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研究生:何俊霖
研究生(外文):Chun-Lin He
論文名稱:黑鯛性轉變機制之研究:DMRT1與性類固醇激素受器基因之表現
論文名稱(外文):The Mechanism of Sex Change in the Protandrous Black Porgy (Acanthopagrus schlegeli) : Gene Expression of DMRT1 and Sex Steroid Receptors
指導教授:張清風張清風引用關係
指導教授(外文):Ching-Fong Chang
學位類別:碩士
校院名稱:國立海洋大學
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:2002
畢業學年度:90
語文別:中文
論文頁數:134
中文關鍵詞:黑鯛雄性素受器雌性素受器性別決定基因性轉變性別決定同步定量PCR
外文關鍵詞:black porgyandrogen receptorestrogen receptorsex determining genesex changesex determinationReal-time PCRDMRT1
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本研究之目的在探討性別相關基因DMRT1與性類固醇激素受器在黑鯛 (Acanthopagrus schlegeli) 性轉變過程中,參與生殖腺分化與發育之分子作用機制。自黑鯛精巢組織選殖出DMRT1與雄性素受器 (androge receptor, AR) 之cDNA,轉譯為胺基酸序列後發現,DMRT1相較於其他物種只有在DM domain具有很高的相似性 (91~100%),在組織分布上DMRT1在生殖腺相關組織的表現顯示出明顯組織特異性,主要表現在精巢與儲精囊。而AR經轉譯為胺基酸後與其他物種之AR序列比較,在不同結構分區上以DNA binding domain與ligand binding domain具有較高的相似性。組織分布方面AR在各個組織中均有表現,但以精巢、儲精囊表現量最高。DMRT1在黑鯛精巢的表現,在性轉變過程中並沒有顯著的差異,然而在第三年齡性轉變確定,生殖腺發育完全後DMRT1的表現量明顯降低。黑鯛性類固醇激素受器 (ERα、AR) 在不同生殖期表現,整體而言精巢的表現量均比卵巢要高,而隨著性轉變過程精巢ERα、AR的表現量逐漸降低,而卵巢ERα的表現量則逐漸增加,以上結果說明雌性激素,如estradiol-17β (E2)在黑鯛性轉變過程中扮演重要的調控角色。經由注射高、低劑量 (50 ng/g 與1.5 μg/g BW) 的性類固醇激素處理,對於黑鯛生殖腺DMRT1之表現均沒有顯著的影響。而testosterone (T) 對於精巢ERα的表現則有抑制的效用,而E2與T則會促進精巢AR的表現,11-ketotestosterone (11-KT) 則不會調控任何基因的表現。此外,注射LHRH analog會造成精巢DMRT1與ERα表現量顯著上升。長時間投餵E2誘導性轉變過程,會導致精巢與卵巢DMRT1表現量降低。而投餵E2對於ERα在精巢與卵巢的表現則分別有抑制與促進的效用,投餵E2則會抑制AR在精巢的表現。由本實驗結果說明DMRT1在黑鯛性轉變過程扮演重要的分子調控角色,其表現模式具有一般性別決定基因在性別確定後即不再表現的特性,故本研究推論DMRT1可能是黑鯛性別決定基因 (sex-determining gene)。然而根據ERα、AR表現的形式,以及受外源性因子調控後表現之影響,證實E2在黑鯛性轉變過程的確是一個關鍵性的調控因子。
The objectives were to investigate the molecular mechanism of sex related gene - DMRT1, and sex steroid receptors (ERα and AR) involved in the gonadal differentiation and development in the process of sex change of the protandrous black porgy (Acanthopagrus schlegeli).
DMRT1 and AR cDNA were cloned from testicular tissue. DMRT1 had a high homology (89~100%) in the DM-domain, but with low similarity outside the DM-domain as compared to other teleosts. Tissue expression of DMRT1 had a gonad-specific pattern, which was mainly found in testis and seminal vesicle. In comparison of the AR with other species, DNA binding domain (DBD) and ligand binding domain (LBD) showed high homology (DBD:70~98% ; LBD:60~96%). AR had a widespread expression, showing high expression in testis and seminal vesicle.
There was no significant difference in the expression of DMRT1 in testis of 1~2 year age, but the DMRT1 was significantly decreased in 3-year-old males. The expression of sex steroid receptors (ERαand AR) in ovarian tissue was much weaker than testiscular tissue in bisexual gonad. ERαtranscripts in ovary increased gradually in the process of sex change; while, ERαand AR transcripts in testis decreased.
Different doses of sex steroids by injection had no effect on gonadal DMRT1 expression, whereas E2 (estradiol-17β) could inhibit ERαexpression in testis. E2 and testosterone (T) could stimulate AR expression in testis but 11-ketotestosterone (11-KT) didn’t regrulate the expression DMRT1、ERαand AR transcripts. Besides, in vivo injection of LHRH analog stimulated the expression of DMRT1 and ERαin testis. Long-term oral administration of E2 reduced the expression of DMRT1 both in testis and ovary. Fed-E2 also inhibited the expression of ERαin testis but increased ERαtranscripts in ovary but oral administration of E2 decreased AR transcripts in testis.
It is concluded that DMRT1 was an important molecular modulator in the process of sex change in black porgy. The expression of DMRT1 significantly declined in 3-year-old males and female. It was suggested that DMRT1 may be the sex determining gene in black porgy. According to the expression pattern of ERαand AR, E2 was further indicated as a critical regulator in the sex change of black porgy.
中文摘要…………………………………………………………………… I
英文摘要………………………………………………………………… III
圖表目錄…………………………………………………………………VIII
壹、前言
一、動物的性別……………………………………………………………1
二、魚類的性別分化………………………………………………………2
三、性別決定因子…………………………………………………………3
四、魚類性別決定基因……………………………………………………5
五、性類固醇激素與性別分化……………………………………………7
六、性類固醇激素受器……………………………………………………9
七、性類固醇激素之相關研究……………………………………………11
八、研究源起………………………………………………………………15
九、研究目的………………………………………………………………17
貳、實驗材料與方法
一、實驗魚種及採樣組織…………………………………………………18
二、實驗設計………………………………………………………………18
(一)、選殖DMRT1與AR基因………………………………………………18
(二)、黑鯛DMRT1與AR基因之組織分佈…………………………………19
(三)、比較不同性別轉變階段雌雄DMRT1、ERα與AR之表現………… 20
(四)、短時間注射不同劑量之性類固醇激素、LHRH analog 對DMRT1、
ERα與AR表現量之影響……………………………………………21
(五)、長時間投餵含有E2飼料對黑鯛生殖腺DMRT1、ERα與AR表現
之影響…………………………………………………………… 21
三、實驗方法………………………………………………………………22
(一)、全量核醣核酸 (Total RNA) 之純化……………………………22
(二)、反轉錄-聚合酶連鎖反應 (RT-PCR)…………………………… 23
(三)、DMRT1基因引子……………………………………………………25
(四)、雌性素受器 (ERα) 基因引子……………………………………26
(五)、雄性素受器 (AR) 基因引子…………………………………… 26
(六)、Rapid Amplification of cDNA Ends (RACE)…………………27
(七)、萃取質體DNA (DNA extraction)……………………………… 29
(八)、南方轉漬雜交法 (Southern blotting hybridization)…… 31
(九)、Absolute Real-time quantitation PCR………………………32
四、實驗藥品及分析試劑…………………………………………………36
(一)、菌種、質體……………………………………………………… 36
(二)、一般化學藥品…………………………………………………… 36
(三)、酵素及生化分析試劑…………………………………………… 37
(四)、實驗藥品及溶液配方…………………………………………… 38
五、統計分析………………………………………………………………40
參、結果
一、黑鯛DMRT1與雄性素受器 (AR) 之基因選殖……………………… 41
(一)、黑鯛DMRT1基因選殖及序列分析…………………………………41
(二)、黑鯛雄性素受器 (AR) 基因選殖及序列分析………………… 42
二、黑鯛DMRT1與AR之組織分布………………………………………… 43
(一)、黑鯛DMRT1基因組織分布…………………………………………43
(二)、黑鯛雄性素受器 (AR) 組織分布……………………………… 44
三、絕對定量Real-time PCR系統之建立……………………………… 44
(一)、黑鯛DMRT1、ERα與AR之Real-time PCR引子設計………………44
(二)、Real-time PCR反應CT值之決定…………………………………44
(三)、絕對定量標準曲線之建立……………………………………… 45
四、黑鯛不同性轉變時期生殖腺DMRT1、ERα和AR之表現量之改變
情形……………………………………………………………………45
(一)、比較不同性轉變階段生殖腺DMRT1的表現………………………45
(二)、比較不同性轉變階段生殖腺ERα的表現…………………………46
(三)、比較不同性轉變階段生殖腺AR的表現………………………… 47
五、注射性類固醇激素對於黑鯛生殖腺DMRT1、ERα與AR基因表現之
影響……………………………………………………………………48
(一)、注射不同劑量性類固醇激素對黑鯛生殖腺DMRT1基因表現之
影響……………………………………………………………… 48
(二)、注射不同劑量性類固醇激素對黑鯛生殖腺ERα基因表現之
影響……………………………………………………………… 48
(三)、注射不同劑量性類固醇激素對黑鯛生殖腺AR基因表現之
影響……………………………………………………………… 49
六、注射LHRH analog對於黑鯛生殖腺DMRT1、ERα與AR基因表現之
影響……………………………………………………………………49
(一)、注射LHRH analog對黑鯛生殖腺DMRT1基因表現之影響……… 49
(二)、注射LHRH analog對黑鯛生殖腺ERα基因表現之影響………… 49
(三)、注射LHRH analog對黑鯛生殖腺AR基因表現之影響……………49
七、長時間投餵E2飼料對黑鯛生殖腺DMRT1、ERα與AR表現之
影響……………………………………………………………………50
(一)、投餵E2 (6 mg/kg feed) 過程,1.5與4.5個月對黑鯛生殖腺
DMRT1基因表現之影響……………………………………………50
(二)、投餵E2 (6 mg/kg feed) 過程,1.5與4.5個月對黑鯛生殖腺
ERα基因表現之影響………………………………………………50
(三)、投餵E2 (6 mg/kg feed) 過程,1.5與4.5個月對黑鯛生殖腺
AR基因表現之影響……………………………………………… 51
肆、討論
一、黑鯛DMRT1基因選殖………………………………………………… 52
二、黑鯛雄性素受器基因之選殖…………………………………………53
三、黑鯛DMRT1和AR之表現……………………………………………… 56
四、絕對定量Real-time PCR分析系統之建立………………………… 58
五、黑鯛DMRT1與性類固醇激素受器基因在不同生殖期表現之
意義……………….………………………………………………… 60
六、DMRT1與性類固醇激素受器基因表現之調控……………………… 64
伍、結論…………………………………………………………………… 69
陸、參考文獻……………………………………………………………… 72
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