affected in the early stages of adaptation. © 2016 European Society For Evolutionary Biology. Journal of Evolutionary Biology © 2016 European Society For Evolutionary Biology.

Evolution of a new sense for wind in flying phasmids? Afferents and interneurons

NASA Astrophysics Data System (ADS)

Hustert, Reinhold; Klug, Rebecca

2009-12-01

The evolution of winged stick insects (phasmids) from secondarily wingless ancestors was proposed in recent studies. We explored the cuticle of flying phasmids for wind sensors that could be involved in their flight control, comparable to those known for locusts. Surprisingly, wind-sensitive hairs (wsH) occur on the palps of mouthparts and on the antennae of the winged phasmid Sipyloidea sipylus which can fly in tethered position only when air currents blow over the mouthparts. The present study describes the morphology and major functional properties of these “new” wsH with soft and bulging hair bases which are different from the beaker-like hair bases of the wsH on the cerci of phasmids and the wsH described in other insects. The most sensitive wsH of antennae and palps respond with phasic-tonic afferents to air currents exceeding 0.2 ms-1. The fields of wsH on one side of the animal respond mainly to ventral, lateral, and frontal wind on the ipsilateral side of the head. Afferent inputs from the wsH converge but also diverge to a group of specific interneurons at their branches in the suboesophageal ganglion and can send their integrated input from wsH fields of the palps and antennae to the thoracic central nervous system. Response types of individual wsH-interneurons are either phasic or phasic-tonic to air puffs or constant air currents and also, the receptive fields of individual interneurons differ. We conclude that the “new” wsH system and its interneurons mainly serve to maintain flight activity in airborne phasmids and also, the “new” wsH must have emerged together with the integrating interneurons during the evolution from wingless to the recent winged forms of phasmids.

Confocal imaging of butterfly tissue.

PubMed

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.

Flapping wing aerodynamics: from insects to vertebrates.

PubMed

Chin, Diana D; Lentink, David

2016-04-01

More than a million insects and approximately 11,000 vertebrates utilize flapping wings to fly. However, flapping flight has only been studied in a few of these species, so many challenges remain in understanding this form of locomotion. Five key aerodynamic mechanisms have been identified for insect flight. Among these is the leading edge vortex, which is a convergent solution to avoid stall for insects, bats and birds. The roles of the other mechanisms - added mass, clap and fling, rotational circulation and wing-wake interactions - have not yet been thoroughly studied in the context of vertebrate flight. Further challenges to understanding bat and bird flight are posed by the complex, dynamic wing morphologies of these species and the more turbulent airflow generated by their wings compared with that observed during insect flight. Nevertheless, three dimensionless numbers that combine key flow, morphological and kinematic parameters - the Reynolds number, Rossby number and advance ratio - govern flapping wing aerodynamics for both insects and vertebrates. These numbers can thus be used to organize an integrative framework for studying and comparing animal flapping flight. Here, we provide a roadmap for developing such a framework, highlighting the aerodynamic mechanisms that remain to be quantified and compared across species. Ultimately, incorporating complex flight maneuvers, environmental effects and developmental stages into this framework will also be essential to advancing our understanding of the biomechanics, movement ecology and evolution of animal flight. © 2016. Published by The Company of Biologists Ltd.

Ontogeny of lift and drag production in ground birds.

PubMed

Heers, Ashley M; Tobalske, Bret W; Dial, Kenneth P

2011-03-01

The juvenile period is often a crucial interval for selective pressure on locomotor ability. Although flight is central to avian biology, little is known about factors that limit flight performance during development. To improve understanding of flight ontogeny, we used a propeller (revolving wing) model to test how wing shape and feather structure influence aerodynamic performance during development in the precocial chukar partridge (Alectoris chukar, 4 to >100 days post hatching). We spun wings in mid-downstroke posture and measured lift (L) and drag (D) using a force plate upon which the propeller assembly was mounted. Our findings demonstrate a clear relationship between feather morphology and aerodynamic performance. Independent of size and velocity, older wings with stiffer and more asymmetrical feathers, high numbers of barbicels and a high degree of overlap between barbules generate greater L and L:D ratios than younger wings with flexible, relatively symmetrical and less cohesive feathers. The gradual transition from immature feathers and drag-based performance to more mature feathers and lift-based performance appears to coincide with ontogenetic transitions in locomotor capacity. Younger birds engage in behaviors that require little aerodynamic force and that allow D to contribute to weight support, whereas older birds may expand their behavioral repertoire by flapping with higher tip velocities and generating greater L. Incipient wings are, therefore, uniquely but immediately functional and provide flight-incapable juveniles with access to three-dimensional environments and refugia. Such access may have conferred selective advantages to theropods with protowings during the evolution of avian flight.

Properties of sunspot cycles and hemispheric wings since the 19th century

NASA Astrophysics Data System (ADS)

Leussu, Raisa; Usoskin, Ilya G.; Arlt, Rainer; Mursula, Kalevi

2016-08-01

Aims: The latitudinal evolution of sunspot emergence over the course of the solar cycle, the so-called butterfly diagram, is a fundamental property of the solar dynamo. Here we present a study of the butterfly diagram of sunspot group occurrence for cycles 7-10 and 11-23 using data from a recently digitized sunspot drawings by Samuel Heinrich Schwabe in 1825-1867, and from RGO/USAF/NOAA(SOON) compilation of sunspot groups in 1874-2015. Methods: We developed a new, robust method of hemispheric wing separation based on an analysis of long gaps in sunspot group occurrence in different latitude bands. The method makes it possible to ascribe each sunspot group to a certain wing (solar cycle and hemisphere), and separate the old and new cycle during their overlap. This allows for an improved study of solar cycles compared to the common way of separating the cycles. Results: We separated each hemispheric wing of the butterfly diagram and analysed them with respect to the number of groups appearing in each wing, their lengths, hemispheric differences, and overlaps. Conclusions: The overlaps of successive wings were found to be systematically longer in the northern hemisphere for cycles 7-10, but in the southern hemisphere for cycles 16-22. The occurrence of sunspot groups depicts a systematic long-term variation between the two hemispheres. During Schwabe time, the hemispheric asymmetry was north-dominated during cycle 9 and south-dominated during cycle 10.

The role of Broad in the development of Tribolium castaneum: implications for the evolution of the holometabolous insect pupa.

PubMed

Suzuki, Yuichiro; Truman, James W; Riddiford, Lynn M

2008-02-01

The evolution of complete metamorphosis in insects is a key innovation that has led to the successful diversification of holometabolous insects, yet the origin of the pupa remains an enigma. Here, we analyzed the expression of the pupal specifier gene broad (br), and the effect on br of isoform-specific, double-stranded RNA-mediated silencing, in a basal holometabolous insect, the beetle Tribolium castaneum. All five isoforms are weakly expressed during the penultimate instar and highly expressed during the prepupal period of the final instar. Application of hydroprene, a juvenile hormone analog, during the penultimate instar caused a repeat of the penultimate br expression patterns, and the formation of supernumerary larvae. Use of dsRNA against the br core region, or against a pair of either the br-Z2 or br-Z3 isoform with the br-Z1 or br-Z4 isoform, produced mobile animals with well-differentiated adult-like appendages, but which retained larval-like urogomphi and epidermis. Disruption of either the br-Z2 or the br-Z3 isoform caused the formation of shorter wings. Disruption of both br-Z1 and br-Z4 caused the appearance of pupal traits in the adults, but disruption of br-Z5 had no morphological effect. Our findings show that the br isoform functions are broadly conserved within the Holometabola and suggest that evolution of br isoform expression may have played an important role in the evolution of the pupa in holometabolous insects.

Rhodnius prolixus and Rhodnius robustus-like (Hemiptera, Reduviidae) wing asymmetry under controlled conditions of population density and feeding frequency.

PubMed

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.

Color-pattern analysis of eyespots in butterfly wings: a critical examination of morphogen gradient models.

PubMed

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.

Feedback and Feeding in the Context of Galaxy Evolution with SPICA: Direct Characterisation of Molecular Outflows and Inflows

NASA Astrophysics Data System (ADS)

González-Alfonso, E.; Armus, L.; Carrera, F. J.; Charmandaris, V.; Efstathiou, A.; Egami, E.; Fernández-Ontiveros, J. A.; Fischer, J.; Granato, G. L.; Gruppioni, C.; Hatziminaoglou, E.; Imanishi, M.; Isobe, N.; Kaneda, H.; Koziel-Wierzbowska, D.; Malkan, M. A.; Martín-Pintado, J.; Mateos, S.; Matsuhara, H.; Miniutti, G.; Nakagawa, T.; Pozzi, F.; Rico-Villas, F.; Rodighiero, G.; Roelfsema, P.; Spinoglio, L.; Spoon, H. W. W.; Sturm, E.; van der Tak, F.; Vignali, C.; Wang, L.

2017-11-01

A far-infrared observatory such as the SPace Infrared telescope for Cosmology and Astrophysics, with its unprecedented spectroscopic sensitivity, would unveil the role of feedback in galaxy evolution during the last 10 Gyr of the Universe (z = 1.5-2), through the use of far- and mid-infrared molecular and ionic fine structure lines that trace outflowing and infalling gas. Outflowing gas is identified in the far-infrared through P-Cygni line shapes and absorption blueshifted wings in molecular lines with high dipolar moments, and through emission line wings of fine-structure lines of ionised gas. We quantify the detectability of galaxy-scale massive molecular and ionised outflows as a function of redshift in AGN-dominated, starburst-dominated, and main-sequence galaxies, explore the detectability of metal-rich inflows in the local Universe, and describe the most significant synergies with other current and future observatories that will measure feedback in galaxies via complementary tracers at other wavelengths.

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.

Aerodynamics of wing-assisted incline running in birds.

PubMed

Tobalske, Bret W; Dial, Kenneth P

2007-05-01

Wing-assisted incline running (WAIR) is a form of locomotion in which a bird flaps its wings to aid its hindlimbs in climbing a slope. WAIR is used for escape in ground birds, and the ontogeny of this behavior in precocial birds has been suggested to represent a model analogous to transitional adaptive states during the evolution of powered avian flight. To begin to reveal the aerodynamics of flap-running, we used digital particle image velocimetry (DPIV) and measured air velocity, vorticity, circulation and added mass in the wake of chukar partridge Alectoris chukar as they engaged in WAIR (incline 65-85 degrees; N=7 birds) and ascending flight (85 degrees, N=2). To estimate lift and impulse, we coupled our DPIV data with three-dimensional wing kinematics from a companion study. The ontogeny of lift production was evaluated using three age classes: baby birds incapable of flight [6-8 days post hatching (d.p.h.)] and volant juveniles (25-28 days) and adults (45+ days). All three age classes of birds, including baby birds with partially emerged, symmetrical wing feathers, generated circulation with their wings and exhibited a wake structure that consisted of discrete vortex rings shed once per downstroke. Impulse of the vortex rings during WAIR was directed 45+/-5 degrees relative to horizontal and 21+/-4 degrees relative to the substrate. Absolute values of circulation in vortex cores and induced velocity increased with increasing age. Normalized circulation was similar among all ages in WAIR but 67% greater in adults during flight compared with flap-running. Estimated lift during WAIR was 6.6% of body weight in babies and between 63 and 86% of body weight in juveniles and adults. During flight, average lift was 110% of body weight. Our results reveal for the first time that lift from the wings, rather than wing inertia or profile drag, is primarily responsible for accelerating the body toward the substrate during WAIR, and that partially developed wings, not yet capable of flight, can produce useful lift during WAIR. We predict that neuromuscular control or power output, rather than external wing morphology, constrain the onset of flight ability during development in birds.

The Stretched Lens Array (SLA): A Low-Risk, Cost-Effective Array Offering Wing-Level Performance of 180 W/KG and 300 W/M2 at 300 VDC

NASA Technical Reports Server (NTRS)

ONeill, Mark; Piszczor, Michael F.; Eskenazi, Michael I.; McDanal, A. J.; George, Patrick J.; Botke, Matthew M.; Brandhorst, Henry W.; Edwards, David L.; Jaster, Paul A.; Lyons, Valerie J. (Technical Monitor)

2002-01-01

At IECEC 2001, our team presented a paper on the new stretched lens array (SLA), including its evolution from the successful SCARLET array on the NASA/JPL Deep Space 1 spacecraft. Since that conference, the SLA team has made significant advances in the SLA technology, including component-level improvements, array-level optimization, space environment exposure testing, and prototype hardware fabrication and evaluation. This paper describes the evolved version of the SLA, highlighting recent improvements in the lens, solar cell, photovoltaic receiver, rigid panel structure, and complete solar array wing.

Computational fluid dynamics applications at McDonnel Douglas

NASA Technical Reports Server (NTRS)

Hakkinen, R. J.

1987-01-01

Representative examples are presented of applications and development of advanced Computational Fluid Dynamics (CFD) codes for aerodynamic design at the McDonnell Douglas Corporation (MDC). Transonic potential and Euler codes, interactively coupled with boundary layer computation, and solutions of slender-layer Navier-Stokes approximation are applied to aircraft wing/body calculations. An optimization procedure using evolution theory is described in the context of transonic wing design. Euler methods are presented for analysis of hypersonic configurations, and helicopter rotors in hover and forward flight. Several of these projects were accepted for access to the Numerical Aerodynamic Simulation (NAS) facility at the NASA-Ames Research Center.

NASAs VESGEN: Systems Analysis of Vascular Phenotypes from Stress and Other Signaling Pathways Using GeneLab.

NASA Technical Reports Server (NTRS)

Parsons-Wingerter, Patricia A.; Weitzel, Alexander; Vyas, Ruchi J.; Murray, Matthew C.; Wyatt, Sarah E.

2016-01-01

One fundamental requirement shared by humans with all higher terrestrial life forms, including insect wings, higher land plants and other vertebrates, is a complex, fractally branching vascular system. NASA's VESsel GENeration Analysis (VESGEN) software maps and quantifies vascular trees, networks, and tree-network composites according to weighted physiological rules such as vessel connectivity, tapering and bifurcational branching. According to fluid dynamics, successful vascular transport requires a complex distributed system of highly regulated laminar flow. Microvascular branching rules within vertebrates, dicot leaves and the other organisms therefore display many similarities. One unifying perspective is that vascular patterning offers a useful readout that necessarily integrates complex molecular signaling pathways. VESGEN has elucidated changes in vascular pattern resulting from inflammatory, stress response, developmental and other signaling within numerous tissues and major model organisms studied for Space Biology. For a new VESGEN systems approach, we analyzed differential gene expression in leaves of Arabidopsis thaliana reported by GeneLab (GLDS-7) for spaceflight. Vascular-related changes in leaf gene expression were identified that can potentially be phenocopied by mutants in ground-based experiments. To link transcriptional, protein and other molecular change with phenotype, alterations in the Euclidean and dynamic dimensions (x,y,t) of vascular patterns for Arabidopsis leaves and other model species are being co-localized with signaling patterns of single molecular expression analyzed as information dimensions (i,j,k,...). Previously, Drosophila microarray data returned from space suggested significant changes in genes related to wing venation development that include EGF, Notch, Hedghog, Wingless and Dpp signaling. Phenotypes of increasingly abnormal ectopic wing venation in the (non-spaceflight) Drosophila wing generated by overexpression of a Notch antagonist were analyzed by VESGEN. Other VESGEN research applications include the mouse retina, GI and coronary vessels, avian placental analogs and translational studies in the astronaut retina related to health challenges for long-duration missions.

Imperfect chemical female mimicry in males of the ant Cardiocondyla obscurior

NASA Astrophysics Data System (ADS)

Cremer, Sylvia; D'Ettorre, Patrizia; Drijfhout, Falko P.; Sledge, Matthew F.; Turillazzi, Stefano; Heinze, Jürgen

2008-11-01

Winged and wingless males coexist in the ant Cardiocondyla obscurior. Wingless (“ergatoid”) males never leave their maternal colony and fight remorselessly among each other for the access to emerging females. The peaceful winged males disperse after about 10 days, but beforehand also mate in the nest. In the first 5 days of their life, winged males perform a chemical female mimicry that protects them against attack and even makes them sexually attractive to ergatoid males. When older, the chemical profile of winged males no longer matches that of virgin females; nevertheless, they are still tolerated, which so far has been puzzling. Contrasting this general pattern, we have identified a single aberrant colony in which all winged males were attacked and killed by the ergatoid males. A comparative analysis of the morphology and chemical profile of these untypical attacked winged males and the tolerated males from several normal colonies revealed that normal old males are still performing some chemical mimicry to the virgin queens, though less perfect than in their young ages. The anomalous attacked winged males, on the other hand, had a very different odour to the females. Our study thus exemplifies that the analysis of rare malfunctioning can add valuable insight on functioning under normal conditions and allows the conclusion that older winged males from normal colonies of the ant C. obscurior are guarded through an imperfect chemical female mimicry, still close enough to protect against attacks by the wingless fighters yet dissimilar enough not to elicit their sexual interest.

Geometric Morphometrics of Nine Field Isolates of Aedes aegypti with Different Resistance Levels to Lambda-Cyhalothrin and Relative Fitness of One Artificially Selected for Resistance

PubMed Central

Jaramillo-O., Nicolás; Fonseca-González, Idalyd; Chaverra-Rodríguez, Duverney

2014-01-01

Aedes aegypti, a mosquito closely associated with humans, is the principal vector of dengue virus which currently infects about 400 million people worldwide. Because there is no way to prevent infection, public health policies focus on vector control; but insecticide-resistance threatens them. However, most insecticide-resistant mosquito populations exhibit fitness costs in absence of insecticides, although these costs vary. Research on components of fitness that vary with insecticide-resistance can help to develop policies for effective integrated management and control. We investigated the relationships in wing size, wing shape, and natural resistance levels to lambda-cyhalothrin of nine field isolates. Also we chose one of these isolates to select in lab for resistance to the insecticide. The main life-traits parameters were assessed to investigate the possible fitness cost and its association with wing size and shape. We found that wing shape, more than wing size, was strongly correlated with resistance levels to lambda-cyhalothrin in field isolates, but founder effects of culture in the laboratory seem to change wing shape (and also wing size) more easily than artificial selection for resistance to that insecticide. Moreover, significant fitness costs were observed in response to insecticide-resistance as proved by the diminished fecundity and survival of females in the selected line and the reversion to susceptibility in 20 generations of the non-selected line. As a practical consequence, we think, mosquito control programs could benefit from this knowledge in implementing efficient strategies to prevent the evolution of resistance. In particular, the knowledge of reversion to susceptibility is important because it can help in planning better strategies of insecticide use to keep useful the few insecticide-molecules currently available. PMID:24801598

Genome-wide introgression among distantly related Heliconius butterfly species.

PubMed

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.

Morphological outcomes of gynandromorphism in Lycaeides butterflies (Lepidoptera: Lycaenidae).

PubMed

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.

Identifying the Flow Physics and Modeling Transient Forces on Two-Dimensional Wings

DTIC Science & Technology

2016-09-02

MODELS USING EDMD (a) ( b ) (c) (d) ( e ) (f) (g) (h... Model EDMD Model , β = 0.5 EDMD Model , optimal β ( b ) Model order 5 10 15 20 25 L im it c y c le f re q u e n c y 0.12 0.125 0.13 0.135 0.14 0.145...GP and EDMD nonlinear models in predicting the evolution of POD coefficients for transitional flow past a cylinder, showing (a) time evolution and ( b

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.

Smart wing rotation and trailing-edge vortices enable high frequency mosquito flight

PubMed Central

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.

Wind-tunnel investigation of aerodynamic efficiency of three planar elliptical wings with curvature of quarter-chord line

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.

Differential expression of genes in the alate and apterous morphs of the brown citrus aphid, Toxoptera citricida

PubMed Central

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

A Wingless and Notch double-repression mechanism regulates G1–S transition in the Drosophila wing

PubMed Central

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

PubMed Central

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

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.

Replication of butterfly wing and natural lotus leaf structures by nanoimprint on silica sol-gel films.

PubMed

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.

Distribution and predictors of wing shape and size variability in three sister species of solitary bees

PubMed Central

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

Wakes of lifting and non-lifting bodies: 1. Instabilities & turbulence in the wake of a delta wing. 2. Control of three-dimensional phase dynamics in the wake of a cylinder

NASA Astrophysics Data System (ADS)

Miller, Gregory Dennis

1997-06-01

In the first part of this work, we study the instabilities and turbulent structures in the wake of a delta wing, using extensive flow visualization, hot wire anemometry, and DPIV. We employ a novel free-flight technique in water, coupled with an image processing technique, to study the evolution of the long-wavelength instability of the primary vortex pair. Although secondary vortical structures have received little attention to date, we find that the 'braid wake' vorticity between the vortex pair imposes small lengthscale turbulence around the principal vortices, as well as influence the development of a 'curtain' of vorticity left far above the descending vortex pair. We study the long-wavelength instability of the trailing vortex pair by measuring growth rate and wavelength of the instability directly, and we also measure all of the critical parameters of the vortices (i.e. vortex core radius, vorticity distribution, axial velocity distribution, spacing and circulation), which provide what appears to be the first complete comparison to the theory describing the instability. We find excellent agreement between measured and theoretical growth rates and wavelengths. In the second part of the work, we have devised a method to control the spanwise end conditions and patterns in the wake of a cylinder using 'end suction', which is both continuously-variable and admits transient control. Classical steady-state patterns, such as parallel or oblique shedding, or the 'chevron' patterns, are simply induced. The wake, at a given Reynolds number (Re), is receptive to a continuous range of oblique shedding angles (θ), rather than to discrete angles, and there is excellent agreement with the 'cos θ' formula for oblique-shedding frequencies. We show that the laminar shedding regime exists up to Re of 194, and that the immense disparity among reported critical Re for wake transition (Re = 140-190) can be explained in terms of spanwise end contamination. Our transient experiments have resulted in the discovery of new phenomena such as 'phase shocks' and 'phase expansions', which have excellent agreement with predictions from a Ginzburg- Landau wake model (collaboration with Peter Monkewitz, Lausanne).

Phenotypic Divergence among West European Populations of Reed Bunting Emberiza schoeniclus: The Effects of Migratory and Foraging Behaviours

PubMed Central

Neto, Júlio M.; Gordinho, Luís; Belda, Eduardo J.; Marín, Marcial; Monrós, Juan S.; Fearon, Peter; Crates, Ross

2013-01-01

Divergent selection and local adaptation are responsible for many phenotypic differences between populations, potentially leading to speciation through the evolution of reproductive barriers. Here we evaluated the morphometric divergence among west European populations of Reed Bunting in order to determine the extent of local adaptation relative to two important selection pressures often associated with speciation in birds: migration and diet. We show that, as expected by theory, migratory E. s. schoeniclus had longer and more pointed wings and a slightly smaller body mass than the resident subspecies, with the exception of E. s. lusitanica, which despite having rounder wings was the smallest of all subspecies. Tail length, however, did not vary according to the expectation (shorter tails in migrants) probably because it is strongly correlated with wing length and might take longer to evolve. E. s. witherbyi, which feed on insects hiding inside reed stems during the winter, had a very thick, stubby bill. In contrast, northern populations, which feed on seeds, had thinner bills. Despite being much smaller, the southern E. s. lusitanica had a significantly thicker, longer bill than migratory E. s. schoeniclus, whereas birds from the UK population had significantly shorter, thinner bills. Geometric morphometric analyses revealed that the southern subspecies have a more convex culmen than E. s. schoeniclus, and E. s. lusitanica differs from the nominate subspecies in bill shape to a greater extent than in linear bill measurements, especially in males. Birds with a more convex culmen are thought to exert a greater strength at the bill tip, which is in agreement with their feeding technique. Overall, the three subspecies occurring in Western Europe differ in a variety of traits following the patterns predicted from their migratory and foraging behaviours, strongly suggesting that these birds have became locally adapted through natural selection. PMID:23667594

Fossil evidence for key innovations in the evolution of insect diversity.

PubMed

Nicholson, David B; Ross, Andrew J; Mayhew, Peter J

2014-10-22

Explaining the taxonomic richness of the insects, comprising over half of all described species, is a major challenge in evolutionary biology. Previously, several evolutionary novelties (key innovations) have been posited to contribute to that richness, including the insect bauplan, wings, wing folding and complete metamorphosis, but evidence over their relative importance and modes of action is sparse and equivocal. Here, a new dataset on the first and last occurrences of fossil hexapod (insects and close relatives) families is used to show that basal families of winged insects (Palaeoptera, e.g. dragonflies) show higher origination and extinction rates in the fossil record than basal wingless groups (Apterygota, e.g. silverfish). Origination and extinction rates were maintained at levels similar to Palaeoptera in the more derived Polyneoptera (e.g. cockroaches) and Paraneoptera (e.g. true bugs), but extinction rates subsequently reduced in the very rich group of insects with complete metamorphosis (Holometabola, e.g. beetles). Holometabola show evidence of a recent slow-down in their high net diversification rate, whereas other winged taxa continue to diversify at constant but low rates. These data suggest that wings and complete metamorphosis have had the most effect on family-level insect macroevolution, and point to specific mechanisms by which they have influenced insect diversity through time. © 2014 The Author(s) Published by the Royal Society. All rights reserved.

Camouflage through an active choice of a resting spot and body orientation in moths.

PubMed

Kang, C-K; Moon, J-Y; Lee, S-I; Jablonski, P G

2012-09-01

Cryptic colour patterns in prey are classical examples of adaptations to avoid predation, but we still know little about behaviours that reinforce the match between animal body and the background. For example, moths avoid predators by matching their colour patterns with the background. Active choice of a species-specific body orientation has been suggested as an important function of body positioning behaviour performed by moths after landing on the bark. However, the contribution of this behaviour to moths' crypticity has not been directly measured. From observations of geometrid moths, Hypomecis roboraria and Jankowskia fuscaria, we determined that the positioning behaviour, which consists of walking and turning the body while repeatedly lifting and lowering the wings, resulted in new resting spots and body orientations in J. fuscaria and in new resting spots in H. roboraria. The body positioning behaviour of the two species significantly decreased the probability of visual detection by humans, who viewed photographs of the moths taken before and after the positioning behaviour. This implies that body positioning significantly increases the camouflage effect provided by moth's cryptic colour pattern regardless of whether the behaviour involves a new body orientation or not. Our study demonstrates that the evolution of morphological adaptations, such as colour pattern of moths, cannot be fully understood without taking into account a behavioural phenotype that coevolved with the morphology for increasing the adaptive value of the morphological trait. © 2012 The Authors. Journal of Evolutionary Biology © 2012 European Society For Evolutionary Biology.

Origin, development, and evolution of butterfly eyespots.

PubMed

Monteiro, Antónia

2015-01-07

This article reviews the latest developments in our understanding of the origin, development, and evolution of nymphalid butterfly eyespots. Recent contributions to this field include insights into the evolutionary and developmental origin of eyespots and their ancestral deployment on the wing, the evolution of eyespot number and eyespot sexual dimorphism, and the identification of genes affecting eyespot development and black pigmentation. I also compare features of old and more recently proposed models of eyespot development and propose a schematic for the genetic regulatory architecture of eyespots. Using this schematic I propose two hypotheses for why we observe limits to morphological diversity across these serially homologous traits.

Knowledge Discovery for Transonic Regional-Jet Wing through Multidisciplinary Design Exploration

NASA Astrophysics Data System (ADS)

Chiba, Kazuhisa; Obayashi, Shigeru; Morino, Hiroyuki

Data mining is an important facet of solving multi-objective optimization problem. Because it is one of the effective manner to discover the design knowledge in the multi-objective optimization problem which obtains large data. In the present study, data mining has been performed for a large-scale and real-world multidisciplinary design optimization (MDO) to provide knowledge regarding the design space. The MDO among aerodynamics, structures, and aeroelasticity of the regional-jet wing was carried out using high-fidelity evaluation models on the adaptive range multi-objective genetic algorithm. As a result, nine non-dominated solutions were generated and used for tradeoff analysis among three objectives. All solutions evaluated during the evolution were analyzed for the tradeoffs and influence of design variables using a self-organizing map to extract key features of the design space. Although the MDO results showed the inverted gull-wings as non-dominated solutions, one of the key features found by data mining was the non-gull wing geometry. When this knowledge was applied to one optimum solution, the resulting design was found to have better performance compared with the original geometry designed in the conventional manner.

Spatially resolved chemical analysis of cicada wings using laser-ablation electrospray ionization (LAESI) imaging mass spectrometry (IMS).

PubMed

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

PubMed Central

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

Ontogeny of lift and drag production in ground birds

PubMed Central

Heers, Ashley M.; Tobalske, Bret W.; Dial, Kenneth P.

2011-01-01

The juvenile period is often a crucial interval for selective pressure on locomotor ability. Although flight is central to avian biology, little is known about factors that limit flight performance during development. To improve understanding of flight ontogeny, we used a propeller (revolving wing) model to test how wing shape and feather structure influence aerodynamic performance during development in the precocial chukar partridge (Alectoris chukar, 4 to >100 days post hatching). We spun wings in mid-downstroke posture and measured lift (L) and drag (D) using a force plate upon which the propeller assembly was mounted. Our findings demonstrate a clear relationship between feather morphology and aerodynamic performance. Independent of size and velocity, older wings with stiffer and more asymmetrical feathers, high numbers of barbicels and a high degree of overlap between barbules generate greater L and L:D ratios than younger wings with flexible, relatively symmetrical and less cohesive feathers. The gradual transition from immature feathers and drag-based performance to more mature feathers and lift-based performance appears to coincide with ontogenetic transitions in locomotor capacity. Younger birds engage in behaviors that require little aerodynamic force and that allow D to contribute to weight support, whereas older birds may expand their behavioral repertoire by flapping with higher tip velocities and generating greater L. Incipient wings are, therefore, uniquely but immediately functional and provide flight-incapable juveniles with access to three-dimensional environments and refugia. Such access may have conferred selective advantages to theropods with protowings during the evolution of avian flight. PMID:21307057

Chromosomal rearrangements maintain a polymorphic supergene controlling butterfly mimicry

PubMed Central

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

First discovery of fossil winged seeds of Pinus L. (family Pinaceae) from the Indian Cenozoic and its palaeobiogeographic significance

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.

Energy Conservation and the Promotion of Legionella pneumophila Growth: The Probable Role of Heat Exchangers in a Nosocomial Outbreak.

PubMed

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.

Selection of Valid Reference Genes for Reverse Transcription Quantitative PCR Analysis in Heliconius numata (Lepidoptera: Nymphalidae)

PubMed Central

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

Characterising the Phenotypic Diversity of Papilio dardanus Wing Patterns Using an Extensive Museum Collection

PubMed Central

Thompson, Martin J.; Timmermans, Martijn J. T. N.

2014-01-01

The history of 20th Century evolutionary biology can be followed through the study of mimetic butterflies. From the initial findings of discontinuous polymorphism through the debates regarding the evolution of mimicry and the step-size of evolutionary change, to the studies on supergene evolution and molecular characterisation of butterfly genomes, mimetic butterflies have been at the heart of evolutionary thought for over 100 years. During this time, few species have received as much attention and in-depth study as Papilio dardanus. To assist all aspects of mimicry research, we present a complete data-derived overview of the extent of polymorphism within this species. Using historical samples permanently held by the NHM London, we document the extent of phenotypic variation and characterise the diversity present in each of the subspecies and how it varies across Africa. We also demonstrate an association between “imperfect” mimetic forms and the transitional race formed in the area where Eastern and Western African populations meet around Lake Victoria. We present a novel portal for access to this collection, www.mimeticbutterflies.org, allowing remote access to this unique repository. It is hoped that this online resource can act as a nucleus for the sharing and dissemination of other collections databases and imagery connected with mimetic butterflies. PMID:24837717

Wing patterning gene redefines the mimetic history of Heliconius butterflies.

PubMed

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.

Bone laminarity in the avian forelimb skeleton and its relationship to flight mode: testing functional interpretations.

PubMed

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.

Bioinspired spring origami

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.

Proprioceptive input patterns elevator activity in the locust flight system.

PubMed

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.

Automatic identification of bird targets with radar via patterns produced by wing flapping.

PubMed

Zaugg, Serge; Saporta, Gilbert; van Loon, Emiel; Schmaljohann, Heiko; Liechti, Felix

2008-09-06

Bird identification with radar is important for bird migration research, environmental impact assessments (e.g. wind farms), aircraft security and radar meteorology. In a study on bird migration, radar signals from birds, insects and ground clutter were recorded. Signals from birds show a typical pattern due to wing flapping. The data were labelled by experts into the four classes BIRD, INSECT, CLUTTER and UFO (unidentifiable signals). We present a classification algorithm aimed at automatic recognition of bird targets. Variables related to signal intensity and wing flapping pattern were extracted (via continuous wavelet transform). We used support vector classifiers to build predictive models. We estimated classification performance via cross validation on four datasets. When data from the same dataset were used for training and testing the classifier, the classification performance was extremely to moderately high. When data from one dataset were used for training and the three remaining datasets were used as test sets, the performance was lower but still extremely to moderately high. This shows that the method generalizes well across different locations or times. Our method provides a substantial gain of time when birds must be identified in large collections of radar signals and it represents the first substantial step in developing a real time bird identification radar system. We provide some guidelines and ideas for future research.

Ectopic KNOX Expression Affects Plant Development by Altering Tissue Cell Polarity and Identity[OPEN

PubMed Central

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

Ultraviolet and optical observations of metal deficient red giants and chromospheric models

NASA Technical Reports Server (NTRS)

Duprele, A. K.; Avrett, E. H.; Hartmann, L.; Smith, G.

1984-01-01

Three metal deficient field stars were observed in the ultraviolet and optical spectral regions: HD 165195, HD 110281, and HD 232078. High dispersion spectra near H alpha, and low dispersion, long wavelength IUE spectra were obtained. The H alpha profiles have strong asymmetric emission with absorption cores that are frequently asymmetric. The surface flux of Mg II lines is similar to that of luminous Pop I stars in spite of the lower metal abundance. Semi-empirical atmospheric models suggest that the characteristic emission in the wings of the H alpha line can arise within static chromospheres. Radial expansion gives an asymmetric, blue-shifted H alpha core accompanied by greater emission in the red line wing than the blue wing. Wind models with extended atmospheres suggest mass loss rates - 2 billion M/yr. Thus H alpha provides no evidence that steady mass loss is substantial enough to significantly affect the evolution of stars on the red giant branch of globular clusters.

Sumoylation Modulates the Activity of Spalt-like Proteins during Wing Development in Drosophila*

PubMed Central

Sánchez, Jonatan; Talamillo, Ana; Lopitz-Otsoa, Fernando; Pérez, Coralia; Hjerpe, Roland; Sutherland, James D.; Herboso, Leire; Rodríguez, Manuel S.; Barrio, Rosa

2010-01-01

The Spalt-like family of zinc finger transcription factors is conserved throughout evolution and is involved in fundamental processes during development and during embryonic stem cell maintenance. Although human SALL1 is modified by SUMO-1 in vitro, it is not known whether this post-translational modification plays a role in regulating the activity of this family of transcription factors. Here, we show that the Drosophila Spalt transcription factors are modified by sumoylation. This modification influences their nuclear localization and capacity to induce vein formation through the regulation of target genes during wing development. Furthermore, spalt genes interact genetically with the sumoylation machinery to repress vein formation in intervein regions and to attain the wing final size. Our results suggest a new level of regulation of Sall activity in vivo during animal development through post-translational modification by sumoylation. The evolutionary conservation of this family of transcription factors suggests a functional role for sumoylation in vertebrate Sall members. PMID:20562097

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.

A large-scale, in vivo transcription factor screen defines bivalent chromatin as a key property of regulatory factors mediating Drosophila wing development

PubMed Central

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

Efficiency of lift production in flapping and gliding flight of swifts.

PubMed

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

PubMed Central

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

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.

Superfast high-resolution absolute 3D recovery of a stabilized flapping flight process.

PubMed

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.

Inwardly rectifying potassium channels influence Drosophila wing morphogenesis by regulating Dpp release.

PubMed

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.

Proteome-wide association studies identify biochemical modules associated with a wing-size phenotype in Drosophila melanogaster.

PubMed

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.

Restructuring the rotor analysis program C-60

NASA Technical Reports Server (NTRS)

1985-01-01

The continuing evolution of the rotary wing industry demands increasing analytical capabilities. To keep up with this demand, software must be structured to accommodate change. The approach discussed for meeting this demand is to restructure an existing analysis. The motivational factors, basic principles, application techniques, and practical lessons from experience with this restructuring effort are reviewed.

Life-history evolution in ants: the case of Cardiocondyla

PubMed Central

2017-01-01

Ants are important components of most terrestrial habitats, and a better knowledge of the diversity of their life histories is essential to understand many aspects of ecosystem functioning. The myrmicine genus Cardiocondyla shows a wide range of colony structures, reproductive behaviours, queen and male lifespans, and habitat use. Reconstructing the evolutionary pathways of individual and social phenotypic traits suggests that the ancestral life history of Cardiocondyla was characterized by the presence of multiple, short-lived queens in small-sized colonies and a male polyphenism with winged dispersers and wingless fighters, which engage in lethal combat over female sexuals within their natal nests. Single queening, queen polyphenism, the loss of winged males and tolerance among wingless males appear to be derived traits that evolved with changes in nesting habits, colony size and the spread from tropical to seasonal environments. The aim of this review is to bring together the information on life-history evolution in Cardiocondyla and to highlight the suitability of this genus for functional genomic studies of adaptation, phenotypic plasticity, senescence, invasiveness and other key life-history traits of ants. PMID:28298341

Red & black or black & white? Phylogeny of the Araschnia butterflies (Lepidoptera: Nymphalidae) and evolution of seasonal polyphenism.

PubMed

Fric, Z; Konvicka, M; Zrzavy, J

2004-03-01

Phylogeny of the butterfly genera Araschnia, Mynes, Symbrenthia and Brensymthia (Lepidoptera: Nymphalidae: Nymphalini) is reconstructed, based on 140 morphological and ecological characters. The resulting tree shows that Araschnia is a sister group of the clade including Symbrenthia, Mynes and Brensymthia (Symbrenthia is paraphyletic in the respect of remaining genera; Symbrenthia hippalus is a derived species of Mynes). The species-level relationships within Araschnia are robustly supported as follows: (A. davidis (prorsoides ((zhangi doris) (dohertyi (levana burejana))))). Analysis of the wing colour-pattern characters linked with the seasonal polyphenism in the Araschnia species suggests that the black and white coloration of the long-day (summer) generation is apomorphic. Biogeographically, the origin of polyphenism in Araschnia predates the dispersal of some Araschnia species towards the Palaearctic temperate zone, and the ecological cause of the polyphenism itself is then probably not linked with thermoregulation. The possible mimetic/cryptic scenarios for the origin of Araschnia polyphenism are discussed.

Wing morphology variations in a natural population of Phlebotomus tobbi Adler and Theodor 1930.

PubMed

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.

Unified aeroacoustics analysis for high speed turboprop aerodynamics and noise. Volume 2: Development of theory for wing shielding

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.

Scaling the Drosophila Wing: TOR-Dependent Target Gene Access by the Hippo Pathway Transducer Yorkie

PubMed Central

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

Positive selection of a duplicated UV-sensitive visual pigment coincides with wing pigment evolution in Heliconius butterflies

PubMed Central

Briscoe, Adriana D.; Bybee, Seth M.; Bernard, Gary D.; Yuan, Furong; Sison-Mangus, Marilou P.; Reed, Robert D.; Warren, Andrew D.; Llorente-Bousquets, Jorge; Chiao, Chuan-Chin

2010-01-01

The butterfly Heliconius erato can see from the UV to the red part of the light spectrum with color vision proven from 440 to 640 nm. Its eye is known to contain three visual pigments, rhodopsins, produced by an 11-cis-3-hydroxyretinal chromophore together with long wavelength (LWRh), blue (BRh) and UV (UVRh1) opsins. We now find that H. erato has a second UV opsin mRNA (UVRh2)—a previously undescribed duplication of this gene among Lepidoptera. To investigate its evolutionary origin, we screened eye cDNAs from 14 butterfly species in the subfamily Heliconiinae and found both copies only among Heliconius. Phylogeny-based tests of selection indicate positive selection of UVRh2 following duplication, and some of the positively selected sites correspond to vertebrate visual pigment spectral tuning residues. Epi-microspectrophotometry reveals two UV-absorbing rhodopsins in the H. erato eye with λmax = 355 nm and 398 nm. Along with the additional UV opsin, Heliconius have also evolved 3-hydroxy-DL-kynurenine (3-OHK)-based yellow wing pigments not found in close relatives. Visual models of how butterflies perceive wing color variation indicate this has resulted in an expansion of the number of distinguishable yellow colors on Heliconius wings. Functional diversification of the UV-sensitive visual pigments may help explain why the yellow wing pigments of Heliconius are so colorful in the UV range compared to the yellow pigments of close relatives lacking the UV opsin duplicate. PMID:20133601

The Effect of Diet Quality and Wing Morph on Male and Female Reproductive Investment in a Nuptial Feeding Ground Cricket

PubMed Central

Hall, Matthew D.; Bussière, Luc F.; Brooks, Robert

2008-01-01

A common approach in the study of life-history trade-off evolution is to manipulate the nutrient content of diets during the life of an individual in order observe how the acquisition of resources influences the relationship between reproduction, lifespan and other life-history parameters such as dispersal. Here, we manipulate the quality of diet that replicate laboratory populations received as a thorough test of how diet quality influences the life-history trade-offs associated with reproductive investment in a nuptial feeding Australian ground cricket (Pteronemobius sp.). In this species, both males and females make significant contributions to the production of offspring, as males provide a nuptial gift by allowing females to chew on a modified tibial spur during copulation and feed directing on their haemolymph. Individuals also have two distinct wing morphs, a short-winged flightless morph and a long-winged morph that has the ability to disperse. By manipulating the quality of diet over seven generations, we found that the reproductive investment of males and females were affected differently by the diet quality treatment and wing morph of the individual. We discuss the broader implications of these findings including the differences in how males and females balance current and future reproductive effort in nuptial feeding insects, the changing nature of sexual selection when diets vary, and how the life-history trade-offs associated with the ability to disperse are expected to differ among populations. PMID:18927614

Positive selection of a duplicated UV-sensitive visual pigment coincides with wing pigment evolution in Heliconius butterflies.

PubMed

Briscoe, Adriana D; Bybee, Seth M; Bernard, Gary D; Yuan, Furong; Sison-Mangus, Marilou P; Reed, Robert D; Warren, Andrew D; Llorente-Bousquets, Jorge; Chiao, Chuan-Chin

2010-02-23

The butterfly Heliconius erato can see from the UV to the red part of the light spectrum with color vision proven from 440 to 640 nm. Its eye is known to contain three visual pigments, rhodopsins, produced by an 11-cis-3-hydroxyretinal chromophore together with long wavelength (LWRh), blue (BRh) and UV (UVRh1) opsins. We now find that H. erato has a second UV opsin mRNA (UVRh2)-a previously undescribed duplication of this gene among Lepidoptera. To investigate its evolutionary origin, we screened eye cDNAs from 14 butterfly species in the subfamily Heliconiinae and found both copies only among Heliconius. Phylogeny-based tests of selection indicate positive selection of UVRh2 following duplication, and some of the positively selected sites correspond to vertebrate visual pigment spectral tuning residues. Epi-microspectrophotometry reveals two UV-absorbing rhodopsins in the H. erato eye with lambda(max) = 355 nm and 398 nm. Along with the additional UV opsin, Heliconius have also evolved 3-hydroxy-DL-kynurenine (3-OHK)-based yellow wing pigments not found in close relatives. Visual models of how butterflies perceive wing color variation indicate this has resulted in an expansion of the number of distinguishable yellow colors on Heliconius wings. Functional diversification of the UV-sensitive visual pigments may help explain why the yellow wing pigments of Heliconius are so colorful in the UV range compared to the yellow pigments of close relatives lacking the UV opsin duplicate.

Transition from wing to leg forces during landing in birds.

PubMed

Provini, Pauline; Tobalske, Bret W; Crandell, Kristen E; Abourachid, Anick

2014-08-01

Transitions to and from the air are critical for aerial locomotion and likely shaped the evolution of flying animals. Research on take-off demonstrates that legs generate greater body accelerations compared with wings, and thereby contribute more to initial flight velocity. Here, we explored coordination between wings and legs in two species with different wingbeat styles, and quantified force production of these modules during the final phase of landing. We used the same birds that we had previously studied during take-off: zebra finch (Taeniopygia guttata, N=4) and diamond dove (Geopelia cuneata, N=3). We measured kinematics using high-speed video, aerodynamics using particle image velocimetry, and ground-reaction forces using a perch mounted on a force plate. In contrast with the first three wingbeats of take-off, the final four wingbeats during landing featured ~2 times greater force production. Thus, wings contribute proportionally more to changes in velocity during the last phase of landing compared with the initial phase of take-off. The two species touched down at the same velocity (~1 m s(-1)), but they exhibited significant differences in the timing of their final wingbeat relative to touchdown. The ratio of average wing force to peak leg force was greater in diamond doves than in zebra finches. Peak ground reaction forces during landing were ~50% of those during take-off, consistent with the birds being motivated to control landing. Likewise, estimations of mechanical energy flux for both species indicate that wings produce 3-10 times more mechanical work within the final wingbeats of flight compared with the kinetic energy of the body absorbed by legs during ground contact. © 2014. Published by The Company of Biologists Ltd.

Transition from leg to wing forces during take-off in birds.

PubMed

Provini, Pauline; Tobalske, Bret W; Crandell, Kristen E; Abourachid, Anick

2012-12-01

Take-off mechanics are fundamental to the ecology and evolution of flying animals. Recent research has revealed that initial take-off velocity in birds is driven mostly by hindlimb forces. However, the contribution of the wings during the transition to air is unknown. To investigate this transition, we integrated measurements of both leg and wing forces during take-off and the first three wingbeats in zebra finch (Taeniopygia guttata, body mass 15 g, N=7) and diamond dove (Geopelia cuneata, body mass 50 g, N=3). We measured ground reaction forces produced by the hindlimbs using a perch mounted on a force plate, whole-body and wing kinematics using high-speed video, and aerodynamic forces using particle image velocimetry (PIV). Take-off performance was generally similar between species. When birds were perched, an acceleration peak produced by the legs contributed to 85±1% of the whole-body resultant acceleration in finch and 77±6% in dove. At lift-off, coincident with the start of the first downstroke, the percentage of hindlimb contribution to initial flight velocity was 93.6±0.6% in finch and 95.2±0.4% in dove. In finch, the first wingbeat produced 57.9±3.4% of the lift created during subsequent wingbeats compared with 62.5±2.2% in dove. Advance ratios were <0.5 in both species, even when taking self-convection of shed vortices into account, so it was likely that wing-wake interactions dominated aerodynamics during wingbeats 2 and 3. These results underscore the relatively low contribution of the wings to initial take-off, and reveal a novel transitional role for the first wingbeat in terms of force production.

Kinematics and aerodynamics of avian upstrokes during slow flight.

PubMed

Crandell, Kristen E; Tobalske, Bret W

2015-08-01

Slow flight is extremely energetically costly per unit time, yet highly important for takeoff and survival. However, at slow speeds it is presently thought that most birds do not produce beneficial aerodynamic forces during the entire wingbeat: instead they fold or flex their wings during upstroke, prompting the long-standing prediction that the upstroke produces trivial forces. There is increasing evidence that the upstroke contributes to force production, but the aerodynamic and kinematic mechanisms remain unknown. Here, we examined the wingbeat cycle of two species: the diamond dove (Geopelia cuneata) and zebra finch (Taeniopygia guttata), which exhibit different upstroke styles - a wingtip-reversal and flexed-wing upstroke, respectively. We used a combination of particle image velocimetry and near-wake streamline measures alongside detailed 3D kinematics. We show that during the middle of the wingtip-reversal upstroke, the hand-wing has a high angular velocity (15.3±0.8 deg ms(-1)) and translational speed (8.4±0.6 m s(-1)). The flexed-wing upstroke, in contrast, has low wingtip speed during mid-upstroke. Instead, later in the stroke cycle, during the transition from upstroke to downstroke, it exhibits higher angular velocities (45.5±13.8 deg ms(-1)) and translational speeds (11.0±1.9 m s(-1)). Aerodynamically, the wingtip-reversal upstroke imparts momentum to the wake, with entrained air shed backward (visible as circulation of 14.4±0.09 m(2) s(-1)). In contrast, the flexed-wing upstroke imparts minimal momentum. Clap and peel in the dove enhances the time course for circulation production on the wings, and provides new evidence of convergent evolution on time-varying aerodynamic mechanisms during flapping in insects and birds. © 2015. Published by The Company of Biologists Ltd.

Evolution of avian flight: muscles and constraints on performance

PubMed Central

2016-01-01

Competing hypotheses about evolutionary origins of flight are the ‘fundamental wing-stroke’ and ‘directed aerial descent’ hypotheses. Support for the fundamental wing-stroke hypothesis is that extant birds use flapping of their wings to climb even before they are able to fly; there are no reported examples of incrementally increasing use of wing movements in gliding transitioning to flapping. An open question is whether locomotor styles must evolve initially for efficiency or if they might instead arrive due to efficacy. The proximal muscles of the avian wing output work and power for flight, and new research is exploring functions of the distal muscles in relation to dynamic changes in wing shape. It will be useful to test the relative contributions of the muscles of the forearm compared with inertial and aerodynamic loading of the wing upon dynamic morphing. Body size has dramatic effects upon flight performance. New research has revealed that mass-specific muscle power declines with increasing body mass among species. This explains the constraints associated with being large. Hummingbirds are the only species that can sustain hovering. Their ability to generate force, work and power appears to be limited by time for activation and deactivation within their wingbeats of high frequency. Most small birds use flap-bounding flight, and this flight style may offer an energetic advantage over continuous flapping during fast flight or during flight into a headwind. The use of flap-bounding during slow flight remains enigmatic. Flap-bounding birds do not appear to be constrained to use their primary flight muscles in a fixed manner. To improve understanding of the functional significance of flap-bounding, the energetic costs and the relative use of alternative styles by a given species in nature merit study. This article is part of the themed issue ‘Moving in a moving medium: new perspectives on flight’. PMID:27528773

Vorticity Transport on a Flexible Wing in Stall Flutter

NASA Astrophysics Data System (ADS)

Akkala, James; Buchholz, James; Farnsworth, John; McLaughlin, Thomas

2014-11-01

The circulation budget within dynamic stall vortices was investigated on a flexible NACA 0018 wing model of aspect ratio 6 undergoing stall flutter. The wing had an initial angle of attack of 6 degrees, Reynolds number of 1 . 5 ×105 and large-amplitude, primarily torsional, limit cycle oscillations were observed at a reduced frequency of k = πfc / U = 0 . 1 . Phase-locked stereo PIV measurements were obtained at multiple chordwise planes around the 62.5% and 75% spanwise locations to characterize the flow field within thin volumetric regions over the suction surface. Transient surface pressure measurements were used to estimate boundary vorticity flux. Recent analyses on plunging and rotating wings indicates that the magnitude of the pressure-gradient-driven boundary flux of secondary vorticity is a significant fraction of the magnitude of the convective flux from the separated leading-edge shear layer, suggesting that the secondary vorticity plays a significant role in regulating the strength of the primary vortex. This phenomenon is examined in the present case, and the physical mechanisms governing the growth and evolution of the dynamic stall vortices are explored. This work was supported by the Air Force Office of Scientific Research through the Flow Interactions and Control Program monitored by Dr. Douglas Smith, and through the 2014 AFOSR/ASEE Summer Faculty Fellowship Program (JA and JB).

Characterisation of cuticular nanostructures on surfaces of insects by atomic force microscopy: mining evolution for smart structures

NASA Astrophysics Data System (ADS)

Watson, Gregory S.; Blach, Jolanta A.

2002-11-01

The optical properties of insect nano-structures have been extensively studied. In particular, nano-scale ordered arrays have been reported from studies of the corneal surfaces of some insects and of insect wings showing anti-reflective properties. These arrays have been ascribed to evolutionary adaptation and survival value arising from increased visual capacity and better camouflage against predators. In this study we show that the Atomic Force Microscope (AFM) can effectively reveal and quantify the three dimensional structures of nano-arrays on moth eyes and cicada wings. It is also shown that the arrays present an ideal surface for in situ characterisation of the AFM probe/tip. In addition, a new structure is presented which has been discovered on a termite wing. The structure is similar to that found on the cicada wing, but has a much larger 'lattice parameter' for the ordered array. The function(s) of the array is unknown at present. It could be effective as an anti-reflective coating, but would then be active in the infra-red region of the light spectrum. Alternatively, it may confer evolutionary advantage by virtue of its mechanical strength, or it may improve the aerodynamics of flying. The study demonstrates that natural selection may be a rich source of 'smart' structures.

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 ...

Ice Roughness and Thickness Evolution on a Swept NACA 0012 Airfoil

NASA Technical Reports Server (NTRS)

McClain, Stephen T.; Vargas, Mario; Tsao, Jen-Ching

2017-01-01

Several recent studies have been performed in the Icing Research Tunnel (IRT) at NASA Glenn Research Center focusing on the evolution, spatial variations, and proper scaling of ice roughness on airfoils without sweep exposed to icing conditions employed in classical roughness studies. For this study, experiments were performed in the IRT to investigate the ice roughness and thickness evolution on a 91.44-cm (36-in.) chord NACA 0012 airfoil, swept at 30-deg with 0deg angle of attack, and exposed to both Appendix C and Appendix O (SLD) icing conditions. The ice accretion event times used in the study were less than the time required to form substantially three-dimensional structures, such as scallops, on the airfoil surface. Following each ice accretion event, the iced airfoils were scanned using a ROMER Absolute Arm laser-scanning system. The resulting point clouds were then analyzed using the self-organizing map approach of McClain and Kreeger to determine the spatial roughness variations along the surfaces of the iced airfoils. The resulting measurements demonstrate linearly increasing roughness and thickness parameters with ice accretion time. Further, when compared to dimensionless or scaled results from unswept airfoil investigations, the results of this investigation indicate that the mechanisms for early stage roughness and thickness formation on swept wings are similar to those for unswept wings.

A second new species of ice crawlers from China (Insecta: Grylloblattodea), with thorax evolution and the prediction of potential distribution.

PubMed

Bai, Ming; Jarvis, Karl; Wang, Shu-Yong; Song, Ke-Qing; Wang, Yan-Ping; Wang, Zhi-Liang; Li, Wen-Zhu; Wang, Wei; Yang, Xing-Ke

2010-09-22

Modern grylloblattids are one of the least diverse of the modern insect orders. The thorax changes in morphology might be associated with the changes of the function of the forelegs, wing loss, changes in behavior and adaptation to habitat. As temperature is the main barrier for migration of modern grylloblattids, the range of each species is extremely limited. The potential distribution areas of grylloblattids remain unclear. A second new species of ice crawlers (Insecta: Grylloblattodea), Grylloblattella cheni Bai, Wang et Yang sp. nov., is described from China. The distribution map and key to species of Grylloblattella are given. A comparison of the thorax of extant and extinct Grylloblattodea is presented, with an emphasis on the pronotum using geometric morphometric analysis, which may reflect thorax adaptation and the evolution of Grylloblattodea. Potential global distribution of grylloblattids is inferred. Highly diversified pronota of extinct Grylloblattodea may reflect diverse habitats and niches. The relatively homogeneous pronota of modern grylloblattids might be explained by two hypotheses: synapomorphy or convergent evolution. Most fossils of Grylloblattodea contain an obviously longer meso- and metathorax than prothorax. The length of the meso- and metathorax of modern grylloblattids is normally shorter than the prothorax. This may be associated with the wing loss, which is accompanied by muscle reduction and changes to the thoracic skeleton system. Threats to grylloblattids and several conservation comments are also provided.

Analysis of four achaete-scute homologs in Bombyx mori reveals new viewpoints of the evolution and functions of this gene family

PubMed Central

Zhou, Qingxiang; Zhang, Tianyi; Xu, Weihua; Yu, Linlin; Yi, Yongzhu; Zhang, Zhifang

2008-01-01

Background achaete-scute complexe (AS-C) has been widely studied at genetic, developmental and evolutional levels. Genes of this family encode proteins containing a highly conserved bHLH domain, which take part in the regulation of the development of central nervous system and peripheral nervous system. Many AS-C homologs have been isolated from various vertebrates and invertebrates. Also, AS-C genes are duplicated during the evolution of Diptera. Functions besides neural development controlling have also been found in Drosophila AS-C genes. Results We cloned four achaete-scute homologs (ASH) from the lepidopteran model organism Bombyx mori, including three proneural genes and one neural precursor gene. Proteins encoded by them contained the characteristic bHLH domain and the three proneural ones were also found to have the C-terminal conserved motif. These genes regulated promoter activity through the Class A E-boxes in vitro. Though both Bm-ASH and Drosophila AS-C have four members, they are not in one by one corresponding relationships. Results of RT-PCR and real-time PCR showed that Bm-ASH genes were expressed in different larval tissues, and had well-regulated expressional profiles during the development of embryo and wing/wing disc. Conclusion There are four achaete-scute homologs in Bombyx mori, the second insect having four AS-C genes so far, and these genes have multiple functions in silkworm life cycle. AS-C gene duplication in insects occurs after or parallel to, but not before the taxonomic order formation during evolution. PMID:18321391

Analysis of four achaete-scute homologs in Bombyx mori reveals new viewpoints of the evolution and functions of this gene family.

PubMed

Zhou, Qingxiang; Zhang, Tianyi; Xu, Weihua; Yu, Linlin; Yi, Yongzhu; Zhang, Zhifang

2008-03-06

achaete-scute complexe (AS-C) has been widely studied at genetic, developmental and evolutional levels. Genes of this family encode proteins containing a highly conserved bHLH domain, which take part in the regulation of the development of central nervous system and peripheral nervous system. Many AS-C homologs have been isolated from various vertebrates and invertebrates. Also, AS-C genes are duplicated during the evolution of Diptera. Functions besides neural development controlling have also been found in Drosophila AS-C genes. We cloned four achaete-scute homologs (ASH) from the lepidopteran model organism Bombyx mori, including three proneural genes and one neural precursor gene. Proteins encoded by them contained the characteristic bHLH domain and the three proneural ones were also found to have the C-terminal conserved motif. These genes regulated promoter activity through the Class A E-boxes in vitro. Though both Bm-ASH and Drosophila AS-C have four members, they are not in one by one corresponding relationships. Results of RT-PCR and real-time PCR showed that Bm-ASH genes were expressed in different larval tissues, and had well-regulated expressional profiles during the development of embryo and wing/wing disc. There are four achaete-scute homologs in Bombyx mori, the second insect having four AS-C genes so far, and these genes have multiple functions in silkworm life cycle. AS-C gene duplication in insects occurs after or parallel to, but not before the taxonomic order formation during evolution.

Osteological histology of the Pan-Alcidae (Aves, Charadriiformes): correlates of wing-propelled diving and flightlessness.

PubMed

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.

T-Box Genes in Drosophila Limb Development.

PubMed

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.

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.

FijiWingsPolarity: An open source toolkit for semi-automated detection of cell polarity.

PubMed

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.

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.

The Utility of Silos and Bunkers in the Evolution of Kinesiology

ERIC Educational Resources Information Center

Kretchmar, R. Scott

2008-01-01

Silos and bunkers have been allies in the development of kinesiology for nearly 50 years. Silos of specialization allow us to go toe-to-toe with researchers in parent disciplines, compete for grants, and otherwise spread our academic wings. The bunkers of utility and generic movement provide an important degree of legitimacy for a subject matter…

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.

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.

Influence of trap design on upwind flight behavior and capture of female grape berry moth (Lepidoptera: Tortricidae) with a kairomone lure.

PubMed

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.

OBSERVATIONS AND SIMULATIONS OF THE Na i D{sub 1} LINE PROFILES IN AN M-CLASS SOLAR FLARE

DOE Office of Scientific and Technical Information (OSTI.GOV)

Kuridze, D.; Mathioudakis, M.; Jess, D. B.

2016-12-01

We study the temporal evolution of the Na i D{sub 1} line profiles in the M3.9 flare SOL2014-06-11T21:03 UT, using observations at high spectral resolution obtained with the Interferometric Bidimensional Spectrometer instrument on the Dunn Solar Telescope combined with radiative hydrodynamic simulations. Our results show a significant increase in the intensities of the line core and wings during the flare. The analysis of the line profiles from the flare ribbons reveals that the Na i D{sub 1} line has a central reversal with excess emission in the blue wing (blue asymmetry). We combine RADYN and RH simulations to synthesize Na imore » D{sub 1} line profiles of the flaring atmosphere and find good agreement with the observations. Heating with a beam of electrons modifies the radiation field in the flaring atmosphere and excites electrons from the ground state 3s {sup 2}S to the first excited state 3p {sup 2}P, which in turn modifies the relative population of the two states. The change in temperature and the population density of the energy states make the sodium line profile revert from absorption into emission. Furthermore, the rapid changes in temperature break the pressure balance between the different layers of the lower atmosphere, generating upflow/downflow patterns. Analysis of the simulated spectra reveals that the asymmetries of the Na i D{sub 1} flare profile are produced by the velocity gradients in the lower solar atmosphere.« less

Genome sequencing of the winged midge, Parochlus steinenii, from the Antarctic Peninsula

PubMed Central

Kim, Sanghee; Oh, Mijin; Jung, Woongsic; Park, Joonho; Choi, Han-Gu

2017-01-01

Abstract Background: In the Antarctic, only two species of Chironomidae occur naturally—the wingless midge, Belgica antarctica, and the winged midge, Parochlus steinenii. B. antarctica is an extremophile with unusual adaptations. The larvae of B. antarctica are desiccation- and freeze-tolerant and the adults are wingless. Recently, the compact genome of B. antarctica was reported and it is the first Antarctic eukaryote to be sequenced. Although P. steinenii occurs naturally in the Antarctic with B. antarctica, the larvae of P. steinenii are cold-tolerant but not freeze-tolerant and the adults are winged. Differences in adaptations in the Antarctic midges are interesting in terms of evolutionary processes within an extreme environment. Herein, we provide the genome of another Antarctic midge to help elucidate the evolution of these species. Results: The draft genome of P. steinenii had a total size of 138 Mbp, comprising 9513 contigs with an N50 contig size of 34,110 bp, and a GC content of 32.2%. Overall, 13,468 genes were predicted using the MAKER annotation pipeline, and gene ontology classified 10,801 (80.2%) predicted genes to a function. Compared with the assembled genome architecture of B. antarctica, that of P. steinenii was approximately 50 Mbp longer with 6.2-fold more repeat sequences, whereas gene regions were as similarly compact as in B. antarctica. Conclusions: We present an annotated draft genome of the Antarctic midge, P. steinenii. The genomes of P. steinenii and B. antarctica will aid in the elucidation of evolution in harsh environments and provide new resources for functional genomic analyses of the order Diptera. PMID:28327954

Comparison of a Fixed-Wing and Multi-Rotor Uav for Environmental Mapping Applications: a Case Study

NASA Astrophysics Data System (ADS)

Boon, M. A.; Drijfhout, A. P.; Tesfamichael, S.

2017-08-01

The advent and evolution of Unmanned Aerial Vehicles (UAVs) and photogrammetric techniques has provided the possibility for on-demand high-resolution environmental mapping. Orthoimages and three dimensional products such as Digital Surface Models (DSMs) are derived from the UAV imagery which is amongst the most important spatial information tools for environmental planning. The two main types of UAVs in the commercial market are fixed-wing and multi-rotor. Both have their advantages and disadvantages including their suitability for certain applications. Fixed-wing UAVs normally have longer flight endurance capabilities while multi-rotors can provide for stable image capturing and easy vertical take-off and landing. Therefore, the objective of this study is to assess the performance of a fixed-wing versus a multi-rotor UAV for environmental mapping applications by conducting a specific case study. The aerial mapping of the Cors-Air model aircraft field which includes a wetland ecosystem was undertaken on the same day with a Skywalker fixed-wing UAV and a Raven X8 multi-rotor UAV equipped with similar sensor specifications (digital RGB camera) under the same weather conditions. We compared the derived datasets by applying the DTMs for basic environmental mapping purposes such as slope and contour mapping including utilising the orthoimages for identification of anthropogenic disturbances. The ground spatial resolution obtained was slightly higher for the multi-rotor probably due to a slower flight speed and more images. The results in terms of the overall precision of the data was noticeably less accurate for the fixed-wing. In contrast, orthoimages derived from the two systems showed small variations. The multi-rotor imagery provided better representation of vegetation although the fixed-wing data was sufficient for the identification of environmental factors such as anthropogenic disturbances. Differences were observed utilising the respective DTMs for the mapping of the wetland slope and contour mapping including the representation of hydrological features within the wetland. Factors such as cost, maintenance and flight time is in favour of the Skywalker fixed-wing. The multi-rotor on the other hand is more favourable in terms of data accuracy including for precision environmental planning purposes although the quality of the data of the fixed-wing is satisfactory for most environmental mapping applications.

Low-Reynolds Number Aerodynamics of an 8.9 Percent Scale Semispan Swept Wing for Assessment of Icing Effects

NASA Technical Reports Server (NTRS)

Broeren, Andy P.; Woodard, Brian S.; Diebold, Jeffrey M.; Moens, Frederic

2017-01-01

Aerodynamic assessment of icing effects on swept wings is an important component of a larger effort to improve three-dimensional icing simulation capabilities. An understanding of ice-shape geometric fidelity and Reynolds and Mach number effects on the iced-wing aerodynamics is needed to guide the development and validation of ice-accretion simulation tools. To this end, wind-tunnel testing and computational flow simulations were carried out for an 8.9%-scale semispan wing based upon the Common Research Model airplane configuration. The wind-tunnel testing was conducted at the Wichita State University 7 ft x 10 ft Beech wind tunnel from Reynolds numbers of 0.8×10(exp 6) to 2.4×10(exp 6) and corresponding Mach numbers of 0.09 to 0.27. This paper presents the results of initial studies investigating the model mounting configuration, clean-wing aerodynamics and effects of artificial ice roughness. Four different model mounting configurations were considered and a circular splitter plate combined with a streamlined shroud was selected as the baseline geometry for the remainder of the experiments and computational simulations. A detailed study of the clean-wing aerodynamics and stall characteristics was made. In all cases, the flow over the outboard sections of the wing separated as the wing stalled with the inboard sections near the root maintaining attached flow. Computational flow simulations were carried out with the ONERA elsA software that solves the compressible, three-dimensional RANS equations. The computations were carried out in either fully turbulent mode or with natural transition. Better agreement between the experimental and computational results was obtained when considering computations with free transition compared to turbulent solutions. These results indicate that experimental evolution of the clean wing performance coefficients were due to the effect of three-dimensional transition location and that this must be taken into account for future data analysis. This research also confirmed that artificial ice roughness created with rapid-prototype manufacturing methods can generate aerodynamic performance effects comparable to grit roughness of equivalent size when proper care is exercised in design and installation. The conclusions of this combined experimental and computational study contributed directly to the successful implementation of follow-on test campaigns with numerous artificial ice-shape configurations for this 8.9% scale model.

Low-Reynolds Number Aerodynamics of an 8.9 Percent Scale Semispan Swept Wing for Assessment of Icing Effects

NASA Technical Reports Server (NTRS)

Broeren, Andy P.; Woodard, Brian S.; Diebold, Jeffrey M.; Moens, Frederic

2017-01-01

Aerodynamic assessment of icing effects on swept wings is an important component of a larger effort to improve three-dimensional icing simulation capabilities. An understanding of ice-shape geometric fidelity and Reynolds and Mach number effects on the iced-wing aerodynamics is needed to guide the development and validation of ice-accretion simulation tools. To this end, wind-tunnel testing and computational flow simulations were carried out for an 8.9 percent-scale semispan wing based upon the Common Research Model airplane configuration. The wind-tunnel testing was conducted at the Wichita State University 7 by 10 ft Beech wind tunnel from Reynolds numbers of 0.8×10(exp 6) to 2.4×10(exp 6) and corresponding Mach numbers of 0.09 to 0.27. This paper presents the results of initial studies investigating the model mounting configuration, clean-wing aerodynamics and effects of artificial ice roughness. Four different model mounting configurations were considered and a circular splitter plate combined with a streamlined shroud was selected as the baseline geometry for the remainder of the experiments and computational simulations. A detailed study of the clean-wing aerodynamics and stall characteristics was made. In all cases, the flow over the outboard sections of the wing separated as the wing stalled with the inboard sections near the root maintaining attached flow. Computational flow simulations were carried out with the ONERA elsA software that solves the compressible, threedimensional RANS equations. The computations were carried out in either fully turbulent mode or with natural transition. Better agreement between the experimental and computational results was obtained when considering computations with free transition compared to turbulent solutions. These results indicate that experimental evolution of the clean wing performance coefficients were due to the effect of three-dimensional transition location and that this must be taken into account for future data analysis. This research also confirmed that artificial ice roughness created with rapid-prototype manufacturing methods can generate aerodynamic performance effects comparable to grit roughness of equivalent size when proper care is exercised in design and installation. The conclusions of this combined experimental and computational study contributed directly to the successful implementation of follow-on test campaigns with numerous artificial ice-shape configurations for this 8.9 percent scale model.

The evolution of airplanes

NASA Astrophysics Data System (ADS)

Bejan, A.; Charles, J. D.; Lorente, S.

2014-07-01

The prevailing view is that we cannot witness biological evolution because it occurred on a time scale immensely greater than our lifetime. Here, we show that we can witness evolution in our lifetime by watching the evolution of the flying human-and-machine species: the airplane. We document this evolution, and we also predict it based on a physics principle: the constructal law. We show that the airplanes must obey theoretical allometric rules that unite them with the birds and other animals. For example, the larger airplanes are faster, more efficient as vehicles, and have greater range. The engine mass is proportional to the body size: this scaling is analogous to animal design, where the mass of the motive organs (muscle, heart, lung) is proportional to the body size. Large or small, airplanes exhibit a proportionality between wing span and fuselage length, and between fuel load and body size. The animal-design counterparts of these features are evident. The view that emerges is that the evolution phenomenon is broader than biological evolution. The evolution of technology, river basins, and animal design is one phenomenon, and it belongs in physics.

Proteasome, but not autophagy, disruption results in severe eye and wing dysmorphia: a subunit- and regulator-dependent process in Drosophila.

PubMed

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.

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.

The susceptibility of Staphylococcus aureus CIP 65.8 and Pseudomonas aeruginosa ATCC 9721 cells to the bactericidal action of nanostructured Calopteryx haemorrhoidalis damselfly wing surfaces.

PubMed

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.

Ear-body lift and a novel thrust generating mechanism revealed by the complex wake of brown long-eared bats (Plecotus auritus)

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.

Why Small Is Beautiful: Wing Colour Is Free from Thermoregulatory Constraint in the Small Lycaenid Butterfly, Polyommatus icarus

PubMed Central

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

Why Small Is Beautiful: Wing Colour Is Free from Thermoregulatory Constraint in the Small Lycaenid Butterfly, Polyommatus icarus.

PubMed

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.

Experimental study of wing leading-edge devices for improved maneuver performance of a supercritical maneuvering fighter configuration. [Langley 7- by 10-ft high speed tunnel tests

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.

Diurnal and developmental differences in gene expression between adult dispersing and flightless morphs of the wing polymorphic cricket, Gryllus firmus: Implications for life-history evolution.

PubMed

Zera, Anthony J; Vellichirammal, Neetha Nanoth; Brisson, Jennifer A

2018-04-12

The functional basis of life history adaptation is a key topic of research in life history evolution. Studies of wing-polymorphism in the cricket Gryllus firmus have played a prominent role in this field. However, prior in-depth investigations of morph specialization have primarily focused on a single hormone, juvenile hormone, and a single aspect of intermediary metabolism, the fatty-acid biosynthetic component of lipid metabolism. Moreover, the role of diurnal variation in life history adaptation in G. firmus has been understudied, as is the case for organisms in general. Here, we identify genes whose expression differs consistently between the morphs independent of time-of-day during early adulthood, as well as genes that exhibit a strong pattern of morph-specific diurnal expression. We find strong, consistent, morph-specific differences in the expression of genes involved in endocrine regulation, carbohydrate and lipid metabolism, and immunity - in particular, in the expression of an insulin-like-peptide precursor gene and genes involved in triglyceride production. We also find that the flight-capable morph exhibited a substantially greater number of genes exhibiting diurnal change in gene expression compared with the flightless morph, correlated with the greater circadian change in the hemolymph juvenile titer in the dispersing morph. In fact, diurnal differences in expression within the dispersing morph at different times of the day were significantly greater in magnitude than differences between dispersing and flightless morphs at the same time-of-day. These results provide important baseline information regarding the potential role of variable gene expression on life history specialization in morphs of G. firmus, and the first information on genetically-variable, diurnal change in gene expression, associated with a key life history polymorphism. These results also suggest the existence of prominent morph-specific circadian differences in gene expression in G. firmus, possibly caused by the morph-specific circadian rhythm in the juvenile hormone titer. Copyright © 2018 Elsevier Ltd. All rights reserved.

Determinants of breeding distributions of ducks

USGS Publications Warehouse

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.

Transcriptomic and epigenomic characterization of the developing bat wing

PubMed Central

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

Feeding ecology of northern pintails and green-winged teal wintering in California

USGS Publications Warehouse

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.

Mechanosensation is evolutionarily tuned to locomotor mechanics

PubMed Central

Aiello, Brett R.; Westneat, Mark W.; Hale, Melina E.

2017-01-01

The biomechanics of animal limbs has evolved to meet the functional demands for movement associated with different behaviors and environments. Effective movement relies not only on limb mechanics but also on appropriate mechanosensory feedback. By comparing sensory ability and mechanics within a phylogenetic framework, we show that peripheral mechanosensation has evolved with limb biomechanics, evolutionarily tuning the neuromechanical system to its functional demands. We examined sensory physiology and mechanics of the pectoral fins, forelimb homologs, in the fish family Labridae. Labrid fishes exhibit extraordinary morphological and behavioral diversity and use pectoral fin-based propulsion with fins ranging in shape from high aspect ratio (AR) wing-like fins to low AR paddle-like fins. Phylogenetic character analysis demonstrates that high AR fins evolved independently multiple times in this group. Four pairs of species were examined; each included a plesiomorphic low AR and a high AR species. Within each species pair, the high AR species demonstrated significantly stiffer fin rays in comparison with the low AR species. Afferent sensory nerve activity was recorded during fin ray bending. In all cases, afferents of stiffer fins were more sensitive at lower displacement amplitudes, demonstrating mechanosensory tuning to fin mechanics and a consistent pattern of correlated evolution. We suggest that these data provide a clear example of parallel evolution in a complex neuromechanical system, with a strong link between multiple phenotypic characters: pectoral fin shape, swimming behavior, fin ray stiffness, and mechanosensory sensitivity. PMID:28396411

Nutritional physiology of life-history trade-offs: how food protein-carbohydrate content influences life-history traits in the wing-polymorphic cricket Gryllus firmus.

PubMed

Clark, Rebecca M; Zera, Anthony J; Behmer, Spencer T

2015-01-15

Although life-history trade-offs result from the differential acquisition and allocation of nutritional resources to competing physiological functions, many aspects of this topic remain poorly understood. Wing-polymorphic insects, which possess alternative morphs that trade off allocation to flight capability versus early reproduction, provide a good model system for exploring this topic. In this study, we used the wing-polymorphic cricket Gryllus firmus to test how expression of the flight capability versus reproduction trade-off was modified across a heterogeneous protein-carbohydrate nutritional landscape. Newly molted adult female long- and short-winged crickets were given one of 13 diets with different concentrations and ratios of protein and digestible carbohydrate; for each cricket, we measured consumption patterns, growth and allocation to reproduction (ovary mass) versus flight muscle maintenance (flight muscle mass and somatic lipid stores). Feeding responses in both morphs were influenced more by total macronutrient concentration than by protein-carbohydrate ratio, except at high-macronutrient concentration, where protein-carbohydrate balance was important. Mass gain tended to be greatest on protein-biased diets for both morphs, but was consistently lower across all diets for long-winged females. When long-winged females were fed high-carbohydrate foods, they accumulated greater somatic lipid stores; on high-protein foods, they accumulated greater somatic protein stores. Food protein-carbohydrate content also affected short-winged females (selected for early reproductive onset), which showed dramatic increases in ovary size, including ovarian stores of lipid and protein, on protein-biased foods. This is the first study to show how the concentration and ratio of dietary protein and carbohydrate affects consumption and allocation to key physiological features associated with the reproduction-dispersal life-history trade-off. © 2015. Published by The Company of Biologists Ltd.

The proteins encoded by the Drosophila Planar Polarity Effector genes inturned, fuzzy and fritz interact physically and can re-pattern the accumulation of “upstream” Planar Cell Polarity proteins

PubMed Central

Wang, Ying; Yan, Jie; Lee, Haeryun; Lu, Qiuheng; Adler, Paul N.

2014-01-01

The frizzled/starry night pathway regulates planar cell polarity in a wide variety of tissues in many types of animals. It was discovered and has been most intensively studied in the Drosophila wing where it controls the formation of the array of distally pointing hairs that cover the wing. The pathway does this by restricting the activation of the cytoskeleton to the distal edge of wing cells. This results in hairs initiating at the distal edge and growing in the distal direction. All of the proteins encoded by genes in the pathway accumulate asymmetrically in wing cells. The pathway is a hierarchy with the Planar Cell Polarity (PCP) genes (aka the core genes) functioning as a group upstream of the Planar Polarity Effector (PPE) genes which in turn function as a group upstream of multiple wing hairs. Upstream proteins, such as Frizzled accumulate on either the distal and/or proximal edges of wing cells. Downstream PPE proteins accumulate on the proximal edge under the instruction of the upstream proteins. A variety of types of data support this hierarchy, however, we have found that when over expressed the PPE proteins can alter both the subcellular location and level of accumulation of the upstream proteins. Thus, the epistatic relationship is context dependent. We further show that the PPE proteins interact physically and can modulate the accumulation of each other in wing cells. We also find that over expression of Frtz results in a marked delay in hair initiation suggesting that it has a separate role/activity in regulating the cytoskeleton that is not shared by other members of the group. PMID:25072625

Phoretic Carrying Capacity of Flying Southern Pine Beetles (Coleoptera: Scolytidae)

Treesearch

John C. Moser

1976-01-01

Mites do not have wings, but in their course of evolution many species have developed an association with insects, using them as a vehicle of distribution. Occasionally they cover the host so completely that the insect cannot fly. The literature is replete with these observations. Except for a single speculation (Fronk 1947), there are no reports as to how many mites...

Natural Selection and Evolution: Using Multimedia Slide Shows to Emphasize the Role of Genetic Variation

ERIC Educational Resources Information Center

Malone, Molly

2012-01-01

Most middle school students comprehend that organisms have adaptations that enable their survival and that successful adaptations prevail in a population over time. Yet they often miss that those bird beaks, moth-wing colors, or whatever traits are the result of random, normal genetic variations that just happen to confer a negative, neutral, or…

Pigeons trade efficiency for stability in response to level of challenge during confined flight.

PubMed

Williams, C David; Biewener, Andrew A

2015-03-17

Individuals traversing challenging obstacles are faced with a decision: they can adopt traversal strategies that minimally disrupt their normal locomotion patterns or they can adopt strategies that substantially alter their gait, conferring new advantages and disadvantages. We flew pigeons (Columba livia) through an array of vertical obstacles in a flight arena, presenting them with this choice. The pigeons selected either a strategy involving only a slight pause in the normal wing beat cycle, or a wings-folded posture granting reduced efficiency but greater stability should a misjudgment lead to collision. The more stable but less efficient flight strategy was not used to traverse easy obstacles with wide gaps for passage but came to dominate the postures used as obstacle challenge increased with narrower gaps and there was a greater chance of a collision. These results indicate that birds weigh potential obstacle negotiation strategies and estimate task difficulty during locomotor pattern selection.

Pigeons trade efficiency for stability in response to level of challenge during confined flight

PubMed Central

Williams, C. David; Biewener, Andrew A.

2015-01-01

Individuals traversing challenging obstacles are faced with a decision: they can adopt traversal strategies that minimally disrupt their normal locomotion patterns or they can adopt strategies that substantially alter their gait, conferring new advantages and disadvantages. We flew pigeons (Columba livia) through an array of vertical obstacles in a flight arena, presenting them with this choice. The pigeons selected either a strategy involving only a slight pause in the normal wing beat cycle, or a wings-folded posture granting reduced efficiency but greater stability should a misjudgment lead to collision. The more stable but less efficient flight strategy was not used to traverse easy obstacles with wide gaps for passage but came to dominate the postures used as obstacle challenge increased with narrower gaps and there was a greater chance of a collision. These results indicate that birds weigh potential obstacle negotiation strategies and estimate task difficulty during locomotor pattern selection. PMID:25733863

The Regulation of Aldehyde Oxidase in Imaginal Wing Discs of Drosophila Hybrids: Evidence for cis- and trans-Acting Control Elements

PubMed Central

Sprey, Th. E.; Kuhn, David T.

1987-01-01

The aldehyde oxidase (Aldox) distribution pattern was determined for wing discs of partial hybrids between D. melanogaster and D. simulans. In these animals the regulation of Aldox activity is not uniform over the disc epithelium as both cis-dominant and trans -acting control were evident in different regions of the disc. The Aldox expression was shown to be regulated by loci on the X chromosome, 2L and 3R of D. melanogaster and 2R and 3R of D. simulans. PMID:17246366

Computer program for prediction of the deposition of material released from fixed and rotary wing aircraft

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.

Complex constraints on allometry revealed by artificial selection on the wing of Drosophila melanogaster

PubMed Central

Bolstad, Geir H.; Cassara, Jason A.; Márquez, Eladio; Hansen, Thomas F.; van der Linde, Kim; Houle, David; Pélabon, Christophe

2015-01-01

Precise exponential scaling with size is a fundamental aspect of phenotypic variation. These allometric power laws are often invariant across taxa and have long been hypothesized to reflect developmental constraints. Here we test this hypothesis by investigating the evolutionary potential of an allometric scaling relationship in drosophilid wing shape that is nearly invariant across 111 species separated by at least 50 million years of evolution. In only 26 generations of artificial selection in a population of Drosophila melanogaster, we were able to drive the allometric slope to the outer range of those found among the 111 sampled species. This response was rapidly lost when selection was suspended. Only a small proportion of this reversal could be explained by breakup of linkage disequilibrium, and direct selection on wing shape is also unlikely to explain the reversal, because the more divergent wing shapes produced by selection on the allometric intercept did not revert. We hypothesize that the reversal was instead caused by internal selection arising from pleiotropic links to unknown traits. Our results also suggest that the observed selection response in the allometric slope was due to a component expressed late in larval development and that variation in earlier development did not respond to selection. Together, these results are consistent with a role for pleiotropic constraints in explaining the remarkable evolutionary stability of allometric scaling. PMID:26371319

UV photoreceptors and UV-yellow wing pigments in Heliconius butterflies allow a color signal to serve both mimicry and intraspecific communication.

PubMed

Bybee, Seth M; Yuan, Furong; Ramstetter, Monica D; Llorente-Bousquets, Jorge; Reed, Robert D; Osorio, Daniel; Briscoe, Adriana D

2012-01-01

Mimetic wing coloration evolves in butterflies in the context of predator confusion. Unless butterfly eyes have adaptations for discriminating mimetic color variation, mimicry also carries a risk of confusion for the butterflies themselves. Heliconius butterfly eyes, which express recently duplicated ultraviolet (UV) opsins, have such an adaptation. To examine bird and butterfly color vision as sources of selection on butterfly coloration, we studied yellow wing pigmentation in the tribe Heliconiini. We confirmed, using reflectance and mass spectrometry, that only Heliconius use 3-hydroxy-DL-kynurenine (3-OHK), which looks yellow to humans but reflects both UV- and long-wavelength light, whereas butterflies in related genera have chemically unknown yellow pigments mostly lacking UV reflectance. Modeling of these color signals reveals that the two UV photoreceptors of Heliconius are better suited to separating 3-OHK from non-3-OHK spectra compared with the photoreceptors of related genera or birds. The co-occurrence of potentially enhanced UV vision and a UV-reflecting yellow wing pigment could allow unpalatable Heliconius private intraspecific communication in the presence of mimics. Our results are the best available evidence for the correlated evolution of a color signal and color vision. They also suggest that predator visual systems are error prone in the context of mimicry. © 2011 by The University of Chicago.

Muscle activation patterns and motor anatomy of Anna's hummingbirds Calypte anna and zebra finches Taeniopygia guttata.

PubMed

Donovan, Edward R; Keeney, Brooke K; Kung, Eric; Makan, Sirish; Wild, J Martin; Altshuler, Douglas L

2013-01-01

Flying animals exhibit profound transformations in anatomy, physiology, and neural architecture. Although much is known about adaptations in the avian skeleton and musculature, less is known about neuroanatomy and motor unit integration for bird flight. Hummingbirds are among the most maneuverable and specialized of vertebrate fliers, and two unusual neuromuscular features have been previously reported: (1) the pectoralis major has a unique distribution pattern of motor end plates (MEPs) compared with all other birds and (2) electromyograms (EMGs) from the hummingbird's pectoral muscles, the pectoralis major and the supracoracoideus, show activation bursts composed of one or a few spikes that appear to have a very consistent pattern. Here, we place these findings in a broader context by comparing the MEPs, EMGs, and organization of the spinal motor neuron pools of flight muscles of Anna's hummingbird Calypte anna, zebra finches Taeniopygia guttata, and, for MEPs, several other species. The previously shown MEP pattern of the hummingbird pectoralis major is not shared with its closest taxonomic relative, the swift, and appears to be unique to hummingbirds. MEP arrangements in previously undocumented wing muscles show patterns that differ somewhat from other avian muscles. In the parallel-fibered strap muscles of the shoulder, MEP patterns appear to relate to muscle length, with the smallest muscles having fibers that span the entire muscle. MEP patterns in pennate distal wing muscles were the same regardless of size, with tightly clustered bands in the middle portion of the muscle, not evenly distributed bands over the muscle's entire length. Muscle activations were examined during slow forward flight in both species, during hovering in hummingbirds, and during slow ascents in zebra finches. The EMG bursts of a wing muscle, the pronator superficialis, were highly variable in peak number, size, and distribution across wingbeats for both species. In the pectoralis major, although the individual EMG bursts were much shorter in duration in hummingbirds relative to zebra finches, the variables describing the normalized amplitude and area of the activation bursts were otherwise indistinguishable between taxa during these flight modes. However, the degree of variation in the time intervals between EMG peaks was much lower in hummingbirds, which is a plausible explanation for the "patterned" EMG signals reported previously.

Biomimetic propulsion under random heaving conditions, using active pitch control

NASA Astrophysics Data System (ADS)

Politis, Gerasimos; Politis, Konstantinos

2014-05-01

Marine mammals travel long distances by utilizing and transforming wave energy to thrust through proper control of their caudal fin. On the other hand, manmade ships traveling in a wavy sea store large amounts of wave energy in the form of kinetic energy for heaving, pitching, rolling and other ship motions. A natural way to extract this energy and transform it to useful propulsive thrust is by using a biomimetic wing. The aim of this paper is to show how an actively pitched biomimetic wing could achieve this goal when it performs a random heaving motion. More specifically, we consider a biomimetic wing traveling with a given translational velocity in an infinitely extended fluid and performing a random heaving motion with a given energy spectrum which corresponds to a given sea state. A formula is invented by which the instantaneous pitch angle of the wing is determined using the heaving data of the current and past time steps. Simulations are then performed for a biomimetic wing at different heave energy spectra, using an indirect Source-Doublet 3-D-BEM, together with a time stepping algorithm capable to track the random motion of the wing. A nonlinear pressure type Kutta condition is applied at the trailing edge of the wing. With a mollifier-based filtering technique, the 3-D unsteady rollup pattern created by the random motion of the wing is calculated without any simplifying assumptions regarding its geometry. Calculated unsteady forces, moments and useful power, show that the proposed active pitch control always results in thrust producing motions, with significant propulsive power production and considerable beneficial stabilizing action to ship motions. Calculation of the power required to set the pitch angle prove it to be a very small percentage of the useful power and thus making the practical application of the device very tractable.

A tool for developing an automatic insect identification system based on wing outlines

PubMed Central

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

Bioinspired spring origami.

PubMed

Faber, Jakob A; Arrieta, Andres F; Studart, André R

2018-03-23

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. Copyright © 2018 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works.

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.

Flow pattern similarities in the near wake of three bird species suggest a common role for unsteady aerodynamic effects in lift generation

PubMed Central

Krishnan, Krishnamoorthy; Ben-Gida, Hadar; Kirchhefer, Adam J.; Kopp, Gregory A.; Guglielmo, Christopher G.

2017-01-01

Analysis of the aerodynamics of flapping wings has yielded a general understanding of how birds generate lift and thrust during flight. However, the role of unsteady aerodynamics in avian flight due to the flapping motion still holds open questions in respect to performance and efficiency. We studied the flight of three distinctive bird species: western sandpiper (Calidris mauri), European starling (Sturnus vulgaris) and American robin (Turdus migratorius) using long-duration, time-resolved particle image velocimetry, to better characterize and advance our understanding of how birds use unsteady flow features to enhance their aerodynamic performances during flapping flight. We show that during transitions between downstroke and upstroke phases of the wing cycle, the near wake-flow structures vary and generate unique sets of vortices. These structures appear as quadruple layers of concentrated vorticity aligned at an angle with respect to the horizon (named ‘double branch’). They occur where the circulation gradient changes sign, which implies that the forces exerted by the flapping wings of birds are modified during the transition phases. The flow patterns are similar in (non-dimensional) size and magnitude for the different birds suggesting that there are common mechanisms operating during flapping flight across species. These flow patterns occur at the same phase where drag reduction of about 5% per cycle and lift enhancement were observed in our prior studies. We propose that these flow structures should be considered in wake flow models that seek to account for the contribution of unsteady flow to lift and drag. PMID:28163881

Distal-less regulates eyespot patterns and melanization in Bicyclus butterflies.

PubMed

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.

Reflection characterization of nano-sized dielectric structure in Morpho butterfly wings

NASA Astrophysics Data System (ADS)

Zhu, Dong

2017-10-01

Morpho butterflies living in Central and South America are well-known for their structural-colored blue wings. The blue coloring originates from the interaction of light with nano-sized dielectric structures that are equipped on the external surface of scales covering over their wings. The high-accuracy nonstandard finite-difference time domain (NS-FDTD) method is used to investigate the reflection characterization from the nanostructures. In the NS-FDTD calculation, a computational model is built to mimic the actual tree-like multilayered structures wherever possible using the hyperbolic tangent functions. It is generally known that both multilayer interference and diffraction grating phenomena can occur when light enters the nano-sized multilayered structure. To answer the question that which phenomenon is mainly responsible for the blue coloring, the NS-FDTD calculation is performed under various incidence angles at wavelengths from 360 to 500 nm. The calculated results at one incident wavelength under different incidence angles are visualized in a two-dimensional mapping image, where horizontal and vertical axes are incidence and reflection angles, respectively. The images demonstrate a remarkable transition from a ring-like pattern at shorter wavelengths to a retro-reflection pattern at longer wavelengths. To clarify the origin of the pattern transition, the model is separated into several simpler parts and compared their mapping images with the theoretical diffraction calculations. It can be concluded that the blue coloring at longer wavelengths is mainly caused by the cooperation of multilayer interference and retro-reflection while the effect of diffraction grating is predominant at shorter wavelengths.

An illustrated checklist of the genus Elymnias Hübner, 1818 (Nymphalidae, Satyrinae)

PubMed Central

Wei, Chia-Hsuan; Lohman, David J.; Peggie, Djunijanti; Yen, Shen-Horn

2017-01-01

Abstract We review the genus Elymnias Hübner, 1818, a morphologically diverse satyrine butterfly clade involved in multifarious Batesian mimicry relationships throughout Asia and Africa. A variety of different model species are mimicked, and many Elymnias species are sexually dimorphic mimics, with males and females resembling different model species. We revise species and subspecies delimitations in light of an integrative taxonomic investigation using external morphology, male and female genital morphology, and a multi-locus molecular phylogeny. There is little interspecific genitalic variation among species in this group, and previous taxonomists therefore relied almost entirely on wing patterns. Our molecular phylogenetic analysis reveals several examples of polymorphism or wing pattern divergence within a single species currently classified as two or more different species. We also found examples of wing pattern convergence among disparate lineages that mimic the same widespread model species. Frequently, two or more phenotypically similar species were classified as a single species. This comprehensive checklist reviews all names associated with Elymnias to align its taxonomy with the evolutionary history of the group. All available information on nomenclature, type localities, repositories of type specimens, and geographical distributions is summarized, and images of adult specimens and genitalia are provided along with distribution maps of all species and selected subspecies. We identify 2 species incertae sedis, establish 15 monophyletic species groups (including 1 species unplaced in any species group), and make 49 taxonomic changes, including 35 new synonyms, 7 new combinations (2 of which have new status), 1 resurrected combination, 1 resurrected subspecies, and 7 status changes. PMID:28769686

A Middle Triassic thoracopterid from China highlights the evolutionary origin of overwater gliding in early ray-finned fishes

PubMed Central

Xu, Guang-Hui; Zhao, Li-Jun; Shen, Chen-Chen

2015-01-01

Gliding adaptations in thoracopterid flying fishes represent a remarkable case of convergent evolution of overwater gliding strategy with modern exocoetid flying fishes, but the evolutionary origin of this strategy was poorly known in the thoracopterids because of lack of transitional forms. Until recently, all thoracopterids, from the Late Triassic of Austria and Italy and the Middle Triassic of South China, were highly specialized ‘four-winged’ gliders in having wing-like paired fins and an asymmetrical caudal fin with the lower caudal lobe notably larger than the upper lobe. Here, we show that the new genus Wushaichthys and the previously alleged ‘peltopleurid’ Peripeltopleurus, from the Middle Triassic (Ladinian, 235–242 Ma) of South China and near the Ladinian/Anisian boundary of southern Switzerland and northern Italy, respectively, represent the most primitive and oldest known thoracopterids. Wushaichthys, the most basal thoracopterid, shows certain derived features of this group in the skull. Peripeltopleurus shows a condition intermediate between Wushaichthys and Thoracopterus in having a slightly asymmetrical caudal fin but still lacking wing-like paired fins. Phylogenetic studies suggest that the evolution of overwater gliding of thoracopterids was gradual in nature; a four-stage adaption following the ‘cranial specialization–asymmetrical caudal fin–enlarged paired fins–scale reduction’ sequence has been recognized in thoracopterid evolution. Moreover, Wushaichthys and Peripeltopleurus bear hooklets on the anal fin of supposed males, resembling those of modern viviparious teleosts. Early thoracopterids probably had evolved a live-bearing reproductive strategy. PMID:25568155

A Second New Species of Ice Crawlers from China (Insecta: Grylloblattodea), with Thorax Evolution and the Prediction of Potential Distribution

PubMed Central

Bai, Ming; Jarvis, Karl; Wang, Shu-Yong; Song, Ke-Qing; Wang, Yan-Ping; Wang, Zhi-Liang; Li, Wen-Zhu; Wang, Wei; Yang, Xing-Ke

2010-01-01

Modern grylloblattids are one of the least diverse of the modern insect orders. The thorax changes in morphology might be associated with the changes of the function of the forelegs, wing loss, changes in behavior and adaptation to habitat. As temperature is the main barrier for migration of modern grylloblattids, the range of each species is extremely limited. The potential distribution areas of grylloblattids remain unclear. A second new species of ice crawlers (Insecta: Grylloblattodea), Grylloblattella cheni Bai, Wang et Yang sp. nov., is described from China. The distribution map and key to species of Grylloblattella are given. A comparison of the thorax of extant and extinct Grylloblattodea is presented, with an emphasis on the pronotum using geometric morphometric analysis, which may reflect thorax adaptation and the evolution of Grylloblattodea. Potential global distribution of grylloblattids is inferred. Highly diversified pronota of extinct Grylloblattodea may reflect diverse habitats and niches. The relatively homogeneous pronota of modern grylloblattids might be explained by two hypotheses: synapomorphy or convergent evolution. Most fossils of Grylloblattodea contain an obviously longer meso- and metathorax than prothorax. The length of the meso- and metathorax of modern grylloblattids is normally shorter than the prothorax. This may be associated with the wing loss, which is accompanied by muscle reduction and changes to the thoracic skeleton system. Threats to grylloblattids and several conservation comments are also provided. PMID:20877572

Hox genes, digit identities and the theropod/bird transition.

PubMed

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

A Nonstationary Markov Model Detects Directional Evolution in Hymenopteran Morphology.

PubMed

Klopfstein, Seraina; Vilhelmsen, Lars; Ronquist, Fredrik

2015-11-01

Directional evolution has played an important role in shaping the morphological, ecological, and molecular diversity of life. However, standard substitution models assume stationarity of the evolutionary process over the time scale examined, thus impeding the study of directionality. Here we explore a simple, nonstationary model of evolution for discrete data, which assumes that the state frequencies at the root differ from the equilibrium frequencies of the homogeneous evolutionary process along the rest of the tree (i.e., the process is nonstationary, nonreversible, but homogeneous). Within this framework, we develop a Bayesian approach for testing directional versus stationary evolution using a reversible-jump algorithm. Simulations show that when only data from extant taxa are available, the success in inferring directionality is strongly dependent on the evolutionary rate, the shape of the tree, the relative branch lengths, and the number of taxa. Given suitable evolutionary rates (0.1-0.5 expected substitutions between root and tips), accounting for directionality improves tree inference and often allows correct rooting of the tree without the use of an outgroup. As an empirical test, we apply our method to study directional evolution in hymenopteran morphology. We focus on three character systems: wing veins, muscles, and sclerites. We find strong support for a trend toward loss of wing veins and muscles, while stationarity cannot be ruled out for sclerites. Adding fossil and time information in a total-evidence dating approach, we show that accounting for directionality results in more precise estimates not only of the ancestral state at the root of the tree, but also of the divergence times. Our model relaxes the assumption of stationarity and reversibility by adding a minimum of additional parameters, and is thus well suited to studying the nature of the evolutionary process in data sets of limited size, such as morphology and ecology. © The Author(s) 2015. Published by Oxford University Press, on behalf of the Society of Systematic Biologists.

Proteasome, but Not Autophagy, Disruption Results in Severe Eye and Wing Dysmorphia: A Subunit- and Regulator-Dependent Process in Drosophila

PubMed Central

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

Comparative analysis of the development of wing-flapping and flight in the fowl.

PubMed

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.

Genome sequencing of the winged midge, Parochlus steinenii, from the Antarctic Peninsula.

PubMed

Kim, Sanghee; Oh, Mijin; Jung, Woongsic; Park, Joonho; Choi, Han-Gu; Shin, Seung Chul

2017-03-01

In the Antarctic, only two species of Chironomidae occur naturally-the wingless midge, Belgica antarctica , and the winged midge, Parochlus steinenii . B. antarctica is an extremophile with unusual adaptations. The larvae of B. antarctica are desiccation- and freeze-tolerant and the adults are wingless. Recently, the compact genome of B. antarctica was reported and it is the first Antarctic eukaryote to be sequenced. Although P. steinenii occurs naturally in the Antarctic with B. antarctica , the larvae of P. steinenii are cold-tolerant but not freeze-tolerant and the adults are winged. Differences in adaptations in the Antarctic midges are interesting in terms of evolutionary processes within an extreme environment. Herein, we provide the genome of another Antarctic midge to help elucidate the evolution of these species. The draft genome of P. steinenii had a total size of 138 Mbp, comprising 9513 contigs with an N50 contig size of 34,110 bp, and a GC content of 32.2%. Overall, 13,468 genes were predicted using the MAKER annotation pipeline, and gene ontology classified 10,801 (80.2%) predicted genes to a function. Compared with the assembled genome architecture of B. antarctica , that of P. steinenii was approximately 50 Mbp longer with 6.2-fold more repeat sequences, whereas gene regions were as similarly compact as in B. antarctica . We present an annotated draft genome of the Antarctic midge, P. steinenii . The genomes of P. steinenii and B. antarctica will aid in the elucidation of evolution in harsh environments and provide new resources for functional genomic analyses of the order Diptera. © The Authors 2017. Published by Oxford University Press.

Evidence of selection on phenotypic plasticity and cost of plasticity in response to host-feeding sources in the major Chagas disease vector Triatoma infestans.

PubMed

Nattero, Julieta; Leonhard, Gustavo; Gürtler, Ricardo E; Crocco, Liliana B

2015-12-01

Phenotypic plasticity is the ability of a genotype to display alternative phenotypes in different environments. Understanding how plasticity evolves and the factors that favor and constrain its evolution have attracted great interest. We investigated whether selection on phenotypic plasticity and costs of plasticity affect head and wing morphology in response to host-feeding sources in the major Chagas disease vector Triatoma infestans. Full-sib families were assigned to blood-feeding on either live pigeons or guinea pigs throughout their lives. We measured diet-induced phenotypic plasticity on wing and head size and shape; characterized selection on phenotypic plasticity for female and male fecundity rates, and evaluated costs of plasticity. Wing size and shape variables exhibited significant differences in phenotypic plasticity associated with host-feeding source in female and male bugs. Evidence of selection on phenotypic plasticity was detected in head size and shape for guinea pig-fed females. A lower female fecundity rate was detected in more plastic families for traits that showed selection on plasticity. These results provide insights into the morphological phenotypic plasticity of T. infestans, documenting fitness advantages of head size and shape for females fed on guinea pigs. This vector species showed measurable benefits of responding plastically to environmental variation rather than adopting a fixed development plan. The presence of cost of plasticity suggests constraints on the evolution of plasticity. Our study indicates that females fed on guinea pigs (and perhaps on other suitable mammalian hosts) have greater chances of evolving under selection on phenotypic plasticity subject to some constraints. Copyright © 2015 Elsevier B.V. All rights reserved.

The accelerated build-up of the red sequence in high-redshift galaxy clusters

NASA Astrophysics Data System (ADS)

Cerulo, P.; Couch, W. J.; Lidman, C.; Demarco, R.; Huertas-Company, M.; Mei, S.; Sánchez-Janssen, R.; Barrientos, L. F.; Muñoz, R. P.

2016-04-01

We analyse the evolution of the red sequence in a sample of galaxy clusters at redshifts 0.8 < z < 1.5 taken from the HAWK-I Cluster Survey (HCS). The comparison with the low-redshift (0.04 < z < 0.08) sample of the WIde-field Nearby Galaxy-cluster Survey (WINGS) and other literature results shows that the slope and intrinsic scatter of the cluster red sequence have undergone little evolution since z = 1.5. We find that the luminous-to-faint ratio and the slope of the faint end of the luminosity distribution of the HCS red sequence are consistent with those measured in WINGS, implying that there is no deficit of red galaxies at magnitudes fainter than M_V^{ast } at high redshifts. We find that the most massive HCS clusters host a population of bright red sequence galaxies at MV < -22.0 mag, which are not observed in low-mass clusters. Interestingly, we also note the presence of a population of very bright (MV < -23.0 mag) and massive (log (M*/M⊙) > 11.5) red sequence galaxies in the WINGS clusters, which do not include only the brightest cluster galaxies and which are not present in the HCS clusters, suggesting that they formed at epochs later than z = 0.8. The comparison with the luminosity distribution of a sample of passive red sequence galaxies drawn from the COSMOS/UltraVISTA field in the photometric redshift range 0.8 < zphot < 1.5 shows that the red sequence in clusters is more developed at the faint end, suggesting that halo mass plays an important role in setting the time-scales for the build-up of the red sequence.