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1 46 6 Vol.46, No OCEANOLOGIA ET LIMNOLOGIA SINICA Nov., 2015 E 2 MT (Oreochromis niloticus) * ( ; ),, PCR, (E 2 MT) GH GHR1 IGF-I mrna : E 2 MT, E 2, MT ; E 2 GH GHR1 IGF-I mrna (P<0.05), GH GHR1 mrna ; MT GH GHR1 IGF-I mrna (P<0.05), GH mrna, GHR1 mrna, GHR1 (P<0.05), mrna,, ; E 2 ; MT; ; Q956 doi: /hyhz , (Growth hormone receptor, GHR) mrna, GHR 5 (Goodyer et al, 2001) (17β-estradiol, E 2 ) GHR mrna (Domené et al, 1994), GHR mrna (Gabrielsson et al, 1995) GHR, (Testosterone, T) GHR mrna (Zung et al, 1999), T GH, (Dihydrotestosterone, DHT),, E 2 (Borski et al, 1996),,, (Mandiki et al, 2004, 2005) GH IGF-I (Wehrenberg et al, 1992),,,, *, ;, 2012ZD07,,, mxl@hzu.edu.cn : , :

2 , (E 2 MT) ( GH GHR1 IGF-I) mrna 1 1.1, (, 2010), (PS); (17β-estradiol, E 2 ) (17α-methyltestosterone, MT) Sigma (Sigma, America),, 50μg/g, ,, ;, h, Real-time PCR E 2 MT ( GH GHR1 IGF-I) mrna (, 2010) , : HSI(%)=100 / (1) K(%)=100 (g) / (cm) 3 (2) AGR W (%) = 100 (W 2 W 1 ) / W 1 (3) AGR L (%) = 100 (L 2 L 1 ) / L 1 (4) SGR W (%/d) =100 [(lnw 2 lnw 1 )] / (ΔT) (5) SGR L (%/d) =100 [(lnl 2 lnl 1 )] / (ΔT) (6) SGR W (%) = 100 ( SGR W SGR W ) / SGR W (7) L 1 : T 1, L 2 : T 2, W 1 : T 1, W 2 : T 2, ΔT=T 2 T 1 (8) Real-time PCR (E 2 MT) GH GHR IGF-I mrna, 1 Real-time PCR (, 2010), E 2 MT 42, E 2 (P<0.05), (P>0.05); MT (P<0.05), (P>0.05)( 1) 表 1 荧光实时定量 PCR 分析所用引物 Tab.1 Primers for real-time PCR β-actin-f 5 -GGTGGGTATGGGTCAGAAAGA-3 β-actin-r 5 -GCTGTCGTGAAGGAGTAG-3 GH-F 5 -ATTATCAAAGTCTGGGAGGC-3 GH-R 5 -GGTAGGTCTCCACCTTGTGC-3 GHR1-F 5 -TAAGAAAGAGCCTCCTACCA-3 GHR1-R 3 -ACTGTCGCTGAATGTCCAAT-5 IGF-I-F 5 -TCTGTGGAGAGCGAGGCTTT-3 IGF-I-R 5 -CACGTGACCGCCTTGCA-3 1 (a., b. ) Fig.1 Effects of steroid hormones on body weights of female (a) and male (b) : ± (n=20) (P<0.05) E 2 MT, 2

3 6 : E 2 MT (Oreochromis niloticus) 1489, E 2 MT (K) (P>0.05), K(P>0.05); (GSI) (P<0.05), GSI (P>0.05); MT (HSI) (P<0.05), MT HSI HSI, E 2 HSI(P<0.05), HSI (P>0.05); E 2 (AGR W ), 24.16%, AGR W, MT AGR W 14.09% 11.20%( 2) 表 2 生长参数 ( 平均值 ± 标准误, n=20) Tab.2 Growth parameters (mean values±s.d., n=20) K(%) GSI(%) HSI(%) AGR W (%) 3.36±0.03 a 0.16±0.14 a 1.61±0.60 a ±0.31 a 0.33±0.14 b 1.54±0.54 a E ±0.21 a 0.18±0.11 a 2.02±1.00 a ±0.16 a 0.29±0.12 b 2.16±0.63 b MT 3.34±1.10 a 0.17±0.09 a 2.35±0.80 b ±0.06 a 0.26±0.11 b 1.55±0.51 a (P<0.05) 2.2 E 2 MT GH mrna E 2 GH mrna (P<0.05), GH mrna (P>0.05); MT GH mrna (P<0.05), GH mrna ( 2) (P<0.05), ( 3) 3 GHR1 mrna Fig.3 Real-time PCR analysis on liver GHR mrna levels in 2 GH mrna Fig.2 Real-time PCR analysis on pituitary GH mrna levels in : ± (n=5) (P<0.05) GHR1 mrna E 2 (P<0.05), GHR1 mrna ; MT, GHR1 mrna IGF-I mrna E 2 IGF-I mrna (P<0.05); MT, IGF-I mrna (P<0.05), E 2 IGF-I MT (P<0.05)( 4) 2.3 E 2 MT GH mrna

25±0.31 a 0.33±0.14 b 1.54±0.54 a 367.20 E 2 3.61±0.21 a 0.18±0.11 a 2.02±1.00 a 434.95 3.49±0.16 a 0.29±0.12 b 2.16±0.63 b 455.90 MT 3.34±1.10 a 0.17±0.09 a 2.35±0.80 b 488.22 3.16±0.06 a 0.26±0.

4 IGF-I mrna Fig.4 Real-time PCR analysis on liver IGF-I mrna levels in E 2 MT, GH mrna E 2 6h, GH mrna, (P<0.05);, MT 12h, GH mrna, (P<0.05)( 5) GHR1 mrna GHR1 mrna E 2 12h GHR1 mrna (P<0.05), 24h (P<0.05); MT 12h GHR1 mrna (P<0.05), 24h GHR1, 24h GHR1 (P<0.05)( 6) IGF-I mrna IGF-I mrna E 2 24h, IGF-I mrna, (P<0.05); MT 24h IGF-I mrna (P<0.05), IGF-I mrna ( 7) 3, E 2 MT, E 2 MT, MT 5 GH mrna Fig.5 Real-time PCR analysis on pituitary GH mrna levels in a., b., ( GH GHR1 IGF-I) mrna ; E 2 GH GHR1 IGF-I, (GSI),,,,,, (Mandiki et al, 2004, 2005; Meinhardt et al, 2006),

05); MT 24h IGF-I mrna (P<0.05), IGF-I mrna ( 7) 3, E 2 MT, E 2 MT, MT 5 GH mrna Fig.5 Real-time PCR analysis on pituitary GH mrna levels in a., b.

5 6 : E 2 MT (Oreochromis niloticus) GHR1 mrna Fig.6 Real-time PCR analysis on liver GHR mrna levels in a., b.,, (Cox et al, 2005) (MT),, T, (Riley et al, 2002), (MT), : (1) MT, ; (2) MT GHR GH ; (3) MT mrna, GH IGF-I, GHR T, GHR mrna (Yu et al, 1996), GHR mrna (Gabrielsson et al, 1995), GHR MT, GHR mrna 7 IGF-I mrna Fig.7 Real-time PCR analysis on liver IGF-I mrna levels in a., b. (, 2006), MT, GHR mrna,, MT GHR mrna MT, IGF-I mrna, T 11-KT, GH, IGF-I IGFBP-3, IGF-I, GH ; GH, IGFBP-3 (Larsen et al, 2004) E 2 GH,, E 2 (Somatotropin Release Inhibiting Factor, SRIF) GH ;, E 2 GH

mrna (Gabrielsson et al, 1995), GHR MT, GHR mrna 7 IGF-I mrna Fig.7 Real-time PCR analysis on liver IGF-I mrna levels in a., b.

6 (Growth Hormone Releasing Hormone, GHRH) GH, GH (Holloway et al, 1997; Zou et al, 1997) ( E 2 ), IGF-I,, E 2 IGF-I, IGF-I mrna, (Borski et al, 1996) E 2, GHR mrna (Yu et al, 1996;, 2006), E 2 GH mrna, GH E 2 GHR1 IGF-I mrna,, ;, E 2,, E 2 GH,, ;, E 2 GH GHR mrna,,,,,,, (CSH) (Oreochromis niloticus)., 41(2): ,,, , 52(6): Borski R J, Tsai W, DeMott-Friberg R et al, Regulation of somatic growth and the somatotropic axis by gonadal steroids: primary effect on insulin-like growth factor I gene expression and secretion. Endocrinology, 137(8): Cox R M, John-Alder H B, Testosterone has opposite effects on male growth in lizards (Sceloporus spp.) with opposite patterns of sexual size dimorphism. J Exp Biol, 208: Domené H M, Marin G, Sztein J et al, Estradiol inhibits growth hormone receptor gene expression in rabbit liver. Mol Cellul Endocrinol, 103(1 2): Gabrielsson B G, Carmignac D F, Flavell D M et al, Steroid regulation of growth hormone (GH) receptor and GH-binding protein messenger ribonucleic acids in the rat. Endocrinology, 136(1): Goodyer C G, Zogopoulos G, Schwartzbauer G et al, Organization and evolution of the human growth hormone receptor gene 5 -flanking region. Endocrinology, 142(5): Holloway A C, Leatherland J F, Effect of gonadal steroid hormones on plasma growth hormone concentrations in sexually immature rainbow trout, Oncorhynchus mykiss. Gen Comp Endocrinol, 105(2): Larsen D A, Shimizu M, Cooper K A et al, Androgen effects on plasma GH, IGF-I, and 41-kDa IGFBP in coho salmon (Oncorhynchus kisutch). Gen Comp Endocrinol, 139(1): Mandiki S N M, Babiak I, Bopopi J M et al, Effects of sex steroids and their inhibitors on endocrine parameters and gender growth differences in Eurasian perch (Perca fluviatilis) juveniles. Steroids, 70(2): Mandiki S N M, Houbart M, Babiak I et al, Are sex steroids involved in the sexual growth Dimorphism in Eurasian perch juveniles?. Physiol Behav, 80(5): Meinhardt U J, Ho Ken K Y, Modulation of growth hormone action by sex steroids. Clin Endocrinol, 65(4): Riley L G, Richman III N H, Hirano T et al, Activation of the growth hormone/insulin-like growth factor axis by treatment with 17 α-methyltestosterone and seawater rearing in the tilapia, Oreochromis mossambicus. Gen Comp Endocrinol, 127(3): Wehrenberg W B, Giustina A, Basic counterpoint: mechanisms and pathways of gonadal steroid modulation of growth hormone secretion. Endocr Rev, 13(2): Yu Y M, Domené H M, Stzein J et al, Developmental changes and differential regulation by testosterone and estradiol of growth hormone receptor expression in the rabbit. Eur J Endocrinol, 135(5): Zou J J, Trudeau V L, Cui Z et al, Estradiol stimulates growth hormone production in female goldfish. Gen Comp Endocrinol, 106(1): Zung A, Phillip M, Chalew S A et al, Testosterone effect on growth and growth mediators of the GH-IGF-I axis in the liver and epiphyseal growth plate of juvenile rats. J Mol Endocrinol, 23(2):

., 52(6): 1096 1106 Borski R J, Tsai W, DeMott-Friberg R et al, 1996.

7 6 : E 2 MT (Oreochromis niloticus) 1493 STEROID HORMONES (E 2 AND MT) DISPLAYED DIFFERENCE IN SEX FOR NILE TILAPIA OREOCHROMIS NILOTICUS MA Xi-Lan 1, ZHANG Yong 2, CHEN Yong-Zhi 1, ZHOU Li-Bin 1 (1. Department of Life Science, Huizhou University and Institute of Biotechnology, Huizhou , China; 2. State Key Laboratory of Biocontrol, Institute of Aquatic Economic Animals and Guangdong Provincial Key Laboratory for Aquatic Economic Animals, School of Life Sciences, Sun Yat-Sen University, Guangzhou , China) Abstract Steroid hormone affects the growth of fish sexual dimorphism, and the mechanism remains unclear. Nile tilapia (Oreochromis niloticus) is a fresh water fish with obvious sexual dimorphism. The effects of steroid hormone on the growth and expressions of GH in pituitary, GHR and IGF-I mrna in liver of were studied. Two experiments in a long and short period were designated and carried out. In the long-period experiment, 200 fish individuals in similar body size were randomly assigned into the control group [intraperitoneal injection (i. p.) with fish physiological saline, PS], the E 2 group (i. p. with E 2, 50μg/g body weight), and the MT group (i. p. with MT, 50μg/g body weight), and fed for 70 days under the same natural conditions. Result shows that E 2 significantly promoted female growth and enhanced female GH, GHR1, and IGF-I mrna expressions (P<0.05), but for males not in growth nor GH, GHR1 mrna expressions. However, MT boosted significantly the growth of both sexes. MT improved GH for males, and GHR1 expressions for both sexes but greater in male than in female (P<0.05). Therefore, steroid hormones affected the growth and the genes expression in different ways in sex, which may become a reason for the dimorphism of the fish. Key words (Oreochromis niloticus); E 2 ; MT; sexual dimorphism; gene expression

State Key Laboratory of Biocontrol, Institute of Aquatic Economic Animals and Guangdong Provincial Key Laboratory for Aquatic Economic Animals, School of Life Sciences, Sun Yat-Sen University,

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.