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1 HEREDITAS (Beijing) , 33(7): ISSN 综述 DOI: /SP.J 鲟形目鱼类内分泌激素的分子生物学研究曹宏, 周荣家, 鲟鱼是处于软骨鱼类向硬骨鱼类过渡的一类古老物种, 被认为是研究脊椎动物起源和演化的重要物种之一目前, 绝大多数种类的鲟鱼都处于濒危状态, 因此, 采用各种手段来保证其物种的繁衍刻不容缓但是到目前为止, 关于鲟鱼的生长和生殖调控的分子机制相关信息, 依然十分有限文章就近年来对鲟鱼的各类内分泌激素基因的研究进展作一概述, 为进一步深入研究鲟鱼的生长和生殖调控的分子机制提供有益的参考资料鲟鱼 ; 内分泌激素 ; 生长 ; 生殖 Molecular biological study on hormones in Acipenseriformes CAO Hong, ZHOU Rong-Jia College of Life Sciences, Wuhan University, Wuhan , China Abstract: Sturgeons belong to the Acipenseriformes, which provide an ideal model for evolutionary studies due to their unique characters. At present, many species of Acipenseriformes are very rare and near extinct. It is urgent to protect these species. However, data on molecular mechanisms of their growth and reproduction regulation are still limited. Hormones are important factors involved in these processes. In this paper, we summarized recent research progresses in the hormones in sturgeon, which will provide valuable information for further studies on molecular mechanisms of growth, sexual development, and reproduction regulation in Acipenseriformes. Keywords: Acipenseriformes; hormone; growth; reproduction (Acipenseriformes),, ( 4.5~3.5 ),, (Chondrostei),,,,,, (Acipenser sinensis),,,,,,,,,, 收稿日期 : ; 修回日期 : 基金项目 : ( ) (973 ) ( 2010CB126306) 作者简介 :,, Tel: ; regancao@ihb.ac.cn 通讯作者 :,,, rjzhou@whu.edu.cn 网络出版时间 : :38:47 URL:

2 708 HEREDITAS (Beijing) ,, ; [1] 20 70,,,,,, [2], [1], [3], [4], [5],,, [6~8], (Acipenser gueldenstaedtii) (Acipenser baeri), (Acipenser transmontanus),,,,, 1 (Growth hormone, GH), (Insulin-like growth factor, IGF) GH,,,, [9] GH GH (Growth hormone receptor, GHR), GHR GHR,,, GH 80, Farmer [10],,, Yasuda [11] 2 GH, 3,, GH, GH GH, 63%~76%, GH 42%~ 63% Din [12] GH cdna, 980 bp, 642 bp, , GH mrna -I (IGF-I), cdna 445 bp, 396 bp, 132, 44, IGF-I,,, mrna GH, 1 5, IGF-I mrna Cao [6] GH, GH cdna 945 bp, 642 bp, 214, 1, GH mrna Western blotting, 1, GH, 2, GH, 1~2, 3, GH,, GH Liao [7] GHR (Chinese sturgeon GHR, csghr) csghr cdna csghr N- (FGDFS) ( )

3 7 : 709 Box 1 Box (CHO) csghr csghr, csghr 2.1(Spi 1.2) (seabream ) (seabream GH, sbgh), sbgh, CHO csghr, GH GHR csghr (Growth hormone binding protein, GHBP) csghbp, csghbp, csghr Asp Glu csghr, Asp Ala csghr, csghr, csghbp csghr GHR, Tarpey [13], GHR GH, GHR GHR GHR, GH, GHR, Fukamachi [14] (Huso dauricus) GHR, cdna bp, bp, 571, GHR (Protopterus dolloi) GHR, GHR, (Somatolactin receptor, SLR) SLR GHR,,, SLR, GHR SLR, GHR SLR 2 (Gonadotropin, GTH),, (Follicle-stimulating hormone, FSH), (Luteinizing hormone, LH) Moberg [15] FSHβ LHβ, GTH I GTH II,,, FSHβ LHβ,, LHβ FSHβ Quérat [16] FSHβ LHβ, FSHβ cdna bp, 128, 20 ; LHβ cdna 570 bp, 137, 22, FSHβ, FSHβ FSHβ ; LHβ LHβ, 4 ; 2.5, FSHβ LHβ Hurvitz [17] GTHα FSHβ LHβ, FSHβ mrna, LHβ mrna ;, LHβ mrna FSHβ mrna, 1, FSHβ mrna, 2, FSHβ mrna, 4 [18] (Acipenser schrenkii) 2 β, FSHβ, 99.1%;, 34.5% LHβ 98.3%;

4 710 HEREDITAS (Beijing) %,, 42.8% FSHβ LHβ Cao [8] GTHα FSHβ LHβ,, Western blotting 3 3, GTHα (Pars intermedia, PI) (Rostral pars distalis, RPD) (Proximal pars distalis, PPD); FSHβ PPD ; LHβ PI PPD RPD, FSHβ LHβ 1~5, Western blotting FSHβ, PPD RPD,, 3 (Somatolactin, SL) (Growth hormone, GH) (Prolactin, PRL), [19] SL Amemiya [20] SL cdna 881 bp, 696 bp, 232, , SL (Melanocyte stimulating hormone, MSH) SL, cdna bp, 696 bp, RT-PCR, SL, 3 ( ) 4, (Somatostatin, SS) ,, (SSR),, 1995, Nishii [21], GH Kim [22] (Scaphirhynchus albus), SS, Trabucchi [23] 2 SS, SS-14 [Pro(2)] SS-14,, 2 SS cdna, 116, 3 SS-14, PSS1 111, 3 [Pro(2)] SS-14, PSS2 PSS2 PSS2 67%, PSS1 ; PSS2 PSS2 mrna Adrio [24], SS, SS,, SS, SS, Li [25] SS, 2, AsSS1 AsSS2, 2, AsSS1, AsSS2,

5 7 : 711, AsSS2 mrna, 5,, AsSS2 AsSS1 5 (Proopiomelanocortin, POMC) γ- (γ-melanocyte stimulating hormone, γ-msh) ACTH ( α-msh (α-, α-melanocyte stimulating hormone) CLIP (, Corticotropin-like intermediate lobe peptide)) γ- (γ-lipotropic hormone, γ-lph) β- (β-melanocyte stimulating hormone, β-msh) β- (β-endorphin) Alrubaian [26] POMC cdna, 2 POMC, POMC A POMC B 2 POMC 45, 26 Danielson [27] (Polyodon spathula) 2 POMC, γ-msh, 6 Quérat [16] (Thyroid-stimulating hormone, TSH), cdna bp, 143, 20 TSH, TSH TSH FSHβ, : FSHβ, TSH, LHβ 7,,,, ;,,,, [28, 29],,,,,, 参考文献 (References): [1] 粦,, 玕,.., 1985, 9(2): [2],,,,.., 1985, (3): [3],,,.., 1985, (4): [4],,,,,,,,,,,.., 1985, (1): 1 5. [5],,,,.., 2004, 28(6): [6] Cao H, Zhou RJ, Wei QW, Li CJ, Gui JF. Molecular characterization of the growth hormone in Chinese sturgeon including expression during embryogenesis and early larval stages. J Appl Ichthyol, 2011, 27(2): [7] Liao ZY, Chen XL, Wu MJ. Molecular cloning and functional analysis of Chinese sturgeon (Acipenser sinensis) growth hormone receptor. Sci China Ser C, 2009, 52(10): [8] Cao H, Zhou L, Zhang YZ, Wei QW, Chen XH, Gui JF.

6 712 HEREDITAS (Beijing) Molecular characterization of Chinese sturgeon gonadotropins and cellular distribution in pituitaries of mature and immature individuals. Mol Cell Endocrinol, 2009, 303(1 2): [9] Tschöp M, Smiley DL, Heiman ML. Ghrelin induces adiposity in rodents. Nature, 2000, 497(19): [10] Farmer SW, Hayashida T, Papkoff H, Polenov AL. Characteristics of growth hormone isolated from sturgeon (Acipenser gueldenstaedtii) pituitaries. Endocrinology, 1981, 108(2): [11] Yasuda A, Yamaguchi K, Noso T, Papkoff H, Polenov AL, Nicoll CS, Kawauchi H. The complete amino acid sequence of growth hormone from sturgeon (Acipencer gueldenstaedtii). Biochim Biophys Acta, 1992, 1120(3): [12] Din SY, Hurvitz A, Goldberg D, Jackson K, Levavi-Sivan B, Degani G. Cloning of Russian sturgeon (Acipenser gueldenstaedtii) growth hormone and insulin-like growth factor I and their expression in male and female fish during the first period of growth. J Endocrinol Invest, 2008, 31(3): [13] Tarpey JF, Nicoll CS. Characterization of hepatic growth hormone binding sites in two fish species, Gillichthys mirabilis (Teleostei) and Acipenser transmontanus (Chondrostei). Gen Comp Endocrinol, 1985, 60(1): [14] Fukamachi S, Meyer A. Evolution of receptors for growth hormone and somatolactin in fish and land vertebrates: lessons from the lungfish and sturgeon orthologues. J Mol Evol, 2007, 65(4): [15] Moberg GP, Watson JG, Doroshov S, Papkoff H, Pavlick RJ Jr. Physiological evidence for two sturgeon gonadotrophins in Acipenser transmontanus. Aquaculture, 1995, 135(1 3): [16] Quérat B, Sellouk A, Salmon C. Phylogenetic analysis of the vertebrate glycoprotein hormone family including new sequences of sturgeon (Acipenser baeri) β subunits of the two gonadotropins and the thyroid-stimulating hormone. Biol Reprod, 2000, 63(1): [17] Avshalom H, Gad D, Doron G, Svetlana YD, Karen J, Berta LS. Cloning of FSHβ, LHβ, and glycoprotein α subunits from the Russian sturgeon (Acipenser gueldenstaedtii), β-subunit mrna expression, gonad development, and steroid levels in immature fish. Gen Comp Endocrinol, 2005, 140(1): [18],,,,,. β cdna., 2006, 52(2): [19] Ono M, Takayama Y, Rand-Weaver M, Sukata S, Yasunaga T, Noso T, Kawauchi H. cdna cloning of somatolactin, a pituitary protein related to growth hormone and prolactin. Proc Natl Acad Sci USA, 1990, 87(11): [20] Amemiya Y, Sogabe Y, Nozaki M, Takahashi A, Kawauchi H. Somatolactin in the white sturgeon and African lungfish and its evolutionary significance. Gen Comp Endocrinol, 1999, 114(2): [21] Nishii M, Movérus B, Bukovskaya OS, Takahashi A, Kawauchi H. Isolation and characterization of [Pro 2 ] somatostatin-14 and melanotropins from Russian sturgeon, Acipenser gueldenstaedti Brandt. Gen Comp Endocrinol, 1995, 99(1): [22] Kim JB, Gadsbøll V, Whittaker J, Barton BA, Conlon JM. Gastroenteropancreatic hormones (insulin, glucagon, somatostatin, and multiple forms of PYY) from the pallid sturgeon, Scaphirhynchus albus (Acipenseriformes). Gen Comp Endocrinol, 2000, 120(3): [23] Trabucchi M, Tostivint H, Lihrmann I, Sollars C, Vallarino M, Dores RM, Vaudry H. Polygenic expression of somatostatin in the sturgeon Acipenser transmontanus: molecular cloning and distribution of the mrnas encoding two somatostatin precursors. J Comp Neurol, 2002, 443(4): [24] Adrio F, Anadón R, Rodríguez-Moldes I. Distribution of somatostatin immunoreactive neurons and fibres in the central nervous system of a chondrostean, the Siberian sturgeon (Acipenser baeri). Brain Res, 2008, 1209: [25] Li CJ, Wei QW, Zhou L, Cao H, Zhang Y, Gui JF. Molecular and expression characterization of two somatostatin genes in the Chinese sturgeon, Acipenser sinensis. Comp Biochem Physiol A, 2009, 154(1): [26] Alrubaian J, Danielson P, Fitzpatrick M, Schreck C, Dores RM. Cloning of a second proopiomelanocortin cdna from the pituitary of the sturgeon, Acipenser transmontanus. Peptides, 1999, 20(4): [27] Danielson PB, Alrubaian J, Muller M, Redding JM, Dores RM. Duplication of the POMC gene in the paddlefish (Polyodon spathula): analysis of γ-msh, ACTH, and β-endorphin regions of ray-finned fish POMC. Gen Comp Endocrinol, 1999, 116(2): [28] Pakarinen P, Kimura S, El-Gehani F, Pelliniemi LJ, Huhtaniemi I. Pituitary hormones are not required for sexual differentiation of male mice: phenotype of the T/ebp/Nkx2.1 null mutant mice. Endocrinology, 2002, 143(11): [29] Rao CV, Lei ZM. Consequences of targeted inactivation of LH receptors. Mol Cell Endocrinol, 2002, 187(1 2):

22 Animal Husbandry & Veterinary Medicine 2011 Vol. 43 No % 3% 3% /% % 6. 67% 1. 71% 1. 11% 0. 14% 0.

22 Animal Husbandry & Veterinary Medicine 2011 Vol. 43 No % 3% 3% /% % 6. 67% 1. 71% 1. 11% 0. 14% 0. 2011 43 5 21 3 1 1* 1 2 3 1 1. 2. 3. 78. 68±2. 95 g 28 300 3 5 3% 3 1 ALT TP ALB AST 0. 05 P>0. 05 ALT TP ALB P>0. 05 AST 0. 05 GLOB P>0. 05 2 INS 0. 05 TSH T 3 0. 05 P>0. 05 GLU GH T4 IGF-I P>0. 05 S816.