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研究生:潘咸安
研究生(外文):Hsien-An Pan
論文名稱:DAZL在顆粒黃體細胞的表現及功能分析
論文名稱(外文):Expression and functional analysis of DAZL in granulosa-lutein cells
指導教授:蔡少正郭保麟郭保麟引用關係
指導教授(外文):Shaw-Jenq TsaiPao-Lin Kuo
學位類別:博士
校院名稱:國立成功大學
系所名稱:臨床醫學研究所
學門:醫藥衛生學門
學類:醫學學類
論文種類:學術論文
論文出版年:2008
畢業學年度:96
語文別:英文
論文頁數:124
中文關鍵詞:DAZL細胞凋亡DAZAP1固醇類生合成顆粒黃體細胞CDC25黃體
外文關鍵詞:DAZLapoptosissteroidogenesisgranulose-lutein cellcorpus luteumCDC25DAZAP1
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DAZL (Deleted in Azoospermia Like)基因是DAZ(Deleted in Azoospermia)基因家族的成員之一,在這個基因家族中包含有BOULE、DAZ和DAZL三個基因,BOULE是最起始的成員,之後再逐漸演化成DAZL和DAZ二個基因。DAZL基因在許多物種的生殖功能上扮演重要的角色,若將老鼠的dazl基因除去,會造成公鼠及母鼠都不孕。在果蠅若將DAZL之同源基因boule除去,雄性果蠅將完全沒有精蟲。DAZL蛋白質表現在不同物種的生殖細胞,在Caenorhabditis elegans,破壞DAZL的表現會使卵子細胞的減數分裂停止,在Xenopus DAZL是早期優勢生殖細胞分化與生殖細胞由內皮層開始移轉所必需,至於在小鼠方面,喪失DAZL同源基因的功能會造成男性與女性生殖細胞的缺失。利用免疫組織染色可以發現在人類男性睪丸中DAZL表現在各個不同時期的精子細胞,從精原細胞到減數分裂時期的精細胞到成熟的精細胞。而在人類女性的卵子形成過程中可以發現DAZL也表現在卵母細胞及發育中的卵子,更進一步在初級濾泡中的顆粒細胞、成熟濾泡的內膜細胞和由本研究中發現黃體內的顆粒黃體細胞也可以看到DAZL的表現。黃體是由優勢濾泡在排卵作用之後所發育形成的,它是卵巢在後排卵時期產生固醇類荷爾蒙的主要來源,黃體是濾泡發育的最終形態,也是卵巢維持早期懷孕成功的重要內分泌構造,由此可見DAZL在女性生殖的調控扮演重要的角色。黃體分泌黃體素是維持早期懷孕的重要因素,而藉由文獻及本研究報告知道DAZL在黃體化顆粒細胞及黃體細胞都有表現,於是推測DAZL應可調控黃體細胞的分泌黃體素的功能,因此本研究探討DAZL在顆粒黃體細胞扮演調控黃體素生成的機制。藉由瑩光免疫染色法,基因轉殖技術,及西方點漬方法,比較DAZL濃度的多寡是否會造成固醇類合成過程中一些重要的改變,經確認黃體素的分泌顯然和DAZL的表現呈正相關。但令人訝異的是,我們在DAZL量降低時,並未觀察到速率決定步驟酶StAR RNA量及蛋白質量的變化,於是我們將轉殖DAZL的顆粒黃體細胞做了細胞存活及凋亡的實驗,令人振奮地發現,DAZL蛋白質量的多寡和細胞凋亡呈正相關而這結果和逐日累計的黃體素量亦呈正相關,至此我們證實DAZL的存在的確會影響黃體素的分泌量改變,但不是經由影響固醇類生合成的路徑,而是藉由影響顆粒黃體細胞凋亡的機制。
The DAZ (deleted in azoospermia) gene family consists of three members in human beings: DAZ on Yq11.2, DAZL(DAZ-like) on 3p25, and BOULE on 2q33. The DAZ gene is present only on the Y chromosomes of great apes and Old World monkeys. The autosomal DAZL gene is present in all vertebrates studied while BOULE orthologs have been isolated from Drosophila and Caenorhabditis elegans in addition to mice and humans. It is believed that the DAZ gene arose from the transposition, repeat amplification, and pruning of the autosomal gene DAZL approximately 40 million years ago, during primate evolution. DAZL is unique in terms of its tissue distribution patterns among DAZ gene family members, considering its expression not only in germ cells but also in somatic cells. In the female reproductive system, DAZL is expressed in the fetal oogonium, adult oocytes, granulosa cells, and theca interna. In earlier reports, we showed evidence that DAZL protein is expressed in granulosa-lutein cells from different phases of human corpus luteum (CL), and in pre-implantation embryo of mice. Because DAZL is abundantly detected in early and mid-stage human corpus luteum, a possible but as-yet-unknown role for DAZL in modulating luteal function has been suggested. We aim to investigate whether DAZL could affect the luteal function by regulating the steroidogenic capacity and survival of luteal cells. Human granulosa-lutein cells (GLCs) were obtained from women undergoing assisted reproduction treatment at the IVF center. Immunohistochemical stain and Western blot were used to confirm and semi-quantify the DAZL expression. Transient transfection and siRNA knockdown approaches were used to investigate gain- or loss- of-function of DAZL. The major product of luteal cells, progesterone, was measured by a competitive ELISA procedure. Finally the DAZL anti-apoptotic effect was measured by TUNEL assay, flow cytometry and cell viability test.
DAZL expression was demonstrated in human GLCs. Progesterone production is positively correlated to the DAZL expression amount but the expression of several steroidogenic enzymes was not affected by levels of DAZL. Overexpression of DAZL increased cell survival while knocking down DAZL increased apoptosis of GLC. The present study provides the first evidence that the amount of DAZL expression has an anti-apoptotic effect and ensures the progesterone secretion to maintain the luteal cell function.
Table of Contents ------------------------------------------------------ i
List of Tables ---------------------------------------------------------------- iv
List of Figures --------------------------------------------------------------- v
Abstract --------------------------------------------------------------------- vii
中文摘要 ------------------------------------------------------------------- ix
Chapter 1 Literature Review
DAZ family ------------------------------------------------------------------ 1
DAZL ------------------------------------------------------------------------- 4
Steroidogenesis -------------------------------------------------------------- 7
DAZAP1 --------------------------------------------------------------------- 9
CDC25 ---------------------------------------------------------------------- 13
Overall subject ------------------------------------------------------------- 16
Chapter 2 Materials and Methods
Collection of ovarian tissue ----------------------------------------------- 17
Generation of anti-DAZL antibody -------------------------------------- 17
Immunohistochemical staining of corpus luteum --------------------- 18
Total RNA isolation and cDNA synthesis of corpus luteum --------- 19
Preparation of native and competitor RNA ----------------------------- 19
Standard curve QC-RT-PCR --------------------------------------------- 21
Reagents --------------------------------------------------------------------- 22
Preparation of GLCs and primary culture ------------------------------ 23
Granulosa – lutein cells Culture and Treatments ---------------------- 24
RNA isolation and RT-PCR of GLCs ----------------------------------- 25
Immunohistochemical staining of GLCs ------------------------------- 26
SDS-PAGE and Western immunoblotting of GLCs ------------------ 27
TdT-mediated dUTP Nick-End Labeling (TUNEL) Assay --------- 28
Detection of apoptotic nuclei -------------------------------------------- 29
Progesterone assays ------------------------------------------------------- 29
Generation of anti-DAZAP1 antibodies -------------------------------- 30
Co-immunoprecipitation (IP) -------------------------------------------- 30
Western blot for CDC25A, -B, -C --------------------------------------- 31
RNA-immunoprecipitation (RNA-IP) ---------------------------------- 32
Data analysis --------------------------------------------------------------- 35
Chapter 3 Results
(I) Expression of DAZL protein in human corpus luteum ----------- 36
(II) DAZL regulates progesterone biosynthesis through inhibition of apoptosis but not by induction of steroidogenesis in human granulosa-lutein cell -------------------------------------------------- 38
(III) Expression patterns of the DAZAP1and in-vivo interaction of DAZL and DAZAP1 in human ovaries --------------------------- 42
(IV) CDC25 protein expression and interaction with DAZL in human corpus luteum --------------------------------------------------------- 44
Chapter 4 Discussion and Perspective --------------- 46
References ---------------------------------------------------------------- 55
Appendixes
(I) Dazl protein expression in mouse preimplantation embryo ------- 100
(II) Award --------------------------------------------------------------------- 117
作者簡歷 ---------------------------------------------------------------- 118
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