{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,19]],"date-time":"2026-03-19T14:12:41Z","timestamp":1773929561300,"version":"3.50.1"},"reference-count":43,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2026,3,1]],"date-time":"2026-03-01T00:00:00Z","timestamp":1772323200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,3,1]],"date-time":"2026-03-01T00:00:00Z","timestamp":1772323200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2025,12,17]],"date-time":"2025-12-17T00:00:00Z","timestamp":1765929600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100012774","name":"Innovation Fund Denmark","doi-asserted-by":"publisher","award":["2040-00062B"],"award-info":[{"award-number":["2040-00062B"]}],"id":[{"id":"10.13039\/100012774","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Cryobiology"],"published-print":{"date-parts":[[2026,3]]},"DOI":"10.1016\/j.cryobiol.2025.105571","type":"journal-article","created":{"date-parts":[[2025,12,23]],"date-time":"2025-12-23T20:23:52Z","timestamp":1766521432000},"page":"105571","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":0,"special_numbering":"C","title":["Temperature-metabolism relationships above and below the body fluid freezing point in the freeze-tolerant white worm, Enchytraeus albidus"],"prefix":"10.1016","volume":"122","author":[{"ORCID":"https:\/\/orcid.org\/0009-0003-2880-2558","authenticated-orcid":false,"given":"Mathias Engell","family":"Dahl Holmstrup","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2851-4551","authenticated-orcid":false,"given":"Johannes","family":"Overgaard","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9974-0014","authenticated-orcid":false,"given":"Stine","family":"Slotsbo","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"78","reference":[{"key":"10.1016\/j.cryobiol.2025.105571_bib1","series-title":"Advances in Insect Physiology","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/S0065-2806(08)60109-5","article-title":"Frost resistance in insects","author":"Asahina","year":"1970"},{"key":"10.1016\/j.cryobiol.2025.105571_bib2","doi-asserted-by":"crossref","first-page":"201","DOI":"10.1016\/0044-8486(92)90149-F","article-title":"Oligochaetes (Tubificidae and Enchytraeidae) as food in fish rearing: a review and preliminary tests","volume":"102","author":"Bouguenec","year":"1992","journal-title":"Aquaculture"},{"key":"10.1016\/j.cryobiol.2025.105571_bib3","doi-asserted-by":"crossref","first-page":"859","DOI":"10.1242\/jeb.026864","article-title":"Dual roles of glucose in the freeze-tolerant earthworm Dendrobaena octaedra: cryoprotection and fuel for metabolism","volume":"212","author":"Calderon","year":"2009","journal-title":"J. Exp. Biol."},{"key":"10.1016\/j.cryobiol.2025.105571_bib4","series-title":"Temperature Biology of Animals","author":"Cossins","year":"1987"},{"key":"10.1016\/j.cryobiol.2025.105571_bib5","doi-asserted-by":"crossref","first-page":"1961","DOI":"10.1242\/jeb.070268","article-title":"Avoidance and tolerance of freezing in ectothermic vertebrates","volume":"216","author":"Costanzo","year":"2013","journal-title":"J. Exp. Biol."},{"key":"10.1016\/j.cryobiol.2025.105571_bib6","doi-asserted-by":"crossref","DOI":"10.1016\/j.jtherbio.2021.102865","article-title":"Thermal optimum for mass production of the live feed organism Enchytraeus albidus","volume":"97","author":"Dai","year":"2021","journal-title":"J. Therm. Biol."},{"key":"10.1016\/j.cryobiol.2025.105571_bib7","doi-asserted-by":"crossref","first-page":"105","DOI":"10.1007\/s13127-019-00402-6","article-title":"The popular model annelid Enchytraeus albidus is only one species in a complex of seashore white worms (Clitellata, Enchytraeidae)","volume":"19","author":"Ers\u00e9us","year":"2019","journal-title":"Org. Divers. Evol."},{"key":"10.1016\/j.cryobiol.2025.105571_bib8","doi-asserted-by":"crossref","first-page":"16","DOI":"10.1016\/j.aquaculture.2017.08.019","article-title":"Production and nutritional composition of white worms Enchytraeus albidus fed different low-cost feeds","volume":"481","author":"Fairchild","year":"2017","journal-title":"Aquaculture"},{"key":"10.1016\/j.cryobiol.2025.105571_bib9","doi-asserted-by":"crossref","DOI":"10.1242\/jeb.105650","article-title":"Effect of repeated freeze-thaw cycles on geographically different populations of the freeze-tolerant worm Enchytraeus albidus (Oligochaeta)","volume":"217","author":"Fisker","year":"2014","journal-title":"J. Exp. Biol."},{"key":"10.1016\/j.cryobiol.2025.105571_bib10","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1007\/s00360-013-0788-6","article-title":"Roles of carbohydrate reserves for local adaptation to low temperatures in the freeze tolerant oligochaete Enchytraeus albidus","volume":"184","author":"Fisker","year":"2014","journal-title":"J. Comp. Physiol. B"},{"key":"10.1016\/j.cryobiol.2025.105571_bib11","series-title":"Effects of Multiple Stressors on Arctic Soil Organisms","author":"Frandsen","year":"2019"},{"key":"10.1016\/j.cryobiol.2025.105571_bib12","doi-asserted-by":"crossref","DOI":"10.7717\/peerj.5675","article-title":"Body size and temperature effects on standard metabolic rate for determining metabolic scope for activity of the polychaete Hediste (Nereis) diversicolor","volume":"6","author":"Galasso","year":"2018","journal-title":"PeerJ"},{"key":"10.1016\/j.cryobiol.2025.105571_bib13","doi-asserted-by":"crossref","DOI":"10.1016\/j.ecoenv.2020.111005","article-title":"Mercury (Hg2+) interferes with physiological adaptations to freezing in the arctic earthworm Enchytraeus albidus","volume":"204","author":"Gerlich","year":"2020","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"10.1016\/j.cryobiol.2025.105571_bib14","series-title":"Animal Physiology","author":"Hill","year":"2012"},{"key":"10.1016\/j.cryobiol.2025.105571_bib15","first-page":"504","article-title":"Overwintering adaptations in earthworms: the 7th international symposium on earthworm ecology \u00b7 Cardiff \u00b7 Wales \u00b7 2002","volume":"47","author":"Holmstrup","year":"2003","journal-title":"Pedobiologia"},{"key":"10.1016\/j.cryobiol.2025.105571_bib16","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1007\/s003600050213","article-title":"Cryoprotective and osmotic responses to cold acclimation and freezing in freeze-tolerant and freeze-intolerant earthworms","volume":"169","author":"Holmstrup","year":"1999","journal-title":"J. Comp. Physiol. B"},{"key":"10.1016\/j.cryobiol.2025.105571_bib17","doi-asserted-by":"crossref","first-page":"2640","DOI":"10.1016\/j.soilbio.2007.05.018","article-title":"Adaptations to overwintering in the earthworm Dendrobaena octaedra: genetic differences in glucose mobilisation and freeze tolerance","volume":"39","author":"Holmstrup","year":"2007","journal-title":"Soil Biol. Biochem."},{"key":"10.1016\/j.cryobiol.2025.105571_bib18","first-page":"192","article-title":"Analysis of heat and cold tolerance of a freeze-tolerant soil invertebrate distributed from temperate to Arctic regions: evidence of selection for extreme cold tolerance","volume":"B","author":"Holmstrup","year":"2022","journal-title":"J. Comp. Physiol."},{"key":"10.1016\/j.cryobiol.2025.105571_bib19","doi-asserted-by":"crossref","first-page":"1","DOI":"10.3920\/JIFF2021.0106","article-title":"A new strategy in rearing of European flounder: using live Enchytraeus albidus to enhance juvenile growth","volume":"8","author":"Holmstrup","year":"2022","journal-title":"Journal of Insects as Food and Feed"},{"key":"10.1016\/j.cryobiol.2025.105571_bib20","doi-asserted-by":"crossref","first-page":"345","DOI":"10.1002\/jez.10082","article-title":"Energy and water conservation in frozen vs. supercooled larvae of the goldenrod gall fly, Eurosta solidaginis (fitch) (Diptera: Tephritidae)","volume":"292","author":"Irwin","year":"2002","journal-title":"J. Exp. Zool."},{"key":"10.1016\/j.cryobiol.2025.105571_bib21","series-title":"Mass Cultivation of Invertebrates : Biology and Methods","author":"Ivleva","year":"1973"},{"key":"10.1016\/j.cryobiol.2025.105571_bib22","doi-asserted-by":"crossref","first-page":"258","DOI":"10.2307\/1539211","article-title":"Histology and metabolism of frozen intertidal animals","volume":"116","author":"Kanwisher","year":"1959","journal-title":"Biol. Bull."},{"key":"10.1016\/j.cryobiol.2025.105571_bib23","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1146\/annurev.physiol.60.1.55","article-title":"Biological ice nucleation and ice distribution in cold-hardy ectothermic animals","volume":"60","author":"Lee","year":"1998","journal-title":"Annu. Rev. Physiol."},{"key":"10.1016\/j.cryobiol.2025.105571_bib24","doi-asserted-by":"crossref","DOI":"10.1016\/j.aquaculture.2022.738586","article-title":"First feed matters: the first diet of larval fish programmes growth, survival, and metabolism of larval ballan wrasse (Labrus bergylta)","volume":"561","author":"Malzahn","year":"2022","journal-title":"Aquaculture"},{"key":"10.1016\/j.cryobiol.2025.105571_bib25","doi-asserted-by":"crossref","first-page":"114","DOI":"10.1016\/j.aquaculture.2008.10.004","article-title":"Viability of subitaneous eggs of the copepod, Acartia tonsa (Dana), following exposure to various cryoprotectants and hypersaline water","volume":"287","author":"Ohs","year":"2009","journal-title":"Aquaculture"},{"key":"10.1016\/j.cryobiol.2025.105571_bib26","first-page":"146","article-title":"Temperature, humidity and inoculative freezing are all signals for glucose accumulation in the freeze tolerant earthworm Dendrobaena octaedra","volume":"A","author":"Overgaard","year":"2007","journal-title":"Comparative Biochemistry and Physiology A-molecular & Integrative Physiology - COMP BIOCHEM PHYSIOL PT"},{"key":"10.1016\/j.cryobiol.2025.105571_bib27","doi-asserted-by":"crossref","first-page":"187","DOI":"10.1146\/annurev-physiol-022516-034142","article-title":"The integrative physiology of insect chill tolerance","volume":"79","author":"Overgaard","year":"2017","journal-title":"Annu. Rev. Physiol."},{"key":"10.1016\/j.cryobiol.2025.105571_bib28","first-page":"2732","article-title":"Soil salinity increases survival of freezing in the enchytraeid Enchytraeus albidus","volume":"216","author":"Patricio Silva","year":"2013","journal-title":"J. Exp. Biol."},{"issue":"Suppl 5","key":"10.1016\/j.cryobiol.2025.105571_bib29","doi-asserted-by":"crossref","first-page":"26","DOI":"10.1093\/humrep\/15.suppl_5.26","article-title":"Aspects of natural cold tolerance in ectothermic animals","volume":"15","author":"Raml\u00f8v","year":"2000","journal-title":"Hum. Reprod."},{"key":"10.1016\/j.cryobiol.2025.105571_bib30","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1002\/jcp.1030420403","article-title":"Studies on the physiology of frozen plants and animals in the arctic","volume":"42","author":"Scholander","year":"1953","journal-title":"J. Cell. Comp. Physiol."},{"key":"10.1016\/j.cryobiol.2025.105571_bib31","doi-asserted-by":"crossref","first-page":"1287","DOI":"10.1242\/jeb.00880","article-title":"Metabolism of the sub-antarctic caterpillar Pringleophaga marioni during cooling, freezing and thawing","volume":"207","author":"Sinclair","year":"2004","journal-title":"J. Exp. Biol."},{"key":"10.1016\/j.cryobiol.2025.105571_bib32","doi-asserted-by":"crossref","first-page":"286","DOI":"10.1016\/j.cryobiol.2008.09.010","article-title":"Freeze tolerance and accumulation of cryoprotectants in the enchytraeid Enchytraeus albidus (Oligochaeta) from Greenland and Europe","volume":"57","author":"Slotsbo","year":"2008","journal-title":"Cryobiology"},{"key":"10.1016\/j.cryobiol.2025.105571_bib33","series-title":"Biochemical Adaptation: Response to Environmental Challenges from Life's Origins to the Anthropocene","author":"Somero","year":"2017"},{"key":"10.1016\/j.cryobiol.2025.105571_bib34","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1152\/physrev.1988.68.1.27","article-title":"Freeze tolerance in animals","volume":"68","author":"Storey","year":"1988","journal-title":"Physiol. Rev."},{"key":"10.1016\/j.cryobiol.2025.105571_bib35","first-page":"365","article-title":"NATURAL FREEZING SURVIVAL IN ANIMALS, annual review of ecology","volume":"27","author":"Storey","year":"1996","journal-title":"Evolution, and Systematics"},{"key":"10.1016\/j.cryobiol.2025.105571_bib36","doi-asserted-by":"crossref","DOI":"10.1242\/jeb.249675","article-title":"Gain of thermal tolerance through acclimation is quicker than the loss by de-acclimation in the freeze-tolerant potworm, Enchytraeus albidus","volume":"228","author":"Touzot","year":"2025","journal-title":"J. Exp. Biol."},{"key":"10.1016\/j.cryobiol.2025.105571_bib37","doi-asserted-by":"crossref","first-page":"1891","DOI":"10.1111\/brv.12425","article-title":"Mechanisms underlying insect freeze tolerance","volume":"93","author":"Toxopeus","year":"2018","journal-title":"Biol. Rev. Camb. Phil. Soc."},{"key":"10.1016\/j.cryobiol.2025.105571_bib38","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/S1164-5563(99)80001-6","article-title":"Respiration activity of Dendrobaena octaedra (Lumbricidae) under constant and diurnally fluctuating temperature regimes in laboratory microcosms","volume":"34","author":"Uvarov","year":"1998","journal-title":"Eur. J. Soil Biol."},{"key":"10.1016\/j.cryobiol.2025.105571_bib39","doi-asserted-by":"crossref","first-page":"461","DOI":"10.1111\/jwas.12210","article-title":"The effects of live and artificial diets on feeding performance of cultured Winter flounder, Pseudopleuronectes americanus, in the wild: survival, feeding, growth, and nucleic acid analyses","volume":"46","author":"Walsh","year":"2015","journal-title":"J. World Aquacult. Soc."},{"key":"10.1016\/j.cryobiol.2025.105571_bib40","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1303\/aez.2006.15","article-title":"Anhydrobiosis in invertebrates","volume":"41","author":"Watanabe","year":"2006","journal-title":"Appl. Entomol. Zool"},{"key":"10.1016\/j.cryobiol.2025.105571_bib41","doi-asserted-by":"crossref","first-page":"799","DOI":"10.1152\/physrev.1985.65.4.799","article-title":"Physiology of cold tolerance in insects","volume":"65","author":"Zachariassen","year":"1985","journal-title":"Physiol. Rev."},{"key":"10.1016\/j.cryobiol.2025.105571_bib42","doi-asserted-by":"crossref","first-page":"180","DOI":"10.1016\/0011-2240(82)90139-0","article-title":"A method for quantitative determination of ice nucleating agents in insect hemolymph","volume":"19","author":"Zachariassen","year":"1982","journal-title":"Cryobiology"},{"key":"10.1016\/j.cryobiol.2025.105571_bib43","doi-asserted-by":"crossref","first-page":"126","DOI":"10.1016\/j.cryobiol.2004.01.004","article-title":"Inorganic ions in cold-hardiness","volume":"48","author":"Zachariassen","year":"2004","journal-title":"Cryobiology"}],"container-title":["Cryobiology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0011224025003773?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0011224025003773?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,3,19]],"date-time":"2026-03-19T11:08:08Z","timestamp":1773918488000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0011224025003773"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,3]]},"references-count":43,"alternative-id":["S0011224025003773"],"URL":"https:\/\/doi.org\/10.1016\/j.cryobiol.2025.105571","relation":{},"ISSN":["0011-2240"],"issn-type":[{"value":"0011-2240","type":"print"}],"subject":[],"published":{"date-parts":[[2026,3]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Temperature-metabolism relationships above and below the body fluid freezing point in the freeze-tolerant white worm, Enchytraeus albidus","name":"articletitle","label":"Article Title"},{"value":"Cryobiology","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.cryobiol.2025.105571","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 Society for Cryobiology.","name":"copyright","label":"Copyright"}],"article-number":"105571"}}