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      Mandible mechanical properties and composition of the larval Glossosoma boltoni (Trichoptera, Insecta)

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          Abstract

          Insect feeding structures, such as mandibles, interact with the ingesta (food or/and substrate) and can be adapted in morphology, composition of material and mechanical properties. The foraging on abrasive ingesta, as on algae covering rocks, is particularly challenging because the mandibles will be prone to wear and structural failure, thus suggesting the presence of mandibular adaptations to accompany this feeding behavior. Adaptations to this are well studied in the mouthparts of molluscs and sea urchins, but for insects there are large gaps in our knowledge. In this study, we investigated the mandibles of a grazing insect, the larvae of the trichopteran Glossosoma boltoni. Using scanning electron microscopy, wear was documented on the mandibles. The highest degree was identified on the medial surface of the sharp mandible tip. Using nanoindentation, the mechanical properties, such as hardness and Young’s modulus, of the medial and lateral mandible cuticles were tested. We found, that the medial cuticle of the tip was significantly softer and more flexible than the lateral one. These findings indicate that a self-sharpening mechanism is present in the mandibles of this species, since the softer medial cuticle is probably abraded faster than the harder lateral one, leading to sharp mandible tips. To investigate the origins of these properties, we visualized the degree of tanning by confocal laser scanning microscopy. The autofluorescence signal related to the mechanical property gradients. The presence of transition and alkaline earth metals by energy dispersive X-ray spectroscopy was also tested. We found Ca, Cl, Cu, Fe, K, Mg, Mn, P, S, Si, and Zn in the cuticle, but the content was very low and did not correlate with the mechanical property values.

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          Design and mechanical properties of insect cuticle.

          Since nearly all adult insects fly, the cuticle has to provide a very efficient and lightweight skeleton. Information is available about the mechanical properties of cuticle-Young's modulus of resilin is about 1 MPa, of soft cuticles about 1 kPa to 50 MPa, of sclerotised cuticles 1-20 GPa; Vicker's Hardness of sclerotised cuticle ranges between 25 and 80 kgf mm(-2); density is 1-1.3 kg m(-3)-and one of its components, chitin nanofibres, the Young's modulus of which is more than 150 GPa. Experiments based on fracture mechanics have not been performed although the layered structure probably provides some toughening. The structural performance of wings and legs has been measured, but our understanding of the importance of buckling is lacking: it can stiffen the structure (by elastic postbuckling in wings, for example) or be a failure mode. We know nothing of fatigue properties (yet, for instance, the insect wing must undergo millions of cycles, flexing or buckling on each cycle). The remarkable mechanical performance and efficiency of cuticle can be analysed and compared with those of other materials using material property charts and material indices. Presented in this paper are four: Young's modulus-density (stiffness per unit weight), specific Young's modulus-specific strength (elastic hinges, elastic energy storage per unit weight), toughness-Young's modulus (fracture resistance under various loading conditions), and hardness (wear resistance). In conjunction with a structural analysis of cuticle these charts help to understand the relevance of microstructure (fibre orientation effects in tendons, joints and sense organs, for example) and shape (including surface structure) of this fibrous composite for a given function. With modern techniques for analysis of structure and material, and emphasis on nanocomposites and self-assembly, insect cuticle should be the archetype for composites at all levels of scale.
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            Insect cuticular sclerotization: a review.

            Different regions of an insect cuticle have different mechanical properties, partly due to different degrees of stabilization and hardening occurring during the process of sclerotization, whereby phenolic material is incorporated into the cuticular proteins. Our understanding of the chemistry of cuticular sclerotization has increased considerably since Mark Pryor in 1940 suggested that enzymatically generated ortho-quinones react with free amino groups, thereby crosslinking the cuticular proteins. The results obtained since then have confirmed the essential features of Pryor's suggestion, and the many observations and experiments, which have been obtained, have led to a detailed and rather complex picture of the sclerotization process, as described in this review. However, many important questions still remain unanswered, especially regarding the precise regional and temporal regulation of the various steps in the process. (c) 2009 Elsevier Ltd. All rights reserved.
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              Functional gradients and heterogeneities in biological materials: Design principles, functions, and bioinspired applications

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                Author and article information

                Contributors
                wencke.krings@uni-hamburg.de
                Journal
                Sci Rep
                Sci Rep
                Scientific Reports
                Nature Publishing Group UK (London )
                2045-2322
                26 February 2024
                26 February 2024
                2024
                : 14
                : 4695
                Affiliations
                [1 ]Department of Electron Microscopy, Institute of Cell and Systems Biology of Animals, Universität Hamburg, ( https://ror.org/00g30e956) Martin-Luther-King-Platz 3, 20146 Hamburg, Germany
                [2 ]Department of Cariology, Endodontology and Periodontology, Universität Leipzig, ( https://ror.org/03s7gtk40) Liebigstraße 12, 04103 Leipzig, Germany
                [3 ]Department of Mammalogy and Palaeoanthropology, Leibniz Institute for the Analysis of Biodiversity Change, ( https://ror.org/03k5bhd83) Martin-Luther-King-Platz 3, 20146 Hamburg, Germany
                [4 ]Department of Functional Morphology and Biomechanics, Zoological Institute, Christian-Albrechts-Universität zu Kiel, ( https://ror.org/04v76ef78) Am Botanischen Garten 1-9, 24118 Kiel, Germany
                Article
                55211
                10.1038/s41598-024-55211-5
                10897335
                38409429
                c4d09ee1-17bf-44bb-a703-ddfbeb1e17c4
                © The Author(s) 2024

                Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.

                History
                : 23 August 2023
                : 21 February 2024
                Funding
                Funded by: FundRef http://dx.doi.org/10.13039/501100001659, Deutsche Forschungsgemeinschaft;
                Award ID: 470833544
                Award Recipient :
                Funded by: Universität Hamburg (1037)
                Categories
                Article
                Custom metadata
                © Springer Nature Limited 2024

                Uncategorized
                cuticle,elemental composition,mechanical properties,feeding,wear,structural biology,biomechanics

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