团头鲂糖皮质激素受体基因组织表达分布及其在应激中的表达变化
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摘要
为了解团头鲂糖皮质激素受体(GR)在应激反应中的调控机制,本研究以皮质醇注射模拟应激事件,采集组织样品,常规检测血糖、血清皮质醇水平;利用荧光定量PCR检测了gr在不同组织中的表达丰度及其参与调控的功能基因的表达变化;并通过常规石蜡切片开展了组织病理学研究。结果显示,gr1在脾脏、鳃、头肾等组织中有较高的表达量,gr2则在肝、垂体、肠等组织中具有较高的表达丰度。应激恢复过程中,下丘脑gr1表达量存在波动,gr2在0 h表达显著上调,gr1/gr2值逐渐增大;垂体中gr1和gr2均呈现先升后降的趋势,gr1/gr2值在2 h达到峰值水平;头肾中gr1表达量有波动,而gr2在0 h和2 h的表达量显著低于其他检测点的表达量,gr1/gr2值逐渐减小;肝脏中gr1的表达量在0 h和2 h的表达量显著高于其他检测点,gr2表达有上下波动的现象,gr1/gr2值有减小的趋势,而pepck表达则出现了显著上调,在2 h达到峰值;皮肤中gr1先升高后降低,gr2在0 h和2 h的表达量显著低于其他检测点,gr1/gr2值在2～8 h维持在较大值,而occ表达呈现先增加后降低的变化趋势;鳃中gr1与gr2均在2 h时发生了显著上调,gr1/gr2值在2～8 h维持在较大值,occ表达峰值出现在2 h。组织学研究显示鳃丝有增生现象,肾间组织中类淋巴细胞增多,其他所检视组织无明显病理改变。应激反应过程中gr在不同组织不同亚型间存在不同的表达变化,以及和不同组织器官中相关功能基因的相关性显示了GR在相关调控中的复杂性,而组织学研究结果则表明了应激反应存在诱发病理变化的风险。
In order to understand the biological function of glucocorticoid receptors(GRs) of Megalobrama amblycephala, we explored the expression characteristics of grs in different tissues and the expression changes of grs and some function genes in the recovery process after stress. Results show that the gr1 was highly expressed in gill, spleen and head kidney, while gr2 was highly expressed in pituitary, liver, intestinal, and so on. During the recovery process, on HPI axis, the expression of gr1 had fluctuations, gr2 significantly up-regulated at the 0 h, and the value of gr1/gr2 was gradually increasing in hypothalamus. The expression of gr1 and gr2 all rose first and then decreased, gr1/gr2 value reached the peak at 2 h in pituitary. The expression of gr1 had fluctuations, whereas gr2 had a low expression at 0 h and 2 h compared with the other time points, gr1/gr2 value gradually decreased in head kidney. In liver, gr1 rose in the early time(0 h and 2 h), and gr2 presented a fluctuating situation, and there was a tendency to reduce the the value of gr1/gr2. At the similar time point the expression of pepck peaked. In gills gr1 rose first, and then decreased, while the expression of gr2 was subdued in the early time(0 h and 2 h), gr1/gr2 value maintained at a large value during 2-8 h, while the occ expression peaked at 2 h. In skin gr1 and gr2 rose significantly in 2 h, gr1/gr2 in 2-8 h also maintained at a large value, and the occ expression rose first and then decreased. Histology research shows that gill filament was hyperplasia, lymphocyte-like cells increased kidney tissues, while other organizations examined had no obvious pathological changes. All the results indicate that GRs have a complex regulatory mechanism in the stress, and the histological analysis shows that the risks of pathological changes come from the stress.
In order to understand the biological function of glucocorticoid receptors(GRs) of Megalobrama amblycephala, we explored the expression characteristics of grs in different tissues and the expression changes of grs and some function genes in the recovery process after stress. Results show that the gr1 was highly expressed in gill, spleen and head kidney, while gr2 was highly expressed in pituitary, liver, intestinal, and so on. During the recovery process, on HPI axis, the expression of gr1 had fluctuations, gr2 significantly up-regulated at the 0 h, and the value of gr1/gr2 was gradually increasing in hypothalamus. The expression of gr1 and gr2 all rose first and then decreased, gr1/gr2 value reached the peak at 2 h in pituitary. The expression of gr1 had fluctuations, whereas gr2 had a low expression at 0 h and 2 h compared with the other time points, gr1/gr2 value gradually decreased in head kidney. In liver, gr1 rose in the early time(0 h and 2 h), and gr2 presented a fluctuating situation, and there was a tendency to reduce the the value of gr1/gr2. At the similar time point the expression of pepck peaked. In gills gr1 rose first, and then decreased, while the expression of gr2 was subdued in the early time(0 h and 2 h), gr1/gr2 value maintained at a large value during 2-8 h, while the occ expression peaked at 2 h. In skin gr1 and gr2 rose significantly in 2 h, gr1/gr2 in 2-8 h also maintained at a large value, and the occ expression rose first and then decreased. Histology research shows that gill filament was hyperplasia, lymphocyte-like cells increased kidney tissues, while other organizations examined had no obvious pathological changes. All the results indicate that GRs have a complex regulatory mechanism in the stress, and the histological analysis shows that the risks of pathological changes come from the stress.
引文
[1]Malandrakis E E,Dadali O,Golomazou E,et al.DNAdamage and differential gene expression associated with physical stress in gilthead seabream(Sparus aurata)[J].General and Comparative Endocrinology,2016,236:98-104.
[2]Schreck C B.Stress and fish reproduction:the roles of allostasis and hormesis[J].General and Comparative Endocrinology,2010,165(3):549-556.
[3]Genova R M,Marchaterre M A,Knapp R,et al.Glucocorticoid and androgen signaling pathways diverge between advertisement calling and non-calling fish[J].Hormones and Behavior,2012,62(4):426-432.
[4]Small B C,Bilodeau A L.Effects of cortisol and stress on channel catfish(Ictalurus punctatus)pathogen susceptibility and lysozyme activity following exposure to Edwardsiella ictaluri[J].General and Comparative Endocrinology,2005,142(1-2):256-262.
[5]刘小玲.鱼类应激反应的研究[J].水利渔业,2007,27(3):1-3.Liu X L.Studies on the emergency responses of fishes[J].Reservoir Fisheries,2007,27(3):1-3(in Chinese).
[6]Duma D,Jewell C M,Cidlowski J A.Multiple glucocorticoid receptor isoforms and mechanisms of post-translational modification[J].Journal of Steroid Biochemistry&Molecular Biology,2006,102(1-5):11-21.
[7]曾灵,顾玮,严军,等.人糖皮质激素受体调控靶基因的生物信息学分析[J].基础医学与临床,2008,28(11):1146-1150.Zeng L,Gu W,Yan J,et al.Bioinformatics analysis of human glucocorticoid receptor regulated target genes[J].Basic&Clinical Medicine,2008,28(11):1146-1150(in Chinese).
[8]Bongiovanni B,Mata-Espinosa D,D'Attilio L,et al.Effect of cortisol and/or DHEA on THP1-derived macrophages infected with Mycobacterium tuberculosis[J].Tuberculosis,2015,95(5):562-569.
[9]Robert O,Boujedidi H,Bigorgne A,et al.Decreased expression of the glucocorticoid receptor-GILZ pathway in Kupffer cells promotes liver inflammation in obese mice[J].Journal of Hepatology,2016,64(4):916-924.
[10]Oh K S,Patel H,Gottschalk R A,et al.Antiinflammatory chromatinscape suggests alternative mechanisms of glucocorticoid receptor action[J].Immunity,2017,47(2):298-309.
[11]Dieken E S,Miesfeld R L.Transcriptional transactivation functions localized to the glucocorticoid receptor Nterminus are necessary for steroid induction of lymphocyte apoptosis[J].Molecular and Cellular Biology,1992,12(2):589-597.
[12]Kitajima T,Ariizumi K,Bergstresser P R,et al.A novel mechanism of glucocorticoid-induced immune suppression:the inhibiton of T cell-mediated terminal maturation of a murine dendritic cell line[J].Journal of Clinical Investigation,1996,98(1):142-147.
[13]Beato M,Herrlich P,Schütz G.Steroid hormone receptors:many actors in search of a plot[J].Cell,1995,83(6):851-857.
[14]Saeij J P J,Verburg-Van Kemenade L B M,Van Muiswinkel W B,et al.Daily handling stress reduces resistance of carp to Trypanoplasma borreli:in vitro modulatory effects of cortisol on leukocyte function and apoptosis[J].Developmental and Comparative Immunology,2003,27(3):233-245.
[15]Gadan K,Marjara I S,Sundh H,et al.Slow release cortisol implants result in impaired innate immune responses and higher infection prevalence following experimental challenge with infectious pancreatic necrosis virus in Atlantic salmon(Salmo salar)parr[J].Fish&Shellfish Immunology,2012,32(5):637-644.
[16]Chasiotis H,Kelly S P.Effect of cortisol on permeability and tight junction protein transcript abundance in primary cultured gill epithelia from stenohaline goldfish and euryhaline trout[J].General and Comparative Endocrinology,2011,172(3):494-504.
[17]Mateus A P,Anjos L,Cardoso J R,et al.Chronic stress impairs the local immune response during cutaneous repair in gilthead sea bream(Sparus aurata,L.)[J].Molecular Immunology,2017,87:267-283.
[18]Vizzini A,Vazzana M,Cammarata M,et al.Peritoneal cavity phagocytes from the teleost sea bass express a glucocorticoid receptor(cloned and sequenced)involved in genomic modulation of the in vitro chemiluminescence response to zymosan[J].General and Comparative Endocrinology,2007,150(1):114-123.
[19]Esteban M A,Rodríguez A,Ayala A G,et al.Effects of high doses of cortisol on innate cellular immune response of seabream(Sparus aurata L.)[J].General and Comparative Endocrinology,2004,137(1):89-98.
[20]Vijayan M M,Raptis S,Sathiyaa R.Cortisol treatment affects glucocorticoid receptor and glucocorticoidresponsive genes in the liver of rainbow trout[J].General and Comparative Endocrinology,2003,132(2):256-263.
[21]Olsen R E,Sundell K,Ring?E,et al.The acute stress response in fed and food deprived Atlantic cod,Gadus morhua L[J].Aquaculture,2008,280(1-4):232-241.
[22]Mazon A F,Huising M O,Taverne-Thiele A J,et al.The first appearance of Rodlet cells in carp(Cyprinus carpio L.)ontogeny and their possible roles during stress and parasite infection[J].Fish&Shellfish Immunology,2007,22(1-2):27-37.
[23]董晶晶,薛春雨,习丙文,等.团头鲂黑素皮质素受体2基因克隆、组织分布和应激后的表达分析[J].江苏农业科学,2014,42(6):20-26.Dong J J,Xue C Y,Xi B W,et al.Gene cloning,tissue distribution and expression after stress of melanocortin receptor 2 in Megalobrama amblycephala[J].Jiangsu Agricultural Sciences,2014,42(6):20-26(in Chinese).
[24]Xue C Y,Xi B W,Ren M C,et al.Molecular cloning,tissue expression of gene Muc2 in blunt snout bream Megalobrama amblycephala and regulation after refeeding[J].Chinese Journal of Oceanology and Limnology,2015,33(2):291-298.
[25]常波,高珊珊,郭琪,等.辽东楤木根皮和人参茎叶总皂苷提取物对过度训练大鼠海马盐皮质类固醇受体MR,糖皮质激素受体GR蛋白表达的影响[J].广州体育学院学报,2014,34(3):94-97,106.Chang B,Gao S S,Guo Q,et al.Empirical study of A.Elata and panax ginseng extracts on the MR,GRprotein expression of hippocampus tissue in the overtraining rat[J].Journal of Guangzhou Sport University,2014,34(3):94-97,106(in Chinese).
[26]万顺伦,徐建军,孙刚.脑内皮质激素受体及其受体前代谢酶的作用[J].生理科学进展,2001,32(2):171-173.Wan S L,Xu J J,Sun G.The actions of glucocorticoid receptor and 11β-hydroxysteroid dehydrogenases in the brain[J].Progress in Physiological Sciences,2001,32(2):171-173(in Chinese).
[27]Small B C,Quiniou S M A.Characterization of two channel catfish,Ictalurus punctatus,glucocorticoid receptors and expression following an acute stressor[J].Comparative Biochemistry and Physiology-Part A:Molecular&Integrative Physiology,2018,216:42-51.
[28]陶争荣,张昊,曾涛,等.鸽子GR基因cDNA克隆与表达分析[J].农业生物技术学报,2016,24(1):83-89.Tao Z R,Zhang H,Zeng T,et al.Molecular cloning and expression analysis of GR gene in pigeon(Columba livia domestica)[J].Journal of Agricultural Biotechnology,2016,24(1):83-89(in Chinese).
[29]Kumai Y,Nesan D,Vijayan M M,et al.Cortisol regulates Na+uptake in zebrafish,Danio rerio,larvae via the glucocorticoid receptor[J].Molecular and Cellular Endocrinology,2012,364(1-2):113-125.
[30]杨玲,何威,李颖.地塞米松给药及撤药后糖皮质激素受体的表达[J].第三军医大学学报,2009,31(21):2086-2089.Yang L,He W,Li Y,et al.Expression of glucocorticoid receptor in HaCaT cells under dexamethasone treatment or after withdrawal[J].Acta Academiae Medicinae Militaris Tertiae,2009,31(21):2086-2089(in Chinese).
[31]程晓刚,粟永萍,罗成基.糖皮质激素受体结构与功能研究进展[J].国外医学分子生物学分册,2003,25(1):29-33.Cheng X G,Su Y P,Luo C J.The structure and function of glucocorticoid receptor[J].Foreign Medical Science Molecular Biology Section,2003,25(1):29-33(in Chinese).
[32]Terova G,Gornati R,Rimoldi S,et al.Quantification of a glucocorticoid receptor in sea bass(Dicentrarchus labrax,L.)reared at high stocking density[J].Gene,2005,357(2):144-151.
[33]Patterson S,Zee J,Wiseman S,et al.Effects of chronic exposure to dietary selenomethionine on the physiological stress response in juvenile white sturgeon(Acipenser transmontanus)[J].Aquatic Toxicology,2017,186:77-86.
[34]Greulich F,Hemmer M C,Rollins D A.There goes the neighborhood:assembly of transcriptional complexes during the regulation of metabolism and inflammation by the glucocorticoid receptor[J].Steroids,2016,114:7-15.
[35]李呈贞.小檗碱治疗肠道炎症的机制研究[J].现代免疫学,2013,33(1):74-76,11.Li C Z.The mechanism of berberine in inflammatory bowel diseas[J].Modern Immunology,2013,33(1):74-76,11(in Chinese).
[36]Duncan A R,Sadowska G B,Stonestreet B S.Ontogeny and the effects of exogenous and endogenous glucocorticoids on tight junction protein expression in ovine cerebral cortices[J].Brain Research,2009,1303:15-25.
[37]Olsen R E,Sundell K,Mayhew T M,et al.Acute stress alters intestinal function of rainbow trout,Oncorhynchus mykiss(Walbaum)[J].Aquaculture,2005,250(1-2):480-495.
[38]Gauberg J,Kolosov D,Kelly S P.Claudin tight junction proteins in rainbow trout(Oncorhynchus mykiss)skin:spatial response to elevated cortisol levels[J].General and Comparative Endocrinology,2017,240:214-226.
[39]Chasiotis H,Wood C M,Kelly S P.Cortisol reduces paracellular permeability and increases occludin abundance in cultured trout gill epithelia[J].Molecular and Cellular Endocrinology,2010,323(2):232-238.
[40]Kelly S P,Chasiotis H.Glucocorticoid and mineralocorticoid receptors regulate paracellular permeability in a primary cultured gill epithelium[J].The Journal of Experimental Biology,2011,214:2308-2318.
[41]Wood C M,Kelly S P,Zhou B S,et al.Cultured gill epithelia as models for the freshwater fish gill[J].Biochimica et Biophysica Acta(BBA)-Biomembranes,2002,1566(1-2):72-83.
[2]Schreck C B.Stress and fish reproduction:the roles of allostasis and hormesis[J].General and Comparative Endocrinology,2010,165(3):549-556.
[3]Genova R M,Marchaterre M A,Knapp R,et al.Glucocorticoid and androgen signaling pathways diverge between advertisement calling and non-calling fish[J].Hormones and Behavior,2012,62(4):426-432.
[4]Small B C,Bilodeau A L.Effects of cortisol and stress on channel catfish(Ictalurus punctatus)pathogen susceptibility and lysozyme activity following exposure to Edwardsiella ictaluri[J].General and Comparative Endocrinology,2005,142(1-2):256-262.
[5]刘小玲.鱼类应激反应的研究[J].水利渔业,2007,27(3):1-3.Liu X L.Studies on the emergency responses of fishes[J].Reservoir Fisheries,2007,27(3):1-3(in Chinese).
[6]Duma D,Jewell C M,Cidlowski J A.Multiple glucocorticoid receptor isoforms and mechanisms of post-translational modification[J].Journal of Steroid Biochemistry&Molecular Biology,2006,102(1-5):11-21.
[7]曾灵,顾玮,严军,等.人糖皮质激素受体调控靶基因的生物信息学分析[J].基础医学与临床,2008,28(11):1146-1150.Zeng L,Gu W,Yan J,et al.Bioinformatics analysis of human glucocorticoid receptor regulated target genes[J].Basic&Clinical Medicine,2008,28(11):1146-1150(in Chinese).
[8]Bongiovanni B,Mata-Espinosa D,D'Attilio L,et al.Effect of cortisol and/or DHEA on THP1-derived macrophages infected with Mycobacterium tuberculosis[J].Tuberculosis,2015,95(5):562-569.
[9]Robert O,Boujedidi H,Bigorgne A,et al.Decreased expression of the glucocorticoid receptor-GILZ pathway in Kupffer cells promotes liver inflammation in obese mice[J].Journal of Hepatology,2016,64(4):916-924.
[10]Oh K S,Patel H,Gottschalk R A,et al.Antiinflammatory chromatinscape suggests alternative mechanisms of glucocorticoid receptor action[J].Immunity,2017,47(2):298-309.
[11]Dieken E S,Miesfeld R L.Transcriptional transactivation functions localized to the glucocorticoid receptor Nterminus are necessary for steroid induction of lymphocyte apoptosis[J].Molecular and Cellular Biology,1992,12(2):589-597.
[12]Kitajima T,Ariizumi K,Bergstresser P R,et al.A novel mechanism of glucocorticoid-induced immune suppression:the inhibiton of T cell-mediated terminal maturation of a murine dendritic cell line[J].Journal of Clinical Investigation,1996,98(1):142-147.
[13]Beato M,Herrlich P,Schütz G.Steroid hormone receptors:many actors in search of a plot[J].Cell,1995,83(6):851-857.
[14]Saeij J P J,Verburg-Van Kemenade L B M,Van Muiswinkel W B,et al.Daily handling stress reduces resistance of carp to Trypanoplasma borreli:in vitro modulatory effects of cortisol on leukocyte function and apoptosis[J].Developmental and Comparative Immunology,2003,27(3):233-245.
[15]Gadan K,Marjara I S,Sundh H,et al.Slow release cortisol implants result in impaired innate immune responses and higher infection prevalence following experimental challenge with infectious pancreatic necrosis virus in Atlantic salmon(Salmo salar)parr[J].Fish&Shellfish Immunology,2012,32(5):637-644.
[16]Chasiotis H,Kelly S P.Effect of cortisol on permeability and tight junction protein transcript abundance in primary cultured gill epithelia from stenohaline goldfish and euryhaline trout[J].General and Comparative Endocrinology,2011,172(3):494-504.
[17]Mateus A P,Anjos L,Cardoso J R,et al.Chronic stress impairs the local immune response during cutaneous repair in gilthead sea bream(Sparus aurata,L.)[J].Molecular Immunology,2017,87:267-283.
[18]Vizzini A,Vazzana M,Cammarata M,et al.Peritoneal cavity phagocytes from the teleost sea bass express a glucocorticoid receptor(cloned and sequenced)involved in genomic modulation of the in vitro chemiluminescence response to zymosan[J].General and Comparative Endocrinology,2007,150(1):114-123.
[19]Esteban M A,Rodríguez A,Ayala A G,et al.Effects of high doses of cortisol on innate cellular immune response of seabream(Sparus aurata L.)[J].General and Comparative Endocrinology,2004,137(1):89-98.
[20]Vijayan M M,Raptis S,Sathiyaa R.Cortisol treatment affects glucocorticoid receptor and glucocorticoidresponsive genes in the liver of rainbow trout[J].General and Comparative Endocrinology,2003,132(2):256-263.
[21]Olsen R E,Sundell K,Ring?E,et al.The acute stress response in fed and food deprived Atlantic cod,Gadus morhua L[J].Aquaculture,2008,280(1-4):232-241.
[22]Mazon A F,Huising M O,Taverne-Thiele A J,et al.The first appearance of Rodlet cells in carp(Cyprinus carpio L.)ontogeny and their possible roles during stress and parasite infection[J].Fish&Shellfish Immunology,2007,22(1-2):27-37.
[23]董晶晶,薛春雨,习丙文,等.团头鲂黑素皮质素受体2基因克隆、组织分布和应激后的表达分析[J].江苏农业科学,2014,42(6):20-26.Dong J J,Xue C Y,Xi B W,et al.Gene cloning,tissue distribution and expression after stress of melanocortin receptor 2 in Megalobrama amblycephala[J].Jiangsu Agricultural Sciences,2014,42(6):20-26(in Chinese).
[24]Xue C Y,Xi B W,Ren M C,et al.Molecular cloning,tissue expression of gene Muc2 in blunt snout bream Megalobrama amblycephala and regulation after refeeding[J].Chinese Journal of Oceanology and Limnology,2015,33(2):291-298.
[25]常波,高珊珊,郭琪,等.辽东楤木根皮和人参茎叶总皂苷提取物对过度训练大鼠海马盐皮质类固醇受体MR,糖皮质激素受体GR蛋白表达的影响[J].广州体育学院学报,2014,34(3):94-97,106.Chang B,Gao S S,Guo Q,et al.Empirical study of A.Elata and panax ginseng extracts on the MR,GRprotein expression of hippocampus tissue in the overtraining rat[J].Journal of Guangzhou Sport University,2014,34(3):94-97,106(in Chinese).
[26]万顺伦,徐建军,孙刚.脑内皮质激素受体及其受体前代谢酶的作用[J].生理科学进展,2001,32(2):171-173.Wan S L,Xu J J,Sun G.The actions of glucocorticoid receptor and 11β-hydroxysteroid dehydrogenases in the brain[J].Progress in Physiological Sciences,2001,32(2):171-173(in Chinese).
[27]Small B C,Quiniou S M A.Characterization of two channel catfish,Ictalurus punctatus,glucocorticoid receptors and expression following an acute stressor[J].Comparative Biochemistry and Physiology-Part A:Molecular&Integrative Physiology,2018,216:42-51.
[28]陶争荣,张昊,曾涛,等.鸽子GR基因cDNA克隆与表达分析[J].农业生物技术学报,2016,24(1):83-89.Tao Z R,Zhang H,Zeng T,et al.Molecular cloning and expression analysis of GR gene in pigeon(Columba livia domestica)[J].Journal of Agricultural Biotechnology,2016,24(1):83-89(in Chinese).
[29]Kumai Y,Nesan D,Vijayan M M,et al.Cortisol regulates Na+uptake in zebrafish,Danio rerio,larvae via the glucocorticoid receptor[J].Molecular and Cellular Endocrinology,2012,364(1-2):113-125.
[30]杨玲,何威,李颖.地塞米松给药及撤药后糖皮质激素受体的表达[J].第三军医大学学报,2009,31(21):2086-2089.Yang L,He W,Li Y,et al.Expression of glucocorticoid receptor in HaCaT cells under dexamethasone treatment or after withdrawal[J].Acta Academiae Medicinae Militaris Tertiae,2009,31(21):2086-2089(in Chinese).
[31]程晓刚,粟永萍,罗成基.糖皮质激素受体结构与功能研究进展[J].国外医学分子生物学分册,2003,25(1):29-33.Cheng X G,Su Y P,Luo C J.The structure and function of glucocorticoid receptor[J].Foreign Medical Science Molecular Biology Section,2003,25(1):29-33(in Chinese).
[32]Terova G,Gornati R,Rimoldi S,et al.Quantification of a glucocorticoid receptor in sea bass(Dicentrarchus labrax,L.)reared at high stocking density[J].Gene,2005,357(2):144-151.
[33]Patterson S,Zee J,Wiseman S,et al.Effects of chronic exposure to dietary selenomethionine on the physiological stress response in juvenile white sturgeon(Acipenser transmontanus)[J].Aquatic Toxicology,2017,186:77-86.
[34]Greulich F,Hemmer M C,Rollins D A.There goes the neighborhood:assembly of transcriptional complexes during the regulation of metabolism and inflammation by the glucocorticoid receptor[J].Steroids,2016,114:7-15.
[35]李呈贞.小檗碱治疗肠道炎症的机制研究[J].现代免疫学,2013,33(1):74-76,11.Li C Z.The mechanism of berberine in inflammatory bowel diseas[J].Modern Immunology,2013,33(1):74-76,11(in Chinese).
[36]Duncan A R,Sadowska G B,Stonestreet B S.Ontogeny and the effects of exogenous and endogenous glucocorticoids on tight junction protein expression in ovine cerebral cortices[J].Brain Research,2009,1303:15-25.
[37]Olsen R E,Sundell K,Mayhew T M,et al.Acute stress alters intestinal function of rainbow trout,Oncorhynchus mykiss(Walbaum)[J].Aquaculture,2005,250(1-2):480-495.
[38]Gauberg J,Kolosov D,Kelly S P.Claudin tight junction proteins in rainbow trout(Oncorhynchus mykiss)skin:spatial response to elevated cortisol levels[J].General and Comparative Endocrinology,2017,240:214-226.
[39]Chasiotis H,Wood C M,Kelly S P.Cortisol reduces paracellular permeability and increases occludin abundance in cultured trout gill epithelia[J].Molecular and Cellular Endocrinology,2010,323(2):232-238.
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