银鲫crh基因的克隆、组织表达谱及其对摄食的影响
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摘要
为研究crh (Corticotropin-releasing hormone)基因对鱼类摄食的调控作用,首次克隆了银鲫(Carassius auratus gibelio) crh基因c DNA全长序列。运用实时荧光定量PCR(RT-qPCR)技术检测crh基因m RNA在不同组织的表达及餐前、餐后和禁食对其表达的影响。结果显示,银鲫crh基因的c DNA序列全长为920 bp,其中,5’-UTR为53 bp,3’-UTR为378 bp,开放阅读框(Open reading frame,ORF)为489 bp。推导银鲫crh基因的ORF区编码蛋白由162个氨基酸组成,其中,含有11个氨基酸的保守区,24个氨基酸的信号肽,41个氨基酸的成熟肽。氨基酸序列的多重比较分析显示,银鲫crh基因与金鱼(Carassiusauratus)的同源性为99%,与鲤(Cyprinuscarpio)的同源性为96%。荧光定量PCR表达分析显示,银鲫crh基因在下丘脑中的表达量最高,其次是端脑和心脏。crh基因的表达量在餐前与餐后没有显著变化(P>0.05),在禁食第5天、第7天出现极显著降低(P<0.01),而在复投喂后第9天、第11天出现极显著升高(P<0.01)。以上结果显示,crh基因在银鲫摄食调控方面可能扮演着重要角色。
To elaborate the mechanism of corticotropin-releasing hormone(crh) on feeding in gibel carp(Carassius auratus gibelio), we cloned and identified the full-length cDNA sequence of crh for the first time. We also employed real-time quantitative PCR(RT-PCR) to explored crh mRNAon distribution in different tissues and investigated the expression of crh mRNA during postprandial and fasting. Here, a920 bp full-length of crh gene cDNA was obtained, including a 53 bp of 5’-UTR, a 378 bp of 3’-UTR, and a 489 bp of an open reading frame(ORF). The ORF of the gibel carp crh gene contains 162 bp nucleotides, including an 11-amino acid conserved region, a 24-amino acid signal peptide, and a 41-amino acid mature peptide. The amino acid sequence of crh shares a high level of similarity with teleost CRH(99% with Carassius auratus and 96% with Cyprinus carpio). The expression of gibel carp crh was observed in almost all tissues, with the highest expression detected in the hypothalamus, followed by the myelencephalon and heart. There was no significant change in crh gene expression in the fed group compared with the fasted group in gibel carp hypothalamus after a meal(P>0.05). However, the expression level of the crh gene showed a significant decrease after fasting on day 5 and day 7(P<0.01),a significant increase of the crh gene was observed after refeeding on day 9 and day 11. The results suggest that the crh gene may play a critical role in feeding regulation in gibel carp.
To elaborate the mechanism of corticotropin-releasing hormone(crh) on feeding in gibel carp(Carassius auratus gibelio), we cloned and identified the full-length cDNA sequence of crh for the first time. We also employed real-time quantitative PCR(RT-PCR) to explored crh mRNAon distribution in different tissues and investigated the expression of crh mRNA during postprandial and fasting. Here, a920 bp full-length of crh gene cDNA was obtained, including a 53 bp of 5’-UTR, a 378 bp of 3’-UTR, and a 489 bp of an open reading frame(ORF). The ORF of the gibel carp crh gene contains 162 bp nucleotides, including an 11-amino acid conserved region, a 24-amino acid signal peptide, and a 41-amino acid mature peptide. The amino acid sequence of crh shares a high level of similarity with teleost CRH(99% with Carassius auratus and 96% with Cyprinus carpio). The expression of gibel carp crh was observed in almost all tissues, with the highest expression detected in the hypothalamus, followed by the myelencephalon and heart. There was no significant change in crh gene expression in the fed group compared with the fasted group in gibel carp hypothalamus after a meal(P>0.05). However, the expression level of the crh gene showed a significant decrease after fasting on day 5 and day 7(P<0.01),a significant increase of the crh gene was observed after refeeding on day 9 and day 11. The results suggest that the crh gene may play a critical role in feeding regulation in gibel carp.
引文
Bernier NJ,Lin X,Peter RE.Differential expression of corticotropin-releasing factor(CRF)and urotensin Iprecursor genes,and evidence of CRF gene expression regulated by cortisol in goldfish brain.General and Comparative Endocrinology,1999,116(3):461-477
Chabbi A,Ganesh CB.Glucocorticoid synthesis inhibitor metyrapone blocks stress-induced suppression along luteinizing hormone secreting cells-ovary axis in the fish Oreochromis mossambicus.Journal of Experimental Zoology Part A Ecological and Integrative Physiology,2014321(3):125-134
Chandrasekar G,Lauterg,Hauptmann G.Distribution of corticotropin-releasing hormone in the developing zebrafish brain.Journal of Comparative Neurology,2010,505(4):337-351
Chen K,Xi BW,Teng T,et al.Molecular cloning,characterization,and regulation of Megalobrama amblycephala corticotropin-releasing hormone receptor expression.Journal of Fishery Sciences of China,2016,23(4):833-845[陈凯,习丙文,滕涛,等.团头鲂促肾上腺皮质激素释放激素受体基因序列特征及表达分析.中国水产科学,2016,23(4):833-845]
Chrousos GP,Zoumakis E.Milestones in CRH Research.Current Molecular Pharmacology,2017,10(4):259
Doyon C,Leclair J,Trudeau VL,et al.Corticotropin-releasing factor and neuropeptide Y m RNA levels are modified by glucocorticoids in rainbow trout,Oncorhynchus mykiss.General and Comparative Endocrinology,2006,146(2):126-135
Fang X,Teng SQ,Yang JL,et al.Pond high-yield,efficient and healthy aquaculture technology of Carassius auratus gibelio China Fisheries,2017(3):91-92[方旭,滕淑芹,杨建利,等."中科3号"异育银鲫池塘高产高效健康养殖技术.中国水产,2017(3):91-92]
Fuentes EN,Kling P,Einarsdotti IE,et al.Plasma leptin and growth hormone levels in the fine flounder(Paralichthys adspersus)increase gradually during fasting and decline rapidly after refeeding.General and Comparative Endocrinology,2012,177(1):120
Fuzesi T,Daviu N,Wamsteeker CJI,et al.Hypothalamic CRHneurons orchestrate complex behaviours after stress.Nature Communications,2016,7:11937
Graf C,Kuehne C,Panhuysen M,et al.Corticotropin-releasing hormone regulates common target genes with divergent functions in corticotrope and neuronal cells.Molecular and Cellular Endocrinology,2012,362(1-2):29-38
Han J,Liu B,Jiang Z,et al.cDNA cloning of neuropeptide Ygene of turbot and its expression analysis for the strategy of starvation and feeding in industrial culture.Engineering Sciences,2014-09
Hoskins LJ,Volkoff H.The comparative endocrinology of feeding in fish:Insights and challenges.General and Comparative Endocrinology,2012,176(3):327-335
Huising MO,Lm VDA,Metz JR,et al.Corticotropin-releasing factor(CRF)and CRF-binding protein expression in and release from the head kidney of common carp:evolutionary conservation of the adrenal CRF system.Journal of Endocrinology,2007,193(3):349-357
Huising MO,Metz JR,Van SC,et al.Structural characterisation of a cyprinid(Cyprinus carpio L.)CRH,CRH-BP and CRH-R1,and the role of these proteins in the acute stress response.Journal of Molecular Endocrinology,2004,32(3):627-648
Jeong KH,Jacobson L,Widmaier EP,et al.Normal suppression of the reproductive axis following stress in corticotropinreleasing hormone-deficient mice.Endocrinology,1999,140(4):1702-1708
Jonsson E,Kaiya H,Bjornsson BT.Ghrelin decreases food intake in juvenile rainbow trout(Oncorhynchus mykiss)through the central anorexigenic corticotropin-releasing factor system.General and Comparative Endocrinology,2010,166(1):39
Liu L,Liang XF,Li J,et al.Feed intake,feed utilization and feeding-related gene expression response to dietary phytic acid for juvenile grass carp(Ctenopharyngodon idellus).Aquaculture,2014,424-425(2):201-206
Lu RH,Sun JJ,Liang XF,et al.Biological characteristics and functions of Leptin in fish.Acta Hydrobiologica Sinica,2015,39(3):583-589[卢荣华,孙君君,梁旭方,等.鱼类leptin的生物学特性及功能.水生生物学报,2015,39(3):583-589]
Lu W,Dow L,Gumusgoz S,et al.Coexpression of corticotropin-releasing hormone and urotensin i precursor genes in the caudal neurosecretory system of the euryhaline flounder(Platichthys flesus):A possible shared role in peripheral regulation.Endocrinology,2004,145(12):5786
Matsuda K,Hagiwara Y,Shibata H,et al.Ovine corticotropinreleasing hormone(oCRH)exerts an anxiogenic-like action in the goldfish,Carassius auratus.General and Comparative Endocrinology,2013,188(1):118-122
Matsuda K,Maruyama K.Regulation of feeding behavior by pituitary adenylate cyclase-activating polypeptide(PACAP)and vasoactive intestinal polypeptide(VIP)in vertebrates Peptides,2007,28(9):1761-1766
Matsuda K,Miura T,Uchiyama M,et al.Relationship between anorexigenic action of pituitary adenylate cyclase-activating polypeptide(PACAP)and that of corticotropin-releasing hormone(CRH)in the goldfish(Carassius auratus).Peptides,2006,27(7):1820-1826
Matsuda K.Regulation of feeding behavior and psychomotor activity by corticotropin-releasing hormone(CRH)in fish.Frontiers in Neuroscience,2013,7(7):91
Mazon AF,Verburg-van Kemenade BM,Flik G,et al.Corticotropin-releasing hormone-receptor 1(CRH-R1)and CRH-binding protein(CRH-BP)are expressed in the gills and skin of common carp Cyprinus carpio L.and respond to acute stress and infection.Journal of Experimental Biology,2006,209(3):510-517
Meyer A,Vav DPY.From 2R to 3R:Evidence for a fish-specific genome duplication(FSGD).Bioessays,2005,27(9):937-945
Michel M,Page-Mccaw PS,Chen W,et al.Leptin signaling regulates glucose homeostasis,but not adipostasis,in the zebrafish.Proceedings of the National Academy of Sciences of the United States of America,2016,113(11):3084
Mu FS,Miao L,Li MY,et al.Cloning and expression of Leptin gene in Acrossocheilus fasciatus fasciatus during fasting and refeeding.Oceanologia et Limnologia Sinica,2017,48(4):822-829[穆方申,苗亮,李明云,等.光唇鱼(Acrossocheilus fasciatus)Leptin基因克隆及禁食-再投喂对其表达的影响.海洋与湖沼,2017,48(4):822-829]
Mu XS.Gene cloning,specific expression in different tissues and recombinant protein preparation of 4 biomarkers in Scophthalmus maximus.Master′s Thesis of Ocean University of China,2013,91-92[穆小生.大菱鲆4种生物标志物基因的克隆、组织特异性表达研究及重组蛋白制备.中国海洋大学硕士研究生学位论文,2013,91-92]
Qin XH,Wu YJ,Shi XT,et al.Effects of light intensity on the hatching rate of fertilized Schizothorax chongi eggs and on the growth and feeding of larvae.Journal of Hydroecology,2017,38(5):97-102[秦希获,邬玉娇,石小涛,等.光照对细鳞裂腹鱼受精卵孵化率及仔鱼生长、摄食的影响.水生态学杂志,2017,38(5):97-102]
Rabasa C,Dickson S L.Impact of stress on metabolism and energy balance.Current Opinion in Behavioral Sciences,2016,9(9):71-77
Rayatpour A,Ghasemi M,Radahmadi M,et al.Effect of intraparaventricular administration of corticotropin releasing hormone on food intake in food-deprived rats.Journal of Isfahan Medical School,2017,35(436):70-775
Shen JJ,LüWQ.Effects of thyroid hormone on CRH gene expression in Paralichthys olivaceus at early development stage.Journal of Southern Agriculture,2017,48(2):323-327[沈娇娇,吕为群.褐牙鲆早期发育阶段CRH基因表达及受甲状腺激素的影响.南方农业学报,2017,48(2):323-327]
Simpson EH,Balsam PD.Behavioral neuroscience of motivation Springer International Publishing.2016:1-12
Sundstrom G,Larsson TA,Brenner S,et al.Evolution of the neuropeptide Y family:New genes by chromosome duplications in early vertebrates and in teleost fishes.General and Comparative Endocrinology,2008,155(3):705
Tort L.Stress and immune modulation in fish.Developmental and Comparative Immunology,2011,35(12):1366
Vale W,Spiess J,Rivier C,et al.Characterization of a 41-residue ovine hypothalamic peptide that stimulates secretion of corticotropin and beta-endorphin.Science,1981,213(4514):1394
Vamvakopoulos NC,Chrousos GP.Evidence of direct estrogenic regulation of human corticotropin-releasing hormone gene expression.Journal of Clinical Investigation,2013,92(4):1896
Volkoff H.The neuroendocrine regulation of food intake in fish:A review of current knowledge.Frontiers in Neuroscience,2016,10:540
Wang GW,Shao BP,Ding YP.The orexigenic role and mechanism of neuropeptide Y.Chinese Journal of Animal Nutrition,2016(2):385-391[王国文,邵宝平,丁艳平.神经肽Y的促摄食作用及其调控机制.动物营养学报,2016(2):385-391]
Wang T,Zhou C,Yuan D,et al.Schizothorax prenanti,corticotropin-releasing hormone(CRH):Molecular cloning,tissue expression,and the function of feeding regulation.Fish Physiology and Biochemistry,2014,40(5):1407-1415
Wang Y,Lu DQ,Li WP,et al.The effect of acute handling stress on the stress-related neuroendocrine factor in Coilia nasus.Journal of Fisheries of China,2014,38(6):803-812[王宇,卢丹琪,李伟萍,等.急性操作胁迫对刀鲚应激反应相关神经内分泌因子的影响.水产学报,2014,38(6):803-812]
Yin XP,Su JM,Pu L,et al.Research on the effect of corticotropin-releasing hormone in stress reaction in the fishes.Chinese Journal of Fisheries,2014(3):57-61[殷新平,苏建明,蒲龄,等.促肾上腺皮质激素释放激素在鱼类应激反应中的作用.水产学杂志,2014(3):57-61]
Yin XP.Cloning and expression of CRH,CRH-R1 from Ctenopharyngodon idella.Master′s Thesis of Hunan Agricultural University,2015,42-43[殷新平.草鱼CRH、CRH-R1基因的全长克隆及表达分析.湖南农业大学硕士研究生学位论文,2015,42-43]
Yin ZY,Xue M,Zheng YH,et al.Effect of yeast extract as feeding stimulants on diet preference of Gibel carp.Feed Industry,2017,38(8):4-8[尹子煜,薛敏,郑银桦,等.酵母提取物在异育银鲫饲料中促摄食作用的研究.饲料工业,2017,38(8):4-8]
You CH,Lu YB,Wang SQ,et al.Cloning of gene encoding corticotrophin-releasing hormone(CRH)and its mRNAexpression after shallow water stress in Siganus canaliculatus.Journal of Shantou University(Natural Science),2014,29(2):43-51[游翠红,卢玉标,王树启,等.黄斑蓝子鱼CRH基因的克隆及其在浅水应激后的m RNA表达.汕头大学学报(自然科学版),2014,29(2):43-51]
Zhang H,Chen H,Zhang Y,et al.Molecular cloning,characterization and expression profiles of multiple leptin genes and a leptin receptor gene in orange-spotted grouper(Epinephelus coioides).General and Comparative Endocrinology,2013,181(1):295-305
Zhang X,Tang N,Qi J,et al.CCK reduces the food intake mainly through CCK1R in Siberian sturgeon(Acipenser baerii Brandt).Scientific Reports,2017,7(1):12413
Zhang YZ,Zheng ZM,Hua JQ,et al.Bio-floc nutrition and nitrogen uptake by shrimp and silver prussian carp in polyculture brackish water.Journal of Ningbo University,2016,29(3):29-33[张亚卓,郑忠明,华建权,等.低盐度条件下生物絮团的营养组分及凡纳滨对虾和银鲫对其摄食效率的研究.宁波大学学报(理工版),2016,29(3):29-33]
Zhou C,Zhang X,Liu T,et al.Schizothorax davidi ghrelin:cDNA cloning,tissue distribution and indication for its stimulatory character in food intake.Gene,2014,534(1):72
Zhou C,Zheng J,Luo L,et al.Evidence that ghrelin,may be associated with the food intake of gibel carp(Carassius auratus gibelio).Fish Physiology and Biochemistry,2016,42(6):1-10
Zhou M,Di R,Hu WP.Different expression of CRH and POMCgenes in reproductive related tissues in small-tail han sheep and sunite sheep.China Herbivore Science,2017,37(4):1-4[周梅,狄冉,胡文萍,等.小尾寒羊和苏尼特羊CRH和POMC基因在繁殖相关组织中的差异性表达.中国草食动物科学,2017,37(4):1-4]
Chabbi A,Ganesh CB.Glucocorticoid synthesis inhibitor metyrapone blocks stress-induced suppression along luteinizing hormone secreting cells-ovary axis in the fish Oreochromis mossambicus.Journal of Experimental Zoology Part A Ecological and Integrative Physiology,2014321(3):125-134
Chandrasekar G,Lauterg,Hauptmann G.Distribution of corticotropin-releasing hormone in the developing zebrafish brain.Journal of Comparative Neurology,2010,505(4):337-351
Chen K,Xi BW,Teng T,et al.Molecular cloning,characterization,and regulation of Megalobrama amblycephala corticotropin-releasing hormone receptor expression.Journal of Fishery Sciences of China,2016,23(4):833-845[陈凯,习丙文,滕涛,等.团头鲂促肾上腺皮质激素释放激素受体基因序列特征及表达分析.中国水产科学,2016,23(4):833-845]
Chrousos GP,Zoumakis E.Milestones in CRH Research.Current Molecular Pharmacology,2017,10(4):259
Doyon C,Leclair J,Trudeau VL,et al.Corticotropin-releasing factor and neuropeptide Y m RNA levels are modified by glucocorticoids in rainbow trout,Oncorhynchus mykiss.General and Comparative Endocrinology,2006,146(2):126-135
Fang X,Teng SQ,Yang JL,et al.Pond high-yield,efficient and healthy aquaculture technology of Carassius auratus gibelio China Fisheries,2017(3):91-92[方旭,滕淑芹,杨建利,等."中科3号"异育银鲫池塘高产高效健康养殖技术.中国水产,2017(3):91-92]
Fuentes EN,Kling P,Einarsdotti IE,et al.Plasma leptin and growth hormone levels in the fine flounder(Paralichthys adspersus)increase gradually during fasting and decline rapidly after refeeding.General and Comparative Endocrinology,2012,177(1):120
Fuzesi T,Daviu N,Wamsteeker CJI,et al.Hypothalamic CRHneurons orchestrate complex behaviours after stress.Nature Communications,2016,7:11937
Graf C,Kuehne C,Panhuysen M,et al.Corticotropin-releasing hormone regulates common target genes with divergent functions in corticotrope and neuronal cells.Molecular and Cellular Endocrinology,2012,362(1-2):29-38
Han J,Liu B,Jiang Z,et al.cDNA cloning of neuropeptide Ygene of turbot and its expression analysis for the strategy of starvation and feeding in industrial culture.Engineering Sciences,2014-09
Hoskins LJ,Volkoff H.The comparative endocrinology of feeding in fish:Insights and challenges.General and Comparative Endocrinology,2012,176(3):327-335
Huising MO,Lm VDA,Metz JR,et al.Corticotropin-releasing factor(CRF)and CRF-binding protein expression in and release from the head kidney of common carp:evolutionary conservation of the adrenal CRF system.Journal of Endocrinology,2007,193(3):349-357
Huising MO,Metz JR,Van SC,et al.Structural characterisation of a cyprinid(Cyprinus carpio L.)CRH,CRH-BP and CRH-R1,and the role of these proteins in the acute stress response.Journal of Molecular Endocrinology,2004,32(3):627-648
Jeong KH,Jacobson L,Widmaier EP,et al.Normal suppression of the reproductive axis following stress in corticotropinreleasing hormone-deficient mice.Endocrinology,1999,140(4):1702-1708
Jonsson E,Kaiya H,Bjornsson BT.Ghrelin decreases food intake in juvenile rainbow trout(Oncorhynchus mykiss)through the central anorexigenic corticotropin-releasing factor system.General and Comparative Endocrinology,2010,166(1):39
Liu L,Liang XF,Li J,et al.Feed intake,feed utilization and feeding-related gene expression response to dietary phytic acid for juvenile grass carp(Ctenopharyngodon idellus).Aquaculture,2014,424-425(2):201-206
Lu RH,Sun JJ,Liang XF,et al.Biological characteristics and functions of Leptin in fish.Acta Hydrobiologica Sinica,2015,39(3):583-589[卢荣华,孙君君,梁旭方,等.鱼类leptin的生物学特性及功能.水生生物学报,2015,39(3):583-589]
Lu W,Dow L,Gumusgoz S,et al.Coexpression of corticotropin-releasing hormone and urotensin i precursor genes in the caudal neurosecretory system of the euryhaline flounder(Platichthys flesus):A possible shared role in peripheral regulation.Endocrinology,2004,145(12):5786
Matsuda K,Hagiwara Y,Shibata H,et al.Ovine corticotropinreleasing hormone(oCRH)exerts an anxiogenic-like action in the goldfish,Carassius auratus.General and Comparative Endocrinology,2013,188(1):118-122
Matsuda K,Maruyama K.Regulation of feeding behavior by pituitary adenylate cyclase-activating polypeptide(PACAP)and vasoactive intestinal polypeptide(VIP)in vertebrates Peptides,2007,28(9):1761-1766
Matsuda K,Miura T,Uchiyama M,et al.Relationship between anorexigenic action of pituitary adenylate cyclase-activating polypeptide(PACAP)and that of corticotropin-releasing hormone(CRH)in the goldfish(Carassius auratus).Peptides,2006,27(7):1820-1826
Matsuda K.Regulation of feeding behavior and psychomotor activity by corticotropin-releasing hormone(CRH)in fish.Frontiers in Neuroscience,2013,7(7):91
Mazon AF,Verburg-van Kemenade BM,Flik G,et al.Corticotropin-releasing hormone-receptor 1(CRH-R1)and CRH-binding protein(CRH-BP)are expressed in the gills and skin of common carp Cyprinus carpio L.and respond to acute stress and infection.Journal of Experimental Biology,2006,209(3):510-517
Meyer A,Vav DPY.From 2R to 3R:Evidence for a fish-specific genome duplication(FSGD).Bioessays,2005,27(9):937-945
Michel M,Page-Mccaw PS,Chen W,et al.Leptin signaling regulates glucose homeostasis,but not adipostasis,in the zebrafish.Proceedings of the National Academy of Sciences of the United States of America,2016,113(11):3084
Mu FS,Miao L,Li MY,et al.Cloning and expression of Leptin gene in Acrossocheilus fasciatus fasciatus during fasting and refeeding.Oceanologia et Limnologia Sinica,2017,48(4):822-829[穆方申,苗亮,李明云,等.光唇鱼(Acrossocheilus fasciatus)Leptin基因克隆及禁食-再投喂对其表达的影响.海洋与湖沼,2017,48(4):822-829]
Mu XS.Gene cloning,specific expression in different tissues and recombinant protein preparation of 4 biomarkers in Scophthalmus maximus.Master′s Thesis of Ocean University of China,2013,91-92[穆小生.大菱鲆4种生物标志物基因的克隆、组织特异性表达研究及重组蛋白制备.中国海洋大学硕士研究生学位论文,2013,91-92]
Qin XH,Wu YJ,Shi XT,et al.Effects of light intensity on the hatching rate of fertilized Schizothorax chongi eggs and on the growth and feeding of larvae.Journal of Hydroecology,2017,38(5):97-102[秦希获,邬玉娇,石小涛,等.光照对细鳞裂腹鱼受精卵孵化率及仔鱼生长、摄食的影响.水生态学杂志,2017,38(5):97-102]
Rabasa C,Dickson S L.Impact of stress on metabolism and energy balance.Current Opinion in Behavioral Sciences,2016,9(9):71-77
Rayatpour A,Ghasemi M,Radahmadi M,et al.Effect of intraparaventricular administration of corticotropin releasing hormone on food intake in food-deprived rats.Journal of Isfahan Medical School,2017,35(436):70-775
Shen JJ,LüWQ.Effects of thyroid hormone on CRH gene expression in Paralichthys olivaceus at early development stage.Journal of Southern Agriculture,2017,48(2):323-327[沈娇娇,吕为群.褐牙鲆早期发育阶段CRH基因表达及受甲状腺激素的影响.南方农业学报,2017,48(2):323-327]
Simpson EH,Balsam PD.Behavioral neuroscience of motivation Springer International Publishing.2016:1-12
Sundstrom G,Larsson TA,Brenner S,et al.Evolution of the neuropeptide Y family:New genes by chromosome duplications in early vertebrates and in teleost fishes.General and Comparative Endocrinology,2008,155(3):705
Tort L.Stress and immune modulation in fish.Developmental and Comparative Immunology,2011,35(12):1366
Vale W,Spiess J,Rivier C,et al.Characterization of a 41-residue ovine hypothalamic peptide that stimulates secretion of corticotropin and beta-endorphin.Science,1981,213(4514):1394
Vamvakopoulos NC,Chrousos GP.Evidence of direct estrogenic regulation of human corticotropin-releasing hormone gene expression.Journal of Clinical Investigation,2013,92(4):1896
Volkoff H.The neuroendocrine regulation of food intake in fish:A review of current knowledge.Frontiers in Neuroscience,2016,10:540
Wang GW,Shao BP,Ding YP.The orexigenic role and mechanism of neuropeptide Y.Chinese Journal of Animal Nutrition,2016(2):385-391[王国文,邵宝平,丁艳平.神经肽Y的促摄食作用及其调控机制.动物营养学报,2016(2):385-391]
Wang T,Zhou C,Yuan D,et al.Schizothorax prenanti,corticotropin-releasing hormone(CRH):Molecular cloning,tissue expression,and the function of feeding regulation.Fish Physiology and Biochemistry,2014,40(5):1407-1415
Wang Y,Lu DQ,Li WP,et al.The effect of acute handling stress on the stress-related neuroendocrine factor in Coilia nasus.Journal of Fisheries of China,2014,38(6):803-812[王宇,卢丹琪,李伟萍,等.急性操作胁迫对刀鲚应激反应相关神经内分泌因子的影响.水产学报,2014,38(6):803-812]
Yin XP,Su JM,Pu L,et al.Research on the effect of corticotropin-releasing hormone in stress reaction in the fishes.Chinese Journal of Fisheries,2014(3):57-61[殷新平,苏建明,蒲龄,等.促肾上腺皮质激素释放激素在鱼类应激反应中的作用.水产学杂志,2014(3):57-61]
Yin XP.Cloning and expression of CRH,CRH-R1 from Ctenopharyngodon idella.Master′s Thesis of Hunan Agricultural University,2015,42-43[殷新平.草鱼CRH、CRH-R1基因的全长克隆及表达分析.湖南农业大学硕士研究生学位论文,2015,42-43]
Yin ZY,Xue M,Zheng YH,et al.Effect of yeast extract as feeding stimulants on diet preference of Gibel carp.Feed Industry,2017,38(8):4-8[尹子煜,薛敏,郑银桦,等.酵母提取物在异育银鲫饲料中促摄食作用的研究.饲料工业,2017,38(8):4-8]
You CH,Lu YB,Wang SQ,et al.Cloning of gene encoding corticotrophin-releasing hormone(CRH)and its mRNAexpression after shallow water stress in Siganus canaliculatus.Journal of Shantou University(Natural Science),2014,29(2):43-51[游翠红,卢玉标,王树启,等.黄斑蓝子鱼CRH基因的克隆及其在浅水应激后的m RNA表达.汕头大学学报(自然科学版),2014,29(2):43-51]
Zhang H,Chen H,Zhang Y,et al.Molecular cloning,characterization and expression profiles of multiple leptin genes and a leptin receptor gene in orange-spotted grouper(Epinephelus coioides).General and Comparative Endocrinology,2013,181(1):295-305
Zhang X,Tang N,Qi J,et al.CCK reduces the food intake mainly through CCK1R in Siberian sturgeon(Acipenser baerii Brandt).Scientific Reports,2017,7(1):12413
Zhang YZ,Zheng ZM,Hua JQ,et al.Bio-floc nutrition and nitrogen uptake by shrimp and silver prussian carp in polyculture brackish water.Journal of Ningbo University,2016,29(3):29-33[张亚卓,郑忠明,华建权,等.低盐度条件下生物絮团的营养组分及凡纳滨对虾和银鲫对其摄食效率的研究.宁波大学学报(理工版),2016,29(3):29-33]
Zhou C,Zhang X,Liu T,et al.Schizothorax davidi ghrelin:cDNA cloning,tissue distribution and indication for its stimulatory character in food intake.Gene,2014,534(1):72
Zhou C,Zheng J,Luo L,et al.Evidence that ghrelin,may be associated with the food intake of gibel carp(Carassius auratus gibelio).Fish Physiology and Biochemistry,2016,42(6):1-10
Zhou M,Di R,Hu WP.Different expression of CRH and POMCgenes in reproductive related tissues in small-tail han sheep and sunite sheep.China Herbivore Science,2017,37(4):1-4[周梅,狄冉,胡文萍,等.小尾寒羊和苏尼特羊CRH和POMC基因在繁殖相关组织中的差异性表达.中国草食动物科学,2017,37(4):1-4]