Role of Soft Matter in the Sandwich Vein of Dragonfly Wing in Its Configuration and Aerodynamic Behaviors

The microstructure of the main longitudinal veins of the dragonfly wing and the aerodynamic behaviors of the wing were investigated in this paper.The microstructure of longitudinal vein presents two circumferential chitin layers and a protein-fiber soft layer.The dragonfly wing is corrugated due to the spatial arrangement of longitudinal veins.It was found that the corrugation angle could significantly influence the lift/drag ratio across a range of attack angles by the wind tunnel experiments.The results of the finite element analysis indicate that the protein soft layer of vein facilitates the change of the corrugation angle by allowing substantial relative twisting deformation between two neighboring veins,which is not possible in veins without a soft sandwich layer.     

Effects of Dragonfly Wing Structure on the Dynamic Performances

The configurations of dragonfly wings, including the corrugations of the chordwise cross-section, the microstructure of the longitudinal veins and membrane, were comprehensively investigated using the Environmental Scanning Electron Microscopy (ESEM). Based on the experimental results reported previously, the multi-scale and multi-dimensional models with different structural features of dragonfly wing were created, and the biological dynamic behaviors of wing models were discussed through the Finite Element Method (FEM). The results demonstrate that the effects of different structural features on dynamic behaviors of dragonfly wing such as natural frequency/modal, bending/torsional deformation, reaction force/torque are very significant. The corrugations of dragonfly wing along the chordwise can observably improve the flapping frequency because of the greater structural stiffness of wings. In updated model, the novel sandwich microstructure of the longitudinal veins remarkably improves the torsional deformation of dragonfly wing while it has a little effect on the flapping frequency and bending deformation. These integrated structural features can adjust the deformation of wing oneself, therefore the flow field around the wings can be controlled adaptively. The fact is that the flights of dragonfly wing with sandwich microstructure of longitudinal veins are more efficient and intelligent.     

The Role of Soft Vein Joints in Dragonfly Flight

Dragonflies are excellent flyers among insects and their flight ability is closely related to the architecture and material properties of their wings.The veins are main structure components of a dragonfly wing,which are found to be connected by resilin with high elasticity at some joints.A three-dimensional (3D) finite element model of dragonfly wing considering the soft vein joints is developed,with some simplifications.Passive deformation under aerodynamic loads and active flapping motion of the wing are both studied.The functions of soft vein joints in dragonfly flight are concluded.In passive deformation,the chordwise flexibility is improved by soft vein joints and the wing is cambered under loads,increasing the action area with air.In active flapping,the wing rigidity in spanwise direction is maintained to achieve the required amplitude.As a result,both the passive deformation and the active control of flapping work well in dragonfly flight.The present study may also inspire the design of biomimetic Flapping Micro Air Vehicles (FMAVs).     

Morphology of hindwing veins in the shield bug Graphosoma italicum (Heteroptera: Pentatomidae)

Light, fluorescence, and electron microscopy were applied to cross sections and -breakage and whole-mount preparations of the anterior hindwing vein of the shield bug Graphosoma italicum. These analyses were complemented by investigations of the basal part of the forewing Corium and Clavus. The integration of structural, histological, and fluorescence data revealed a complex arrangement of both rigid and elastic structures in the wall of wing veins and provided insights into the constitution of transition zones between rigid and elastic regions. Beneath the exocuticular layers, which are continuous with the dorsal and ventral cuticle of the wing membrane, the lumen of the veins is encompassed by a mesocuticular layer, an internal circular exocuticular layer, and an internal longitudinal endocuticular layer. Separate parallel lumina within the anterior longitudinal vein of the hindwing, arranged side-by-side rostro-caudally, suggest that several veins have fused in the phylogenetic context of vein reduction in the pentatomid hindwing. Gradual structural transition zones and resilin enrichment between sclerotized layers of the vein wall and along the edges of the claval furrow are interpreted as mechanical adaptations to enhance the reliability and durability of the mechanically stressed wing veins.     

现存蜉蝣翅基纵脉走向及愈合模式(昆虫纲：蜉蝣目)

有翅昆虫翅基纵脉的走向及愈合模式在系统发育重建中占有重要地位。然而,现存蜉蝣翅基纵脉的走向及愈合状况在大部分种类变化极大,无法推测其原始状况,只在极少数种类保留有部分可见残迹。中国拟短丝蜉Siphluriscus chinensis的翅基保留有独立的亚前缘脉弓、部分中脉M和肘脉Cu主干以及前中脉MA及径分脉Rs的走向痕迹。据此并结合红斑蜉Ephemera rufomaculata 和大网脉蜉Chromarcys magnifica翅基的相关特征,本文提出了蜉蝣目主要纵脉基部走向及愈合的基本模式,其要点有：中脉主干在基部与径脉主干独自发出后先接近或愈合后又分离、它们各自分成两支后的前中脉及径分脉又先愈合再分离、肘脉始终独立。这种中脉与径脉先接近或愈合后分离的模式非常接近新翅类的情况而与蜻蜓很不相同(在蜻蜓,中脉与肘脉在基部愈合) 。亚前缘脉弓的作用相信是加强了因翅基骨板发达而相互远离的纵脉间的连结作用。这个假说也可以来解释蜻蜓复杂脉相的形成原因。     

The Wettability and Mechanism of Geometric Non-Smooth Structure of Dragonfly Wing Surface

Scanning electron microscope and optical contact angle measuring instruments were used to investigate the microstructure and wettability of geometric non-smooth structure of dragonfly wing surface. Results show that the geometric non-smooth structure of dragonfly wing surface is one part of epicuticle, some organic solvents can effectively dissolve the main ingredient of non-smooth structure. The hydrophobicity of dragonfly wing surface is induced by the co-coupling of the non-smooth structure and the waxy layer covering.     

Oogenesis and egg-shell formation in Aspiculuris tetraptera Schulz (Nematoda: Oxyuroidea).

The ovary of Aspiculuris tetraptera has a prominent terminal cap cell. This is considered to be part of the ovarian epithelium. Oogonia detach from the short rachis and increase in size from 6 to 60 microns; accumulating hyaline granules, shell granules and glycogen. The hyaline granules persist in the eff cytoplasm after shell formation has been completed and are considered to be lipoprotein yolk. The shell granules contribute to the non-chitin fraction of the chitinous layer. A classification of the cytoplasmic inclusions of the nematode oocyte is proposed. Upon fertilization a vitelline membrane is formed which constitutes the vitelline layer of the egg-shell. The chitinous layer is secreted in the perivitelline space, between the vitelline layer and the egg oolemma. Upon completion of chitinous layer synthesis, the egg cytoplasm contracts away from its inner surface. The material of the lipid layer is secreted at the surface of the egg cytoplasm and adheres to the inner surface of the chitinous layer. During secretion of the chitinous and lipid layers by the egg cytoplasm, the uterine cells secrete the unit membrane-like external uterine layer and the crystalline internal uterine layer. A complex system of interconnecting spaces develops in the internal uterine layer. This system is open to the exterior via breaks in the external uterine layer. There is no direct involvement of the uterine cells in the formation of this structure.     

Oogenesis and egg shell formation in Heterakis gallinarum (Nematoda)

The structure of the developing ova and egg shell formation of Heterakis gallinarum has been described. The oogonia are small, undifferentiated cells which are arranged around a central cytoplasmic rachis. The oogonia and young oocytes are in cytoplasmic continuity with the rachis and it is suggested that the rachis may influence synchronous development of the oocytes. The oocytes contain two types of granule; refringent, which give rise to the ascaroside layer of the egg shell, and another kind which appear to be a type of yolk for the developing egg. After fertilization the spermatozoon produces numerous ribosomes; a second unit membrane appears beneath the oolemma, and the chitinous layer of the shell forms between the oolemma and this inner membrane. The refringent granules later produce the ascaroside layer of the shell between the chitinous layer and the inner membrane. The outermost layer of the shell is produced from material secreted by the cells of the uterus.     

Envelope structure of four gliding filamentous cyanobacteria.

The cell walls of four gliding filamentous Oscillatoriaceae species comprising three different genera were studied by freeze substitution, freeze fracturing, and negative staining. In all species, the multilayered gram-negative cell wall is covered with a complex external double layer. The first layer is a tetragonal crystalline S-layer anchored on the outer membrane. The second array is formed by parallel, helically arranged surface fibrils with diameters of 8 to 12 nm. These fibrils have a serrated appearance in cross sections. In all cases, the orientation of the surface fibrils correlates with the sense of revolution of the filaments during gliding, i.e., clockwise in both Phormidium strains and counterclockwise in Oscillatoria princeps and Lyngbya aeruginosa. The lack of longitudinal corrugations or contractions of the surface fibrils and the identical appearances of motile and nonmotile filaments suggest that this structure plays a passive screw thread role in gliding. It is hypothesized that the necessary propulsive force is generated by shear forces between the surface fibrils and the continuing flow of secreted extracellular slime. Furthermore, the so-called junctional pores seem to be the extrusion sites of the slime. In motile cells, these pores exhibit a different staining behavior than that seen in nonmotile ones. In the former, the channels of the pores are filled with electron-dense material, whereas in the latter, the channels appear comparatively empty, highly contrasting the peptidoglycan. Finally, the presence of regular surface structures in other gliding prokaryotes is considered an indication that comparable structures are general features of the cell walls of gliding microbes.     

The distribution of chitin in larval shells of the bivalve mollusk Mytilus galloprovincialis

The insoluble matrix of larval shells of the marine bivalve mollusk Mytilus galloprovincialis is investigated by confocal laser scanning microscopy using a GFP fusion protein with a chitin-binding domain for labeling of chitinous structures. We show that chitinous material is present in the larval shell, presumably as a chitin-protein complex. We further show that the structure of the chitinous material changes with the development of the larvae. We conclude from the presence of characteristic chitinous structures in certain shell regions that chitin fulfills an important function in the formation and functionality of larval bivalve shells.     

Resilin-bearing wing vein joints in the dragonfly Epiophlebia superstes

In this study, we compared the dorsal and ventral patterns of three vein joint types and three types of resilin patches in the wings of the dragonfly Epiophlebia superstes. The joint types were classified according to their general structure and the resilin patch types according to their arrangement at joints and in the adjacent wing membrane. Resilin patches are found in both dorsal and ventral pleat valleys of the corrugated wings of E. superstes, which results in different patterns of resilin distribution on the dorsal and ventral sides of the wing. In addition to its probable function in conferring flexibility to stressed joints, resilin may also have a damping function. Our results suggest that resilin patches in the leading edge may be loaded in compression, whereas in the trailing area, they may be involved in angle widening and thus loaded in tension. Possible adaptations to the deformability of different areas of the wing, e.g. during the process of camber formation, are discussed.     

Use of a Digital Image Correlation Technique for Measuring the Material Properties of Beetle Wing

Beetle wings are very specialized flight organs consisting of the veins and membranes.Therefore it is necessary from abionic view to investigate the material properties of a beetle wing experimentally.In the present study,we have used a DigitalImage Correlation (DIC) technique to measure the elastic modulus of a beetle wing membrane.Specimens were prepared bycarefully cutting a beetle hind wing into 3.0 mm by 7.0 mm segments (the gage length was 5 mm).We used a scanning electronmicroscope for a precise measurement of the thickness of the beetle wing membrane.The specimen was attached to a designedfixture to induce a uniform displacement by means of a micromanipulator.We used an ARAMISTM system based on the digitalimage correlation technique to measure the corresponding displacement of a specimen.The thickness of the beetle wing variedat different points of the membrane.The elastic modulus differed in relation to the membrane arrangement showing a structuralanisotropy;the elastic modulus in the chordwise direction is approximately 2.65 GPa,which is three times larger than the elasticmodulus in the spanwise direction of 0.84 GPa.As a result,the digital image correlation-based ARAMIS system was suc-cessfully used to measure the elastic modulus of a beetle wing.In addition to membrane’s elastic modulus,we considered thePoisson’s ratio of the membrane and measured the elastic modulus of a vein using an Instron universal tensile machine.Theresult reveals the Poisson’s ratio is nearly zero and the elastic modulus of a vein is about 11 GPa.     

Homology and function in the wings of Heteroptera

Abstract. The homology of veins and other wing characters in Heteroptera is reviewed in the light of palaeontology and new functional studies. A cladogram is given for the higher taxa of Hemiptera. It is probable that the vannus is an autapomorphy of Auchenorrhyncha+Heteropteroidea; that the leading edge vein of heteropteran fore- and hindwings is C+Sc; that Rs cannot be distinguished from R; that the hamus is part of M; that the glochis is a secondary structure. The difficulty of defining a vein is stressed. The functional significance of the hemielytron, cuneal fracture and longitudinal flexion lines is discussed. A preliminary ground-plan for Heteroptera wings is presented.     

Expression pattern of glutamine synthetase marks transition from collecting into conducting hepatic veins.

The expression of glutamine synthetase (GS) is confined to a rim of hepatocytes surrounding the efferent hepatic veins in all mammalian species investigated. In rat liver, a two- to three-cell thick layer of GS-positive (GS(+)) hepatocytes uniformly surrounds the two to four terminal branching generations of the hepatic vein that collect blood from sinusoids (central veins). With increasing diameter of the efferent vessel, this multilayered rim of GS(+) hepatocytes becomes confined to patches surrounding the decreasing number of central vein outlets. The remaining part of the wall of these sublobar hepatic veins is bordered by a one-cell thick layer of GS(+) hepatocytes. Around still larger veins, this single-cell layer of GS(+) hepatocytes gradually disappears. The expression pattern of GS is therefore a convenient biological parameter to delimit sinusoidal draining ("collecting") from nondraining ("conducting") surfaces in the wall of the efferent hepatic vessels. The hepatocytes surrounding a single tree of central veins together form a compound liver lobule. (J Histochem Cytochem 47:1507-1511, 1999)     

brinker and optomotor-blind act coordinately to initiate development of the L5 wing vein primordium in Drosophila

The stereotyped pattern of Drosophila wing veins is determined by the action of two morphogens, Hedgehog (Hh) and Decapentaplegic (Dpp), which act sequentially to organize growth and patterning along the anterior-posterior axis of the wing primordium. An important unresolved question is how positional information established by these morphogen gradients is translated into localized development of morphological structures such as wing veins in precise locations. In the current study, we examine the mechanism by which two broadly expressed Dpp signaling target genes, optomotor-blind (omb) and brinker (brk), collaborate to initiate formation of the fifth longitudinal (L5) wing vein. omb is broadly expressed at the center of the wing disc in a pattern complementary to that of brk, which is expressed in the lateral regions of the disc and represses omb expression. We show that a border between omb and brk expression domains is necessary and sufficient for inducing L5 development in the posterior regions. Mosaic analysis indicates that brk-expressing cells produce a short-range signal that can induce vein formation in adjacent omb-expressing cells. This induction of the L5 primordium is mediated by abrupt, which is expressed in a narrow stripe of cells along the brk/omb border and plays a key role in organizing gene expression in the L5 primordium. Similarly, in the anterior region of the wing, brk helps define the position of the L2 vein in combination with another Dpp target gene, spalt. The similar mechanisms responsible for the induction of L5 and L2 development reveal how boundaries set by dosage-sensitive responses to a long-range morphogen specify distinct vein fates at precise locations.     

Insights into the molecular mechanisms underlying diversified wing venation among insects

Insect wings are great resources for studying morphological diversities in nature as well as in fossil records. Among them, variation in wing venation is one of the most characteristic features of insect species. Venation is therefore, undeniably a key factor of species-specific functional traits of the wings; however, the mechanism underlying wing vein formation among insects largely remains unexplored. Our knowledge of the genetic basis of wing development is solely restricted to Drosophila melanogaster. A critical step in wing vein development in Drosophila is the activation of the decapentaplegic (Dpp)/bone morphogenetic protein (BMP) signalling pathway during pupal stages. A key mechanism is the directional transport of Dpp from the longitudinal veins into the posterior crossvein by BMP-binding proteins, resulting in redistribution of Dpp that reflects wing vein patterns. Recent works on the sawfly Athalia rosae, of the order Hymenoptera, also suggested that the Dpp transport system is required to specify fore- and hindwing vein patterns. Given that Dpp redistribution via transport is likely to be a key mechanism for establishing wing vein patterns, this raises the interesting possibility that distinct wing vein patterns are generated, based on where Dpp is transported. Experimental evidence in Drosophila suggests that the direction of Dpp transport is regulated by prepatterned positional information. These observations lead to the postulation that Dpp generates diversified insect wing vein patterns through species-specific positional information of its directional transport. Extension of these observations in some winged insects will provide further insights into the mechanisms underlying diversified wing venation among insects.     

Peculiarities of structural-functional organization of the motor neuropil in the dragonfly thoracic ganglia

The study considers structural-functional relations in motor neuropil of the thoracic ganglia in dragonflies-insects capable of performing very complex and fast maneuvering in flight. The motor neuropil in dragonflies was shown to be more differentiated than in less mobile insects, while its motor nuclei are more outlined and approached to each other. There were revealed dendrites of the leg muscle motoneurons (intermediate nucleus), running to the anterior and posterior nuclei that contain dendrites of the wing muscle motoneurons. A possible role of such a dendrite approaching is discussed for close functional cooperation of wing and leg muscles essential for dragonflies to catch a large prey in flight by using their legs. Peculiarities of structural organization of the wing muscle motoneurons in dragonflies and locusts are considered to suggest the greater functional capabilities of motoneurons in the dragonfly motor apparatus.     

Changing spatial patterns of DNA replication in the developing wing of Drosophila

Using an antibody against bromodeoxyuridine we have analyzed the distribution of S-phase nuclei in the wing disc of Drosophila as the larval disc transforms into the adult wing during metamorphosis. On the basis of the timing of replication three cell populations can be distinguished: the cells of the presumptive wing margin, the precursor cells of the longitudinal veins, and those of the intervein regions. In each of these populations the cell cycle is first arrested and later resumes at a specific time, so that at each developmental time point a characteristic spatial pattern of S-phase nuclei is seen. An interpretation of these changing patterns in terms of vein formation, compartments, and neural development is offered.