{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,1,9]],"date-time":"2026-01-09T23:45:01Z","timestamp":1768002301904,"version":"3.49.0"},"reference-count":91,"publisher":"Public Library of Science (PLoS)","issue":"2","license":[{"start":{"date-parts":[[2025,2,13]],"date-time":"2025-02-13T00:00:00Z","timestamp":1739404800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/"}],"funder":[{"DOI":"10.13039\/100015704","name":"Bonneville Power Administration","doi-asserted-by":"publisher","award":["project 1993-056-00"],"award-info":[{"award-number":["project 1993-056-00"]}],"id":[{"id":"10.13039\/100015704","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.plosone.org"],"crossmark-restriction":false},"short-container-title":["PLoS ONE"],"abstract":"<jats:p>Most steelhead trout hatcheries increase growth rate during rearing to produce and release yearling smolts for harvest augmentation, but natural steelhead exhibit variable age of smoltification, so this common rearing practice may not be ideal for programs focused on recovering imperiled wild stocks; therefore, it is important to investigate and compare alternative hatchery rearing methods that promote life history diversity. Over six consecutive years, the Winthrop National Fish Hatchery on the Methow River, WA reared and released paired groups of age-1 (S1) and age-2 (S2) steelhead smolts. To understand how the two rearing methods affected developmental ontogeny and life-history, fish were sampled prior to hatchery release for factors associated with smoltification (size, gill Na+\/K+ ATPase activity, and a qualitative smolt phenotype) and sexual maturation (sex, pituitary and testis mRNA transcripts, gonadosomatic index, and plasma 11-ketotestosterone). Our objectives were to quantify levels of smoltification and male maturation during hatchery rearing, combine metrics to estimate residualism (failure to migrate upon release), and compare the treatments by sex. Overall, S2 rearing produced 7.8% more smolts and 44-fold (4.4 vs. 0.1%) more precociously mature males than S1 rearing. Conversely, S1 rearing produced 31.6% more residuals than S2 rearing. While the proportion of total male residuals was comparable between treatments, the S1 treatment produced approximately five-fold more female residuals (20.6 vs. 4.2%). Because residuals contribute minimally to adult returns and the number of returning adult females is critical to the success of salmonid supplementation efforts, developing rearing techniques that maximize migration in females is a management priority. Physiological assessments are useful for characterizing and quantifying the effects and risks of different hatchery rearing regimes on steelhead life-history, in addition to providing sex-specific guidance to inform and optimize conservation management goals in supplementation programs.<\/jats:p>","DOI":"10.1371\/journal.pone.0315016","type":"journal-article","created":{"date-parts":[[2025,2,13]],"date-time":"2025-02-13T18:29:13Z","timestamp":1739471353000},"page":"e0315016","update-policy":"https:\/\/doi.org\/10.1371\/journal.pone.corrections_policy","source":"Crossref","is-referenced-by-count":1,"title":["Age at release affects developmental physiology and sex-specific phenotypic diversity of hatchery steelhead trout (Oncorhynchus mykiss)"],"prefix":"10.1371","volume":"20","author":[{"ORCID":"https:\/\/orcid.org\/0009-0009-0557-7865","authenticated-orcid":true,"given":"Mollie A.","family":"Middleton","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5862-1665","authenticated-orcid":true,"given":"Donald A.","family":"Larsen","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Christopher P.","family":"Tatara","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Barry A.","family":"Berejikian","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Chris R.","family":"Pasley","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Jon T.","family":"Dickey","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Penny","family":"Swanson","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"340","published-online":{"date-parts":[[2025,2,13]]},"reference":[{"key":"pone.0315016.ref001","first-page":"1","article-title":"The life histories of the steelhead rainbow trout (Salmo gairdneri gairdneri) and silver salmon (Oncorhynchus kisutch), with special reference to Waddell Creek, California, and recommendations regarding their management","volume":"98","author":"L Shapovalov","year":"1954","journal-title":"Calif Dep Fish Game Fish Bull"},{"issue":"5","key":"pone.0315016.ref002","doi-asserted-by":"crossref","first-page":"1035","DOI":"10.1111\/1365-2656.12212","article-title":"Life-history diversity and its importance to population stability and persistence of a migratory fish: steelhead in two large North American watersheds","volume":"83","author":"JW Moore","year":"2014","journal-title":"J Anim Ecol"},{"key":"pone.0315016.ref003","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1139\/cjfas-2014-0192","article-title":"Anadromy and residency in steelhead and rainbow trout (Oncorhynchus mykiss): a review of the processes and patterns","volume":"72","author":"NW Kendall","year":"2015","journal-title":"Can J Fish Aquat Sci"},{"key":"pone.0315016.ref004","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1023\/A:1023355114612","article-title":"Genetic comparison between sympatric anadromous steelhead and freshwater resident rainbow trout in British Columbia, Canada","volume":"4","author":"MF Docker","year":"2003","journal-title":"Conserv Genet"},{"key":"pone.0315016.ref005","doi-asserted-by":"crossref","first-page":"539","DOI":"10.1111\/j.1600-0633.2007.00248.x","article-title":"Population structure and partial anadromy in Oncorhynchus mykiss from Kamchatka: relevance for conservation strategies around the Pacific Rim","volume":"16","author":"MV McPhee","year":"2007","journal-title":"Ecol Freshw Fish"},{"key":"pone.0315016.ref006","doi-asserted-by":"crossref","first-page":"421","DOI":"10.1007\/s11160-005-0802-5","article-title":"Anadromy and the marine migrations of Pacific salmon and trout: Rounsefell revisited","volume":"14","author":"TP Quinn","year":"2004","journal-title":"Rev Fish Biol Fish"},{"issue":"5","key":"pone.0315016.ref007","doi-asserted-by":"crossref","first-page":"701","DOI":"10.1139\/cjfas-2012-0457","article-title":"Resident rainbow trout produce anadromous offspring in a large interior watershed","volume":"70","author":"II Courter","year":"2013","journal-title":"Can J Fish Aquat Sci"},{"key":"pone.0315016.ref008","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1577\/1548-8675(1994)014<0077:AALOSS>2.3.CO;2","article-title":"Age and length of steelhead smolts from the mid-Columbia River basin, Washington","volume":"14","author":"CM Peven","year":"1994","journal-title":"N Am J Fish Manag"},{"key":"pone.0315016.ref009","first-page":"275","volume-title":"Fish Physiology,The physiology of developing fish, Part B, Viviparity and posthatching juveniles","author":"WS Hoar","year":"1988"},{"issue":"1","key":"pone.0315016.ref010","doi-asserted-by":"crossref","first-page":"84","DOI":"10.1111\/j.1365-294X.2007.03510.x","article-title":"Evolutionary responses by native species to major anthropogenic changes to their ecosystems: Pacific salmon in the Columbia River hydropower system","volume":"17","author":"RS Waples","year":"2008","journal-title":"Mol Ecol"},{"key":"pone.0315016.ref011","doi-asserted-by":"crossref","first-page":"609","DOI":"10.1038\/nature09060","article-title":"Population diversity and the portfolio effect in an exploited species","volume":"465","author":"DE Schindler","year":"2010","journal-title":"Nature"},{"key":"pone.0315016.ref012","doi-asserted-by":"crossref","first-page":"222","DOI":"10.1111\/j.1752-4571.2008.00025.x","article-title":"A review of quantitative genetic components of fitness in salmonids: implications for adaptation to future change","volume":"1","author":"SM Carlson","year":"2008","journal-title":"Evol Appl"},{"key":"pone.0315016.ref013","unstructured":"Ruzycki JR, Clarke LR, Flesher MW, Carmichael RW, Eddy DL. Performance of progeny from steelhead and rainbow trout crosses. USFWS (US Fish and Wildlife Service) Lower Snake River compensation plan, final report. Contract number 1411-08-J009 (2007\u20132008). ODFW (Oregon Department of Fish and Wildlife), Salem, Oregon. 2009."},{"key":"pone.0315016.ref014","doi-asserted-by":"crossref","first-page":"772","DOI":"10.1080\/02755947.2012.686953","article-title":"Should I stay or should I go? The influence of genetic origin on emigration behavior and physiology of resident and anadromous juvenile Oncorhynchus mykiss","volume":"32","author":"SA Hayes","year":"2012","journal-title":"N Am J Fish Manag"},{"issue":"2","key":"pone.0315016.ref015","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1080\/00028487.2013.862181","article-title":"Maternal control over offspring life history in a partially anadromous species, Oncorhynchus mykiss","volume":"143","author":"BA Berejikian","year":"2014","journal-title":"Trans Am Fish Soc"},{"issue":"5","key":"pone.0315016.ref016","doi-asserted-by":"crossref","first-page":"1263","DOI":"10.1577\/T09-146.1","article-title":"Smolt transformation in two California steelhead populations: effects of temporal variability in growth","volume":"139","author":"MP Beakes","year":"2010","journal-title":"Trans Am Fish Soc"},{"key":"pone.0315016.ref017","doi-asserted-by":"crossref","first-page":"343","DOI":"10.1007\/s10641-011-9921-0","article-title":"Individual condition and stream temperature influence early maturation of rainbow and steelhead trout, Oncorhynchus mykiss","volume":"93","author":"JR McMillan","year":"2012","journal-title":"Environ Biol Fish"},{"issue":"2","key":"pone.0315016.ref018","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1080\/00028487.2013.849617","article-title":"Growth-mediated life history traits of steelhead reveal phenotypic divergence and plastic response to temperature","volume":"143","author":"K Doctor","year":"2014","journal-title":"Trans Am Fish Soc"},{"key":"pone.0315016.ref019","doi-asserted-by":"crossref","first-page":"689","DOI":"10.1007\/s11160-014-9344-z","article-title":"Ecological and evolutionary patterns of freshwater maturation in Pacific and Atlantic salmonines","volume":"24","author":"MR Sloat","year":"2014","journal-title":"Rev Fish Biol Fish"},{"key":"pone.0315016.ref020","doi-asserted-by":"crossref","first-page":"181","DOI":"10.1098\/rspb.1990.0122","article-title":"A polygenic model for the evolution and maintenance of conditional strategies","volume":"242","author":"WN Hazel","year":"1990","journal-title":"Proc R Soc Lond B Biol Sci"},{"key":"pone.0315016.ref021","doi-asserted-by":"crossref","first-page":"421","DOI":"10.1038\/hdy.2010.166","article-title":"Old wine in new bottles: reaction norms in salmonid fishes","volume":"106","author":"JA Hutchings","year":"2011","journal-title":"Heredity"},{"key":"pone.0315016.ref022","volume-title":"Salmon without rivers: a history of the Pacific salmon crisis.:","author":"J. Lichatowich","year":"1999","edition":"1"},{"key":"pone.0315016.ref023","first-page":"383","volume-title":"Ecological and genetic implications of aquaculture activitiesMethods and technologies in fish biology and fisheries","author":"RS Waples","year":"2007"},{"key":"pone.0315016.ref024","doi-asserted-by":"crossref","first-page":"1018","DOI":"10.1139\/f03-087","article-title":"Interactions between hatchery and wild salmonids in streams: differences in biology and evidence for competition","volume":"60","author":"ED Weber","year":"2003","journal-title":"Can J Fish Aquat Sci"},{"key":"pone.0315016.ref025","doi-asserted-by":"crossref","first-page":"577","DOI":"10.1139\/f04-019","article-title":"Differences in juvenile phenotypes and survival between hatchery stocks and a natural population provide evidence for modified selection due to captive breeding","volume":"61","author":"KE Kostow","year":"2004","journal-title":"Can J Fish Aquat Sci"},{"key":"pone.0315016.ref026","first-page":"61","volume-title":"Advances in marine biology","author":"KA Naish","year":"2007"},{"issue":"4","key":"pone.0315016.ref027","doi-asserted-by":"crossref","first-page":"535","DOI":"10.1111\/j.1752-4571.2008.00036.x","article-title":"How well can captive breeding programs conserve biodiversity? A review of salmonids","volume":"1","author":"DJ Fraser","year":"2008","journal-title":"Evol Appl"},{"issue":"2","key":"pone.0315016.ref028","doi-asserted-by":"crossref","first-page":"342","DOI":"10.1111\/j.1752-4571.2008.00026.x","article-title":"Fitness of hatchery-reared salmonids in the wild","volume":"1","author":"H Araki","year":"2008","journal-title":"Evol Appl"},{"key":"pone.0315016.ref029","doi-asserted-by":"crossref","first-page":"10676","DOI":"10.1038\/ncomms10676","article-title":"A single generation of domestication heritably alters the expression of hundreds of genes","volume":"7","author":"MR Christie","year":"2016","journal-title":"Nat Comm"},{"key":"pone.0315016.ref030","doi-asserted-by":"crossref","first-page":"273","DOI":"10.1139\/cjfas-2016-0175","article-title":"Rearing strategies alter patterns of size-selective mortality and heritable size variation in steelhead trout (Oncorhynchus mykiss)","volume":"74","author":"BA Berejikian","year":"2017","journal-title":"Can J Fish Aquat Sci"},{"key":"pone.0315016.ref031","first-page":"27","volume-title":"Common strategies of anadromous and catadromous fishes, American Fisheries Society, Symposium 1","author":"RG Randall","year":"1987"},{"key":"pone.0315016.ref032","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1007\/s10641-011-9788-0","article-title":"Development of natural growth regimes for hatchery-reared steelhead to reduce residualism, fitness loss, and negative ecological interactions","volume":"94","author":"BA Berejikian","year":"2012","journal-title":"Environ Biol Fish"},{"issue":"4","key":"pone.0315016.ref033","doi-asserted-by":"crossref","first-page":"717","DOI":"10.1577\/1548-8675(2001)021<0717:CORTSD>2.0.CO;2","article-title":"Comparisons of run timing, spatial distribution, and length of wild and newly established hatchery populations of steelhead in Forks Creek, Washington","volume":"21","author":"G Mackey","year":"2001","journal-title":"N Am J Fish Manag"},{"issue":"2","key":"pone.0315016.ref034","doi-asserted-by":"crossref","first-page":"446","DOI":"10.1111\/j.1523-1739.2005.00515.x","article-title":"Nonrandom, size- and timing-biased breeding in a hatchery population of steelhead trout","volume":"19","author":"JE McLean","year":"2005","journal-title":"Conserv Biol"},{"key":"pone.0315016.ref035","doi-asserted-by":"crossref","first-page":"905","DOI":"10.1080\/02755947.2012.711269","article-title":"Residualization of hatchery steelhead: a meta-analysis of hatchery practices","volume":"32","author":"SJ Hausch","year":"2012","journal-title":"N Am J Fish Manag"},{"issue":"3","key":"pone.0315016.ref036","doi-asserted-by":"crossref","first-page":"468","DOI":"10.1002\/nafm.10284","article-title":"Age at release, size, and maturation status influence residualism in hatchery steelhead","volume":"39","author":"CP Tatara","year":"2019","journal-title":"N Am J Fish Manag"},{"key":"pone.0315016.ref037","volume-title":"Effect of steelhead smolt size on residualism and adult return rates. USFWS (US Fish and Wildlife Service) Lower Snake River compensation plan","author":"FE Partridge","year":"1986"},{"key":"pone.0315016.ref038","doi-asserted-by":"crossref","first-page":"1355","DOI":"10.1577\/M05-220.1","article-title":"Growth modulation during juvenile rearing can reduce rates of residualism in the progeny of wild steelhead broodstock.","volume":"27","author":"CS Sharpe","year":"2007","journal-title":"N Am J Fish Manag"},{"key":"pone.0315016.ref039","doi-asserted-by":"crossref","first-page":"230","DOI":"10.1577\/1548-8659(1997)126<0230:EORHRS>2.3.CO;2","article-title":"Effects of residual hatchery-reared steelhead on growth of wild rainbow trout and spring chinook salmon","volume":"126","author":"GA McMichael","year":"1997","journal-title":"Trans Am Fish Soc"},{"key":"pone.0315016.ref040","doi-asserted-by":"crossref","first-page":"948","DOI":"10.1577\/1548-8675(1999)019<0948:BIAHRS>2.0.CO;2","article-title":"Behavioral interactions among hatchery-reared steelhead smolts and wild Oncorhynchus mykiss in natural stream","volume":"19","author":"GA McMichael","year":"1999","journal-title":"N Am J Fish Manag"},{"key":"pone.0315016.ref041","doi-asserted-by":"crossref","first-page":"943","DOI":"10.1577\/1548-8675(2001)021<0943:UMORHS>2.0.CO;2","article-title":"Upstream movement of residual hatchery steelhead into areas containing bull trout and cutthroat trout","volume":"21","author":"GA McMichael","year":"2001","journal-title":"N Am J Fish Manag"},{"issue":"s1","key":"pone.0315016.ref042","doi-asserted-by":"crossref","first-page":"122","DOI":"10.1111\/j.0022-1112.2004.00562.x","article-title":"Implications of domestication and rearing conditions for the behavior of cultivated fishes","volume":"65","author":"FA Huntingford","year":"2004","journal-title":"J Fish Biol"},{"key":"pone.0315016.ref043","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1007\/s10641-011-9906-z","article-title":"Mechanisms influencing competition between hatchery and wild juvenile anadromous Pacific salmonids in fresh water and their relative competitive abilities","volume":"94","author":"CP Tatara","year":"2012","journal-title":"Environ Biol Fish"},{"key":"pone.0315016.ref044","doi-asserted-by":"crossref","first-page":"1100","DOI":"10.1080\/02755947.2013.824938","article-title":"Ecological and demographic costs of releasing nonmigratory juvenile hatchery steelhead in the Methow River, Washingto","volume":"33","author":"CG Snow","year":"2013","journal-title":"n. N Am J Fish Manag"},{"key":"pone.0315016.ref045","first-page":"199","volume-title":"Fish Physiology, Vol. 32, Euryhaline fishes","author":"SD McCormick","year":"2013"},{"key":"pone.0315016.ref046","first-page":"211","article-title":"Preparatory physiological adaptations for marine life of salmonids: osmoregulation, growth, and metabolism","volume":"1","author":"SD McCormick","year":"1987","journal-title":"Am Fish Soc Symp"},{"key":"pone.0315016.ref047","doi-asserted-by":"crossref","first-page":"285","DOI":"10.3354\/meps335285","article-title":"Maturation response of salmonids to changing developmental opportunities","volume":"335","author":"JE Thorpe","year":"2007","journal-title":"Mar Ecol Prog Ser"},{"issue":"4","key":"pone.0315016.ref048","doi-asserted-by":"crossref","first-page":"663","DOI":"10.1080\/00028487.2017.1296492","article-title":"Use of morphological and physiological indices to characterize life history diversity in juvenile hatchery winter-run steelhead","volume":"146","author":"DA Larsen","year":"2017","journal-title":"Trans Am Fish Soc"},{"key":"pone.0315016.ref049","doi-asserted-by":"crossref","first-page":"30","DOI":"10.1016\/j.ygcen.2019.05.010","article-title":"Evaluation of endocrine and transcriptomic markers of male maturation in winter-run steelhead trout (Oncorhynchus mykiss)","volume":"281","author":"MA Middleton","year":"2019","journal-title":"Gen Comp Endocrinol"},{"issue":"5","key":"pone.0315016.ref050","doi-asserted-by":"crossref","first-page":"506","DOI":"10.1139\/f03-042","article-title":"Energy allocation among developmental stages, age groups, and types of Atlantic salmon (Salmo salar) spawners","volume":"60","author":"N Jonsson","year":"2003","journal-title":"Can J Fish Aquat Sci"},{"issue":"17","key":"pone.0315016.ref051","doi-asserted-by":"crossref","first-page":"e13809","DOI":"10.14814\/phy2.13809","article-title":"Sexual maturation and smoltification in domesticated Atlantic salmon (Salmo salar L.)\u2013is there a developmental conflict?","volume":"6","author":"PG Fjelldal","year":"2018","journal-title":"Physiol Rep"},{"key":"pone.0315016.ref052","doi-asserted-by":"crossref","first-page":"390","DOI":"10.1016\/j.ygcen.2009.02.013","article-title":"Spermatogenesis in fish","volume":"165","author":"RW Schulz","year":"2010","journal-title":"Gen Comp Endocrinol"},{"issue":"9","key":"pone.0315016.ref053","doi-asserted-by":"crossref","first-page":"3257","DOI":"10.1210\/en.2010-1469","article-title":"Proteolytically activated, recombinant anti-Mullerian hormone inhibits androgen secretion, proliferation, and differentiation of spermatogonia in adult zebrafish testis organ cultures","volume":"152","author":"KS Skaar","year":"2011","journal-title":"Endocrinology"},{"issue":"10","key":"pone.0315016.ref054","doi-asserted-by":"crossref","first-page":"3804","DOI":"10.1210\/en.2015-1157","article-title":"Fsh stimulates spermatogonial proliferation and differentiation in zebrafish via Igf3","volume":"156","author":"RH N\u00f3brega","year":"2015","journal-title":"Endocrinology"},{"key":"pone.0315016.ref055","doi-asserted-by":"crossref","first-page":"413","DOI":"10.1006\/gcen.1998.7222","article-title":"Growth hormone (GH) and gonadotropin subunit gene expression and pituitary and plasma changes during spermatogenesis and oogenesis in rainbow trout (Oncorhynchus mykiss)","volume":"113","author":"JM Gomez","year":"1999","journal-title":"Gen Comp Endocrinol"},{"key":"pone.0315016.ref056","unstructured":"USFWS (US Fish and Wildlife Service). Hatchery and genetic management plan (HGMP) for summer steelhead (Oncorhynchus mykiss) at Winthrop National Fish Hatchery. USFWS, Leavenworth Fisheries Complex, Leavenworth, Washington. 2009."},{"key":"pone.0315016.ref057","unstructured":"Bjornn TC, Ringe RR. Evaluation of two-year rearing program for steelhead trout at Dworshak NFH. Technical Report 84\u20137 to US Department of the Army, Corps of Engineers, Walla Walla District, WA. Contract number DACW 68-79-C-0044. Idaho Cooperative Fish and Wildlife Research Unit, University of Idaho, Moscow, ID. 1984."},{"key":"pone.0315016.ref058","doi-asserted-by":"crossref","first-page":"700","DOI":"10.1080\/02755947.2017.1317676","article-title":"Age and method of release affect migratory performance of hatchery steelhead","volume":"37","author":"CP Tatara","year":"2017","journal-title":"N Am J Fish Manag"},{"key":"pone.0315016.ref059","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/0044-8486(82)90004-7","article-title":"Morphological indices of development progress in the parr-smolt coho salmon, Oncorhynchus kisutch","volume":"28","author":"A Gorbman","year":"1982","journal-title":"Aquaculture"},{"key":"pone.0315016.ref060","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1139\/f82-027","article-title":"A simplified preparation for adenosine triphosphate determination in gill tissue","volume":"39","author":"WS Zaugg","year":"1982","journal-title":"Can J Fish Aquat Sci"},{"key":"pone.0315016.ref061","doi-asserted-by":"crossref","first-page":"656","DOI":"10.1139\/f93-075","article-title":"Methods for nonlethal gill biopsy and measurement of Na+, K+ -ATPase activity","volume":"50","author":"SD McCormick","year":"1993","journal-title":"Can J Fish Aquat Sci"},{"key":"pone.0315016.ref062","doi-asserted-by":"crossref","first-page":"401","DOI":"10.1016\/j.ygcen.2003.10.010","article-title":"A quantitative real-time RT-PCR assay for salmon IGF-I mRNA, and its application in the study of GH regulation of IGF-I gene expression in primary culture of salmon hepatocytes","volume":"135","author":"AL Pierce","year":"2004","journal-title":"Gen Comp Endocrinol"},{"key":"pone.0315016.ref063","first-page":"195","article-title":"Sexual steroids during puberty in male African catfish (Clarius gariepinus): serum levels and gonadotropin-stimulated testicular secretion in vitro","volume":"164","author":"RW Schulz","year":"1994","journal-title":"Comp Biochem Physiol Part B"},{"key":"pone.0315016.ref064","first-page":"229","article-title":"Enzyme immunoassay for 11-ketotestosterone using acetylcholinesterase as label: application to the measurement of 11-ketotestosterone in plasma of Siberian sturgeon","volume":"108C","author":"B Cuisset","year":"1994","journal-title":"Comp Biochem Physiol Part C"},{"issue":"6","key":"pone.0315016.ref065","doi-asserted-by":"crossref","first-page":"1","DOI":"10.18637\/jss.v032.i06","article-title":"mixtools: An R package for analyzing finite mixture models","volume":"32","author":"T Benaglia","year":"2009","journal-title":"J Stat Soft"},{"key":"pone.0315016.ref066","volume-title":"An {R} companion to applied regression","author":"J Fox","edition":"3"},{"key":"pone.0315016.ref067","doi-asserted-by":"crossref","article-title":"agricolae: Statistical procedures for agricultural research","author":"F. de Mendiburu","DOI":"10.32614\/CRAN.package.agricolae"},{"key":"pone.0315016.ref068","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1080\/10641260490280313","article-title":"Population structure of Columbia River basin chinook salmon and steelhead trout","volume":"12","author":"EL Brannon","year":"2004","journal-title":"Rev Fish Sci"},{"issue":"2","key":"pone.0315016.ref069","doi-asserted-by":"crossref","first-page":"375","DOI":"10.1139\/f07-003","article-title":"The growing degree-day and fish size-at-age: the overlooked metric","volume":"64","author":"AB Neuheimer","year":"2007","journal-title":"Can J Aquat Fish Sci"},{"key":"pone.0315016.ref070","doi-asserted-by":"crossref","first-page":"491","DOI":"10.1139\/cjfas-2013-0366","article-title":"Individual condition, standard metabolic rate, and rearing temperature influence steelhead and rainbow trout (Oncorhynchus mykiss) life histories","volume":"71","author":"MR Sloat","year":"2014","journal-title":"Can J Fish Aquat Sci"},{"issue":"s1","key":"pone.0315016.ref071","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1139\/d98-011","article-title":"Movement, migration, and smolting of Atlantic salmon (Salmo salar)","volume":"55","author":"SD McCormick","year":"1998","journal-title":"Can J Fish Aquat Sci"},{"issue":"4","key":"pone.0315016.ref072","doi-asserted-by":"crossref","first-page":"310","DOI":"10.1577\/1548-8640(1997)059<0310:EOSLAR>2.3.CO;2","article-title":"Effect of smolt length at release on adult returns of hatchery-reared winter steelhead","volume":"59","author":"JM Tipping","year":"1997","journal-title":"Prog Fish-Cult"},{"issue":"1","key":"pone.0315016.ref073","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1080\/15222055.2013.837127","article-title":"Hatchery steelhead smolt release size effects on adult production and straying","volume":"76","author":"LR Clarke","year":"2014","journal-title":"N Am J Aquac"},{"key":"pone.0315016.ref074","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1007\/978-94-007-0983-6_27","volume-title":"Developments in environmental biology of fishesGenetics of subpolar fish and invertebrates","author":"TR Seamons","year":"2004"},{"key":"pone.0315016.ref075","doi-asserted-by":"crossref","first-page":"1000","DOI":"10.1139\/cjfas-2019-0255","article-title":"Duration in captivity affects competitive ability and breeding success of male but not female steelhead trout (Oncorhynchus mykiss)","volume":"77","author":"BA Berejikian","year":"2020","journal-title":"Can J Fish Aquat Sci"},{"issue":"3","key":"pone.0315016.ref076","doi-asserted-by":"crossref","first-page":"343","DOI":"10.1080\/15222055.2014.999846","article-title":"Fish hatchery genetic management techniques: integrating theory with implementation","volume":"77","author":"KM Fisch","year":"2015","journal-title":"N Am J Aquac"},{"key":"pone.0315016.ref077","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1007\/s11160-019-09551-5","article-title":"Advancing mate choice studies in salmonids","volume":"29","author":"HL Auld","year":"2019","journal-title":"Rev Fish Biol Fish"},{"issue":"3","key":"pone.0315016.ref078","doi-asserted-by":"crossref","first-page":"209","DOI":"10.1016\/0044-8486(84)90188-1","article-title":"Indices of parr-smolt transformation in juvenile steelhead trout (Salmo gairdneri) undergoing volitional release at Cole Rivers Hatchery","volume":"40","author":"RD Ewing","year":"1984","journal-title":"Oregon. Aquaculture"},{"issue":"2","key":"pone.0315016.ref079","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1577\/A08-001.1","article-title":"Juvenile steelhead release strategies: a comparison of volitional- and forced-release practices","volume":"71","author":"WL Gale","year":"2009","journal-title":"N Am J Aquac"},{"key":"pone.0315016.ref080","doi-asserted-by":"crossref","first-page":"339","DOI":"10.1016\/0044-8486(89)90420-1","article-title":"Migratory behavior of underyearling Oncorhynchus tshawytscha and survival to adulthood as related to prerelease gill (Na+ -K+)-ATPase development","volume":"82","author":"WS Zaugg","year":"1989","journal-title":"Aquaculture"},{"key":"pone.0315016.ref081","doi-asserted-by":"crossref","first-page":"564","DOI":"10.1577\/T08-209.1","article-title":"Examining the conflict between smolting and precocious male maturation in spring (stream-type) Chinook salmon","volume":"139","author":"DA Larsen","year":"2010","journal-title":"Trans Am Fish Soc"},{"key":"pone.0315016.ref082","unstructured":"Tatara CP, Swanson P, Dickey J, Young G, Cooper M, Humling M, et al. Advance Hatchery Reform Research. Annual report to BPA (Bonneville Power Administration). Project number 1993-056-00 (2018). BPA, Portland, OR. 2019."},{"issue":"5","key":"pone.0315016.ref083","doi-asserted-by":"crossref","first-page":"621","DOI":"10.1098\/rsbl.2009.0315","article-title":"Carry-over effect of captive breeding reduces reproductive fitness of wild-born descendants in the wild","volume":"5","author":"H Araki","year":"2009","journal-title":"Biol Lett"},{"issue":"10","key":"pone.0315016.ref084","doi-asserted-by":"crossref","first-page":"e0164801","DOI":"10.1371\/journal.pone.0164801","article-title":"Broodstock history strongly influences natural spawning success in hatchery steelhead (Oncorhynchus mykiss)","volume":"11","author":"MJ Ford","year":"2016","journal-title":"PLoS ONE"},{"issue":"7","key":"pone.0315016.ref085","doi-asserted-by":"crossref","first-page":"1090","DOI":"10.1111\/eva.12089","article-title":"Assessing strategies to minimize unintended fitness consequences of aquaculture on wild populations","volume":"6","author":"ML Baskett","year":"2013","journal-title":"Evol Appl"},{"issue":"10","key":"pone.0315016.ref086","doi-asserted-by":"crossref","first-page":"956","DOI":"10.1111\/eva.12331","article-title":"Effectiveness of managed gene flow in reducing genetic divergence associated with captive breeding","volume":"8","author":"CD Waters","year":"2015","journal-title":"Evol Appl"},{"issue":"6","key":"pone.0315016.ref087","doi-asserted-by":"crossref","first-page":"284","DOI":"10.1002\/fsh.10084","article-title":"Why keep hatcheries? Weighing the economic cost and value of fish production for public use and public trust purposes","volume":"43","author":"JT Trushenski","year":"2018","journal-title":"Fisheries"},{"issue":"1","key":"pone.0315016.ref088","doi-asserted-by":"crossref","first-page":"208","DOI":"10.1080\/00028487.2012.728162","article-title":"Potential effects of changes in temperature and food resources on life history trajectories of juvenile Oncorhynchus mykiss","volume":"142","author":"JR Benjamin","year":"2013","journal-title":"Trans Am Fish Soc"},{"issue":"3","key":"pone.0315016.ref089","doi-asserted-by":"crossref","first-page":"221","DOI":"10.1111\/j.1752-4571.2009.00103.x","article-title":"State-dependent life history models in a changing (and regulated) environment: steelhead in the California Central Valley","volume":"3","author":"WH Satterthwaite","year":"2010","journal-title":"Evol Appl"},{"key":"pone.0315016.ref090","doi-asserted-by":"crossref","first-page":"413","DOI":"10.1007\/s10584-012-0627-x","article-title":"The effects of increased stream temperatures on juvenile steelhead growth in the Yakima River Basin based on projected climate change scenarios","volume":"124","author":"JM Hardiman","year":"2014","journal-title":"Clim Change"},{"issue":"3","key":"pone.0315016.ref091","doi-asserted-by":"crossref","first-page":"373","DOI":"10.1111\/eff.12151","article-title":"Interactions between body mass and water temperature cause energetic bottlenecks in juvenile steelhead","volume":"24","author":"KM Myrvold","year":"2014","journal-title":"Ecol Freshw Fish"}],"container-title":["PLOS ONE"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0315016","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,2,13]],"date-time":"2025-02-13T18:30:10Z","timestamp":1739471410000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0315016"}},"subtitle":[],"editor":[{"given":"Tzong-Yueh","family":"Chen","sequence":"first","affiliation":[],"role":[{"role":"editor","vocabulary":"crossref"}]}],"short-title":[],"issued":{"date-parts":[[2025,2,13]]},"references-count":91,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2025,2,13]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pone.0315016","relation":{},"ISSN":["1932-6203"],"issn-type":[{"value":"1932-6203","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,2,13]]}}}