{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,4]],"date-time":"2026-08-04T08:42:52Z","timestamp":1785832972087,"version":"3.56.0"},"reference-count":83,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2025,8,1]],"date-time":"2025-08-01T00:00:00Z","timestamp":1754006400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2025,8,1]],"date-time":"2025-08-01T00:00:00Z","timestamp":1754006400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2025,5,23]],"date-time":"2025-05-23T00:00:00Z","timestamp":1747958400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001659","name":"German Research Foundation","doi-asserted-by":"publisher","award":["BE3215\/5\\u20131"],"award-info":[{"award-number":["BE3215\/5\\u20131"]}],"id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001665","name":"French National Research Agency","doi-asserted-by":"publisher","award":["AAPG2023"],"award-info":[{"award-number":["AAPG2023"]}],"id":[{"id":"10.13039\/501100001665","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Acta Biomaterialia"],"published-print":{"date-parts":[[2025,8]]},"DOI":"10.1016\/j.actbio.2025.05.046","type":"journal-article","created":{"date-parts":[[2025,5,22]],"date-time":"2025-05-22T02:54:01Z","timestamp":1747882441000},"page":"377-393","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":8,"special_numbering":"C","title":["Exoskeletal cuticle proteins enable Drosophila locomotion"],"prefix":"10.1016","volume":"202","author":[{"given":"Maximilian","family":"G\u00f6pfert","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jing","family":"Yang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0008-9750-9634","authenticated-orcid":false,"given":"Dhyeykumar","family":"Rabadiya","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2970-6894","authenticated-orcid":false,"given":"Dietmar","family":"Riedel","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bernard","family":"Moussian","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9869-1257","authenticated-orcid":false,"given":"Matthias","family":"Behr","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.actbio.2025.05.046_bib0001","doi-asserted-by":"crossref","first-page":"44","DOI":"10.1016\/j.bone.2013.03.010","article-title":"Muscle loading is necessary for the formation of a functional tendon enthesis","volume":"55","author":"Schwartz","year":"2013","journal-title":"Bone"},{"key":"10.1016\/j.actbio.2025.05.046_bib0002","doi-asserted-by":"crossref","DOI":"10.1073\/pnas.2004840117","article-title":"Toward the correction of muscular dystrophy by gene editing","volume":"118","author":"Olson","year":"2021","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.actbio.2025.05.046_bib0003","doi-asserted-by":"crossref","DOI":"10.1155\/2015\/421746","article-title":"Biology of bone tissue: structure, function, and factors that influence bone cells","author":"Florencio-Silva","year":"2015","journal-title":"BioMed Res. Int."},{"key":"10.1016\/j.actbio.2025.05.046_bib0004","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1247\/csf.25.73","article-title":"Mammalian skeletogenesis and extracellular matrix: what can we learn from knockout mice?","volume":"25","author":"Asz\u00f3di","year":"2000","journal-title":"Cell Struct. Funct."},{"key":"10.1016\/j.actbio.2025.05.046_bib0005","doi-asserted-by":"crossref","first-page":"333","DOI":"10.32098\/mltj.03.2014.12","article-title":"The enthesis: a review of the tendon-to-bone insertion","volume":"4","author":"Apostolakos","year":"2014","journal-title":"Muscles Ligaments Tendons J."},{"key":"10.1016\/j.actbio.2025.05.046_bib0006","doi-asserted-by":"crossref","first-page":"e55","DOI":"10.1371\/journal.pgen.0010055","article-title":"Gain-of-function screen for genes that affect Drosophila muscle pattern formation","volume":"1","author":"Staudt","year":"2005","journal-title":"PLoS Genet."},{"key":"10.1016\/j.actbio.2025.05.046_bib0007","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1016\/j.devcel.2004.06.010","article-title":"Muscle building; mechanisms of myotube guidance and attachment site selection","volume":"7","author":"Schnorrer","year":"2004","journal-title":"Dev. Cell"},{"key":"10.1016\/j.actbio.2025.05.046_bib0008","doi-asserted-by":"crossref","first-page":"448","DOI":"10.1016\/S0168-9525(99)01862-4","article-title":"Singling out Drosophila tendon cells: a dialogue between two distinct cell types","volume":"15","author":"Volk","year":"1999","journal-title":"Trends Genet."},{"key":"10.1016\/j.actbio.2025.05.046_bib0009","doi-asserted-by":"crossref","first-page":"8606","DOI":"10.1073\/pnas.94.16.8606","article-title":"Epidermal muscle attachment site-specific target gene expression and interference with myotube guidance in response to ectopic stripe expression in the developing Drosophila epidermis","volume":"94","author":"Vorbr\u00fcggen","year":"1997","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.actbio.2025.05.046_bib0010","doi-asserted-by":"crossref","first-page":"2691","DOI":"10.1101\/gad.11.20.2691","article-title":"The Drosophila neuregulin homolog Vein mediates inductive interactions between myotubes and their epidermal attachment cells","volume":"11","author":"Yarnitzky","year":"1997","journal-title":"Genes Dev."},{"key":"10.1016\/j.actbio.2025.05.046_bib0011","doi-asserted-by":"crossref","first-page":"566","DOI":"10.1016\/j.ceb.2010.08.009","article-title":"Born to run: creating the muscle fiber","volume":"22","author":"Schejter","year":"2010","journal-title":"Curr. Opin. Cell Biol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0012","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pbio.3000057","article-title":"A small proportion of Talin molecules transmit forces at developing muscle attachments in vivo","volume":"17","author":"Lemke","year":"2019","journal-title":"PLoS Biol"},{"key":"10.1016\/j.actbio.2025.05.046_bib0013","doi-asserted-by":"crossref","first-page":"311","DOI":"10.1016\/S1534-5807(02)00265-4","article-title":"Integrins in development","volume":"3","author":"B\u00f6kel","year":"2002","journal-title":"Dev. Cell"},{"key":"10.1016\/j.actbio.2025.05.046_bib0014","doi-asserted-by":"crossref","first-page":"463","DOI":"10.1016\/j.mod.2007.03.005","article-title":"AlphaPS2 integrin-mediated muscle attachment in Drosophila requires the ECM protein thrombospondin","volume":"124","author":"Chanana","year":"2007","journal-title":"Mech. Dev."},{"key":"10.1016\/j.actbio.2025.05.046_bib0015","doi-asserted-by":"crossref","DOI":"10.1016\/j.cub.2021.01.010","article-title":"Mechano-chemical enforcement of tendon apical ECM into nano-filaments during Drosophila flight muscle development","author":"Chu","year":"2021","journal-title":"Curr. Biol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0016","doi-asserted-by":"crossref","DOI":"10.1073\/pnas.2120853119","article-title":"Chitin deacetylases are necessary for insect femur muscle attachment and mobility","volume":"119","author":"Mun","year":"2022","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.actbio.2025.05.046_bib0017","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1006\/dbio.2000.9711","article-title":"Integrins as mediators of morphogenesis in Drosophila","volume":"223","author":"Brown","year":"2000","journal-title":"Dev. Biol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0018","doi-asserted-by":"crossref","first-page":"1086","DOI":"10.1016\/S0960-9822(03)00416-0","article-title":"Shortstop recruits EB1\/APC1 and promotes microtubule assembly at the muscle-tendon junction","volume":"13","author":"Subramanian","year":"2003","journal-title":"Curr. Biol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0019","doi-asserted-by":"crossref","first-page":"633","DOI":"10.1242\/jcs.01619","article-title":"Papillote and piopio: drosophila ZP-domain proteins required for cell adhesion to the apical extracellular matrix and microtubule organization","volume":"118","author":"B\u00f6kel","year":"2005","journal-title":"J. Cell. Sci."},{"key":"10.1016\/j.actbio.2025.05.046_bib0020","doi-asserted-by":"crossref","first-page":"563","DOI":"10.1006\/dbio.1994.1054","article-title":"The development of cellular junctions in the drosophila embryo","volume":"161","author":"Tepass","year":"1994","journal-title":"Dev. Biol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0021","doi-asserted-by":"crossref","DOI":"10.1038\/srep18340","article-title":"Chitinases and imaginal disc growth factors organize the extracellular matrix formation at barrier tissues in insects","volume":"6","author":"Pesch","year":"2016","journal-title":"Sci. Rep."},{"key":"10.1016\/j.actbio.2025.05.046_bib0022","first-page":"1396","article-title":"Double cuticle barrier in two global pests, the whitefly trialeurodes vaporariorum and the bedbug cimex lectularius","volume":"220","author":"Wang","year":"2017","journal-title":"J. Exp. Biol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0023","doi-asserted-by":"crossref","first-page":"10071","DOI":"10.1074\/jbc.M114.614933","article-title":"Obstructor A organizes matrix assembly at the apical cell surface to promote enzymatic cuticle maturation in drosophila","volume":"290","author":"Pesch","year":"2015","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.actbio.2025.05.046_bib0024","doi-asserted-by":"crossref","first-page":"337","DOI":"10.1016\/j.matbio.2007.02.001","article-title":"Assembly of the Drosophila larval exoskeleton requires controlled secretion and shaping of the apical plasma membrane","volume":"26","author":"Moussian","year":"2007","journal-title":"Matrix Biol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0025","series-title":"Insect Molecular Biology and Biochemistry","first-page":"193","article-title":"Chitin metabolism in insects","author":"Muthukrishnan","year":"2012"},{"key":"10.1016\/j.actbio.2025.05.046_bib0026","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1007\/978-981-13-7318-3_5","article-title":"Chitin organizing and modifying enzymes and proteins involved In remodeling of the insect cuticle","volume":"1142","author":"Muthukrishnan","year":"2019","journal-title":"Adv. Exp. Med. Biol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0027","doi-asserted-by":"crossref","first-page":"6985","DOI":"10.1074\/jbc.RA117.001454","article-title":"Group I chitin deacetylases are essential for higher order organization of chitin fibers in beetle cuticle","volume":"293","author":"Noh","year":"2018","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.actbio.2025.05.046_bib0028","doi-asserted-by":"crossref","DOI":"10.1074\/jbc.M116.720581","article-title":"Helicoidal organization of chitin in the cuticle of the migratory locust requires the function of the chitin deacetylase 2 enzyme (LmCDA2)","author":"Yu","year":"2016","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.actbio.2025.05.046_bib0029","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1111\/j.1365-2583.2012.01134.x","article-title":"The transcription factor grainy head and the steroid hormone ecdysone cooperate during differentiation of the skin of Drosophila melanogaster","volume":"21","author":"Gangishetti","year":"2012","journal-title":"Insect Mol. Biol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0030","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1186\/s40694-021-00127-2","article-title":"Deciphering the ChitoCode: fungal chitins and chitosans as functional biopolymers","volume":"8","author":"Cord-Landwehr","year":"2021","journal-title":"Fungal Biol. Biotechnol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0031","doi-asserted-by":"crossref","first-page":"180","DOI":"10.1016\/j.cub.2005.11.074","article-title":"Septate-junction-dependent luminal deposition of Chitin deacetylases restricts tube elongation in the Drosophila trachea","volume":"16","author":"Wang","year":"2006","journal-title":"Curr. Biol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0032","doi-asserted-by":"crossref","first-page":"186","DOI":"10.1016\/j.cub.2005.11.072","article-title":"serpentine and vermiform encode matrix proteins with chitin binding and deacetylation domains that limit tracheal tube length in Drosophila","volume":"16","author":"Luschnig","year":"2006","journal-title":"Curr. Biol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0033","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pgen.1003268","article-title":"Retroactive maintains cuticle integrity by promoting the trafficking of Knickkopf into the procuticle of Tribolium castaneum","volume":"9","author":"Chaudhari","year":"2013","journal-title":"PLoS Genet"},{"key":"10.1016\/j.actbio.2025.05.046_bib0034","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1242\/dev.02177","article-title":"Drosophila Knickkopf and Retroactive are needed for epithelial tube growth and cuticle differentiation through their specific requirement for chitin filament organization","volume":"133","author":"Moussian","year":"2006","journal-title":"Development"},{"key":"10.1016\/j.actbio.2025.05.046_bib0035","doi-asserted-by":"crossref","first-page":"1056","DOI":"10.1002\/dvdy.20389","article-title":"Retroactive, a membrane-anchored extracellular protein related to vertebrate snake neurotoxin-like proteins, is required for cuticle organization in the larva ofDrosophila melanogaster","volume":"233","author":"Moussian","year":"2005","journal-title":"Dev. Dyn."},{"key":"10.1016\/j.actbio.2025.05.046_bib0036","doi-asserted-by":"crossref","first-page":"17028","DOI":"10.1073\/pnas.1112288108","article-title":"Knickkopf protein protects and organizes chitin in the newly synthesized insect exoskeleton","volume":"108","author":"Chaudhari","year":"2011","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.actbio.2025.05.046_bib0037","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0049844","article-title":"Gene families of cuticular proteins analogous to peritrophins (CPAPs) in Tribolium castaneum have diverse functions","volume":"7","author":"Jasrapuria","year":"2012","journal-title":"PLoS ONE"},{"key":"10.1016\/j.actbio.2025.05.046_bib0038","doi-asserted-by":"crossref","first-page":"6827","DOI":"10.1016\/j.febslet.2005.11.021","article-title":"Identification of the novel evolutionary conserved obstructor multigene family in invertebrates","volume":"579","author":"Behr","year":"2005","journal-title":"FEBS Lett."},{"key":"10.1016\/j.actbio.2025.05.046_bib0039","doi-asserted-by":"crossref","first-page":"21396","DOI":"10.1074\/jbc.M112.359984","article-title":"Obstructor-A is required for epithelial extracellular matrix dynamics, exoskeleton function, and tubulogenesis","volume":"287","author":"Petkau","year":"2012","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.actbio.2025.05.046_bib0040","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0067415","article-title":"Control of airway tube diameter and integrity by secreted chitin-binding proteins in Drosophila","volume":"8","author":"Tiklova","year":"2013","journal-title":"PLoS ONE"},{"key":"10.1016\/j.actbio.2025.05.046_bib0041","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pgen.1006548","article-title":"Mechanical control of whole body shape by a single cuticular protein obstructor-E in drosophila melanogaster","volume":"13","author":"Tajiri","year":"2017","journal-title":"PLoS Genet."},{"key":"10.1016\/j.actbio.2025.05.046_bib0042","doi-asserted-by":"crossref","DOI":"10.1016\/j.ibmb.2024.104071","article-title":"The biology of insect chitinases and their roles at chitinous cuticles","author":"Rabadiya","year":"2024","journal-title":"Insect Biochem. Mol. Biol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0043","doi-asserted-by":"crossref","DOI":"10.1038\/s41598-020-70185-w","article-title":"Glycosylhydrolase genes control respiratory tubes sizes and airway stability","volume":"10","author":"Behr","year":"2020","journal-title":"Sci. Rep."},{"key":"10.1016\/j.actbio.2025.05.046_bib0044","doi-asserted-by":"crossref","first-page":"4","DOI":"10.1016\/j.asd.2016.11.002","article-title":"Drosophila Chitinase 2 is expressed in chitin producing organs for cuticle formation","volume":"46","author":"Pesch","year":"2017","journal-title":"Arthropod Struct. Dev."},{"key":"10.1016\/j.actbio.2025.05.046_bib0045","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pgen.1007307","article-title":"A chitinase with two catalytic domains is required for organization of the cuticular extracellular matrix of a beetle","volume":"14","author":"Noh","year":"2018","journal-title":"PLoS Genet."},{"key":"10.1016\/j.actbio.2025.05.046_bib0046","doi-asserted-by":"crossref","first-page":"524","DOI":"10.1016\/j.tcb.2010.06.002","article-title":"From A to Z: apical structures and zona pellucida-domain proteins","volume":"20","author":"Plaza","year":"2010","journal-title":"Trends Cell Biol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0047","doi-asserted-by":"crossref","first-page":"895","DOI":"10.1038\/ncb1049","article-title":"Epithelial tube morphogenesis during Drosophila tracheal development requires Piopio, a luminal ZP protein","volume":"5","author":"Ja\u017awi\u0144ska","year":"2003","journal-title":"Nat. Cell Biol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0048","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.91079","article-title":"The proteolysis of ZP proteins is essential to control cell membrane structure and integrity of developing tracheal tubes in Drosophila","volume":"12","author":"Drees","year":"2023","journal-title":"Elife"},{"key":"10.1016\/j.actbio.2025.05.046_bib0049","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pgen.1007882","article-title":"Conserved function of the matriptase-prostasin proteolytic cascade during epithelial morphogenesis","volume":"15","author":"Drees","year":"2019","journal-title":"PLoS Genet."},{"key":"10.1016\/j.actbio.2025.05.046_bib0050","doi-asserted-by":"crossref","first-page":"1041","DOI":"10.3390\/ma15031041","article-title":"Improving polysaccharide-based chitin\/Chitosan-aerogel materials by learning from genetics and molecular biology","volume":"15","author":"Behr","year":"2022","journal-title":"Materials"},{"key":"10.1016\/j.actbio.2025.05.046_bib0051","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1016\/j.ejcb.2017.12.003","article-title":"The ABC transporter Snu and the extracellular protein snsl cooperate in the formation of the lipid-based inward and outward barrier in the skin of Drosophila","volume":"97","author":"Zuber","year":"2018","journal-title":"Eur. J. Cell Biol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0052","doi-asserted-by":"crossref","DOI":"10.1098\/rspb.2015.2994","article-title":"Regionalization of surface lipids in insects","volume":"283","author":"Wang","year":"2016","journal-title":"Proc. R. Soc. B."},{"key":"10.1016\/j.actbio.2025.05.046_bib0053","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pgen.1008363","article-title":"Dysfunction of Oskyddad causes harlequin-type ichthyosis-like defects in Drosophila melanogaster","volume":"16","author":"Wang","year":"2020","journal-title":"PLoS Genet."},{"key":"10.1016\/j.actbio.2025.05.046_bib0054","doi-asserted-by":"crossref","first-page":"423","DOI":"10.1016\/j.devcel.2005.07.012","article-title":"A transient luminal chitinous matrix is required to model epithelial tube diameter in the Drosophila trachea","volume":"9","author":"Tonning","year":"2005","journal-title":"Dev. Cell"},{"key":"10.1016\/j.actbio.2025.05.046_bib0055","doi-asserted-by":"crossref","DOI":"10.1016\/j.devcel.2017.06.004","article-title":"Basement membrane manipulation in drosophila wing discs affects dpp retention but not growth mechanoregulation","volume":"42","author":"Ma","year":"2017","journal-title":"Dev. Cell"},{"key":"10.1016\/j.actbio.2025.05.046_bib0056","doi-asserted-by":"crossref","first-page":"62","DOI":"10.1016\/j.ydbio.2014.12.038","article-title":"The many faces of cell adhesion during Drosophila muscle development","volume":"401","author":"Maartens","year":"2015","journal-title":"Dev. Biol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0057","doi-asserted-by":"crossref","first-page":"42","DOI":"10.1016\/j.yjmcc.2020.02.008","article-title":"Matrix metalloproteinases regulate ECM accumulation but not larval heart growth in Drosophila melanogaster","volume":"140","author":"Hughes","year":"2020","journal-title":"J. Mol. Cell. Cardiol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0058","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1101\/gad.308270.117","article-title":"Dendrite regeneration of adult drosophila sensory neurons diminishes with aging and is inhibited by epidermal-derived matrix metalloproteinase 2","volume":"32","author":"DeVault","year":"2018","journal-title":"Genes Dev."},{"key":"10.1016\/j.actbio.2025.05.046_bib0059","doi-asserted-by":"crossref","first-page":"218","DOI":"10.1016\/j.isci.2019.05.027","article-title":"Degradation of extracellular matrix by matrix metalloproteinase 2 is essential for the establishment of the blood-brain barrier in Drosophila","volume":"16","author":"Kanda","year":"2019","journal-title":"iScience"},{"key":"10.1016\/j.actbio.2025.05.046_bib0060","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1016\/S1097-2765(05)00011-0","article-title":"Drosophila segment borders result from unilateral repression of hedgehog activity by wingless signaling","volume":"6","author":"Piepenburg","year":"2000","journal-title":"Mol. Cell"},{"key":"10.1016\/j.actbio.2025.05.046_bib0061","doi-asserted-by":"crossref","first-page":"790","DOI":"10.1016\/j.devcel.2010.03.015","article-title":"A developmentally regulated two-step process generates a noncentrosomal microtubule network in Drosophila tracheal cells","volume":"18","author":"Brodu","year":"2010","journal-title":"Dev. Cell"},{"key":"10.1016\/j.actbio.2025.05.046_bib0062","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1016\/j.asd.2006.05.003","article-title":"Cuticle differentiation during Drosophila embryogenesis","volume":"35","author":"Moussian","year":"2006","journal-title":"Arthropod Struct. Dev."},{"key":"10.1016\/j.actbio.2025.05.046_bib0063","doi-asserted-by":"crossref","first-page":"441","DOI":"10.1016\/S0040-8166(73)80036-9","article-title":"The formation of the ecdysial droplets and the ecdysial membrane in an insect","volume":"5","author":"Locke","year":"1973","journal-title":"Tissue Cell"},{"key":"10.1016\/j.actbio.2025.05.046_bib0064","doi-asserted-by":"crossref","first-page":"1305","DOI":"10.1083\/jcb.200307089","article-title":"Maintaining epithelial integrity: a function for gigantic spectraplakin isoforms in adherens junctions","volume":"162","author":"R\u00f6per","year":"2003","journal-title":"J. Biol. Cell."},{"key":"10.1016\/j.actbio.2025.05.046_bib0065","doi-asserted-by":"crossref","first-page":"5374","DOI":"10.1038\/s41598-019-41734-9","article-title":"The putative C-type lectin Schlaff ensures epidermal barrier compactness in Drosophila","volume":"9","author":"Zuber","year":"2019","journal-title":"Sci. Rep."},{"key":"10.1016\/j.actbio.2025.05.046_bib0066","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1038\/s42003-020-0751-5","article-title":"Chitin is a functional component of the larval adhesive of barnacles","volume":"3","author":"Aldred","year":"2020","journal-title":"Commun. Biol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0067","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1002\/jmor.10324","article-title":"Involvement of chitin in exoskeleton morphogenesis in Drosophila melanogaster","volume":"264","author":"Moussian","year":"2005","journal-title":"J. Morphol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0068","doi-asserted-by":"crossref","unstructured":"Y. Itakura, H. Wada, S. Inagaki, S. Hayashi, Mechanical control of the insect extracellular matrix nanostructure, 2024.","DOI":"10.1101\/2024.08.20.608778"},{"key":"10.1016\/j.actbio.2025.05.046_bib0069","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0209058","article-title":"Trynity controls epidermal barrier function and respiratory tube maturation in Drosophila by modulating apical extracellular matrix nano-patterning","volume":"13","author":"Itakura","year":"2018","journal-title":"PLoS ONE"},{"key":"10.1016\/j.actbio.2025.05.046_bib0070","doi-asserted-by":"crossref","DOI":"10.1098\/rsif.2017.0310","article-title":"Stiffness distribution in insect cuticle: a continuous or a discontinuous profile?","volume":"14","author":"Rajabi","year":"2017","journal-title":"J. R. Soc. Interface."},{"key":"10.1016\/j.actbio.2025.05.046_bib0071","doi-asserted-by":"crossref","DOI":"10.1098\/rsta.2020.0332","article-title":"The spider cuticle: a remarkable material toolbox for functional diversity","volume":"379","author":"Politi","year":"2021","journal-title":"Phil. Trans. R. Soc. A."},{"key":"10.1016\/j.actbio.2025.05.046_bib0072","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s00339-021-04439-3","article-title":"Biomechanics of insect cuticle: an interdisciplinary experimental challenge","volume":"127","author":"Stamm","year":"2021","journal-title":"Appl. Phys. A"},{"key":"10.1016\/j.actbio.2025.05.046_bib0073","doi-asserted-by":"crossref","first-page":"187","DOI":"10.1016\/j.asd.2004.05.006","article-title":"Design and mechanical properties of insect cuticle","volume":"33","author":"Vincent","year":"2004","journal-title":"Arthropod Struct. Dev."},{"key":"10.1016\/j.actbio.2025.05.046_bib0074","doi-asserted-by":"crossref","unstructured":"S.A. Shaked, S. Weil, R. Manor, E. Aflalo, N. Maman, R. Maria, B. Kruppke, T. Hanke, J. Eichler, B. Ratzker, M. Sokol, A. Sagi, Crusticuls: a Family of cuticular protein that affect the spatial organization of skeletal chitin bundles (2024). 10.2139\/ssrn.4987166.","DOI":"10.2139\/ssrn.4987166"},{"key":"10.1016\/j.actbio.2025.05.046_bib0075","doi-asserted-by":"crossref","DOI":"10.1016\/j.ibmb.2023.104058","article-title":"Advances in understanding insect chitin biosynthesis","volume":"164","author":"Yu","year":"2024","journal-title":"Insect Biochem. Mol. Biol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0076","doi-asserted-by":"crossref","DOI":"10.1088\/1748-3190\/11\/5\/055006","article-title":"Functional adaptation of crustacean exoskeletal elements through structural and compositional diversity: a combined experimental and theoretical study","volume":"11","author":"Fabritius","year":"2016","journal-title":"Bioinspir. Biomim."},{"key":"10.1016\/j.actbio.2025.05.046_bib0077","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1016\/j.jmbbm.2013.04.003","article-title":"Effects of homogenization technique and introduction of interfaces in a multiscale approach to predict the elastic properties of arthropod cuticle","volume":"23","author":"Lhadi","year":"2013","journal-title":"J. Mech. Behav. Biomed. Mater."},{"key":"10.1016\/j.actbio.2025.05.046_bib0078","doi-asserted-by":"crossref","first-page":"3551","DOI":"10.1073\/pnas.1915798117","article-title":"Unique subsite specificity and potential natural function of a chitosan deacetylase from the human pathogen Cryptococcus neoformans","volume":"117","author":"Hembach","year":"2020","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.actbio.2025.05.046_bib0079","doi-asserted-by":"crossref","DOI":"10.1038\/srep38018","article-title":"A chitin deacetylase from the endophytic fungus Pestalotiopsis sp. efficiently inactivates the elicitor activity of chitin oligomers in rice cells","volume":"6","author":"Cord-Landwehr","year":"2016","journal-title":"Sci. Rep."},{"key":"10.1016\/j.actbio.2025.05.046_bib0080","doi-asserted-by":"crossref","first-page":"301","DOI":"10.1111\/imb.12554","article-title":"LmCDA1 organizes the cuticle by chitin deacetylation in Locusta migratoria","volume":"28","author":"Yu","year":"2019","journal-title":"Insect Mol. Biol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0081","doi-asserted-by":"crossref","first-page":"128","DOI":"10.1016\/j.ibmb.2019.05.008","article-title":"The putative chitin deacetylases Serpentine and Vermiform have non-redundant functions during Drosophila wing development","volume":"110","author":"Zhang","year":"2019","journal-title":"Insect Biochem. Mol. Biol."},{"key":"10.1016\/j.actbio.2025.05.046_bib0082","doi-asserted-by":"crossref","first-page":"272","DOI":"10.1186\/s12951-025-03347-5","article-title":"Nanocarrier-mediated transdermal delivery of Lmidgf4 dsRNA expedites biological control of locusts by Beauveria bassiana","volume":"23","author":"Kong","year":"2025","journal-title":"J. Nanobiotechnology"},{"key":"10.1016\/j.actbio.2025.05.046_bib0083","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1186\/s40634-014-0008-7","article-title":"The role of aggrecan in normal and osteoarthritic cartilage","volume":"1","author":"Roughley","year":"2014","journal-title":"J. Exp. Orthop"}],"container-title":["Acta Biomaterialia"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S1742706125003733?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S1742706125003733?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,5,16]],"date-time":"2026-05-16T15:21:05Z","timestamp":1778944865000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S1742706125003733"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,8]]},"references-count":83,"alternative-id":["S1742706125003733"],"URL":"https:\/\/doi.org\/10.1016\/j.actbio.2025.05.046","relation":{},"ISSN":["1742-7061"],"issn-type":[{"value":"1742-7061","type":"print"}],"subject":[],"published":{"date-parts":[[2025,8]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Exoskeletal cuticle proteins enable Drosophila locomotion","name":"articletitle","label":"Article Title"},{"value":"Acta Biomaterialia","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.actbio.2025.05.046","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2025 The Authors. Published by Elsevier Inc. on behalf of Acta Materialia Inc.","name":"copyright","label":"Copyright"}]}}