The comparative morphology of the wings and axillae of selected Heteroptera

The wings of selected species are here treated as functional deformable aerofoils and their morphology described with this in mind. A novel type of non-tubular vein structure in the forewing and hindwing membranes, a 'channel vein', is described. The presence of small, cuticular membrane thickenings in the hindwing is recorded and their possible functions are discussed.
A comparative account of the structure and musculature of the axillae is presented with a view in particular to clarifying some of the associated terminology.
A mechanism for wing folding is described and previously unrecorded adaptations for this purpose seen in Notonecta are described and discussed.     

Functioning of the wings and axillary sclerites of Heteroptera during flight

A high degree of automation of the wingstroke in the Heteroptera has resulted in the simplification of the axillae and associated musculature. The functions of the indirect, asynchronous flight muscles are summarised from previous investigations and original work. Functions are suggested for the tonic muscles but await confirmation by electrophysiological research.
The deformations seen in heteropteran wings can be explained in terms of muscularly generated and aerodynamic forces. Unsteady aerodynamic benefits of such deformations, though unproven, may be of primary importance. Torsion, camber change and transverse flexion are all candidates for the generation of unsteady effects. The 'near clap and peel' seen at pronation has been described in other insects by Ellington (1984a), and it seems likely that it serves the same purpose in Heteroptera.
Heteroptera, like many other insects, show wing-tip flexion. Among several possible functions, inertial stress reduction is shown to be important.     

The function of resilin in beetle wings

This account shows the distribution of elastic elements in hind wings in the scarabaeid Pachnoda marginata and coccinellid Coccinella septempunctata (both Coleoptera). Occurrence of resilin, a rubber-like protein, in some mobile joints together with data on wing unfolding and flight kinematics suggest that resilin in the beetle wing has multiple functions. First, the distribution pattern of resilin in the wing correlates with the particular folding pattern of the wing. Second, our data show that resilin occurs at the places where extra elasticity is needed, for example in wing folds, to prevent material damage during repeated folding and unfolding. Third, resilin provides the wing with elasticity in order to be deformable by aerodynamic forces. This may result in elastic energy storage in the wing.     

Design, function and evolution in the wings of holometabolous insects

The wings of Holometabola are reviewed in functional terms. Two basic types, anteroposteriorly symmetric ('symmetric') and anteroposteriorly asymmetric ('asymmetric'), are recognized and their modes of operation discussed. Most neuropteroids have asymmetric wings, whereas relative anteroposterior symmetry appears plesiomorphic for mecopteroids. Wing coupling has only occurred in lineages with wings of symmetric type, but anteroposterior asymmetry of the wing couple has later developed several times in association with improved, more versatile, flight performance. Coleoptera wings seem to have developed from predecessors of symmetric type, but their morphology is strongly influenced by the need to fold up at rest. Diptera provide an excellent illustration of the evolution of asymmetric, high-performance wings from the symmetric, 'mecopteran' type. Asymmetric and symmetric wings can both be derived from less specialized types. Both are well represented as fossils from the Permian onwards, and these supply further examples of the evolution of asymmetry from the symmetric pattern.     

Homology in Development and the Development of the Homology Concept

Homology is a central concept for Developmental Evolution. HereI argue that homology should be explained within the referenceprocesses of development and evolution; development becauseit is the proximate cause of morphological characters and evolutionbecause it deals with organic transformations and stability.This was already recognized by Hans Spemann in 1915. In a seminalessay "A history and critique of the homology concept" Spemannanalyzed the history and present problems of the homology concept.Here I will continue Spemann's project and analyze some of the20th century contributions to homology. I will end with a fewreflections about the connections between developmental processesand homology and conclude that developmental processes are inherentin (i) the assessment of homology, (ii) the explanation of homology,(iii) the origin of evolutionary innovations (incipient homologues),and (iv) can be considered homologous themselves.     

Hox and wings

In many bilaterian phyla, appendages are morphological traits that characterise the identity of the various body parts. In pterygote insects, wings are dorsal appendages on the thorax. The famous "bithorax" fly created by Ed Lewis is the emblematic example of the role of Hox genes.1 Now, Tomoyasu et al.,2 using classical genetics, transgenesis and RNAi, have examined the function of thoracic Hox genes in the beetle Tribolium castaneum. Beetles have rigid elytra in place of the first pair of wings. Instead of the expected transformation of the elytron into a wing, loss of Hox genes' function leads to the homeotic transformation of the second pair of dorsal appendages, the wings, into elytra. This has important consequences for the way that we see the role of Hox genes in development and evolution.     

Structure and function of the dorsal abdominal gland and defence mechanism in cocoa-capsids (Miridae: Heteroptera)

An account is given of the structure of the dorsal abdominal gland in Distantiella theobroma, Sahlbergella singularis, Bryocoropsis laticollis and Helopeltis corbisieri. The function is discussed, and defence mechanisms in the four species are described.     

Homology modeling and function of trehalose synthase from Pseudomonas putida P06

Trehalose is a non-reducing disaccharide that has wide applications in the food industry and pharmaceutical manufacturing. Trehalose synthase (TreS) from Pseudomonas putida P06 catalyzes the reversible interconversion of maltose and trehalose and may have applications in the food industry. However, the catalytic mechanism of TreS is not well understood. Here, we investigated the structural characteristics of this enzyme by homology modeling. The highly conserved Asp294 residue was identified to be critical for catalytic activity. In addition, flexible docking studies of the enzyme–substrate system were performed to predict the interactions between TreS and its substrate, maltose. Amino acids that interact extensively with the substrate and stabilize the substrate in an orientation suitable for enzyme catalysis were identified. The importance of these residues for catalytic activity was confirmed by the biochemical characterization of the relevant mutants generated by site-directed mutagenesis.     

Phenotypic selection and function of reproductive behavior in the subsocial bug Elasmucha putoni (Heteroptera: Acanthosomatidae)

To investigate the function of maternal care and determinantsof reproductive success in the subsocial bug Elasmucha putoni (Heteroptera: Acanthosomatidae), I used two different approaches,the measurement of phenotypic selection and female-removalexperiments under conditions differing in biotic-environmentalpressure. For two field populations, unattended eggs and youngernymphs consistently suffered severe predation pressure andattendance by parent females greatly enhanced their survival.In contrast, under enemy-excluded conditions, offspring performance was not reduced in broods without parent females, indicatingthat maternal care functions as a physical defense againstpredators. However, the determinant of female reproductivesuccess in E. putoni in the field was not the care behavioralone. Selection gradient analysis showed that early seasonoviposition and larger clutch size, as well as a longer durationof care by a female, was favored during the breeding episode.This study is the first to evaluate phenotypic selection onparental care and other reproductive traits in arthropods.     

Homology and the origin of correspondence

Homology is a natural kind term and a precise account of what homologyis has to come out of theories about the role of homologues in evolution anddevelopment. Definitions of homology are discussed with respect to the questionas to whether they are able to give a non-circular account of thecorrespondenceor sameness referred to by homology. It is argued that standard accounts tiehomology to operational criteria or specific research projects, but are not yetable to offer a concept of homology that does not presuppose a version ofhomology or a comparable notion of sameness. This is the case for phylogeneticdefinitions that trace structures back to the common ancestor as well as fordevelopmental approaches such as Wagner's biological homology concept. Incontrast, molecular homology is able to offer a definition of homology in genesand proteins that explicates homology by reference to more basic notions.Molecular correspondence originates by means of specific features of causalprocesses. It is speculated that further understanding of morphogenesis mightenable biologists to give a theoretically deeper definition of homology alongsimilar lines: an account which makes reference to the concrete mechanisms thatoperate in organisms.     

Support and deformability in insect wings

Coupled investigations of insect wing movements and detailed wing morphology are in progress, and some functional principles underlying wing design are emerging. High speed cine and still photography and stroboscopy indicate that most wings undergo orderly deformation in flight. Common patterns are described and their significance discussed in the light of recent aerodynamic studies.
Many aspects of wing morphology–venational features, relief, thickened areas, flexionlines and vein fractures–may be related to the control of three-dimensional shape while beating. It is usually possible to distinguish areas specialized for deformability, and for support and the limiting of deformation. Some structural adaptations for these roles are described and illustrated.     

Homology and errors

A recent review of the homology concept in cladistics is critiqued in light of the historical literature. Homology as a notion relevant to the recognition of clades remains equivalent to synapomorphy. Some symplesiomorphies are “homologies” inasmuch as they represent synapomorphies of more inclusive taxa; others are complementary character states that do not imply any shared evolutionary history among the taxa that exhibit the state. Undirected character‐state change (as characters optimized on an unrooted tree) is a necessary but not sufficient test of homology, because the addition of a root may alter parsimonious reconstructions. Primary and secondary homology are defended as realistic representations of discovery procedures in comparative biology, recognizable even in Direct Optimization. The epistemological relationship between homology as evidence and common ancestry as explanation is again emphasized. An alternative definition of homology is proposed. © The Willi Hennig Society 2012.     

Origin and development of the wings of Lepidoptera

Summary In Lepidoptera (Pieris rapae) the imaginal wing discs originate by a thickening of the thoracic pleural hypodermis at the close of embryonic life. The tracheoles originate in the endotracheal layer at the close of the first moult and become functional at the close of second moult. They grow very rapidly and reach their limit of penetration in the larval wing at end of third moult. At the beginning of fifth instar they become less abundant degenerate and disappear by absorption in the pupal wing.The permanent system of tracheoles evaginate from the wing trachea in the prepupal stage and become functional during pupal life. The wing passes to the exterior by a drawing back of the hypodermal wing sac. The rapid expansion of the wing causes the withdrawal of hypodermis. The sharp division into larval and pupal stages applies more particularly to the exterior of the body as they follow one another after successive moults, the internal development is a continuous series of transformations, between which there is no sharp line of demarcation. Yet on the whole the form of larva, pupa and imago are kept distinct in adaptation to their separate environments and habits.

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Zusammenfassung Die imaginalen Flügelscheiben nehmen bei Schmetterlingen (Pieris rapae) ihren Ursprung ans einer Verdickung der pleuralen Thoraxhypodermis am Schlusse des Embryonallebens. Die Tracheolen nehmen ihren Ursprung in der endotrachealen Schicht am Schlusse der ersten Häutung und beginnen am Schlusse der zweiten zu funktionieren. Sie wachsen sehr rasch und erreichen die Grenze ihres Eindringens in den Larvenflügel am Ende der dritten Häutung. Am Beginne der fünften werden sie dagegen weniger reichlich, degenerieren und verschwinden durch Absorption im Flügel der Puppe.Das bleibende Tracheolensystem stülpt sich aus der Flügeltrachea im Vorpuppenstadium aus und tritt während des Puppenlebens in Funktion. Der Flügel gelangt durch ein Zurückziehen des hypodermalen Flügelsackes an die Außenwelt. Die rapide Ausdehnung des Flügels veranlaßt das Zurückweichen der Hypodermis. Die scharfe Teilung in Larven- und Puppenstadium findet ihre Anwendung mehr auf das Körperäußere, da das eine auf das andre nach successiven Häutungen folgt. Die innere Entwicklung ist eine kontinuierliche Reihe von Formveränderungen, zwischen denen keine scharfe Trennungslinie existiert. Im ganzen aber bleiben die Larven-, Puppen- und Imagoform je nach Anpassung an ihre verschiedene Umgebung und ihre verschiedenen Lebensgewohnheiten streng getrennt.

     

Homology and misdirection

Hennig (1966) recognized symplesiomorphies as homologies, and that view is logically correct under the concept of homology (homogeny) prevalent among evolutionists since 1870. Nelson and Platnick (1981) instead wanted homology to exclude symplesiomorphies for reasons that they never made clear but which certainly included opposition to Hennig. They and some of their followers, most recently Platnick (2013) and Brower and de Pinna (2013), have continued to advocate that anti‐Hennigian position, often under the slogan “homology equals synapomorphy,” while ironically passing themselves off as cladists and often using ambiguous or falsified citations to pretend that legitimate phylogeneticists think likewise. Such authors have seldom shown much concern for accuracy or logic, with the result that a great deal of print has been wasted. Those problems can be avoided simply by maintaining a Hennigian view and so discarding the purported equivalence of homology and synapomorphy.     

Studies on D-alanine in the Heteroptera*

Four milkweed associated insects were assayed for free D-alanine by gas-liquid chromatographic methods. Oncopeltus fasciatus and Lygaeus kalmii, reared aseptically on milkweed seeds, showed high titres of free D-alanine. Labidomera clivicollis showed traces of the D-isomer whereas none could be detected in Tetraopes tetrophthalmus. O.fasciatus reared aseptically on sunflower seed showed high titres of D-alanine. No D-alanine could be detected in milkweed seed hydrolysates. Several museum specimens of the orders Hemiptera and Homoptera were assayed and all showed high titres of D-alanine.     

Homology and the hierarchy of biological systems

Homology is the similarity between organisms due to common ancestry. Introduced by Richard Owen in 1843 in a paper entitled "Lectures on comparative anatomy and physiology of the invertebrate animals", the concept of homology predates Darwin's "Origin of Species" and has been very influential throughout the history of evolutionary biology. Although homology is the central concept of all comparative biology and provides a logical basis for it, the definition of the term and the criteria of its application remain controversial. Here, I will discuss homology in the context of the hierarchy of biological organization. I will provide insights gained from an exemplary case study in evolutionary developmental biology that indicates the uncoupling of homology at different levels of biological organization. I argue that continuity and hierarchy are separate but equally important issues of homology.