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研究生:黃靜端
研究生(外文):Jing-Duan Huang
論文名稱:黑鯛生殖腺組織及微細構造之特性研究
論文名稱(外文):The studies of histological and ultrastructural characteristics of gonad in the protandrous black porgy, Acanthopagrus schlegeli
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:博士
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:131
中文關鍵詞:性轉變生殖腺構造精子生成卵子生成血管原始生殖表皮細胞
外文關鍵詞:sex changegonadal structurespermatogenesisoogenesisblood vesselsprimordial germ epithelia
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黑鯛(Acanthopagrus schlegeli)屬雄性先熟型的雌雄同體魚,在第一、二個繁殖季節都表現雄性功能,進入第三個繁殖季節才有部份魚會進行性轉變成為雌魚,本論文利用光學組織切片技術、掃描及穿透式電子顯微鏡技術研究黑鯛生殖腺在個體發生學上型態的變化及精、卵巢組織內生殖細胞與體細胞的發育及微細構造變化。
根據黑鯛生殖週期及生殖腺組織特性,我們將黑鯛每年的生殖週期分成四個不同階段:繁殖期(spawning period)為每年十二月至隔年三月,繁殖後期(post-spawning period )四月至五月,靜止期(resting period)為六月至九月,繁殖前期(pre-spawning period)為十至十一月。
四個月齡(七月份)的黑鯛生殖腺開始發現卵巢腔,此時只能發現少數幾顆生殖細胞,六個月齡(九月份)的黑鯛生殖腺才開始在卵巢腔旁及生殖腺底端發現原始生殖細胞,精原細胞和卵原細胞無法從光學顯微鏡下分辨。七個月齡(十月份)的黑鯛生殖腺內精巢組織開始增生,可以發現各種不同階段的生殖細胞,在卵巢腔一側發現少數的卵母細胞。
黑鯛雄性生殖細胞的發育可分為精原細胞(spermatogonia)、精母細胞(spermatocytes)、精細胞(spermatids)及精子(spermatozoa)四個主要階段。從精母細胞進入精細胞時,發生一系列細胞型態變化,包括細胞質丟棄,遠端小體(distal centriole)及近端小體(proximal centriole)移至細胞質側邊,鞭毛(9+2構造)從遠端小體延伸出細胞外,隨後細胞核旋轉,遠端小體及近端小體陷入細胞核凹陷內。
雌性生殖細胞及體細胞組織由卵巢腔一側的原始生殖表皮細胞(primordial germ epithelia)開始分化增生。在第一次繁殖季節後,卵巢腔內的卵巢薄板開始增生。黑鯛卵細胞的發育階段分為四期:卵原細胞期(oogonia phase)、染色質-核仁及周邊核仁期(chromatin-nucleolus and perinucleolar phase)、皮質小粒及卵黃堆積期(cortical alveoli and vitellogenic phase)、成熟及排卵期(maturation and ovulation phase)。卵原細胞期時間非常短,在未進行性轉變前,卵細胞都停滯在周邊核仁期階段。卵細胞進入成熟階段時,zona radiata層逐漸增厚,並形成Z1、Z2及Z3三層不同構造。卵細胞亦伸出微絨毛(microvilli),穿過zona radiata層,進入濾泡層的granulosa cells,使得granulosa cells的細胞質呈現電子極化現象。
在兩次繁殖期間的兩性階段,精、卵巢組織的比例及其個別的發育狀態有各種不同的類型;在不同魚齡,不同發育階段,雌雄生殖組織及體細胞組織會有不同消長特性。
黑鯛生殖腺的精巢組織與卵巢組織由結締組織層加以分隔,結締組織層主要由膠原纖維(collagen)、彈性纖維(elastin)及平滑肌細胞等組成,隨著生殖腺生殖細胞消長及性轉變過程中的變化,結締組織組成亦發生變化。
黑鯛生殖腺內之血管組織對雌、雄兩性生殖腺組織之消長扮演重要角色,排空的精巢組織顯露許多血管構造,每一卵細胞皆與一組血管連接,其微血管分支再包覆於卵細胞的濾泡細胞層外。尚未變性的雄魚即使在非繁殖期間,精巢組織萎縮,但其血管系統及精巢小葉構造仍完整健全;即將進行性轉變或已發生性轉變為雌魚者,其生殖腺內精巢組織退化成痕跡物,也無健全的血管系統及精巢小葉構造存在。
綜合本論文的發現,黑鯛自然性轉變過程中,生殖腺內生殖細胞組織間、生殖細胞與體細胞組織間及體細胞與體細胞組織間,隨著個體發生、分化、成熟不同階段皆有不同的細胞特徵與組織特性。本研究論文對日後進行黑鯛生殖調控、自然性轉變機制及人為性轉變調控研究上,在生殖腺之細胞學及組織學提供非常重要之基礎資料。
Black porgy, Acanthopagrus schlegeli, is a marine protandrous hermaphrodite. They are functional males at the first and the second spawning seasons, while part of them start to sexually change and become functional females at the third spawning season. The objectives were to investigate the ontogenic pattern of the gonadal structures and the development and ultrastructural changes of the germ cells and somatic tissues in both testes and ovary by light microscopy, scanning and transmission electron microscopy.
According to the observations, the reproductive season of black porgy was defined into four stages: post-spawning period (from April to May), resting period (from June to September), pre-spawning period (from October to November) and spawning period (from December to March).
The ovarian cavities began to be found in the gonads of four-month-old (July) fish. Very few germ cells could be found during this stage. Generally spermatogonia and oogonia could not be found until six-month-old (September). They located in the ventral part and inside the ovarian cavity of the gonad, respectively. Spermatogonia could not be distinguished from oogonia by light microscopy. Zone of connective tissues, constituting part of the wall of ovarian cavity, separated the testicular tissues from the ovarian tissues. The proliferation of testicular tissues started in October and a few oocytes inside the ovarian cavity also could be found.
Development of male germ cells was classified into four stages: spermatogonia, spermatocytes, spermatids and spermatozoa. Male germ cells were arrested in the spermatogonia stage during the resting period. Spermiogenesis included a series of cytological changes: casting of spermatocyte cytoplasm, the 9+2 flagellum structure developing and elongating from the distal centriole, nucleus rotating and the diplosome invaging into nucleus.
Both female germ cells and somatic tissues differentiated and proliferated from primordial germ epithelia, which were found in one side of ovarian cavity. After the first spawning spawning, ovarian lamella, the unit of ovary, began to develop. Development of female germ cells was classified into four stages: oogonia phase, chromatin-nucleolus and perinucleolar phase, cortical alveoli and vitellogenic phase and maturation and ovulation phase. Oogonia phase was very short. Female germ cells were nearly arrested in the perinucleolar phase before fish proceeded to sexually change. Granulosa cells surrounded the perinucleolar oocytes and directly contacted with them. When fish proceeded to sexually change, perinucleolar oocytes started to go into vitellogenesis in the prespawning period. Then zona radiata began to appear between granulosa cells and oocyte. Following the processes of the maturation of oocytes, zona radiata grew into three zones: Z1, Z2 and Z3. Microvilli, elongating from oocytes and penetrating through the zona radiata, entered into granulosa cells. Consequently, the cytoplasm of granulosa cells became electron polarization.
Blood vessel system played an important role in dynamic variation of the gonadal structures. Main artery and vein appeared in the zone of ovary. Each oocyte was connected with a single group of capillaries, which were then distributed on the surface of the follicular layer. Lots of small blood vessels could be seen within the partially emptied testis after spermiation. The lobular structures and blood vessel system were still intact in the regressed testis of male fish in the resting period. On the contrary, they became disintegrated in the residues of testis of the female fish.
In conclusion, these findings provide the important and fundamental cytological and histological basis for the further studies of ultrastructural changes in bisexual gonad and mechanism of natural and controlled sex change.
目 錄
中文摘要 ------------------------------------------------------------- I
英文摘要 ------------------------------------------------------------- IV
目錄 ------------------------------------------------------------------- VI
第一章 緒論 ------------------------------------------------------- 1
一、魚類精巢結構及精子生成過程
二、魚類卵巢結構及卵生成過程
三、雌雄同體魚生殖腺結構特徵
四、魚類生殖腺中的結締組織
第二章 黑鯛生殖腺發育型態之變化 ------------------------ 15
第三章 黑鯛精巢結構與精子生成微細構造 --------------- 36
第四章 黑鯛卵巢結構與卵子生成微細構造 --------- ----- 65
第五章 黑鯛生殖腺中的血管系統與結締組織 ------ ----- 107
第六章 結論 ------------------------------------------------------ 126
參考文獻 ------------------------------------------------------------ 128
謝辭 ------------------------------------------------------------------ 131
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