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研究生:林倩如
研究生(外文):Lin, Chien-Ju
論文名稱:在下視丘-腦下垂體-生殖腺主軸影響下的腦部變化
論文名稱(外文):The Brain Changes under Control of Hypothalamus-Pituitary-Gonad Axis in Protandrous Black Porgy, Acanthopagrus schlegelii
指導教授:張清風張清風引用關係
指導教授(外文):Chang, Ching-Fong
口試委員:黃鵬鵬Dufour, Sylvie廖欽峯周信佑吳貫忠張清風
口試委員(外文):Dufour, Sylvie Chang, Ching-Fong
口試日期:2015-07-30
學位類別:博士
校院名稱:國立臺灣海洋大學
系所名稱:水產養殖學系
學門:農業科學學門
學類:漁業學類
論文種類:學術論文
論文出版年:2016
畢業學年度:103
語文別:中文
論文頁數:92
中文關鍵詞:黑鯛促性腺素釋放激素促性腺素釋放激素受體促性腺激素性類固醇環化酵素雌二醇腦下垂體細胞腦部發育神經新生作用神經性類固醇合成作用
外文關鍵詞:Black porgyGnRHGnRH receptorGTHSex steroidsaromataseestradiolPituitary cellsBrain developmentNeurogenenesisNeurosteroidogenesis
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下視丘-腦下垂體-生殖腺主軸 (Hypothalamus-Pituitary-Gonad axis, HPG axis) 在生殖生理中扮演重要角色,neuropeptide、gonadotropin、性類固醇激素皆參與其中,透過對HPG主軸中各影響因子的分析,進一步瞭解黑鯛腦中生殖內分泌的調控機制。
根據本研究的結果發現 (1) 黑鯛有兩型的GnRH-R,gnrh-r1、gnrh-r2在腦下垂體、生殖腺均有表現。從季節性變化的分析結果推測,黑鯛生殖腺gnrh-r1與gnrh-r2的表現受生殖腺發育狀態影響,應與黑鯛的性轉變以及卵巢成長有關,腦下垂體上gnrh-r1可能參與在性轉變過程中的調控。(2) 黑鯛腦下垂體gnrh-r1的表現量遠大於gnrh-r2,且激素能刺激gnrh-r1表現增加,而gnrh-r2的表現不受影響。在發育期間或季節性採樣中,腦下垂體gnrh-r2的表現都沒有太大變化。(3) 黑鯛腦下垂體gnrh-r1對生殖生理的調控較為重要,應扮演比gnrh-r2更重要的角色;而生殖腺gnrh-r1與gnrh-r2的表現應同樣重要,均參與生殖內分泌和生殖腺功能的調控。(4) 分析第一繁殖季前生殖腺分化至發育期間腦組織中的基因表現,腦下垂體gthα、fshβ、lhβ應參與繁殖季之前的調控,而與生殖腺性別分化較無關聯性。(5) 腦組織中sbgnrh與cyp19a1b應參與在生殖腺性別分化時期以及第一繁殖季節的調控。(6) 在生殖腺分化期間,可能也發生了腦的性別分化,且應以下視丘、前腦為主進行調控作用。(7) 在生殖腺分化期間,黑鯛腦中神經性類固醇合成作用及神經新生作用的活性均上升。(8) E2能促進神經性類固醇合成作用及神經新生作用的活性,而AI顯著抑制細胞增生活性,顯示E2在黑鯛早期腦發育過程中扮演促進神經新生的角色。(9) 生殖腺摘除後,腦組織中基因表現、細胞數目、免疫染色活性等皆與控制組沒有顯著差異,顯示發生在生殖腺分化時期的腦中神經類固醇合成作用和神經新生作用的波峰現象,與性別或生殖腺無關,應是受到腦部內生性雌二醇的調控。
The objectives of the presents study were to investigate the brain differences in protandrous black porgy at the time of gonadal differentiation, development and sex change. The regulatory factors involved in the Hypothalamus-Pituitary-Gonad axis, including neuropeptides, gonadotropin, and sex steroids, were analyzed.
The present data demonstrates that (1) Two forms of GnRH-R were detected in black porgy, with differential expression and regulation in the pituitary, testicular and ovarian tissues. gnrh-r1 but not gnrh-r2 expression was higher in the pituitary during the spawning period as compared to the prespawning. The expression profiles of both forms of GnRH-R were variable in the gonads according to the gonad stage and season. (2) In pituitary, sex steroids (E2 and T) increased the expression of gnrh-r1, but not gnrh-r2. No significant differences were found in gnrh-r2 from the prespawning to the spawning period. (3) gnrh-r1 may play a major role in the pituitary, while both forms may be involved in the regulation of gonadal functions and further ovarian development at the time of sex change. (4) The gene expressions of gthα, fshβ, and lhβ in pituitary may be involved in the regulation of the first spawning season rather than sex differentiation period. (5) sbgnrh and cyp19a1b may involved in achieving gonadal sex differentiation, and may also in the regulation of the first spawning season in black porgy. (6) Brain sex differentiation is organized during gonadal sex differentiation period in black porgy, and suggests that the may regulate in forebrain and hypothalamus mainly. (7) The peaks in brain neurosteroidogenesis and neurogenesis activities occur at the time of gonad differentiation in the black porgy. (8) Exogenous E2 stimulated neurosteroidogenesis and neurogenesis, while AI decreased brain cell proliferation. These findings support the hypothesis that E2 plays important roles in the induction of early brain development in black porgy. (9) No significant differences in gene expression and brain cell numbers were found between control and castrated fish. These data further support that the peak in early brain activity is controlled by locally produced brain E2, and is independent of the gonads.
摘要 I
Abstract II
目錄 III
圖表目錄 V

緒論 1

第一章 黑鯛腦下垂體和生殖腺GnRH-R的表現 6
摘要 6
1-1 前言 7
1-2 材料與方法 9
1-2.1 實驗魚種及採樣組織 9
1-2.2 實驗設計 9
1-2.3 腦下垂體初級細胞培養 (Pituitary primary cell culture) 10
1-2.4 Quantitative Real-time PCR 11
1-2.5 統計分析 11
1-3 結果 11
1-3.1 黑鯛繁殖季節與gnrh-r的基因表現 11
1-3.2 gnrh-r在雄魚與雌魚腦下垂體中的基因表現 12
1-3.3 活體注射對gnrh-r表現量之影響 12
1-3.4 腦下垂體細胞培養短時間處理不同劑量之LHRH analog和性類固醇激素對gnrh-r1與gnrh-r2表現量之影響 13
1-4 討論 13
圖 17

第二章 腦下垂體及各腦區內與GnRH-R相關基因之表現 29
摘要 29
2-1 前言 30
2-2 材料與方法 32
2-2.1 實驗魚種及採樣組織 32
2-2.2 實驗設計 32
2-2.3 Quantitative Real-time PCR 33
2-2.4 統計分析 33
2-3 結果 33
2-3.1 第一繁殖季前腦下垂體gnrh-r1與gonadotropin次單元的基因表現 33
2-3.2 第一繁殖季前各腦區gnrh的基因表現 33
2-3.3 第一繁殖季前各腦區cyp19a1b的基因表現 34
2-3.4 生殖腺性別分化期間E2對腦下垂體gnrh-r1與gonadotropin次單元基因的調控 34
2-3.5 生殖腺性別分化期間E2對各腦區gnrh基因的調控 35
2-3.6 生殖腺性別分化期間E2對各腦區cyp19a1b基因的調控 35
2-4 討論 35
圖 39

第三章 生殖腺性別分化期間腦中類固醇生成與神經新生 50
摘要 50
3-1 前言 51
3-2 材料與方法 53
3-2.1 實驗魚種及採樣組織 53
3-2.2 實驗設計 53
3-2.3 腦組織中類固醇含量的檢測 54
3-2.4 組織切片 54
3-2.5 Quantitative Real-time PCR 55
3-2.6 幼齡黑鯛的生殖腺摘除手術 55
3-2.7 統計分析 55
3-3 結果 56
3-3.1 生殖腺分化期間幼齡黑鯛腦部的變化 56
3-3.2 E2對生殖腺分化期間幼齡黑鯛腦部的調控 57
3-3.3 生殖腺分化期間幼齡黑鯛腦部變化與生殖腺的關係 57
3-4 討論 58
圖 62

第四章 結論 71

參考文獻 73
附錄 著作目錄 92
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