Wetting Characteristics of Insect Wing Surfaces

Biological tiny structures have been observed on many kinds of surfaces such as lotus leaves,which have an effect on thecoloration of Morpho butterflies and enhance the hydrophobicity of natural surfaces.We investigated the micro-scale andnano-scale structures on the wing surfaces of insects and found that the hierarchical multiple roughness structures help in enhancingthe hydrophobicity.After examining 10 orders and 24 species of flying Pterygotan insects,we found that micro-scaleand nano-scale structures typically exist on both the upper and lower wing surfaces of flying insects.The tiny structures such asdenticle or setae on the insect wings enhance the hydrophobicity,thereby enabling the wings to be cleaned more easily.And thehydrophobic insect wings undergo a transition from Cassie to Wenzel states at pitch/size ratio of about 20.In order to examinethe wetting characteristics on a rough surface,a biomimetic surface with micro-scale pillars is fabricated on a silicon wafer,which exhibits the same behavior as the insect wing,with the Cassie-Wenzel transition occurring consistently around apitch/width value of 20.     

基于四种金龟的昆虫后翅关节骨片三维形态复杂性研究

【目的】昆虫的翅非常精巧与灵活,翅脉及翅关节的形态及功能长久以来受到众多领域科学家的广泛关注。由于历史条件的限制,昆虫翅的研究主要集中在翅脉,即使少量的有关翅关节形态的研究也主要是停留在二维形态数据分析的层面上。更重要的是,各骨片内部形态结构还未见报道。本研究的目的就是为了重建翅关节骨片内部和外部复杂的三维形态结构,全面呈现利用传统形态学方法无法获得的形态学信息,进而深入探究昆虫翅的形态与功能的关系。【方法】本文利用显微CT对鞘翅目4种金龟进行了扫描,通过计算机三维重建技术,对折叠和展开状态时后翅关节各个骨片（第1, 2和3腋片及中片）的内部和外部的三维形态进行研究,展示和分析昆虫翅关节内部与外部形态结构和空间运动的复杂性。【结果】翅关节骨片的三维重建模型及虚拟切面图展示了其复杂的外部形态,主要表现在表面曲率的不均匀变化和部分结构的互相遮挡两个方面。前者主要表现骨片表面具有突起、沟槽、弯折以及外长物等。后者指各骨片均呈现了不同程度的弯折,有的弯折还会互相接触,最终形成筒状结构,这样不可避免造成部分结构被遮挡或包裹。三维重建模型的断层图显示了翅关节骨片并非是实心的结构,而是分为两层：靠近表皮的为高度骨化的外骨骼,而靠近骨片核心则为疏松的类似海绵状结构。本文还展示了各个骨片在后翅折叠状和展开状态下的空间位置,并对所研究的4个科的翅关节骨片的三维形态进行了比较。【结论】翅关节骨片具有复杂的内部和外部形态结构。关节骨片的内部海绵结构和外层强烈骨化的双层结构,可能与其尽量减小骨片的重量和节约运动能量,同时又尽量保持骨片的刚性结构的形态适应策略有关。此类形态适应在材料学、空气动力学等领域具有重要的仿生学意义。     

Structural Characteristics of Allomyrina Dichotoma Beetle＇s Hind Wings for Flapping Wing Micro Air Vehicle

In this study, we present a complete structural analysis ofAllomyrina dichotoma beetle＇s hind wings by investigating their static and dynamic characteristics. The wing was subjected to the static loading to determine its overall flexural stiffness. Dy- namic characteristics such as natural frequency, mode shape, and damping ratio of vibration modes in the operating frequency range were determined using a Bruel ＆ Kjaer fast Fourier transform analyzer along with a laser sensor. The static and dynamic characteristics of natural Allomyrina dichotoma beetle＇s hind wings were compared to those of a fabricated artificial wing. The results indicate that natural frequencies of the natural wing were significantly correlated to the wing surface area density that was defined as the wing mass divided by the hind wing surface area. Moreover, the bending behaviors of the natural wing and artificial wing were similar to that of a cantilever beam. Furthermore, the flexural stiffness of the artificial wing was a little higher than that of the natural one whereas the natural frequency of the natural wing was close to that of the artificial wing. These results provide important information for the biomimetic design of insect-scale artificial wings, with which highly ma- neuverable and efficient micro air vehicles can be designed.     

Effects of Methanol on Wettability of the Non-Smooth Surface on Butterfly Wing

The contact angles of distilled water and methanol solution on the wings of butterflies were determined by a visual contact angle measuring system. The scale structures of the wings were observed using scanning electron microscopy, The influence of the scale micro- and ultra-structure on the wettability was investigated. Results show that the contact angle of distilled water on the wing surfaces varies from 134.0° to 159.2°. High hydrophobicity is found in six species with contact angles greater than 150°. The wing surfaces of some species are not only hydrophobic but also resist the wetting by methanol solution with 55% concentration. Only two species in Parnassius can not resist the wetting because the micro-structure （spindle-like shape） and ultra-structure （pinnule-like shape） of the wing scales are remarkably different from that of other species. The concentration of methanol solution for the occurrence of spreading/wetting on the wing surfaces of different species varies from 70% to 95%. After wetting by methanol solution for 10 min, the distilled water contact angle on the wing surface increases by 0.8°-2.1°, showing the promotion of capacity against wetting by distilled water.     

Aerodynamic performance of a hovering hawkmoth with flexible wings: a computational approach

Insect wings are deformable structures that change shape passively and dynamically owing to inertial and aerodynamic forces during flight. It is still unclear how the three-dimensional and passive change of wing kinematics owing to inherent wing flexibility contributes to unsteady aerodynamics and energetics in insect flapping flight. Here, we perform a systematic fluid-structure interaction based analysis on the aerodynamic performance of a hovering hawkmoth, Manduca, with an integrated computational model of a hovering insect with rigid and flexible wings. Aerodynamic performance of flapping wings with passive deformation or prescribed deformation is evaluated in terms of aerodynamic force, power and efficiency. Our results reveal that wing flexibility can increase downwash in wake and hence aerodynamic force: first, a dynamic wing bending is observed, which delays the breakdown of leading edge vortex near the wing tip, responsible for augmenting the aerodynamic force-production; second, a combination of the dynamic change of wing bending and twist favourably modifies the wing kinematics in the distal area, which leads to the aerodynamic force enhancement immediately before stroke reversal. Moreover, an increase in hovering efficiency of the flexible wing is achieved as a result of the wing twist. An extensive study of wing stiffness effect on aerodynamic performance is further conducted through a tuning of Young's modulus and thickness, indicating that insect wing structures may be optimized not only in terms of aerodynamic performance but also dependent on many factors, such as the wing strength, the circulation capability of wing veins and the control of wing movements.     

Wing flexibility enhances load-lifting capacity in bumblebees

The effect of wing flexibility on aerodynamic force production has emerged as a central question in insect flight research. However, physical and computational models have yielded conflicting results regarding whether wing deformations enhance or diminish flight forces. By experimentally stiffening the wings of live bumblebees, we demonstrate that wing flexibility affects aerodynamic force production in a natural behavioural context. Bumblebee wings were artificially stiffened in vivo by applying a micro-splint to a single flexible vein joint, and the bees were subjected to load-lifting tests. Bees with stiffened wings showed an 8.6 per cent reduction in maximum vertical aerodynamic force production, which cannot be accounted for by changes in gross wing kinematics, as stroke amplitude and flapping frequency were unchanged. Our results reveal that flexible wing design and the resulting passive deformations enhance vertical force production and load-lifting capacity in bumblebees, locomotory traits with important ecological implications.     

Differences in wing morphology between juvenile and adult European Turtle Doves Streptopelia turtur: implications for migration and predator escape

Behaviour has direct links to wing morphology in bird species. Many studies have postulated migration to be one of the most important forces of selection acting on wing morphology, particularly in relation to wing pointedness. Studies in passerines have found that adults have longer and more pointed wings than juveniles, especially in migratory species. We analysed differences in wing morphology between age groups of the European Turtle Dove, a non‐passerine migratory species that benefits from rounded wings during their daily activity, due to its ground‐feeding behaviour and acrobatic flight style. Our results show that adults of this species have longer but more rounded wings than juveniles. This suggests that in this species wing morphology in juveniles is selected to facilitate the first migration, whereas other selection forces (e.g. flight manoeuvrability) are more important after the first moult. These data also explain why juveniles are not as adept at escaping from predators or hunters as adults.     

Variations in the arrangement of sensory bristles along butterfly wing margins

The surfaces of insect wings exhibit numerous sensilla, which have been suggested to have a behavioral function. Some evidence suggests that the sensory bristles along the wing margin of lepidopteran insects (butterflies and moths) are involved in the regulation of wing movement. We investigated the arrangement of sensory bristles along the wing margins of 62 species of papilionoid butterflies, using light-microscopic examination of mounts of whole wings after removing the scales surrounding the bristles. In the majority of the wings examined, bristles were located on the ventral wing surfaces and were continuously distributed along the wing margins, except in the vicinity of the wing bases. In some wings, bristles were also located on the dorsal wing surfaces, and were continuously or discontinuously distributed along the wing margins of different species. In a minority of the species studied, we observed bristle distribution in the vicinity of the wing base, discontinuous bristle distribution on both the dorsal and ventral wing surfaces, or an absence of bristles along the wing margins. This variation in the arrangement of bristles along the wing margins is discussed in relation to the reception and transmission of sensory information in the wings.     

Surface Laser Scanning of Fossil Insect Wings

Primary homologization of wing venation is of crucial importance in taxonomic studies of fossil and recent insects, with implications for large-scale phylogenies. Homologization is usually based on relative relief of veins (with an insect ground plan of alternating concave and convex vein sectors). However, this method has led to divergent interpretations, notably because vein relief can be attenuated in fossil material or because wings were originally flat. In order to interpret better vein relief in fossil insect wings, we tested the application of non-contact laser scanning. This method enables high resolution three-dimensional (3-D) data visualization of a surface, and produces high quality images of fossil insect wings. These images facilitate and improve interpretation of the homologization of wing venation. In addition, because the surface information is digitised in three axes (X, Y, Z), the data may be processed for a wide range of surface characteristics, and may be easily and widely distributed electronically. Finally, this method permits users to reconstruct accurately the fossils and opens the field of biomechanical interpretation using numerical modelling methods.     

Comparative and functional morphology of wing coupling structures in Trichoptera: Annulipalpia

Several orders of morphologically four‐winged insects have evolved mechanisms that enforce a union between the mesothoracic and metathoracic wings (forewings and hindwings) during the wing beat cycle. Such mechanisms result in a morphologically tetrapterous insect flying as if it were functionally dipterous, and these mechanisms have been described for several insect orders. The caddisfly suborders Annulipalpia and Integripalpia (Trichoptera) each have evolved a wing coupling apparatus, with at least three systems having evolved within the suborder Annulipalpia. The comparative and inferred functional morphology of the putative wing coupling mechanisms is described for the annulipalpian families Hydropsychidae (subfamilies Macronematinae and Hydropsychinae), Polycentropodidae and Ecnomidae, and a novel form‐functional complex putatively involved with at‐rest forewing‐forewing coupling is described for Hydropsychidae: Smicrideinae. It is proposed that the morphology of the wing coupling apparatuses of Hydropsychinae and Macronematinae are apomorphies for those clades. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.     

A revised interpretation of the wing base structure in Odonata

Abstract Homology of the wing base structure in the Odonata is highly controversial, and many different interpretations of homology have been proposed. In extreme cases, two independent origins of insect wings have been suggested, based on comparative morphology between the odonate and other pterygote wing bases. Difficulties in establishing homology of the wing base structures between Odonata and other Pterygota result mainly from their extreme differences in morphology and function. In the present paper, we establish homology of the wing base structures between Neoptera, Ephemeroptera and Odonata using highly conservative and unambiguously identifiable characters (the basal wing hinge and subcostal veins) as principal landmarks. Homology of the odonate wing base structure with those of Ephemeroptera and Neoptera can be identified reliably. Based on this interpretation, the ancestral condition of the insect wing base structure is discussed.     

Resilin in dragonfly and damselfly wings and its implications for wing flexibility

Although there is mounting evidence that passive mechanical dynamics of insect wings play an integral role in insect flight, our understanding of the structural details underlying insect wing flexibility remains incomplete. Here, we use comparative morphological and mechanical techniques to illuminate the function and diversity of two mechanisms within Odonata wings presumed to affect dynamic wing deformations: flexible resilin vein‐joints and cuticular spikes. Mechanical tests show that joints with more resilin have lower rotational stiffness and deform more in response to a load applied to an intact wing. Morphological studies of 12 species of Odonata reveal that resilin joints and cuticular spikes are widespread taxonomically, yet both traits display a striking degree of morphological and functional diversity that follows taxonomically distinct patterns. Interestingly, damselfly wings (suborder Zygoptera) are mainly characterized by vein‐joints that are double‐sided (containing resilin both dorsally and ventrally), whereas dragonfly wings (suborder Epiprocta) are largely characterized by single‐sided vein‐joints (containing resilin either ventrally or dorsally, but not both). The functional significance and diversity of resilin joints and cuticular spikes could yield insight into the evolutionary relationship between form and function of wings, as well as revealing basic principles of insect wing mechanical design. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc.     

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

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.     

The development of butterfly settling posture: the role of predators, climate, hostplant-habitat and phylogeny

A model is described that explains the evolution of adult butterfly settling posture. The role of climate and the activity of butterflies is fundamental in determining the effectiveness of primary and secondary visual defence systems based on wing colour and pattern. Both dorsal (open winged) and lateral (closed wing) postures can be regarded as evolutionarily conservative; common postures are found within almost all sub-families. Lateral postures tend to be associated with cool temperate and arctic climates. Conditions that are marginal for activity place an emphasis on the development of underside primary defence, uncompromised by secondary visual defences. In turn, any primary defensive function of the upper surface is reduced but secondary defence may be emphasized. There is interaction between segregation of functions on different wing surfaces and the development of lateral posture. The use of predictable hostplant-habitat structures reduces the role of visual communication in mate-location and facilitates the adoption of a lateral resting posture and a cryptic underside wing pattern. When hostplant-habitat structures are unpredictable but a lateral posture is adopted, primary defence is more important than visual communication for the resting insect. Adjustments of wing morphology facilitate postural conservatism, allowing species to retain particular postures in new hostplant-habitats and climatic zones.     

Hox基因与昆虫翅的特化

自从1978年E.B. Lewis描述了著名的果蝇双胸突变体（bithorax）以来,大量的比较发育遗传学研究为我们揭示了形态进化的遗传基础,从而使形态进化研究进入了一个新的时代。同时,Hox基因的研究也成为这一领域的焦点。本文综述了昆虫翅的起源及其特化类群翅的发育遗传学研究的最新进展。一般认为,原始的有翅昆虫胸腹部多附肢（包括翅）; 之后不同的体节受到了不同Hox的抑制,形成两对翅以及前后翅的分化; Ubx的不同表达导致了前后翅的分化,并且Ubx负责识别后翅。我们选择翅特化最为显著的3个类群——鞘翅目（T2鞘翅）、双翅目（T3平衡棒）和捻翅目（T2平衡棒）,结合Hox的表达情况讨论了翅的特化机理。目前已知双翅目和鞘翅目的翅的控制模式存在巨大差异,两种模式的比较研究对于理解翅的形态进化具有重要的意义。但是对捻翅目昆虫的研究则很少。     

Directional and stabilizing selection on wing size and shape in migrant and resident monarch butterflies, Danaus plexippus (L.), in Cuba

The majority of migrant monarchs (Danaus plexippus) from the eastern USA and south‐eastern Canada migrate to Mexico; however, some of them migrate to Cuba. Cuban migrants hatch in south‐east Canada and eastern USA, and then engage in a southern trip of 4000 km to this Caribbean island. In Cuba, these migrants encounter resident monarchs, which do not migrate, and instead move between plant patches looking for nectar, mating partners and host plants. These differences in flight behaviour between migrant and resident Cuban monarchs may have resulted in different selective pressures in the wing size and shape. Two modes of selection were tested, directional and stabilizing. In addition, wing condition was compared between these two groups. Monarchs were collected for 4 years in Cuba and classified as resident or migrant using two independent techniques: Thin‐layer chromatography and stable hydrogen and stable carbon isotope measurements. Wing size was measured and wing condition was rated in the butterflies. Fourier analysis and wing angular measurements were used to assess wing shape differences. Migrants have significantly longer wings than residents, thus supporting the action of directional selection on wing size. In addition, directional selection acts on wing shape; that is, migrant females differ significantly from resident females in their wing angles. However, the results do not support the action of stabilizing selection: there was no significant variance between migrant and resident monarchs in their wing size or shape. Also, migrant females and males differed in wing condition as a result of differences in flight behaviour. In conclusion, eastern North American monarchs offer a good opportunity to study the selective pressures of migration on wing morphology and how different migratory routes and behaviours are linked to wing morphology and condition. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 92 , 605–616.     

Morphological variation associated with dispersal capacity in a tree‐killing bark beetle Dendroctonus ponderosae Hopkins

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The Microstructures of Butterfly Wing Scales in Northeast of China

There are billions of tiny scales on the butterfly wings, which array regularly as the tiles on the roof. Such tilts can form various colors of the wing and afford the species many abilities to survive and propagate. Morphological experiments on the wing scales of six butterfly species living in northeast of China were conducted. By the optics microscope; the form, geometry dimension and array of the scales were observed generally. By using scanning electron microscope （SEM）, the 2D scanning and measurement were carried out and the surface micro configurations of scales were observed. The dimension and microstructure characteristics of the cross section of single scale were achieved through transmission electron microscope （TEM）. Finally, by using 3D software, three 3D models were described and the 3D visual effect was achieved. This work can put forward a basic method for the future study on the morphology of biological microstructure.     

Functional morphology and structural characteristics of the hind wings of the bamboo weevil Cyrtotrachelus buqueti (Coleoptera,Curculionidae)

Research data of the microstructure and surface morphology of insect wings have been used to help design micro air vehicles (MAV) and coating materials. The present study aimed to examine the microstructure and morphology of the hind wings of Cyrtotrachelus buqueti using inverted fluorescence microscopy (IFM), scanning electron microscopy (SEM), and a mechanical testing system. IFM was used to investigate the distribution of resilin in the hind wing, and SEM was performed to assess the functional characteristics and cross-sectional microstructure of the wings. Moreover, mechanical properties regarding the intersecting location of folding lines and the bending zone (BZ) were examined. Resilin, a rubber-like protein, was found in several mobile joints and in veins walls that are connected to the wing membranes. Taken together, structural data, unfolding motions, and results of tensile testing suggest two conclusions on resilin in the hind wing of C. buqueti: firstly, the resilin distribution is likely associated with specific folding mechanisms of the hind wings, and secondly, resilin occurs at positions where additional elasticity is needed, such as in the bending zone, in order to prevent structural damage during repeated folding and unfolding of the hind wings. The functional significance of resilin joints may shed light on the evolutionary relationship between morphological and structural hind wing properties.     

Longitudinal data reveal ontogenetic changes in the wing morphology of a long‐distance migratory bird

In migratory bird species, juveniles normally have shorter and more rounded wings than adults. The causes of this age‐specific difference in wing morphology, however, are largely unknown. Here, we used longitudinal data collected over 3 years from a Pied Flycatcher Ficedula hypoleuca population to assess whether age‐related differences in wing morphology are a consequence of ontogenetic changes or of selection favouring birds with longer and more pointed wings. Our study provides evidence of ontogenetic changes in wing length and shape, whereby birds grow longer and more pointed wings as they grow older. Age‐dependent variation is likely to be adaptive and may partly explain age differences in spring migration phenology and breeding success.