{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,6]],"date-time":"2026-03-06T12:35:07Z","timestamp":1772800507467,"version":"3.50.1"},"reference-count":105,"publisher":"Oxford University Press (OUP)","issue":"11","license":[{"start":{"date-parts":[[2024,8,29]],"date-time":"2024-08-29T00:00:00Z","timestamp":1724889600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001782","name":"University of Melbourne","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100001782","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Australian Government stipend"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2024,11,6]]},"abstract":"<jats:title>Abstract<\/jats:title>\n               <jats:p>Ecdysteroids are major hormones in insects and control molting, growth, reproduction, physiology, and behavior. The biosynthesis of ecdysteroids such as 20-hydroxyecdysone (20E) from dietary sterols is well characterized, but ecdysteroid catabolism is poorly understood. Ecdysteroid kinases (EcKs) mediate the reversible phosphorylation of ecdysteroids, which has been implicated in ecdysteroid recycling during embryogenesis and reproduction in various insects. However, to date, only 2 EcK-encoding genes have been identified, in the silkworm Bombyx mori and the mosquito Anopheles gambiae. Previously, we identified 2 ecdysteroid kinase-like (EcKL) genes\u2014Wallflower (Wall) and Pinkman (pkm)\u2014in the model fruit fly Drosophila melanogaster that are orthologs of the ecdysteroid 22-kinase gene BmEc22K. Here, using gene knockdown, knockout, and misexpression, we explore Wall and pkm's possible functions and genetically test the hypothesis that they encode EcKs. Wall and pkm null mutants are viable and fertile, suggesting that they are not essential for development or reproduction, whereas phenotypes arising from RNAi and somatic CRISPR appear to derive from off-target effects or other artifacts. However, misexpression of Wall results in dramatic phenotypes, including developmental arrest, and defects in trachea, cuticle, and pigmentation. Wall misexpression fails to phenocopy irreversible ecdysteroid catabolism through misexpression of Cyp18a1, suggesting that Wall does not directly inactivate 20E. Additionally, Wall misexpression phenotypes are not attenuated in Cyp18a1 mutants, strongly suggesting that Wall is not an ecdysteroid 26-kinase. We hypothesize that the substrate of Wall in this misexpression experiment and possibly generally is an unknown, atypical ecdysteroid that plays essential roles in Drosophila development, and may highlight aspects of insect endocrinology that are as-yet uncharacterized. We also provide preliminary evidence that CG5644 encodes an ecdysteroid 22-kinase conserved across Diptera.<\/jats:p>","DOI":"10.1093\/g3journal\/jkae204","type":"journal-article","created":{"date-parts":[[2024,8,29]],"date-time":"2024-08-29T22:11:25Z","timestamp":1724969485000},"source":"Crossref","is-referenced-by-count":3,"title":["Genetic characterization of candidate ecdysteroid kinases in <i>Drosophila melanogaster<\/i>"],"prefix":"10.1093","volume":"14","author":[{"given":"Jack L","family":"Scanlan","sequence":"first","affiliation":[{"name":"School of BioSciences, The University of Melbourne , Parkville Campus, Melbourne, Victoria 3010 ,","place":["Australia"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7733-6763","authenticated-orcid":false,"given":"Charles","family":"Robin","sequence":"additional","affiliation":[{"name":"School of BioSciences, The University of Melbourne , Parkville Campus, Melbourne, Victoria 3010 ,","place":["Australia"]}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"286","published-online":{"date-parts":[[2024,8,29]]},"reference":[{"key":"2024110622335965800_jkae204-B1","doi-asserted-by":"publisher","first-page":"e104347","DOI":"10.15252\/embj.2019104347","article-title":"Steroid-dependent switch of OvoL\/Shavenbaby controls self-renewal versus differentiation of intestinal stem cells","volume-title":"EMBO J","author":"Al Hayek","year":"2021"},{"key":"2024110622335965800_jkae204-B2","first-page":"57","article-title":"Staging the metamorphosis of Drosophila melanogaster","volume":"66","author":"Bainbridge","year":"1981","journal-title":"J Embryol Exp Morphol"},{"issue":"6","key":"2024110622335965800_jkae204-B3","doi-asserted-by":"publisher","first-page":"731","DOI":"10.1016\/s0092-8674(03)00420-3","article-title":"The Drosophila orphan nuclear receptor DHR38 mediates an atypical ecdysteroid signaling pathway","volume":"113","author":"Baker","year":"2003","journal-title":"Cell"},{"issue":"11","key":"2024110622335965800_jkae204-B4","doi-asserted-by":"publisher","first-page":"1037","DOI":"10.1016\/s0965-1748(00)00075-8","article-title":"Transcriptional activation of the Drosophila ecdysone receptor by insect and plant ecdysteroids","volume":"30","author":"Baker","year":"2000","journal-title":"Insect Biochem Mol Biol."},{"issue":"6","key":"2024110622335965800_jkae204-B5","doi-asserted-by":"publisher","first-page":"570","DOI":"10.1016\/j.ibmb.2007.03.001","article-title":"Specific transcriptional responses to juvenile hormone and ecdysone in Drosophila","volume":"37","author":"Beckstead","year":"2007","journal-title":"Insect Biochem Mol Biol."},{"issue":"2","key":"2024110622335965800_jkae204-B6","doi-asserted-by":"publisher","first-page":"761","DOI":"10.1534\/genetics.104.026427","article-title":"The BDGP gene disruption project: single transposon insertions associated with 40% of Drosophila genes","volume":"167","author":"Bellen","year":"2004","journal-title":"Genetics"},{"issue":"12","key":"2024110622335965800_jkae204-B7","doi-asserted-by":"publisher","first-page":"3562","DOI":"10.1002\/dvdy.21372","article-title":"The commonly used marker ELAV is transiently expressed in neuroblasts and glial cells in the Drosophila embryonic CNS","volume":"236","author":"Berger","year":"2007","journal-title":"Dev Dyn."},{"issue":"2","key":"2024110622335965800_jkae204-B8","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1002\/arch.940040207","article-title":"Ecdysone metabolism in Pieris brassicae during the feeding last larval instar","volume":"4","author":"Beydon","year":"1987","journal-title":"Arch Insect Biochem Physiol."},{"issue":"11","key":"2024110622335965800_jkae204-B9","doi-asserted-by":"publisher","first-page":"2434","DOI":"10.1242\/dev.088757","article-title":"A versatile platform for creating a comprehensive UAS-ORFeome library in Drosophila","volume":"140","author":"Bischof","year":"2013","journal-title":"Development"},{"issue":"9","key":"2024110622335965800_jkae204-B10","doi-asserted-by":"publisher","first-page":"3312","DOI":"10.1073\/pnas.0611511104","article-title":"An optimized transgenesis system for Drosophila using germ-line-specific phiC31 integrases","volume":"104","author":"Bischof","year":"2007","journal-title":"Proc Natl Acad Sci U S A."},{"issue":"4","key":"2024110622335965800_jkae204-B11","doi-asserted-by":"publisher","first-page":"398","DOI":"10.1016\/j.cbpa.2010.08.025","article-title":"The role of 20-hydroxyecdysone signaling in Drosophila pupal metabolism","volume":"157","author":"Bond","year":"2010","journal-title":"Comparative Biochemistry and Physiology a-Molecular & Integrative Physiology"},{"issue":"4","key":"2024110622335965800_jkae204-B12","doi-asserted-by":"publisher","first-page":"497","DOI":"10.1016\/j.devcel.2006.02.004","article-title":"Systematic analysis of the transcriptional switch inducing migration of border cells","volume":"10","author":"Borghese","year":"2006","journal-title":"Dev Cell."},{"key":"2024110622335965800_jkae204-B13","doi-asserted-by":"publisher","first-page":"dev165654","DOI":"10.1242\/dev.165654","article-title":"Growth control through regulation of insulin signalling by nutrition-activated steroid hormone in Drosophila","volume":"145","author":"Buhler","year":"2018","journal-title":"Development"},{"issue":"3","key":"2024110622335965800_jkae204-B14","doi-asserted-by":"publisher","first-page":"170039","DOI":"10.1098\/rsos.170039","article-title":"Drosophila enhancer-Gal4 lines show ectopic expression during development","volume":"4","author":"Casas-Tint\u00f3","year":"2017","journal-title":"R Soc Open Sci"},{"issue":"3","key":"2024110622335965800_jkae204-B15","doi-asserted-by":"publisher","first-page":"e1011204","DOI":"10.1371\/journal.pgen.1011204","article-title":"Cis-regulatory polymorphism at fiz ecdysone oxidase contributes to polygenic evolutionary response to malnutrition in Drosophila","volume":"20","author":"Cavigliasso","year":"2024","journal-title":"PLoS Genet."},{"issue":"9","key":"2024110622335965800_jkae204-B16","doi-asserted-by":"publisher","first-page":"e290","DOI":"10.1371\/journal.pbio.0040290","article-title":"Shavenbaby couples patterning to epidermal cell shape control","volume":"4","author":"Chanut-Delalande","year":"2006","journal-title":"PLoS Biol"},{"key":"2024110622335965800_jkae204-B17","doi-asserted-by":"publisher","first-page":"1035","DOI":"10.1038\/ncb3052","article-title":"Pri peptides are mediators of ecdysone for the temporal control of development","volume":"16","author":"Chanut-Delalande","year":"2014","journal-title":"Nat Cell Biol."},{"issue":"50","key":"2024110622335965800_jkae204-B18","doi-asserted-by":"publisher","first-page":"17959","DOI":"10.1073\/pnas.1409241111","article-title":"Control of metabolic adaptation to fasting by dILP6-induced insulin signaling in Drosophila oenocytes","volume":"111","author":"Chatterjee","year":"2014","journal-title":"Proc Natl Acad Sci U S A."},{"issue":"2","key":"2024110622335965800_jkae204-B19","doi-asserted-by":"publisher","first-page":"394","DOI":"10.1111\/j.1432-1033.1996.0394r.x","article-title":"Purification and characterisation of haemolymph 3-dehydroecdysone 3\u03b2-reductase in relation to ecdysteroid biosynthesis in the cotton leafworm Spodoptera littoralis","volume":"242","author":"Chen","year":"1996","journal-title":"Eur J Biochem."},{"issue":"6","key":"2024110622335965800_jkae204-B20","doi-asserted-by":"publisher","first-page":"e1006126","DOI":"10.1371\/journal.pgen.1006126","article-title":"Steroid hormone signaling is essential for pheromone production and oenocyte survival","volume":"12","author":"Chiang","year":"2016","journal-title":"PLoS Genet"},{"issue":"3","key":"2024110622335965800_jkae204-B21","doi-asserted-by":"publisher","first-page":"1071","DOI":"10.1534\/genetics.106.066597","article-title":"Cis-regulatory elements in the Accord retrotransposon result in tissue-specific expression of the Drosophila melanogaster insecticide resistance gene Cyp6g1","volume":"175","author":"Chung","year":"2007","journal-title":"Genetics"},{"issue":"20","key":"2024110622335965800_jkae204-B22","doi-asserted-by":"publisher","first-page":"10257","DOI":"10.1002\/ece3.4502","article-title":"Social density, but not sex ratio, drives ecdysteroid hormone provisioning to eggs by female house crickets (Acheta domesticus)","volume":"8","author":"Crocker","year":"2018","journal-title":"Ecol Evol."},{"issue":"3","key":"2024110622335965800_jkae204-B23","doi-asserted-by":"publisher","first-page":"720","DOI":"10.1002\/prot.21357","article-title":"An unsuspected ecdysteroid\/steroid phosphatase activity in the key T-cell regulator, Sts-1: surprising relationship to insect ecdysteroid phosphate phosphatase","volume":"67","author":"Davies","year":"2007","journal-title":"Proteins: Struct. Funct. Bioinf"},{"issue":"5","key":"2024110622335965800_jkae204-B24","doi-asserted-by":"publisher","first-page":"e1005163","DOI":"10.1371\/journal.pgen.1005163","article-title":"Genetic architecture of abdominal pigmentation in Drosophila melanogaster","volume":"11","author":"Dembeck","year":"2015","journal-title":"PLoS Genet"},{"issue":"7150","key":"2024110622335965800_jkae204-B25","doi-asserted-by":"publisher","first-page":"151","DOI":"10.1038\/nature05954","article-title":"A genome-wide transgenic RNAi library for conditional gene inactivation in Drosophila","volume":"448","author":"Dietzl","year":"2007","journal-title":"Nature"},{"issue":"1\u20132","key":"2024110622335965800_jkae204-B26","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1002\/gene.10150","article-title":"GAL4 system in drosophila: a fly geneticist's Swiss army knife","volume":"34","author":"Duffy","year":"2002","journal-title":"Genesis"},{"issue":"7751","key":"2024110622335965800_jkae204-B27","doi-asserted-by":"publisher","first-page":"193","DOI":"10.1038\/s41586-019-1064-z","article-title":"Genetic compensation triggered by mutant mRNA degradation","volume":"568","author":"El-Brolosy","year":"2019","journal-title":"Nature"},{"issue":"7","key":"2024110622335965800_jkae204-B28","doi-asserted-by":"publisher","first-page":"e1006780","DOI":"10.1371\/journal.pgen.1006780","article-title":"Genetic compensation: a phenomenon in search of mechanisms","volume":"13","author":"El-Brolosy","year":"2017","journal-title":"PLoS Genet"},{"issue":"2","key":"2024110622335965800_jkae204-B29","doi-asserted-by":"publisher","first-page":"219","DOI":"10.1016\/0020-1790(88)90027-3","article-title":"Fate of maternally-acquired ecdysteroids in unfertilized eggs of Manduca sexta","volume":"18","author":"Feldlaufer","year":"1988","journal-title":"Insect Biochem"},{"issue":"4","key":"2024110622335965800_jkae204-B30","doi-asserted-by":"crossref","first-page":"240","DOI":"10.1002\/(SICI)1526-968X(200004)26:4<240::AID-GENE40>3.0.CO;2-P","article-title":"Temperature-dependent gene silencing by an expressed inverted repeat in Drosophila","volume":"26","author":"Fortier","year":"2000","journal-title":"Genesis"},{"issue":"6","key":"2024110622335965800_jkae204-B31","doi-asserted-by":"publisher","first-page":"1006","DOI":"10.1101\/gr.081349.108","article-title":"Genomic mapping of binding regions for the Ecdysone receptor protein complex","volume":"19","author":"Gauhar","year":"2009","journal-title":"Genome Res"},{"issue":"4","key":"2024110622335965800_jkae204-B32","doi-asserted-by":"publisher","first-page":"385","DOI":"10.1002\/arch.940010409","article-title":"Metabolism of [3H]-ecdysone in Schistocerca gregaria; formation of ecdysteroid acids together with free and phosphorylated ecdysteroid acetates","volume":"1","author":"Gibson","year":"1984","journal-title":"Arch Insect Biochem Physiol."},{"issue":"4","key":"2024110622335965800_jkae204-B33","doi-asserted-by":"publisher","first-page":"iyac035","DOI":"10.1093\/genetics\/iyac035","article-title":"FlyBase: a guided tour of highlighted features","volume":"220","author":"Gramates","year":"2022","journal-title":"Genetics"},{"key":"2024110622335965800_jkae204-B34","first-page":"189","article-title":"Ovary-specific interaction of ecdysone 22-phosphate with proteins in adult Drosophila melanogaster (Diptera: Drosophilidae)","volume":"92","author":"Grau","year":"1995","journal-title":"Eur J Entomol."},{"issue":"7339","key":"2024110622335965800_jkae204-B35","doi-asserted-by":"publisher","first-page":"473","DOI":"10.1038\/nature09715","article-title":"The developmental transcriptome of Drosophila melanogaster","volume":"471","author":"Graveley","year":"2011","journal-title":"Nature"},{"issue":"1","key":"2024110622335965800_jkae204-B36","doi-asserted-by":"publisher","first-page":"35","DOI":"10.1016\/j.ydbio.2010.09.023","article-title":"CYP18A1, a key enzyme of Drosophila steroid hormone inactivation, is essential for metamorphosis","volume":"349","author":"Guittard","year":"2011","journal-title":"Dev Biol."},{"issue":"3","key":"2024110622335965800_jkae204-B38","doi-asserted-by":"publisher","first-page":"690","DOI":"10.1016\/j.cell.2011.08.047","article-title":"A protein complex network of Drosophila melanogaster","volume":"147","author":"Guruharsha","year":"2011","journal-title":"Cell"},{"key":"2024110622335965800_jkae204-B39","volume-title":"Isolation and Characterization of Ecdysteroid Phosphotransferases","author":"Hilton","year":"2004"},{"issue":"2","key":"2024110622335965800_jkae204-B40","doi-asserted-by":"publisher","first-page":"185","DOI":"10.1016\/0024-3205(85)90422-9","article-title":"Apolar ecdysteroid esters in adult female crickets, Gryllus bimaculatus","volume":"37","author":"Hoffmann","year":"1985","journal-title":"Life Sci"},{"issue":"2","key":"2024110622335965800_jkae204-B41","doi-asserted-by":"publisher","first-page":"533","DOI":"10.1042\/bj2130533","article-title":"Identification of the 22-phosphate esters of ecdysone, 2-deoxyecdysone, 20-hydroxyecdysone and 2-deoxy-20-hydroxyecdysone from newly laid eggs of the desert locust, Schistocerca gregaria","volume":"213","author":"Isaac","year":"1983","journal-title":"Biochem J."},{"key":"2024110622335965800_jkae204-B42","first-page":"340","article-title":"Enzymic hydrolysis of ecdysteroid phosphate during embryogenesis in the desert locust (Schistocerca gregaria)","volume":"50","author":"Isaac","year":"1983","journal-title":"Biochim Biophys Acta."},{"issue":"10","key":"2024110622335965800_jkae204-B43","doi-asserted-by":"publisher","first-page":"e1003843","DOI":"10.1371\/journal.pgen.1003843","article-title":"A novel role for ecdysone in Drosophila conditioned behavior: linking GPCR-mediated non-canonical steroid action to cAMP signaling in the adult brain","volume":"9","author":"Ishimoto","year":"2013","journal-title":"PLoS Genet"},{"key":"2024110622335965800_jkae204-B44","doi-asserted-by":"publisher","first-page":"56","DOI":"10.1016\/j.jinsphys.2015.09.002","article-title":"The genetic basis for mating-induced sex differences in starvation resistance in Drosophila melanogaster","volume":"82","author":"Jang","year":"2015","journal-title":"J Insect Physiol."},{"issue":"2","key":"2024110622335965800_jkae204-B45","doi-asserted-by":"publisher","first-page":"200373","DOI":"10.1098\/rsob.200373","article-title":"Regulation of ecdysone production in Drosophila by neuropeptides and peptide hormones","volume":"11","author":"Kannangara","year":"2021","journal-title":"Open Biol"},{"issue":"4083","key":"2024110622335965800_jkae204-B46","doi-asserted-by":"publisher","first-page":"307","DOI":"10.1126\/science.180.4083.307","article-title":"26-Hydroxyecdysone: new insect molting hormone from the egg of the tobacco hornworm","volume":"180","author":"Kaplanis","year":"1973","journal-title":"Science"},{"key":"2024110622335965800_jkae204-B47","doi-asserted-by":"crossref","first-page":"106","DOI":"10.1016\/B978-0-12-384749-2.10004-4","volume-title":"In: Insect Endocrinology","author":"Lafont","year":"2012"},{"key":"2024110622335965800_jkae204-B48","first-page":"168","volume-title":"In: Proceedings in Life Sciences. Biosynthesis, Metabolism and Mode of Action of Invertebrate Hormones","author":"Lagueux","year":"1984"},{"issue":"D1","key":"2024110622335965800_jkae204-B49","doi-asserted-by":"publisher","first-page":"D809","DOI":"10.1093\/nar\/gkx976","article-title":"FlyAtlas 2: a new version of the Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data","volume":"46","author":"Leader","year":"2018","journal-title":"Nucleic Acids Res"},{"key":"2024110622335965800_jkae204-B50","doi-asserted-by":"publisher","first-page":"33","DOI":"10.1016\/j.ibmb.2014.08.007","article-title":"CYP18A1 regulates tissue-specific steroid hormone inactivation in Bombyx mori","volume":"54","author":"Li","year":"2014","journal-title":"Insect Biochem Mol Biol."},{"key":"2024110622335965800_jkae204-B51","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1016\/j.ibmb.2014.03.014","article-title":"Wolbachia-induced paternal defect in Drosophila is likely by interaction with the juvenile hormone pathway","volume":"49","author":"Liu","year":"2014","journal-title":"Insect Biochem Mol Biol."},{"issue":"1","key":"2024110622335965800_jkae204-B52","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41598-018-27076-y","article-title":"The Drosophila homologue of MEGF8 is essential for early development","volume":"8","author":"Lloyd","year":"2018","journal-title":"Sci Rep."},{"issue":"1","key":"2024110622335965800_jkae204-B53","doi-asserted-by":"publisher","first-page":"63","DOI":"10.1002\/arch.940120106","article-title":"Profiles of free and conjugated ecdysteroids and ecdysteroid acids during pupal\u2013adult development of Manduca sexta","volume":"12","author":"Lozano","year":"1989","journal-title":"Arch Insect Biochem Physiol."},{"issue":"3","key":"2024110622335965800_jkae204-B54","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1002\/arch.10055","article-title":"A possible role of 20-hydroxyecdysone in embryonic development of the silkworm Bombyx mori","volume":"51","author":"Makka","year":"2002","journal-title":"Arch Insect Biochem Physiol."},{"issue":"3","key":"2024110622335965800_jkae204-B55","doi-asserted-by":"crossref","first-page":"310","DOI":"10.1002\/(SICI)1097-0177(199707)209:3<310::AID-AJA6>3.0.CO;2-L","article-title":"GAL4 enhancer traps expressed in the embryo, larval brain, imaginal discs, and ovary of Drosophila","volume":"209","author":"Manseau","year":"1997","journal-title":"Dev Dyn."},{"issue":"7","key":"2024110622335965800_jkae204-B56","doi-asserted-by":"publisher","first-page":"633","DOI":"10.1016\/0022-1910(88)90071-6","article-title":"Embryonic ecdysteroids of Drosophila melanogaster","volume":"34","author":"Mar\u00f3y","year":"1988","journal-title":"J Insect Physiol."},{"issue":"3","key":"2024110622335965800_jkae204-B57","doi-asserted-by":"publisher","first-page":"151","DOI":"10.1159\/000502361","article-title":"Involvement of ecdysone signaling in the expression of the doublesex gene during embryonic development in the silkworm, Bombyx mori","volume":"13","author":"Matsushima","year":"2019","journal-title":"Sex Dev"},{"issue":"22","key":"2024110622335965800_jkae204-B58","doi-asserted-by":"publisher","first-page":"3593","DOI":"10.1080\/15384101.2015.1100772","article-title":"Early-late genes of the ecdysone cascade as models for transcriptional studies","volume":"14","author":"Mazina","year":"2015","journal-title":"Cell Cycle"},{"issue":"3","key":"2024110622335965800_jkae204-B59","doi-asserted-by":"publisher","first-page":"161","DOI":"10.1080\/01688170.1984.10510087","article-title":"Ecdysone metabolism in Locusta migratoria larvae and adults","volume":"7","author":"Modde","year":"1984","journal-title":"Int J Invert Reproduct Dev"},{"key":"2024110622335965800_jkae204-B60","doi-asserted-by":"publisher","first-page":"727","DOI":"10.1093\/icb\/21.3.727","article-title":"Ecdysteroid metabolism in insects","volume":"21","author":"Ohtaki","year":"1981","journal-title":"Integr Comp Biol."},{"issue":"1","key":"2024110622335965800_jkae204-B61","doi-asserted-by":"publisher","first-page":"32","DOI":"10.1016\/j.ydbio.2014.04.004","article-title":"Ecdysone differentially regulates metamorphic timing relative to 20-hydroxyecdysone by antagonizing juvenile hormone in Drosophila melanogaster","volume":"391","author":"Ono","year":"2014","journal-title":"Dev Biol."},{"issue":"1","key":"2024110622335965800_jkae204-B62","doi-asserted-by":"publisher","first-page":"247","DOI":"10.1016\/j.celrep.2016.05.053","article-title":"The insect prothoracic gland as a model for steroid hormone biosynthesis and regulation","volume":"16","author":"Ou","year":"2016","journal-title":"Cell Rep"},{"issue":"1","key":"2024110622335965800_jkae204-B63","doi-asserted-by":"publisher","first-page":"iyad211","DOI":"10.1093\/genetics\/iyad211","article-title":"FlyBase: updates to the Drosophila genes and genomes database","volume":"227","author":"\u00d6zt\u00fcrk-\u00c7olak","year":"2024","journal-title":"Genetics"},{"issue":"8","key":"2024110622335965800_jkae204-B64","doi-asserted-by":"publisher","first-page":"e1002925","DOI":"10.1371\/journal.pgen.1002925","article-title":"Drosophila melanogaster acetyl-CoA-carboxylase sustains a fatty acid\u2013dependent remote signal to waterproof the respiratory system","volume":"8","author":"Parvy","year":"2012","journal-title":"PLoS Genet"},{"issue":"7921","key":"2024110622335965800_jkae204-B65","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1038\/s41586-022-04908-6","article-title":"A male steroid controls female sexual behaviour in the malaria mosquito","volume":"608","author":"Peng","year":"2022","journal-title":"Nature"},{"issue":"7","key":"2024110622335965800_jkae204-B66","doi-asserted-by":"publisher","first-page":"1361","DOI":"10.1242\/jcs.02839","article-title":"Brush border spectrin is required for early endosome recycling in Drosophila","volume":"119","author":"Phillips","year":"2006","journal-title":"J Cell Sci."},{"key":"2024110622335965800_jkae204-B67","first-page":"41","article-title":"Analysis of ecdysteroid conjugates: chromatographic characterization of sulfates, phosphates and glucosides","volume":"92","author":"Pis","year":"1995","journal-title":"Eur J Entomol."},{"issue":"1","key":"2024110622335965800_jkae204-B68","doi-asserted-by":"publisher","first-page":"65","DOI":"10.1016\/0022-1910(83)90107-5","article-title":"Ecdysteroid synthesis by the ring gland of Drosophila melanogaster during late-larval, prepupal and pupal development","volume":"29","author":"Redfern","year":"1983","journal-title":"J Insect Physiol."},{"issue":"8","key":"2024110622335965800_jkae204-B69","doi-asserted-by":"publisher","first-page":"741","DOI":"10.1016\/j.ibmb.2007.02.012","article-title":"Molecular evolution of the insect Halloween family of cytochrome P450s: phylogeny, gene organization and functional conservation","volume":"37","author":"Rewitz","year":"2007","journal-title":"Insect Biochem Mol Biol."},{"issue":"6","key":"2024110622335965800_jkae204-B70","doi-asserted-by":"publisher","first-page":"1256","DOI":"10.1042\/bst0341256","article-title":"The Halloween genes code for cytochrome P450 enzymes mediating synthesis of the insect moulting hormone","volume":"34","author":"Rewitz","year":"2006","journal-title":"Biochem Soc Trans."},{"issue":"6","key":"2024110622335965800_jkae204-B71","doi-asserted-by":"publisher","first-page":"895","DOI":"10.1016\/j.devcel.2010.10.021","article-title":"Steroid hormone inactivation is required during the juvenile\u2013adult transition in Drosophila","volume":"19","author":"Rewitz","year":"2010","journal-title":"Dev Cell."},{"key":"2024110622335965800_jkae204-B72","first-page":"899","volume-title":"The Development of Drosophila melanogaster","author":"Riddiford","year":"1993"},{"issue":"1","key":"2024110622335965800_jkae204-B73","doi-asserted-by":"publisher","first-page":"123","DOI":"10.1242\/dev.042036","article-title":"The Drosophila nuclear receptors DHR3 and betaFTZ-F1 control overlapping developmental responses in late embryos","volume":"137","author":"Ruaud","year":"2010","journal-title":"Development"},{"issue":"3","key":"2024110622335965800_jkae204-B74","doi-asserted-by":"publisher","first-page":"179","DOI":"10.1002\/arch.940100303","article-title":"Mediation of ecdysone synthesis in Manduca sexta by a hemolymph enzyme","volume":"10","author":"Sakurai","year":"1989","journal-title":"Arch Insect Biochem Physiol."},{"issue":"5","key":"2024110622335965800_jkae204-B75","doi-asserted-by":"publisher","first-page":"e0233231","DOI":"10.1371\/journal.pone.0233231","article-title":"Small heat shock proteins determine synapse number and neuronal activity during development","volume":"15","author":"Santana","year":"2020","journal-title":"PLoS One"},{"key":"2024110622335965800_jkae204-B76","doi-asserted-by":"publisher","first-page":"100030","DOI":"10.1016\/j.cris.2022.100030","article-title":"Ecdysteroid kinase-like (EcKL) paralogs confer developmental tolerance to caffeine in Drosophila melanogaster","volume":"2","author":"Scanlan","year":"2022","journal-title":"Curr Res Insect Sci."},{"key":"2024110622335965800_jkae204-B77","doi-asserted-by":"publisher","first-page":"103429","DOI":"10.1016\/j.ibmb.2020.103429","article-title":"Genomic and transcriptomic analyses in Drosophila suggest that the ecdysteroid kinase-like (EcKL) gene family encodes the \u2018detoxification-by-phosphorylation\u2019enzymes of insects","volume":"123","author":"Scanlan","year":"2020","journal-title":"Insect Biochem Mol Biol."},{"issue":"2","key":"2024110622335965800_jkae204-B78","doi-asserted-by":"publisher","first-page":"evae019","DOI":"10.1093\/gbe\/evae019","article-title":"Phylogenomics of the ecdysteroid kinase-like (EcKL) gene family in insects highlights roles in both steroid hormone metabolism and detoxification","volume":"16","author":"Scanlan","year":"2024","journal-title":"Genome Biol Evol."},{"key":"2024110622335965800_jkae204-B79","doi-asserted-by":"publisher","first-page":"103891","DOI":"10.1016\/j.ibmb.2022.103891","article-title":"Rethinking the ecdysteroid source during Drosophila pupal\u2013adult development","volume":"152","author":"Scanlan","year":"2023","journal-title":"Insect Biochem Mol Biol."},{"issue":"4","key":"2024110622335965800_jkae204-B80","doi-asserted-by":"publisher","first-page":"206","DOI":"10.1093\/bfgp\/elr017","article-title":"RNAi-induced off-target effects in Drosophila melanogaster: frequencies and solutions","volume":"10","author":"Seinen","year":"2011","journal-title":"Brief Funct Genomics."},{"issue":"1","key":"2024110622335965800_jkae204-B81","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1002\/arch.940150105","article-title":"P1 gene expression in Drosophila larval fat body: induction by various ecdysteroids","volume":"15","author":"Somm\u00e9-Martin","year":"1990","journal-title":"Arch Insect Biochem Physiol."},{"issue":"7","key":"2024110622335965800_jkae204-B82","doi-asserted-by":"publisher","first-page":"729","DOI":"10.1016\/0020-1790(88)90082-0","article-title":"Conversion of ecdysone and 20-hydroxyecdysone into 3-dehydroecdysteroids is a major pathway in third instar Drosophila melanogaster larvae","volume":"18","author":"Somm\u00e9-Martin","year":"1988","journal-title":"Insect Biochem"},{"issue":"7","key":"2024110622335965800_jkae204-B83","doi-asserted-by":"publisher","first-page":"735","DOI":"10.1016\/0020-1790(88)90083-2","article-title":"Metabolism and biosynthesis of ecdysteroids in the Drosophila development mutant ecd1","volume":"18","author":"Somm\u00e9-Martin","year":"1988","journal-title":"Insect Biochem"},{"issue":"1\u20132","key":"2024110622335965800_jkae204-B84","doi-asserted-by":"publisher","first-page":"25","DOI":"10.1016\/j.mce.2009.03.017","article-title":"Phosphoconjugation and dephosphorylation reactions of steroid hormone in insects","volume":"307","author":"Sonobe","year":"2009","journal-title":"Mol Cell Endocrinol."},{"issue":"40","key":"2024110622335965800_jkae204-B85","doi-asserted-by":"publisher","first-page":"29513","DOI":"10.1074\/jbc.m604035200","article-title":"Purification, kinetic characterization, and molecular cloning of a novel enzyme, ecdysteroid 22-kinase","volume":"281","author":"Sonobe","year":"2006","journal-title":"J Biol Chem."},{"issue":"5","key":"2024110622335965800_jkae204-B86","doi-asserted-by":"publisher","first-page":"503","DOI":"10.2108\/zsj.21.503","article-title":"Ecdysteroids during early embryonic development in silkworm Bombyx mori: metabolism and functions","volume":"21","author":"Sonobe","year":"2004","journal-title":"Zoolog Sci."},{"issue":"3","key":"2024110622335965800_jkae204-B87","doi-asserted-by":"publisher","first-page":"683","DOI":"10.1534\/g3.115.023366","article-title":"Diverse hormone response networks in 41 independent Drosophila cell lines","volume":"6","author":"Stoiber","year":"2016","journal-title":"G3 (Bethesda)"},{"issue":"2","key":"2024110622335965800_jkae204-B88","doi-asserted-by":"publisher","first-page":"200","DOI":"10.1016\/j.devcel.2018.11.030","article-title":"Drosophila HNF4 directs a switch in lipid metabolism that supports the transition to adulthood","volume":"48","author":"Storelli","year":"2019","journal-title":"Dev Cell."},{"issue":"1","key":"2024110622335965800_jkae204-B89","doi-asserted-by":"publisher","first-page":"125","DOI":"10.7150\/ijbs.8.125","article-title":"Molecular cloning and characterization of Ecdysone oxidase and 3-dehydroecdysone-3\u03b1-reductase involved in the ecdysone inactivation pathway of silkworm, Bombyx mori","volume":"8","author":"Sun","year":"2012","journal-title":"Int J Biol Sci."},{"issue":"5","key":"2024110622335965800_jkae204-B90","doi-asserted-by":"publisher","first-page":"e36544","DOI":"10.1371\/journal.pone.0036544","article-title":"A cytochrome P450 conserved in insects is involved in cuticle formation","volume":"7","author":"Sztal","year":"2012","journal-title":"PLoS One"},{"issue":"29","key":"2024110622335965800_jkae204-B91","doi-asserted-by":"publisher","first-page":"26819","DOI":"10.1074\/jbc.m104291200","article-title":"Regulation of ecdysteroid signaling: cloning and characterization of ecdysone oxidase","volume":"276","author":"Takeuchi","year":"2001","journal-title":"J Biol Chem."},{"issue":"7","key":"2024110622335965800_jkae204-B92","doi-asserted-by":"publisher","first-page":"743","DOI":"10.1016\/s0022-1910(02)00099-9","article-title":"Possible involvement of ecdysteroids in embryonic diapause of Locusta migratoria","volume":"48","author":"Tawfik","year":"2002","journal-title":"J Insect Physiol."},{"issue":"1","key":"2024110622335965800_jkae204-B93","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1002\/arch.940040102","article-title":"Metabolism of 26-[C-14]hydroxyecdysone 26-phosphate in the tobacco hornworm, Manduca sexta L., to a new ecdysteroid conjugate: 26-[C-14]hydroxyecdysone 22-glucoside","volume":"4","author":"Thompson","year":"1987","journal-title":"Arch Insect Biochem Physiol."},{"issue":"1","key":"2024110622335965800_jkae204-B94","doi-asserted-by":"publisher","first-page":"8","DOI":"10.1186\/1472-6793-10-8","article-title":"Hypoxia modifies the feeding preferences of Drosophila. Consequences for diet dependent hypoxic survival","volume":"10","author":"Vigne","year":"2010","journal-title":"BMC Physiol"},{"issue":"11","key":"2024110622335965800_jkae204-B95","doi-asserted-by":"publisher","first-page":"3870","DOI":"10.1128\/mcb.20.11.3870-3879.2000","article-title":"Molecular determinants of differential ligand sensitivities of insect ecdysteroid receptors","volume":"20","author":"Wang","year":"2000","journal-title":"Mol Cell Biol."},{"issue":"2","key":"2024110622335965800_jkae204-B96","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1111\/jzo.12226","article-title":"Report on Drosophila melanogaster larvae without functional tracheae","volume":"296","author":"Wang","year":"2015","journal-title":"J Zool."},{"issue":"5","key":"2024110622335965800_jkae204-B97","doi-asserted-by":"publisher","first-page":"605","DOI":"10.1038\/nprot.2009.55","article-title":"Determination of gene expression patterns using high-throughput RNA in situ hybridization to whole-mount Drosophila embryos","volume":"4","author":"Weiszmann","year":"2009","journal-title":"Nat Protoc."},{"issue":"2","key":"2024110622335965800_jkae204-B98","doi-asserted-by":"publisher","first-page":"91","DOI":"10.1006\/pest.2001.2588","article-title":"Species specificity of changes in ecdysteroid metabolism in response to ecdysteroid agonists","volume":"72","author":"Williams","year":"2002","journal-title":"Pestic Biochem Physiol."},{"issue":"1","key":"2024110622335965800_jkae204-B99","doi-asserted-by":"publisher","first-page":"52","DOI":"10.1016\/j.ydbio.2017.08.018","article-title":"Debris buster is a Drosophila scavenger receptor essential for airway physiology","volume":"430","author":"Wingen","year":"2017","journal-title":"Dev Biol."},{"key":"2024110622335965800_jkae204-B100","doi-asserted-by":"publisher","first-page":"103210","DOI":"10.1016\/j.ibmb.2019.103210","article-title":"CYP303A1 has a conserved function in adult eclosion in Locusta migratoria and Drosophila melanogaster","volume":"113","author":"Wu","year":"2019","journal-title":"Insect Biochem Mol Biol."},{"issue":"4","key":"2024110622335965800_jkae204-B101","doi-asserted-by":"publisher","first-page":"811","DOI":"10.1038\/sj.emboj.7600112","article-title":"A lysosomal tetraspanin associated with retinal degeneration identified via a genome-wide screen","volume":"23","author":"Xu","year":"2004","journal-title":"EMBO J"},{"issue":"29","key":"2024110622335965800_jkae204-B102","doi-asserted-by":"publisher","first-page":"26365","DOI":"10.1074\/jbc.m304158200","article-title":"Purification, kinetic characterization, and molecular cloning of a novel enzyme ecdysteroid-phosphate phosphatase","volume":"278","author":"Yamada","year":"2003","journal-title":"J Biol Chem."},{"issue":"2","key":"2024110622335965800_jkae204-B103","doi-asserted-by":"publisher","first-page":"187","DOI":"10.2108\/zsj.22.187","article-title":"Release of ecdysteroid-phosphates from egg yolk granules and their dephosphorylation during early embryonic development in silkworm, Bombyx mori","volume":"22","author":"Yamada","year":"2005","journal-title":"Zoolog Sci."},{"issue":"1","key":"2024110622335965800_jkae204-B104","doi-asserted-by":"publisher","first-page":"497","DOI":"10.1146\/annurev-ento-120811-153608","article-title":"Ecdysone control of developmental transitions: lessons from Drosophila research","volume":"58","author":"Yamanaka","year":"2013","journal-title":"Annu Rev Entomol."},{"issue":"1","key":"2024110622335965800_jkae204-B105","doi-asserted-by":"publisher","first-page":"235","DOI":"10.1016\/0012-1606(80)90511-4","article-title":"Endocrine control of the metamorphosis of the larval muscles in Calliphora erythrocephala (Diptera): in vitro studies of the role of ecdysteroids","volume":"80","author":"Zachary","year":"1980","journal-title":"Dev Biol."},{"issue":"1","key":"2024110622335965800_jkae204-B106","doi-asserted-by":"publisher","first-page":"125","DOI":"10.1016\/j.devcel.2023.11.016","article-title":"Eclosion muscles secrete ecdysteroids to initiate asymmetric intestinal stem cell division in Drosophila","volume":"59","author":"Zhang","year":"2024","journal-title":"Dev Cell."}],"container-title":["G3: Genes, Genomes, Genetics"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/g3journal\/advance-article-pdf\/doi\/10.1093\/g3journal\/jkae204\/59596851\/jkae204.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/g3journal\/article-pdf\/14\/11\/jkae204\/60456387\/jkae204.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/g3journal\/article-pdf\/14\/11\/jkae204\/60456387\/jkae204.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,11,6]],"date-time":"2024-11-06T22:37:05Z","timestamp":1730932625000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/g3journal\/article\/doi\/10.1093\/g3journal\/jkae204\/7745362"}},"subtitle":[],"editor":[{"given":"J","family":"Tennessen","sequence":"additional","affiliation":[],"role":[{"role":"editor","vocabulary":"crossref"}]}],"short-title":[],"issued":{"date-parts":[[2024,8,29]]},"references-count":105,"journal-issue":{"issue":"11","published-online":{"date-parts":[[2024,8,29]]},"published-print":{"date-parts":[[2024,11,6]]}},"URL":"https:\/\/doi.org\/10.1093\/g3journal\/jkae204","relation":{},"ISSN":["2160-1836"],"issn-type":[{"value":"2160-1836","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2024,11]]},"published":{"date-parts":[[2024,8,29]]},"article-number":"jkae204"}}