Attenuation of the cortisol response to stress in female rainbow trout chronically exposed to dietary selenomethionine
► Trout exposed to Se-Met had greater concentration of cortisol compared to controls. ► Transcript abundance of mc2r was greater in trout exposed to Se-Met. ► Trout exposed to Se-Met had a reduced cortisol response to a handling stressor. ► Cortisone concentration was greater in Se-Met exposed trout...
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creator | Wiseman, Steve Thomas, Jith K. McPhee, Landon Hursky, Olesya Raine, Jason C. Pietrock, Michael Giesy, John P. Hecker, Markus Janz, David M. |
description | ► Trout exposed to Se-Met had greater concentration of cortisol compared to controls. ► Transcript abundance of mc2r was greater in trout exposed to Se-Met. ► Trout exposed to Se-Met had a reduced cortisol response to a handling stressor. ► Cortisone concentration was greater in Se-Met exposed trout post-handling stressor.
Selenomethionine (Se-Met) is the major dietary form of selenium (Se). While Se is a required nutrient, it can also influence the physiological stress response because it stimulates greater concentrations of cortisol in blood plasma of exposed fish. However, little is known about the effects of exposure to Se on the ability to cope with a secondary stressor. In the current study, female rainbow trout were exposed to an environmentally relevant dietary concentration (8.47
mg
Se/kg dry mass (dm)) of Se-Met for 126
d, after which time fish were subjected to a 3-min handling stressor and sampled at 2
h and 24
h post-stressor exposure. Concentrations of cortisol, cortisone, glucose, and lactate in blood plasma and concentrations of glycogen and triglycerides in liver and muscle were determined. Abundances of transcripts of proteins involved in corticosteroidogenesis were determined using quantitative RT-PCR. Concentrations of cortisol were significantly greater in blood plasma of trout exposed to Se-Met, relative to control trout sampled prior to the handling stressor. A typical response of cortisol to the handling stressor was observed in the control trout. However, trout exposed to Se-Met were unable to mount a cortisol response to the handling stressor. Concentrations of cortisone, the inactive metabolite of cortisol, were significantly greater following the handling stressor in trout exposed to Se-Met. In trout exposed to Se-Met, transcript abundance of melanocortin 2 receptor (
mc2r) and peripheral benzodiazepine receptor (
pbr) were greater, which is consistent with the conclusion that synthesis of cortisol was greater. However, abundances of transcripts of cytochrome P450 side-chain cleavage (
p450scc) and cytochrome P450 11B1 (
cyp11b1) were not significantly different between controls and Se-Met exposed trout. Exposure to Se-Met affected accumulation and tissue partitioning of glycogen and triglycerides in liver and muscle as concentrations of these energy reserves were greater in muscle, but not liver. Concentrations of glycogen and triglycerides in muscle, but not in liver, were lesser following the handling stressor suggesting that |
doi_str_mv | 10.1016/j.aquatox.2011.09.002 |
format | Article |
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Selenomethionine (Se-Met) is the major dietary form of selenium (Se). While Se is a required nutrient, it can also influence the physiological stress response because it stimulates greater concentrations of cortisol in blood plasma of exposed fish. However, little is known about the effects of exposure to Se on the ability to cope with a secondary stressor. In the current study, female rainbow trout were exposed to an environmentally relevant dietary concentration (8.47
mg
Se/kg dry mass (dm)) of Se-Met for 126
d, after which time fish were subjected to a 3-min handling stressor and sampled at 2
h and 24
h post-stressor exposure. Concentrations of cortisol, cortisone, glucose, and lactate in blood plasma and concentrations of glycogen and triglycerides in liver and muscle were determined. Abundances of transcripts of proteins involved in corticosteroidogenesis were determined using quantitative RT-PCR. Concentrations of cortisol were significantly greater in blood plasma of trout exposed to Se-Met, relative to control trout sampled prior to the handling stressor. A typical response of cortisol to the handling stressor was observed in the control trout. However, trout exposed to Se-Met were unable to mount a cortisol response to the handling stressor. Concentrations of cortisone, the inactive metabolite of cortisol, were significantly greater following the handling stressor in trout exposed to Se-Met. In trout exposed to Se-Met, transcript abundance of melanocortin 2 receptor (
mc2r) and peripheral benzodiazepine receptor (
pbr) were greater, which is consistent with the conclusion that synthesis of cortisol was greater. However, abundances of transcripts of cytochrome P450 side-chain cleavage (
p450scc) and cytochrome P450 11B1 (
cyp11b1) were not significantly different between controls and Se-Met exposed trout. Exposure to Se-Met affected accumulation and tissue partitioning of glycogen and triglycerides in liver and muscle as concentrations of these energy reserves were greater in muscle, but not liver. Concentrations of glycogen and triglycerides in muscle, but not in liver, were lesser following the handling stressor suggesting that the muscle energy reserves are an important source of energy required for recovery from the handling stressor. The results of the study demonstrate that chronic exposure to dietary Se-Met elicits a stress response, but prevents a cortisol response to a secondary handling stressor, most likely due to cortisol inactivation. Moreover, exposure to Se-Met has effects on concentrations of energy reserves that are important for providing the energy necessary to cope with a secondary stressor.</description><identifier>ISSN: 0166-445X</identifier><identifier>EISSN: 1879-1514</identifier><identifier>DOI: 10.1016/j.aquatox.2011.09.002</identifier><identifier>PMID: 21963595</identifier><identifier>CODEN: AQTODG</identifier><language>eng</language><publisher>Amsterdam: Elsevier B.V</publisher><subject>Animal, plant and microbial ecology ; Animals ; Applied ecology ; Biological and medical sciences ; Blood Glucose - drug effects ; Blood Glucose - metabolism ; blood plasma ; chronic exposure ; Corticosteroidogenesis ; Cortisol ; Cortisone ; Cortisone - blood ; cytochrome P-450 ; dietary exposure ; Ecotoxicology, biological effects of pollution ; energy ; Female ; Food Contamination ; Fundamental and applied biological sciences. Psychology ; General aspects ; glucose ; Glycogen - metabolism ; Handling (Psychology) ; Hydrocortisone - blood ; lactation ; liver ; Liver - drug effects ; Liver - metabolism ; MC2R ; Muscles - drug effects ; Muscles - metabolism ; Oncorhynchus mykiss ; Oncorhynchus mykiss - growth & development ; Oncorhynchus mykiss - metabolism ; PBR ; Random Allocation ; Real-Time Polymerase Chain Reaction ; Selenium ; selenomethionine ; Selenomethionine - pharmacokinetics ; Selenomethionine - toxicity ; Stress response ; Stress, Physiological - drug effects ; Stressor ; Toxicity Tests, Chronic ; triacylglycerols ; Triglycerides - metabolism ; trout ; Water Pollutants, Chemical - pharmacokinetics ; Water Pollutants, Chemical - toxicity</subject><ispartof>Aquatic toxicology, 2011-10, Vol.105 (3), p.643-651</ispartof><rights>2011 Elsevier B.V.</rights><rights>2015 INIST-CNRS</rights><rights>Copyright © 2011 Elsevier B.V. All rights reserved.</rights><lds50>peer_reviewed</lds50><woscitedreferencessubscribed>false</woscitedreferencessubscribed><citedby>FETCH-LOGICAL-c450t-b47f76a235658eb2d401bf1f48ed2010450fa6e71d8093bc59c8b3a90a5f93963</citedby><cites>FETCH-LOGICAL-c450t-b47f76a235658eb2d401bf1f48ed2010450fa6e71d8093bc59c8b3a90a5f93963</cites></display><links><openurl>$$Topenurl_article</openurl><openurlfulltext>$$Topenurlfull_article</openurlfulltext><thumbnail>$$Tsyndetics_thumb_exl</thumbnail><linktohtml>$$Uhttps://dx.doi.org/10.1016/j.aquatox.2011.09.002$$EHTML$$P50$$Gelsevier$$H</linktohtml><link.rule.ids>314,778,782,3539,27907,27908,45978</link.rule.ids><backlink>$$Uhttp://pascal-francis.inist.fr/vibad/index.php?action=getRecordDetail&idt=24781931$$DView record in Pascal Francis$$Hfree_for_read</backlink><backlink>$$Uhttps://www.ncbi.nlm.nih.gov/pubmed/21963595$$D View this record in MEDLINE/PubMed$$Hfree_for_read</backlink></links><search><creatorcontrib>Wiseman, Steve</creatorcontrib><creatorcontrib>Thomas, Jith K.</creatorcontrib><creatorcontrib>McPhee, Landon</creatorcontrib><creatorcontrib>Hursky, Olesya</creatorcontrib><creatorcontrib>Raine, Jason C.</creatorcontrib><creatorcontrib>Pietrock, Michael</creatorcontrib><creatorcontrib>Giesy, John P.</creatorcontrib><creatorcontrib>Hecker, Markus</creatorcontrib><creatorcontrib>Janz, David M.</creatorcontrib><title>Attenuation of the cortisol response to stress in female rainbow trout chronically exposed to dietary selenomethionine</title><title>Aquatic toxicology</title><addtitle>Aquat Toxicol</addtitle><description>► Trout exposed to Se-Met had greater concentration of cortisol compared to controls. ► Transcript abundance of mc2r was greater in trout exposed to Se-Met. ► Trout exposed to Se-Met had a reduced cortisol response to a handling stressor. ► Cortisone concentration was greater in Se-Met exposed trout post-handling stressor.
Selenomethionine (Se-Met) is the major dietary form of selenium (Se). While Se is a required nutrient, it can also influence the physiological stress response because it stimulates greater concentrations of cortisol in blood plasma of exposed fish. However, little is known about the effects of exposure to Se on the ability to cope with a secondary stressor. In the current study, female rainbow trout were exposed to an environmentally relevant dietary concentration (8.47
mg
Se/kg dry mass (dm)) of Se-Met for 126
d, after which time fish were subjected to a 3-min handling stressor and sampled at 2
h and 24
h post-stressor exposure. Concentrations of cortisol, cortisone, glucose, and lactate in blood plasma and concentrations of glycogen and triglycerides in liver and muscle were determined. Abundances of transcripts of proteins involved in corticosteroidogenesis were determined using quantitative RT-PCR. Concentrations of cortisol were significantly greater in blood plasma of trout exposed to Se-Met, relative to control trout sampled prior to the handling stressor. A typical response of cortisol to the handling stressor was observed in the control trout. However, trout exposed to Se-Met were unable to mount a cortisol response to the handling stressor. Concentrations of cortisone, the inactive metabolite of cortisol, were significantly greater following the handling stressor in trout exposed to Se-Met. In trout exposed to Se-Met, transcript abundance of melanocortin 2 receptor (
mc2r) and peripheral benzodiazepine receptor (
pbr) were greater, which is consistent with the conclusion that synthesis of cortisol was greater. However, abundances of transcripts of cytochrome P450 side-chain cleavage (
p450scc) and cytochrome P450 11B1 (
cyp11b1) were not significantly different between controls and Se-Met exposed trout. Exposure to Se-Met affected accumulation and tissue partitioning of glycogen and triglycerides in liver and muscle as concentrations of these energy reserves were greater in muscle, but not liver. Concentrations of glycogen and triglycerides in muscle, but not in liver, were lesser following the handling stressor suggesting that the muscle energy reserves are an important source of energy required for recovery from the handling stressor. The results of the study demonstrate that chronic exposure to dietary Se-Met elicits a stress response, but prevents a cortisol response to a secondary handling stressor, most likely due to cortisol inactivation. Moreover, exposure to Se-Met has effects on concentrations of energy reserves that are important for providing the energy necessary to cope with a secondary stressor.</description><subject>Animal, plant and microbial ecology</subject><subject>Animals</subject><subject>Applied ecology</subject><subject>Biological and medical sciences</subject><subject>Blood Glucose - drug effects</subject><subject>Blood Glucose - metabolism</subject><subject>blood plasma</subject><subject>chronic exposure</subject><subject>Corticosteroidogenesis</subject><subject>Cortisol</subject><subject>Cortisone</subject><subject>Cortisone - blood</subject><subject>cytochrome P-450</subject><subject>dietary exposure</subject><subject>Ecotoxicology, biological effects of pollution</subject><subject>energy</subject><subject>Female</subject><subject>Food Contamination</subject><subject>Fundamental and applied biological sciences. Psychology</subject><subject>General aspects</subject><subject>glucose</subject><subject>Glycogen - metabolism</subject><subject>Handling (Psychology)</subject><subject>Hydrocortisone - blood</subject><subject>lactation</subject><subject>liver</subject><subject>Liver - drug effects</subject><subject>Liver - metabolism</subject><subject>MC2R</subject><subject>Muscles - drug effects</subject><subject>Muscles - metabolism</subject><subject>Oncorhynchus mykiss</subject><subject>Oncorhynchus mykiss - growth & development</subject><subject>Oncorhynchus mykiss - metabolism</subject><subject>PBR</subject><subject>Random Allocation</subject><subject>Real-Time Polymerase Chain Reaction</subject><subject>Selenium</subject><subject>selenomethionine</subject><subject>Selenomethionine - pharmacokinetics</subject><subject>Selenomethionine - toxicity</subject><subject>Stress response</subject><subject>Stress, Physiological - drug effects</subject><subject>Stressor</subject><subject>Toxicity Tests, Chronic</subject><subject>triacylglycerols</subject><subject>Triglycerides - metabolism</subject><subject>trout</subject><subject>Water Pollutants, Chemical - pharmacokinetics</subject><subject>Water Pollutants, Chemical - toxicity</subject><issn>0166-445X</issn><issn>1879-1514</issn><fulltext>true</fulltext><rsrctype>article</rsrctype><creationdate>2011</creationdate><recordtype>article</recordtype><sourceid>EIF</sourceid><recordid>eNqNkUuPFCEUhYnROO3oT1DZGFdVXqqgqliZycRXMokLncQdoaiLTacKeoCax7-XTre6VDaE5Dv3cs4h5CWDmgHr3u1qfbPqHO7rBhirQdYAzSOyYUMvKyYYf0w2hesqzsWPM_IspR2U03D5lJw1THatkGJDbi9yRl8GueBpsDRvkZoQs0thphHTPviENAeacnkl6jy1uOgZadTOj-GO5hjWTM02Bu-MnucHivf7kHA6qCaHWccHmnBGHxbM27LHeXxOnlg9J3xxus_J9ccP3y8_V1dfP325vLiqDBeQq5H3tu9004pODDg2Ewc2Wmb5gFOxDQWyusOeTQPIdjRCmmFstQQtrGyLx3Py9jh3H8PNiimrxSWD86w9hjUpyRjrSlr_QUIDkgH0hRRH0sSQUkSr9tEtxaVioA7dqJ06daMO3SiQqgRfdK9OG9ZxwemP6ncZBXhzAnQqSdqovXHpL8f7gcmWFe71kbM6KP0zFub6W9nEAZiQwKEQ748ElmxvHUaVjENvcHIRTVZTcP_47C_nsrsa</recordid><startdate>20111001</startdate><enddate>20111001</enddate><creator>Wiseman, Steve</creator><creator>Thomas, Jith K.</creator><creator>McPhee, Landon</creator><creator>Hursky, Olesya</creator><creator>Raine, Jason C.</creator><creator>Pietrock, Michael</creator><creator>Giesy, John P.</creator><creator>Hecker, Markus</creator><creator>Janz, David M.</creator><general>Elsevier B.V</general><general>Elsevier</general><scope>FBQ</scope><scope>IQODW</scope><scope>CGR</scope><scope>CUY</scope><scope>CVF</scope><scope>ECM</scope><scope>EIF</scope><scope>NPM</scope><scope>AAYXX</scope><scope>CITATION</scope><scope>7X8</scope><scope>7QH</scope><scope>7ST</scope><scope>7TV</scope><scope>7U7</scope><scope>7UA</scope><scope>C1K</scope><scope>F1W</scope><scope>H97</scope><scope>L.G</scope><scope>SOI</scope></search><sort><creationdate>20111001</creationdate><title>Attenuation of the cortisol response to stress in female rainbow trout chronically exposed to dietary selenomethionine</title><author>Wiseman, Steve ; Thomas, Jith K. ; McPhee, Landon ; Hursky, Olesya ; Raine, Jason C. ; Pietrock, Michael ; Giesy, John P. ; Hecker, Markus ; Janz, David M.</author></sort><facets><frbrtype>5</frbrtype><frbrgroupid>cdi_FETCH-LOGICAL-c450t-b47f76a235658eb2d401bf1f48ed2010450fa6e71d8093bc59c8b3a90a5f93963</frbrgroupid><rsrctype>articles</rsrctype><prefilter>articles</prefilter><language>eng</language><creationdate>2011</creationdate><topic>Animal, plant and microbial ecology</topic><topic>Animals</topic><topic>Applied ecology</topic><topic>Biological and medical sciences</topic><topic>Blood Glucose - drug effects</topic><topic>Blood Glucose - metabolism</topic><topic>blood plasma</topic><topic>chronic exposure</topic><topic>Corticosteroidogenesis</topic><topic>Cortisol</topic><topic>Cortisone</topic><topic>Cortisone - blood</topic><topic>cytochrome P-450</topic><topic>dietary exposure</topic><topic>Ecotoxicology, biological effects of pollution</topic><topic>energy</topic><topic>Female</topic><topic>Food Contamination</topic><topic>Fundamental and applied biological sciences. Psychology</topic><topic>General aspects</topic><topic>glucose</topic><topic>Glycogen - metabolism</topic><topic>Handling (Psychology)</topic><topic>Hydrocortisone - blood</topic><topic>lactation</topic><topic>liver</topic><topic>Liver - drug effects</topic><topic>Liver - metabolism</topic><topic>MC2R</topic><topic>Muscles - drug effects</topic><topic>Muscles - metabolism</topic><topic>Oncorhynchus mykiss</topic><topic>Oncorhynchus mykiss - growth & development</topic><topic>Oncorhynchus mykiss - metabolism</topic><topic>PBR</topic><topic>Random Allocation</topic><topic>Real-Time Polymerase Chain Reaction</topic><topic>Selenium</topic><topic>selenomethionine</topic><topic>Selenomethionine - pharmacokinetics</topic><topic>Selenomethionine - toxicity</topic><topic>Stress response</topic><topic>Stress, Physiological - drug effects</topic><topic>Stressor</topic><topic>Toxicity Tests, Chronic</topic><topic>triacylglycerols</topic><topic>Triglycerides - metabolism</topic><topic>trout</topic><topic>Water Pollutants, Chemical - pharmacokinetics</topic><topic>Water Pollutants, Chemical - toxicity</topic><toplevel>peer_reviewed</toplevel><toplevel>online_resources</toplevel><creatorcontrib>Wiseman, Steve</creatorcontrib><creatorcontrib>Thomas, Jith K.</creatorcontrib><creatorcontrib>McPhee, Landon</creatorcontrib><creatorcontrib>Hursky, Olesya</creatorcontrib><creatorcontrib>Raine, Jason C.</creatorcontrib><creatorcontrib>Pietrock, Michael</creatorcontrib><creatorcontrib>Giesy, John P.</creatorcontrib><creatorcontrib>Hecker, Markus</creatorcontrib><creatorcontrib>Janz, David M.</creatorcontrib><collection>AGRIS</collection><collection>Pascal-Francis</collection><collection>Medline</collection><collection>MEDLINE</collection><collection>MEDLINE (Ovid)</collection><collection>MEDLINE</collection><collection>MEDLINE</collection><collection>PubMed</collection><collection>CrossRef</collection><collection>MEDLINE - Academic</collection><collection>Aqualine</collection><collection>Environment Abstracts</collection><collection>Pollution Abstracts</collection><collection>Toxicology Abstracts</collection><collection>Water Resources Abstracts</collection><collection>Environmental Sciences and Pollution Management</collection><collection>ASFA: Aquatic Sciences and Fisheries Abstracts</collection><collection>Aquatic Science & Fisheries Abstracts (ASFA) 3: Aquatic Pollution & Environmental Quality</collection><collection>Aquatic Science & Fisheries Abstracts (ASFA) Professional</collection><collection>Environment Abstracts</collection><jtitle>Aquatic toxicology</jtitle></facets><delivery><delcategory>Remote Search Resource</delcategory><fulltext>fulltext</fulltext></delivery><addata><au>Wiseman, Steve</au><au>Thomas, Jith K.</au><au>McPhee, Landon</au><au>Hursky, Olesya</au><au>Raine, Jason C.</au><au>Pietrock, Michael</au><au>Giesy, John P.</au><au>Hecker, Markus</au><au>Janz, David M.</au><format>journal</format><genre>article</genre><ristype>JOUR</ristype><atitle>Attenuation of the cortisol response to stress in female rainbow trout chronically exposed to dietary selenomethionine</atitle><jtitle>Aquatic toxicology</jtitle><addtitle>Aquat Toxicol</addtitle><date>2011-10-01</date><risdate>2011</risdate><volume>105</volume><issue>3</issue><spage>643</spage><epage>651</epage><pages>643-651</pages><issn>0166-445X</issn><eissn>1879-1514</eissn><coden>AQTODG</coden><abstract>► Trout exposed to Se-Met had greater concentration of cortisol compared to controls. ► Transcript abundance of mc2r was greater in trout exposed to Se-Met. ► Trout exposed to Se-Met had a reduced cortisol response to a handling stressor. ► Cortisone concentration was greater in Se-Met exposed trout post-handling stressor.
Selenomethionine (Se-Met) is the major dietary form of selenium (Se). While Se is a required nutrient, it can also influence the physiological stress response because it stimulates greater concentrations of cortisol in blood plasma of exposed fish. However, little is known about the effects of exposure to Se on the ability to cope with a secondary stressor. In the current study, female rainbow trout were exposed to an environmentally relevant dietary concentration (8.47
mg
Se/kg dry mass (dm)) of Se-Met for 126
d, after which time fish were subjected to a 3-min handling stressor and sampled at 2
h and 24
h post-stressor exposure. Concentrations of cortisol, cortisone, glucose, and lactate in blood plasma and concentrations of glycogen and triglycerides in liver and muscle were determined. Abundances of transcripts of proteins involved in corticosteroidogenesis were determined using quantitative RT-PCR. Concentrations of cortisol were significantly greater in blood plasma of trout exposed to Se-Met, relative to control trout sampled prior to the handling stressor. A typical response of cortisol to the handling stressor was observed in the control trout. However, trout exposed to Se-Met were unable to mount a cortisol response to the handling stressor. Concentrations of cortisone, the inactive metabolite of cortisol, were significantly greater following the handling stressor in trout exposed to Se-Met. In trout exposed to Se-Met, transcript abundance of melanocortin 2 receptor (
mc2r) and peripheral benzodiazepine receptor (
pbr) were greater, which is consistent with the conclusion that synthesis of cortisol was greater. However, abundances of transcripts of cytochrome P450 side-chain cleavage (
p450scc) and cytochrome P450 11B1 (
cyp11b1) were not significantly different between controls and Se-Met exposed trout. Exposure to Se-Met affected accumulation and tissue partitioning of glycogen and triglycerides in liver and muscle as concentrations of these energy reserves were greater in muscle, but not liver. Concentrations of glycogen and triglycerides in muscle, but not in liver, were lesser following the handling stressor suggesting that the muscle energy reserves are an important source of energy required for recovery from the handling stressor. The results of the study demonstrate that chronic exposure to dietary Se-Met elicits a stress response, but prevents a cortisol response to a secondary handling stressor, most likely due to cortisol inactivation. Moreover, exposure to Se-Met has effects on concentrations of energy reserves that are important for providing the energy necessary to cope with a secondary stressor.</abstract><cop>Amsterdam</cop><pub>Elsevier B.V</pub><pmid>21963595</pmid><doi>10.1016/j.aquatox.2011.09.002</doi><tpages>9</tpages></addata></record> |
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subjects | Animal, plant and microbial ecology Animals Applied ecology Biological and medical sciences Blood Glucose - drug effects Blood Glucose - metabolism blood plasma chronic exposure Corticosteroidogenesis Cortisol Cortisone Cortisone - blood cytochrome P-450 dietary exposure Ecotoxicology, biological effects of pollution energy Female Food Contamination Fundamental and applied biological sciences. Psychology General aspects glucose Glycogen - metabolism Handling (Psychology) Hydrocortisone - blood lactation liver Liver - drug effects Liver - metabolism MC2R Muscles - drug effects Muscles - metabolism Oncorhynchus mykiss Oncorhynchus mykiss - growth & development Oncorhynchus mykiss - metabolism PBR Random Allocation Real-Time Polymerase Chain Reaction Selenium selenomethionine Selenomethionine - pharmacokinetics Selenomethionine - toxicity Stress response Stress, Physiological - drug effects Stressor Toxicity Tests, Chronic triacylglycerols Triglycerides - metabolism trout Water Pollutants, Chemical - pharmacokinetics Water Pollutants, Chemical - toxicity |
title | Attenuation of the cortisol response to stress in female rainbow trout chronically exposed to dietary selenomethionine |
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