综述 REVIEW

鱼类雷帕霉素靶蛋白信号通路生物学功能的研究进展

  • 吴龙华 ,
  • 梁化亮 ,
  • 戈贤平 ,
  • 任鸣春
展开
  • 1. 南京农业大学无锡渔业学院, 无锡 214081;
    2. 中国水产科学研究院淡水渔业研究中心, 农业农村部淡水渔业和种质资源利用重点实验室, 无锡 214081
吴龙华(1997-),男,江苏淮安人,硕士研究生,水产动物营养与饲料专业。E-mail:wlhaz1021@163.com

收稿日期: 2021-03-24

  网络出版日期: 2021-10-16

基金资助

国家重点研发计划(2018YFD0900400);现代农业产业技术体系专项——国家大宗淡水鱼产业技术体系(CARS-45);江苏省自然科学基金青年基金项目(BK20200169)

Research Progress on Biological Function of Target of Rapamycin Signaling Pathway in Fish

  • WU Longhua ,
  • LIANG Hualiang ,
  • GE Xianping ,
  • REN Mingchun
Expand
  • 1. Wuxi Fisheries College, Nanjing Agricultural University, Wuxi 214081, China;
    2. Key Laboratory of Freshwater Fisheries and Germplasm Resources Utilization, Ministry of Agriculture and Rural Affairs, Freshwater Fisheries Research Center, Chinese Academy of Fishery Sciences, Wuxi 214081, China

Received date: 2021-03-24

  Online published: 2021-10-16

Supported by

 

摘要

雷帕霉素靶蛋白(TOR)信号通路是营养、化学和运动等因素导致细胞生长和分化的一个关键信号通路。本文介绍了到目前为止TOR信号通路的生物学功能以及其在水产动物中的研究进展,主要包括TOR信号通路对蛋白质合成代谢、糖代谢、脂代谢、摄食调控以及机体免疫、抗氧化调控的影响,同时阐述了胰岛素抵抗机制及精氨酸对团头鲂免疫与抗氧化信号通路的调控机制,以为进一步全面研究TOR信号通路在水产动物中的应用提供参考。

本文引用格式

吴龙华 , 梁化亮 , 戈贤平 , 任鸣春 . 鱼类雷帕霉素靶蛋白信号通路生物学功能的研究进展[J]. 动物营养学报, 2021 , 33(10) : 5486 -5496 . DOI: 10.3969/j.issn.1006-267x.2021.10.010

Abstract

Target of rapamycin (TOR) signaling pathway is a key signaling pathway in cell growth and differentiation caused by nutritional, chemical and motor factors. This study was introduced the biological function of TOR signaling pathway and its research progress in aquatic animals so far, to provide reference for further comprehensive study of TOR signaling pathway in aquatic animals. It mainly included the effects of TOR signaling pathway on the regulation of protein anabolism, glucose metabolism lipid metabolism, feed intake regulation, as well as the regulation of immunity and antioxidant status. It also expounded the mechanism of insulin resistance and the regulation mechanism of arginine in blunt snout bream (Megalobrama ambylcephala) on immune antioxidant signaling pathway.

参考文献

[1] 刘宁, 刘国华, 蔡辉益, 等.营养介导的TOR信号传导研究进展[J].中国畜牧兽医, 2010, 37(6):25-29. LIU N, LIU G H, CAI H Y, et al.Advances in nutrient regulation of tor signal transduction[J].China Animal Husbandry & Veterinary Medicine, 2010, 37(6):25-29.(in Chinese)
[2] SABATINI D M.Twenty-five years of mTOR:uncovering the link from nutrients to growth[J].Proceedings of the National Academy of Sciences of the United States of America, 2017, 114(45):11818-11825.  
[3] WOLFSON R L, SABATINI D M.The dawn of the age of amino acid sensors for the mTORC1 pathway[J].Cell Metabolism, 2017, 26(2):301-309.  
[4] CHANTRANUPONG L, WOLFSON R L, SABATINI D M.Nutrient-sensing mechanisms across evolution[J].Cell, 2015, 161(1):67-83.  
[5] LAPLANTE M, SABATINI D M.mTOR signaling in growth control and disease[J].Cell, 2012, 149(2):274-293.  
[6] WEICHHART T, COSTANTINO G, POGLITSCH M A, et al.The TSC-mTOR signaling pathway regulates the innate inflammatory response[J].Immunity, 2008, 29(4):565-577.  
[7] 邓玉平.亮氨酸对生长中期草鱼生长、肌肉品质和肠道免疫的影响研究[D].硕士学位论文.雅安:四川农业大学, 2014. DENG Y P.Effects of graded levels of dietary leucine on growth, flesh quality, intestinal immunity and antioxidant statusin young grass carp (Ctenopharyngodon idella)[D].Master's Thesis.Ya'an:Sichuan Agricultural University, 2014.(in Chinese)
[8] COTA D, MATTER E K, WOODS S C, et al.The role of hypothalamic mammalian target of rapamycin complex 1 signaling in diet-induced obesity[J].The Journal of Neuroscience, 2008, 28(28):7202-7208.  
[9] BLOUET C, ONO H, SCHWARTZ G J.Mediobasal hypothalamic p70 S6 kinase 1 modulates the control of energy homeostasis[J].Cell Metabolism, 2008, 8(6):459-467.  
[10] 梁晓芳.花鲈利用鱼粉和植物蛋白源的选择性摄食调控机制研究[D].博士学位论文.北京:中国农业科学院, 2017. LIANG X F.Mechanism on feed intake regulation of Lateolabrax japonicas when fishmeal was replaced by plant protein[D].Ph.D.Thesis.Beijing:Chinese Academy of Agricultural Sciences, 2017.(in Chinese)
[11] COTA D, PROULX K, SMITH K A B, et al.Hypothalamic mTOR signaling regulates food intake[J].Science, 2006, 312(5775):927-930.  
[12] HAO S Z, SHARP J W, ROSS-INTA C M, et al.Uncharged tRNA and sensing of amino acid deficiency in mammalian piriform cortex[J].Science, 2005, 307(5716):1776-1778.  
[13] ALBERT V, CORNU M, HALL M N.mTORC1 signaling in Agrp neurons mediates circadian expression of AgRP and NPY but is dispensable for regulation of feeding behavior[J].Biochemical and Biophysical Research Communications, 2015, 464(2):480-486.  
[14] SMITH M A, KATSOURI L, IRVINE E E, et al.Ribosomal S6K1 in POMC and AgRP neurons regulates glucose homeostasis but not feeding behavior in mice[J].Cell Reports, 2015, 11(3):335-343.  
[15] LIU Y Y, LI F N, KONG X F, et al.Signaling pathways related to protein synthesis and amino acid concentration in pig skeletal muscles depend on the dietary protein level, genotype and developmental stages[J].PLoS One, 2015, 10(9):e0138277.
[16] GWINN D M, SHACKELFORD D B, EGAN D F, et al.AMPK phosphorylation of raptor mediates a metabolic checkpoint[J].Molecular Cell, 2008, 30(2):214-226.  
[17] INOKI K, ZHU T Q, GUAN K L.TSC2 mediates cellular energy response to control cell growth and survival[J].Cell, 2003, 115(5):577-590.  
[18] HAISSAGUERRE M, SAUCISSE N, COTA D.Influence of mTOR in energy and metabolic homeostasis[J].Molecular and Cellular Endocrinology, 2014, 397(1/2):67-77.
[19] LAEGER T, REED S D, HENAGAN T M, et al.Leucine acts in the brain to suppress food intake but does not function as a physiological signal of low dietary protein[J].American Journal of Physiology:Regulatory, Integrative and Comparative Physiology, 2014, 307(3):R310-R320.
[20] VON HOLSTEIN-RATHLOU S, BONDURANT L D, PELTEKIAN L, et al.FGF21 mediates endocrine control of simple sugar intake and sweet taste preference by the liver[J].Cell Metabolism, 2016, 23(2):335-343.  
[21] FON TACER K, BOOKOUT A L, DING X S, et al.Research resource:comprehensive expression atlas of the fibroblast growth factor system in adult mouse[J].Molecular Endocrinology, 2010, 24(10):2050-2064.  
[22] MASTER A, NAUMAN A.Molecular mechanisms of protein biosynthesis initiation-biochemical and biomedical implications of a new model of translation enhanced by the RNA hypoxia response element (rHRE)[J].Postepy Biochemii, 2014, 60(1):39-54.
[23] DENNIS P B, JAESCHKE A, SAITOH M, et al.Mammalian TOR:a homeostatic ATP sensor[J].Science, 2001, 294(5544):1102-1105.  
[24] 刘新伟.凡纳滨对虾TOR信号通路及其2个重要成员的功能研究[D].硕士学位论文.青岛:中国科学院大学, 2018. LIU X W.Study on TOR signaling pathway and functional analysis of two important factors in Litopenaeus vannamei[D].Master's Thesis.Qingdao:University of Chinese Academy of Sciences, 2018.(in Chinese)
[25] 孟艳梅.慢性粒细胞白血病患者SHIP基因及4EBP1基因的表达及其意义[D].硕士学位论文.石家庄:河北医科大学, 2014. MENG Y M.The significance and expression of SHIP gene and 4EBP1 gene in chronic myeloid leukemia patients[D].Master's Thesis.Shijiazhuang:Hebei Medical University, 2014.(in Chinese)
[26] 孟兰佳.Bufalin对人食管癌细胞4EBP1活化及凋亡的影响[D].硕士学位论文.石家庄:河北医科大学, 2014. MENG L J.The effect of Bufalin on the activation of 4EBP1 and cell apoptosis in human esophageal cancer cells[D].Master's Thesis.Shijiazhuang:Hebei Medical University, 2014.(in Chinese)
[27] INOKI K, OUYANG H J, LI Y, et al.Signaling by target of rapamycin proteins in cell growth control[J].Microbiology and Molecular Biology Reviews, 2005, 69(1):79-100.  
[28] PERVIN S, SINGH R, HERNANDEZ E, et al.Nitric oxide in physiologic concentrations targets the translational machinery to increase the proliferation of human breast cancer cells:involvement of mammalian target of rapamycin/eIF4E pathway[J].Cancer Research, 2007, 67(1):289-299.  
[29] YUAN C, DING Y, HE Q, et al.L-arginine upregulates the gene expression of target of rapamycin signaling pathway and stimulates protein synthesis in chicken intestinal epithelial cells[J].Poultry Science, 2015, 94(5):1043-1051.  
[30] WU L H, LIANG H L, HAMUNJO C M K, et al.Culture salinity alters dietary protein requirement, whole body composition and nutrients metabolism related genes expression in juvenile genetically improved farmed tilapia (GIFT) (Oreochromis niloticus)[J].Aquaculture, 2021, 531:735961.
[31] LAPLANTE M, SABATINI D M.An emerging role of mTOR in lipid biosynthesis[J].Current Biology, 2009, 19(22):R1046-R1052.
[32] GAGNON A, LAU S, SORISKY A.Rapamycin-sensitive phase of 3T3-L1 preadipocyte differentiation after clonal expansion[J].Journal of Cellular Physiology, 2001, 189(1):14-22.  
[33] ZHANG H H, HUANG J X, DVVEL K, et al.Insulin stimulates adipogenesis through the Akt-TSC2-mTORC1 pathway[J].PLoS One, 2009, 4(7):e6189.
[34] CHAKRABARTI P, ENGLISH T, SHI J, et al.Mammalian target of rapamycin complex 1 suppresses lipolysis, stimulates lipogenesis, and promotes fat storage[J].Diabetes, 2010, 59(4):775-781.  
[35] CARNEVALLI L S, MASUDA K, FRIGERIO F, et al.S6K1 plays a critical role in early adipocyte differentiation[J].Developmental Cell, 2010, 18(5):763-774.  
[36] 周招洪.饲粮能量和精氨酸水平对育肥猪肉品质和脂肪代谢的影响[D].硕士学位论文.雅安:四川农业大学, 2014. ZHOU Z H.Effects of dietary energy and arginine levels on pork quality and lipid metabolism of finishing pigs[D].Master's Thesis.Ya'an:Sichuan Agricultural University, 2014.(in Chinese)
[37] BISPHAM J, GARDNER D S, GNANALINGHAM M G, et al.Maternal nutritional programming of fetal adipose tissue development:differential effects on messenger ribonucleic acid abundance for uncoupling proteins and peroxisome proliferator-activated and prolactin receptors[J].Endocrinology, 2005, 146(9):3943-3949.  
[38] LE BACQUER O, PETROULAKIS E, PAGLIALUNGA S, et al.Elevated sensitivity to diet-induced obesity and insulin resistance in mice lacking 4E-BP1 and 4E-BP2[J].The Journal of Clinical Investigation, 2007, 117(2):387-396.  
[39] UM S H, FRIGERIO F, WATANABE M, et al.Absence of S6K1 protects against age- and diet-induced obesity while enhancing insulin sensitivity[J].Nature, 2004, 431(7005):200-205.  
[40] DVVEL K, YECIES J L, MENON S, et al.Activation of a metabolic gene regulatory network downstream of mTOR complex 1[J].Molecular Cell, 2010, 39(2):171-183.  
[41] TIAN J, GOLDSTEIN J L, BROWN M S.Insulin induction of SREBP-1c in rodent liver requires LXRα-C/EBPβ complex[J].Proceedings of the National Academy of Sciences of the United States of America, 2016, 113(29):8182-8187.  
[42] HAAS J T, MIAO J, CHANDA D, et al.Hepatic insulin signaling is required for obesity-dependent expression of SREBP-1c mRNA but not for feeding-dependent expression[J].Cell Metabolism, 2012, 15(6):873-884.  
[43] 赵迪, 朱燕婷, 史道华.mTOR介导转录因子调控细胞糖脂代谢的研究进展[J].基础医学与临床, 2014, 34(11):1574-1577. ZHAO D, ZHU Y T, SHI D H.Research progress in cell metabolism of glucose and lipid regulated by transcription factors via mTOR[J].Basic & Clinical Medicine, 2014, 34(11):1574-1577.(in Chinese)
[44] KUMAR A, LAWRENCE J C, Jr, JUNG D Y, et al.Fat cell-specific ablation of Rictor in mice impairs insulin-regulated fat cell and whole-body glucose and lipid metabolism[J].Diabetes, 2010, 59(6):1397-1406.  
[45] TANG Y F, WALLACE M, SANCHEZ-GURMACHES J, et al.Adipose tissue mTORC2 regulates ChREBP-driven de novo lipogenesis and hepatic glucose metabolism[J].Nature Communications, 2016, 7:11365.
[46] HAGIWARA A, CORNU M, CYBULSKI N, et al.Hepatic mTORC2 activates glycolysis and lipogenesis through Akt, glucokinase, and SREBP1c[J].Cell Metabolism, 2012, 15(5):725-738.  
[47] SEILIEZ I, MÉDALE F, AGUIRRE P, et al.Postprandial regulation of growth- and metabolism-related factors in zebrafish[J].Zebrafish, 2013, 10(2):237-248.  
[48] DAI W W, PANSERAT S, MENNIGEN J A, et al.Post-prandial regulation of hepatic glucokinase and lipogenesis requires the activation of TORC1 signalling in rainbow trout (Oncorhynchus mykiss)[J].Journal of Experimental Biology, 2013, 216(23):4483-4492.
[49] 黄皓琰.N-氨甲酰谷氨酸在花鲈饲料中的有效性和耐受评价及其调控营养代谢的机制研究[D].硕士学位论文.北京:中国农业科学院, 2019. HUANG H Y.Efficacy and tolerance evaluation of N-carbamylglutamate in Japanese sebass (Lateolabrax japonicas) diet and the related nutrient metabolism regulation mechanism[D].Master's Thesis.Beijing:Chinese Academy of Agricultural Sciences, 2019.(in Chinese)
[50] BRUGAROLAS J B, VAZQUEZ F, REDDY A, et al.TSC2 regulates VEGF through mTOR-dependent and-independent pathways[J].Cancer Cell, 2003, 4(2):147-158.  
[51] HUDSON C C, LIU M, CHIANG G G, et al.Regulation of hypoxia-inducible factor 1alpha expression and function by the mammalian target of rapamycin[J].Molecular and Cellular Biology, 2002, 22(20):7004-7014.  
[52] LAUGHNER E, TAGHAVI P, CHILES K, et al.HER2(neu) signaling increases the rate of hypoxia-inducible factor 1alpha (HIF-1alpha) synthesis:novel mechanism for HIF-1-mediated vascular endothelial growth factor expression[J].Molecular and Cellular Biology, 2001, 21(12):3995-4004.  
[53] CUNNINGHAM J T, RODGERS J T, ARLOW D H, et al.mTOR controls mitochondrial oxidative function through a YY1-PGC-1alpha transcriptional complex[J].Nature, 2007, 450(7170):736-740.  
[54] LEFEBVRE P, CHINETTI G, FRUCHART J C, et al.Sorting out the roles of PPAR alpha in energy metabolism and vascular homeostasis[J].The Journal of Clinical Investigation, 2006, 116(3):571-580.  
[55] KIM K, PYO S, UM S H.S6 kinase 2 deficiency enhances ketone body production and increases peroxisome proliferator-activated receptor alpha activity in the liver[J].Hepatology, 2012, 55(6):1727-1737.  
[56] AL-JADA D N, AHMAD M N.Dietary fat and insulin resistance:a connection through leptin and PPARγ activation[J].Functional Foods in Health and Disease, 2016, 6(6):306-328.  
[57] 栾会玲, 王茉, 翁雨晴, 等.mTOR信号调节糖脂代谢的研究进展[J].海峡药学, 2018, 30(5):5-8. LUAN H L, WANG M, WENG Y Q, et al.Advances in mTOR signal regulation of glycolipid metabolism[J].Strait Pharmaceutical Journal, 2018, 30(5):5-8.(in Chinese)
[58] BRUCE K D, HANSON M A.The developmental origins, mechanisms, and implications of metabolic syndrome[J].The Journal of Nutrition, 2010, 140(3):648-652.  
[59] CHEATHAM B, KAHN C R.Insulin action and the insulin signaling network[J].Endocrine Reviews, 1995, 16(2):117-142.
[60] 唐小红, 樊佳佳, 于凌云, 等.鱼类糖酵解关键酶的研究进展[J].中国农学通报, 2014, 30(2):69-75. TANG X H, FAN J J, YU L Y, et al.Research advances in glycolytic key enzyme of fish[J].Chinese Agricultural Science Bulletin, 2014, 30(2):69-75.(in Chinese)
[61] WULLSCHLEGER S, LOEWITH R, HALL M N.TOR signaling in growth and metabolism[J].Cell, 2006, 124(3):471-484.  
[62] 梁化亮.精氨酸对团头鲂幼鱼生长、营养代谢和免疫功能的影响及作用机制[D].博士学位论文.南京:南京农业大学, 2019. LIANG H L.Effects of dietary arginine on growth, nutrient metabolism and immune capacity involved in fuctional mechanism in juvenile blunt snout bream, Megalobrama amblycephala[D].Ph.D.Thesis.Nanjing:Nanjing Agricultural University, 2019.(in Chinese)
[63] FUHRMANN A, LOPES P, SERENO J, et al.Molecular mechanisms underlying the effects of cyclosporin A and sirolimus on glucose and lipid metabolism in liver, skeletal muscle and adipose tissue in an in vivo rat model[J].Biochemical Pharmacology, 2014, 88(2):216-228.  
[64] 许戈阳, 刘芬婷, 沈哲民, 等.mTOR信号通路在糖代谢中作用[J].生理科学进展, 2015, 46(2):94-98. XU G Y, LIU F T, SHEN Z M, et al.The role of mTOR signaling pathway in glucose metabolism[J].Progress in Physiological Sciences, 2015, 46(2):94-98.(in Chinese)
[65] LANSARD M, PANSERAT S, PLAGNES-JUAN E, et al.Integration of insulin and amino acid signals that regulate hepatic metabolism-related gene expression in rainbow trout:role of TOR[J].Amino Acids, 2010, 39(3):801-810.  
[66] 辛芳, 王雷, 刘梅, 等.水产动物雷帕霉素受体信号通路的研究进展[J].海洋科学, 2016, 40(1):147-154. XIN F, WANG L, LIU M, et al.Mechanistic target of rapamycin signaling in aquatic animals[J].Marine Sciences, 2016, 40(1):147-154.(in Chinese)
[67] LANSARD M, PANSERAT S, PLAGNES-JUAN E, et al.L-leucine, L-methionine, and L-lysine are involved in the regulation of intermediary metabolism-related gene expression in rainbow trout hepatocytes[J].The Journal of Nutrition, 2011, 141(1):75-80.  
[68] 汪福保, 罗莉, 文华, 等.镁对草鱼生长、形体、肝功能和糖代谢的影响[J].淡水渔业, 2011, 41(2):57-62, 68. WANG F B, LUO L, WEN H, et al.Effects of dietary magnesium on the growth, body index, liver function and glucose metabolism of grass carp, Ctenopharyngodon idella[J].Freshwater Fisheries, 2011, 41(2):57-62, 68.(in Chinese)
[69] 陈银涛, 于秉治, 武迪迪.PI3K/Akt/mTOR信号通路及临床相关肿瘤抑制剂[J].中国生物化学与分子生物学报, 2014, 30(10):949-956. CHEN Y T, YU B Z, WU D D.PI3K/Akt/mTOR signaling and the related cancer inhibitors[J].Chinese Journal of Biochemistry and Molecular Biology, 2014, 30(10):949-956.(in Chinese)
[70] 陈洪菊, 屈艺, 母得志.mTOR信号通路的生物学功能[J].生命的化学, 2010, 30(4):555-561. CHEN H J, QU Y, MU D Z.The progress of study on the biological function of mTOR pathway[J].Chemistry of Life, 2010, 30(4):555-561.(in Chinese)
[71] ROMBOUT J H W M, ABELLI L, PICCHIETTI S, et al.Teleost intestinal immunology[J].Fish & Shellfish Immunology, 2011, 31(5):616-626.  
[72] 杨冰贞, 张民, 王克坚.NF-κB信号通路在鱼类先天性免疫中的作用[J].生物技术通报, 2014(1):46-52. YANG B Z, ZHANG M, WANG K J.Role of NF-κB signal pathway in the innate immune system of fish[J].Biotechnology Bulletin, 2014(1):46-52.(in Chinese)
[73] ZHOU Q, JIN M, ELMADA Z C, 等.饲料中不同精氨酸水平对黄鲶幼鱼(黄颡鱼)生长、免疫力以及抗嗜水气单胞菌能力的影响[J].饲料博览, 2015(2):46. ZHOU Q, JIN M, ELMADA Z C, et al.Effects of dietary arginine levels on growth, immune ability and resistance to Aeromonas hydrophilus of juvenile yellow catfish (Pelteobagrus fulvidraco)[J].Feed Review, 2015(2):46.(in Chinese)
[74] 韩凤禄, 张琴, 黄国强, 等.斜带石斑鱼幼鱼的饲料精氨酸需求量[J].中国水产科学, 2016, 23(3):584-593. HAN F L, ZHANG Q, HUANG G Q, et al.Requirement of dietary arginine for juvenile orange-spotted grouper, Epinephelus coioides[J].Journal of Fishery Sciences of China, 2016, 23(3):584-593.(in Chinese)
[75] LI P, YIN Y L, LI D F, et al.Amino acids and immune function[J].The British Journal of Nutrition, 2007, 98(2):237-252.  
[76] 王标.精氨酸对中期草鱼肉质和铜诱导的鳃屏障功能的影响[D].硕士学位论文.雅安:四川农业大学, 2014. WANG B.The effect of dietary arginine supplement on flesh quality and Cu-induced gill barrier function of young grass carp (Ctenopharyngodon idellus)[D].Master's Thesis.Ya'an:Sichuan Agricultural University, 2014.(in Chinese)
[77] LIANG H L, MOKRANI A, JI K, et al.Effects of dietary arginine on intestinal antioxidant status and immunity involved in Nrf2 and NF-κB signaling pathway in juvenile blunt snout bream, Megalobrama amblycephala[J].Fish & Shellfish Immunology, 2018, 82:243-249.
[78] 胡欣欣.γ射线照射对蛋白质结构和功能影响的研究[D].硕士学位论文.济南:山东大学, 2015. HU X X.Studies on the effect of γ-radiation on the conformation and function of protein[D].Master's Thesis.Jinan:Shandong University, 2015.(in Chinese)
[79] ZHANG W B, CHEN Q Y, MAI K S, et al.Effects of dietary α-lipoic acid on the growth and antioxidative responses of juvenile abalone Haliotis discus hannai Ino[J].Aquaculture Research, 2010, 41(11):e781-e787.
[80] TOKUR B, KORKMAZ K.The effects of an iron-catalyzed oxidation system on lipids and proteins of dark muscle fish[J].Food Chemistry, 2007, 104(2):754-760.  
[81] 李文.苯丙氨酸对生长中期草鱼生长性能、肌肉品质和肠道黏膜免疫功能的影响研究[D].硕士学位论文.雅安:四川农业大学, 2014. LI W.The effect of dietary phenylalanine supplement on growth, flesh quality parameters, antioxidant capacity and intestine immune function of young grass carp (Ctenopharyngodon idellus)[D].Master's Thesis.Ya'an:Sichuan Agricultural University, 2014.(in Chinese)
[82] LIANG H L, MOKRANI A, JI K, et al.Dietary leucine modulates growth performance, Nrf2 antioxidant signaling pathway and immune response of juvenile blunt snout bream (Megalobrama amblycephala)[J].Fish & Shellfish Immunology, 2018, 73:57-65.
[83] SEILIEZ I, GABILLARD J C, SKIBA-CASSY S, et al.An in vivo and in vitro assessment of TOR signaling cascade in rainbow trout (Oncorhynchus mykiss)[J].American Journal of Physiology:Regulatory, Integrative and Comparative Physiology, 2008, 295(1):R329-R335.
[84] NUTTALL F Q, SCHWEIM K J, GANNON M C.Effect of orally administered phenylalanine with and without glucose on insulin, glucagon and glucose concentrations[J].Hormone and Metabolic Research, 2006, 38(8):518-523.  
[85] WANG X M, PROUD C G.The mTOR pathway in the control of protein synthesis[J].Physiology, 2006, 21(5):362-369.  
[86] 黄陈翠, 孙健, 吉红, 等.硫辛酸对草鱼脂肪细胞脂质含量及脂代谢相关基因表达的影响[J].淡水渔业, 2020, 50(1):87-92. HUANG C C, SUN J, JI H, et al.Effect of α-lipoic acid on lipid content and lipid metabolism related gene expression in Ctenopharyngodon idellus adipocyte[J].Freshwater Fisheries, 2020, 50(1):87-92.(in Chinese)
[87] 陈冰, 杨继华, 曹俊明, 等.桑叶黄酮对吉富罗非鱼肌肉抗氧化指标及营养组成的影响[J].淡水渔业, 2018, 48(3):90-95. CHEN B, YANG J H, CAO J M, et al.Effects of dietary mulberry leaf flavonoids on muscle antioxidant indices and nutritional compositions of GIFT, Oreochromis niloticus[J].Freshwater Fisheries, 2018, 48(3):90-95.(in Chinese)
文章导航

/