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While stress-response mechanisms activated during and immediately after freezing are well studied, less is known about how freeze-tolerant organisms recover from freezing in the longer term and whether this recovery carries energetic costs. Here, we tracked changes in gene expression and energy stores over the course of 15\u2005days following extracellular freezing using the world's southernmost insect, Belgica antarctica (Diptera: Chironomidae), as a study system. We found that B. antarctica employed a coordinated \u2018emergency\u2019 stress-response system during early recovery (0\u20131\u2005day), which includes the upregulation of genes involved in inhibiting cell death (e.g. Bcl-2, IAP-1) alongside mechanisms involved in damage repair and clearance of cell debris (e.g. heat shock proteins, autophagy, proteasome). Concomitantly, genes involved in ecdysone biosynthesis and juvenile hormone degradation were downregulated through the 1st day of recovery, and several genes involved in cuticle development were downregulated between the 3rd and 15th day of recovery, suggesting that homeostasis may not have been completely restored for 2\u00a0weeks after B. antarctica had thawed. These physiological changes did not lead to a detectable depletion of energy stores, indicating that this emergency response system does not incur significant energy drain. These results suggest that evoking a coordinated stress response with minimal energy usage may be crucial for B. antarctica to persist in cold environments, and that blocking apoptosis and pausing development could provide sufficient time for freezing injury to be fully addressed.<\/jats:p>","DOI":"10.1242\/jeb.252542","type":"journal-article","created":{"date-parts":[[2026,7,14]],"date-time":"2026-07-14T09:48:16Z","timestamp":1784022496000},"source":"Crossref","is-referenced-by-count":1,"title":["Molecular mechanisms and energetic costs of recovery from freezing in the polyextremophile midge\n                    <i>Belgica antarctica<\/i>"],"prefix":"10.1242","volume":"229","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-0020-8939","authenticated-orcid":false,"given":"Cleverson","family":"Lima","sequence":"first","affiliation":[{"name":"University of Kentucky 1 Department of Entomology , , Lexington, KY 40546-0091 ,","place":["USA"]},{"name":"University of Kentucky 2 Department of Biology , , Lexington, KY 40508-0225 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4727-1951","authenticated-orcid":false,"given":"Melise C.","family":"Lecheta","sequence":"additional","affiliation":[{"name":"University of Kentucky 1 Department of Entomology , , Lexington, KY 40546-0091 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0003-6786-3978","authenticated-orcid":false,"given":"Samuel","family":"Cecconi","sequence":"additional","affiliation":[{"name":"University of Kentucky 1 Department of Entomology , , Lexington, KY 40546-0091 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5965-0986","authenticated-orcid":false,"given":"Scott","family":"Hotaling","sequence":"additional","affiliation":[{"name":"Utah State University 3 Department of Watershed Sciences , , Logan, UT 84322 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4801-7579","authenticated-orcid":false,"given":"Paul B.","family":"Frandsen","sequence":"additional","affiliation":[{"name":"Brigham Young University 4 Department of Plant and Wildlife Sciences and Bean Life Science Museum , , Provo, UT 84602 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8497-9903","authenticated-orcid":false,"given":"Peter","family":"Convey","sequence":"additional","affiliation":[{"name":"Natural Environment Research Council 5 British Antarctic Survey , , Cambridge, CB3 0ET ,","place":["UK"]},{"name":"University of Johannesburg 6 Department of Zoology , , Auckland Park 2006 ,","place":["South Africa"]},{"name":"Millennium Institute - Biodiversity of Antarctic and Sub-Antarctic Ecosystems (BASE) 7 , 3425 Santiago ,","place":["Chile"]},{"name":"School of Biosciences, University of Birmingham 8 , Birmingham, B15 2TT ,","place":["UK"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1899-6630","authenticated-orcid":false,"given":"Scott A. L.","family":"Hayward","sequence":"additional","affiliation":[{"name":"School of Biosciences, University of Birmingham 8 , Birmingham, B15 2TT ,","place":["UK"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3847-6031","authenticated-orcid":false,"given":"J. D.","family":"Gantz","sequence":"additional","affiliation":[{"name":"Hendrix College 9 Department of Biology and Health Sciences , , Conway, AR 72032 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuta","family":"Kawarasaki","sequence":"additional","affiliation":[{"name":"Gustavus Adolphus College 10 Department of Biology , , Saint Peter, MN 56082 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5333-5531","authenticated-orcid":false,"given":"Jeramiah J.","family":"Smith","sequence":"additional","affiliation":[{"name":"University of Kentucky 2 Department of Biology , , Lexington, KY 40508-0225 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0963-7457","authenticated-orcid":false,"given":"Nicholas M.","family":"Teets","sequence":"additional","affiliation":[{"name":"University of Kentucky 1 Department of Entomology , , Lexington, KY 40546-0091 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","published-online":{"date-parts":[[2026,8,12]]},"reference":[{"key":"2026081203534043100_JEB252542C1","doi-asserted-by":"publisher","first-page":"542","DOI":"10.1186\/s13071-021-05056-y","article-title":"Functional implication of heat shock protein 70\/90 and tubulin in cold stress of Dermacentor silvarum","volume":"14","author":"Agwunobi","year":"2021","journal-title":"Parasit. 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PhD Thesis, Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije. https:\/\/repositorij.upr.si\/IzpisGradiva.php? lang=env&id=21738\/"},{"key":"2026081203534043100_JEB252542C40","doi-asserted-by":"publisher","first-page":"D353","DOI":"10.1093\/nar\/gkw1092","article-title":"KEGG: new perspectives on genomes, pathways, diseases and drugs","volume":"45","author":"Kanehisa","year":"2017","journal-title":"Nucleic Acids Res."},{"key":"2026081203534043100_JEB252542C41","doi-asserted-by":"publisher","first-page":"753","DOI":"10.1007\/s00300-014-1475-0","article-title":"Wet hibernacula promote inoculative freezing and limit the potential for cryoprotective dehydration in the Antarctic midge, Belgica antarctica","volume":"37","author":"Kawarasaki","year":"2014","journal-title":"Polar Biol."},{"key":"2026081203534043100_JEB252542C42","doi-asserted-by":"publisher","first-page":"933","DOI":"10.1111\/1365-2435.12229","article-title":"Alternative overwintering strategies in an Antarctic midge: freezing vs. cryoprotective dehydration","volume":"28","author":"Kawarasaki","year":"2014","journal-title":"Funct. 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