Analysis of modularity and integration suggests evolution of dragonfly wing venation mainly in response to functional demands

Insect wings show a high variability in wing venation. Selection for function, developmental pathways and phylogeny likely influenced wing vein diversification, however, quantitative data to estimate these influences and their interplay are missing. Here, it is tested how dragonfly wing vein configuration is influenced by functional demands, development, phylogeny and allometry using the concepts of modularity and integration. In an evolutionary context, modules are sets of characters that evolve in relative independence to other characters, while integration refers to a high degree of association between subparts of a structure. Results show allometric and phylogenetic signal in the wing shape variation, however, patterns of integration and modularity are not influenced by these two factors. Overall, dragonfly wings are highly integrated structures with almost no modular signal. Configuration changes in one wing vein or wing area thus influence wing shape as a whole. Moreover, the fore- and hindwings correlate with each other in their evolutionary shape variation supporting biomechanical data of wing interdependence. Despite the overall high degree of evolutionary integration, functional hypotheses of modularity could be confirmed for two wing areas, the arculus–triangle complex at the base of the wing which is responsible for passive wing folding especially during flapping flight and the location of the pterostigma, a coloured wing cell which is more heavy that other wing cells and passively regulates wing pitch as well as critical flight speeds during gliding. Although evolving as distinct modules, these specific vein regions also show high integration and evolve at the same rates like the whole wing which suggests an influence of these structures on the shape evolution of the rest of the wing. Their biomechanical role as passive regulators of wing corrugation and wing pitch suggests that these structures decisively influenced the evolution of advanced modern flight styles and explains their retention once they had evolved early within the lineage Odonatoptera

Form-function relationships in dragonfly mandibles under an evolutionary perspective

© 2017 The Author(s). Functional requirements may constrain phenotypic diversification or foster it. For insect mouthparts, the quantification of the relationship between shape and function in an evolutionary framework remained largely unexplored. Here, the question of a functional influence on phenotypic diversification for dragonfly mandibles is assessed with a large-scale biomechanical analysis covering nearly all anisopteran families, using finite element analysis in combination with geometric morphometrics. A constraining effect of phylogeny could be found for shape, the mandibular mechanical advantage (MA), and certain mechanical joint parameters, while stresses and strains, the majority of joint parameters and size are influenced by shared ancestry. Furthermore, joint mechanics are correlated with neither strain nor mandibular MA and size effects have virtually play no role for shape or mechanical variation. The presence of mandibular strengthening ridges shows no phylogenetic signal except for one ridge peculiar to Libelluloidea, and ridge presence is also not correlated with each other. The results suggest that functional traits are more variable at this taxonomic level and that they are not influenced by shared ancestry. At the same time, the results contradict the widespread idea that mandibular morphology mainly reflects functional demands at least at this taxonomic level. The varying functional factors rather lead to the same mandibular performance as expressed by the MA, which suggests a many-to-one mapping of the investigated parameters onto the same narrow mandibular performance space

A biomechanical analysis of prognathous and orthognathous insect head capsules: Evidence for a many to one mapping of ridge strain to head strain

Insect head shapes are remarkably variable but the influences of these changes on biomechanical performance are unclear. Among “basal” winged insects, such as dragonflies, mayflies, earwigs, and stoneflies, some of the most prominent anatomical changes are the general mouthpart orientation, eye size and the connection of the endoskeleton to the head. Here, we assess these variations for the first time using modern engineering methods including multibody dynamics modelling and finite element analysis in order to quantify and compare their influence on overall head performance. We show that a range of peculiar structures such as the genal/subgenal, epistomal, and circumoccular areas are consistently highly loaded in all species, despite drastically differing morphologies in species with forward projecting (prognathous) and downwards projecting (orthognathous) mouthparts. Sensitivity analyses show that the presence of eyes has a negligible influence on head capsule strain if a circumoccular ridge is present. In contrast, the connection of the dorsal endoskeletal arms to the head capsule especially affects overall head loading in species with downward projecting mouthparts. Analysis of the relative strains between species for each head region reveals that most head regions map onto a similar biomechanical performance space thus showing a many to one mapping of differing forms to the same functional space. Concerted changes in head substructures such as the subgenal area, the endoskeleton and the epistomal area lead to a consistent relative loading pattern in prognathous and orthognathous insects. It appears that biting-chewing loads are managed by a system of strengthening ridges on the head capsule irrespective of the general mouthpart and head orientation. Concerted changes in ridge and endoskeleton configuration allow more radical anatomical changes such as general mouthpart orientation which could be an explanation for the variability of this trait among insects. In an evolutionary context, many to one mapping of diverse forms to similar functions indeed could have fostered the dynamic diversification processes seen in insects

Ultra high-resolution biomechanics suggest that substructures within insect mechanosensors decisively affect their sensitivity

Insect load sensors, called campaniform sensilla (CS), measure strain changes within the cuticle of appendages. This mechanotransduction provides the neuromuscular system with feedback for posture and locomotion. Owing to their diverse morphology and arrangement, CS can encode different strain directions. We used nano-computed tomography and finite-element analysis to investigate how different CS morphologies within one location-the femoral CS field of the leg in the fruit fly Drosophila-interact under load. By investigating the influence of CS substructures' material properties during simulated limb displacement with naturalistic forces, we could show that CS substructures (i.e. socket and collar) influence strain distribution throughout the whole CS field. Altered socket and collar elastic moduli resulted in 5% relative differences in displacement, and the artificial removal of all sockets caused differences greater than 20% in cap displacement. Apparently, CS sockets support the distribution of distal strain to more proximal CS, while collars alter CS displacement more locally. Harder sockets can increase or decrease CS displacement depending on sensor location. Furthermore, high-resolution imaging revealed that sockets are interconnected in subcuticular rows. In summary, the sensitivity of individual CS is dependent on the configuration of other CS and their substructures

Hubungan Kadar Carcinoembryonic Antigen (Cea) Dan Albumin Serum Dengan Lokasi Kanker Kolorektal Studi Kasus Di Rsup Dr. Kariadi

Latar belakang : Kolonoskopi masih menjadi alat deteksi utama untuk mengetahui lokasi kanker kolorektal. Diperlukan pemantuan petanda yang tidak invasif untuk membantu mengetahui lokasi kanker kolorektal. Kadar Carcinoembryonic antigen (CEA) dan albumin pada pasien kanker kolorektal dapat digunakan sebagai petanda lokasi dan prognosis.Tujuan : Untuk mengetahui hubungan kadar carcinoembryonic antigen (CEA) dan albumin serum dengan lokasi kanker kolorektal.Metode Penelitian : ini merupakan penelitian observasional dengan desain belah lintang. Data didapatkan dari rekam medik pasien kanker kolorektal di RSUP Dr. Kariadi dari Januari 2012-Desember 2015. Sejumlah 63 pasien menjadi subyek penelitian. Kadar CEA dan albumin dilihat dari hasil pemeriksaan laboratorium darah sebelum terapi. Lokasi tumor diketahui setelah pasien menjalani prosedur sigmoidoskopi atau kolonoskopi. Uji hipotesis yang digunakan adalah uji korelasi Spearman.Hasil : Subyek penelitian rerata berusia 50,50±13,69 tahun, dengan 49,2% pasien pria dan 50,8% wanita. Sebanyak 55,6% subyek penelitian mengalami peningkatan kadar CEA (>5 ng/mL). Sebanyak 52,4% subyek penelitian memiliki kadar albumin rendah, 46,0% memiliki kadar albumin normal, dan 1,6% memiliki kadar albumin tinggi. Lokasi tumor tersering berada di rektum (77,8%). Kadar CEA tidak berhubungan dengan lokasi tumor(rs = -0,019). Kadar albumin tidak berhubungan dengan lokasi tumor(rs= -0,060). Kadar CEA dan albumin tidak berhubungan dengan lokasi tumor(rs = -0,048).Kesimpulan : Tidak terdapat hubungan kadar CEA dan albumin dengan lokasi kanker kolorektal

A leg model based on anatomical landmarks to study 3D joint kinematics of walking in Drosophila melanogaster

Walking is the most common form of how animals move on land. The model organism Drosophila melanogaster has become increasingly popular for studying how the nervous system controls behavior in general and walking in particular. Despite recent advances in tracking and modeling leg movements of walking Drosophila in 3D, there are still gaps in knowledge about the biomechanics of leg joints due to the tiny size of fruit flies. For instance, the natural alignment of joint rotational axes was largely neglected in previous kinematic analyses. In this study, we therefore present a detailed kinematic leg model in which not only the segment lengths but also the main rotational axes of the joints were derived from anatomical landmarks, namely, the joint condyles. Our model with natural oblique joint axes is able to adapt to the 3D leg postures of straight and forward walking fruit flies with high accuracy. When we compared our model to an orthogonalized version, we observed that our model showed a smaller error as well as differences in the used range of motion (ROM), highlighting the advantages of modeling natural rotational axes alignment for the study of joint kinematics. We further found that the kinematic profiles of front, middle, and hind legs differed in the number of required degrees of freedom as well as their contributions to stepping, time courses of joint angles, and ROM. Our findings provide deeper insights into the joint kinematics of walking in Drosophila, and, additionally, will help to develop dynamical, musculoskeletal, and neuromechanical simulations

Juvenile ecology drives adult morphology in two insect orders

Most animals undergo ecological niche shifts between distinct life phases,but such shifts can result in adaptive conflicts of phenotypic traits. Metamor-phosis can reduce these conflicts by breaking up trait correlations, allowingeach life phase to independently adapt to its ecological niche. This process iscalled adaptive decoupling. It is, however, yet unknown to what extentadaptive decoupling is realized on a macroevolutionary scale in hemimeta-bolous insects and if the degree of adaptive decoupling is correlated with thestrength of ontogenetic niche shifts. It is also unclear whether the degree ofadaptive decoupling is correlated with phenotypic disparity. Here, we quan-tify nymphal and adult trait correlations in 219 species across the wholephylogeny of earwigs and stoneflies to test whether juvenile and adulttraits are decoupled from each other. We demonstrate that adult head mor-phology is largely driven by nymphal ecology, and that adult head shapedisparity has increased with stronger ontogenetic niche shifts in some stone-fly lineages. Our findings implicate that the hemimetabolan metamorphosisin earwigs and stoneflies does not allow for high degrees of adaptive decou-pling, and that high phenotypic disparity can even be realized when theevolution of distinct life phases is coupled

Intersubunit Interactions at Putative Sites of Ethanol Action in the M3 and M4 Domains of the NMDA Receptor GluN1 and GluN2B Subunits

Background and Purpose: The N-methyl-D-aspartate (NMDA) receptor is an important target of alcohol action in the brain. Recent studies in this laboratory have demonstrated that alcohol-sensitive positions in the intersubunit interfaces of the M3 and M4 domains of GluN1 and GluN2A subunits interact with respect to ethanol sensitivity and receptor kinetics, and that alcohol-sensitive positions in the M domains of GluN2A and GluN2B subunits differ. In this study we tested for interactions among alcohol-sensitive positions at the M domain intersubunit interfaces in GluN1/GluN2B NMDA receptors. Experimental Approach: We used whole-cell patch-clamp recording in tsA201 cells expressing tryptophan substitution mutants at ethanol-sensitive positions in the GluN1 and GluN2B NMDA receptor subunits to test for interactions among positions. Key Results: Six pairs of positions in GluN1/GluN2B significantly interacted to regulate ethanol inhibition: Gly638/Met824, Gly638/Leu825, Phe639/Leu825, Phe639/Gly826, Met818/Phe637 and Val820/Phe637. Tryptophan substitution at Met824 or Leu825 in GluN2B did not alter ethanol sensitivity but interacted with positions in the GluN1 M3 domain to regulate ethanol action, whereas tryptophan substitution at Gly638, which is the cognate of an ethanol-sensitive position in GluN2A, did not alter ethanol sensitivity or interact with positions in GluN1. Two and three pairs of positions interacted to regulate glutamate steady-state and peak current EC50, respectively, and one pair interacted with respect to macroscopic desensitization. Conclusions: Despite highly-conserved M domain sequences and similar ethanol sensitivity in the GluN2A and GluN2B subunits, the manner in which these subunits interact with the GluN1 subunit to regulate ethanol sensitivity and receptor kinetics differs

Four myriapod relatives – but who are sisters? No end to debates on relationships among the four major myriapod subgroups

BackgroundPhylogenetic relationships among the myriapod subgroups Chilopoda, Diplopoda, Symphyla and Pauropoda are still not robustly resolved. The first phylogenomic study covering all subgroups resolved phylogenetic relationships congruently to morphological evidence but is in conflict with most previously published phylogenetic trees based on diverse molecular data. Outgroup choice and long-branch attraction effects were stated as possible explanations for these incongruencies. In this study, we addressed these issues by extending the myriapod and outgroup taxon sampling using transcriptome data.ResultsWe generated new transcriptome data of 42 panarthropod species, including all four myriapod subgroups and additional outgroup taxa. Our taxon sampling was complemented by published transcriptome and genome data resulting in a supermatrix covering 59 species. We compiled two data sets, the first with a full coverage of genes per species (292 single-copy protein-coding genes), the second with a less stringent coverage (988 genes). We inferred phylogenetic relationships among myriapods using different data types, tree inference, and quartet computation approaches. Our results unambiguously support monophyletic Mandibulata and Myriapoda. Our analyses clearly showed that there is strong signal for a single unrooted topology, but a sensitivity of the position of the internal root on the choice of outgroups. However, we observe strong evidence for a clade Pauropoda+Symphyla, as well as for a clade Chilopoda+Diplopoda.ConclusionsOur best quartet topology is incongruent with current morphological phylogenies which were supported in another phylogenomic study. AU tests and quartet mapping reject the quartet topology congruent to trees inferred with morphological characters. Moreover, quartet mapping shows that confounding signal present in the data set is sufficient to explain the weak signal for the quartet topology derived from morphological characters. Although outgroup choice affects results, our study could narrow possible trees to derivatives of a single quartet topology. For highly disputed relationships, we propose to apply a series of tests (AU and quartet mapping), since results of such tests allow to narrow down possible relationships and to rule out confounding signal

Location and arrangement of campaniform sensilla in Drosophila melanogaster

Sensory systems provide input to motor networks on the state of the body and environment. One such sensory system in insects is the campaniform sensilla (CS), which detect deformations of the exoskeleton arising from resisted movements or external perturbations. When physical strain is applied to the cuticle, CS external structures are compressed, leading to transduction in an internal sensory neuron. In Drosophila melanogaster, the distribution of CS on the exoskeleton has not been comprehensively described. To investigate CS number, location, spatial arrangement, and potential differences between individuals, we compared the front, middle, and hind legs of multiple flies using scanning electron microscopy. Additionally, we imaged the entire body surface to confirm known CS locations. On the legs, the number and relative arrangement of CS varied between individuals, and single CS of corresponding segments showed characteristic differences between legs. This knowledge is fundamental for studying the relevance of cuticular strain information within the complex neuromuscular networks controlling posture and movement. This comprehensive account of all D. melanogaster CS helps set the stage for experimental investigations into their responsivity, sensitivity, and roles in sensory acquisition and motor control in a light-weight model organism