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研究生:黃仕聰
研究生(外文):Shih-Tsung Huang
論文名稱:點帶石斑(Epinepheluscoioides)生長素釋質(ghrelin)基因之分子選殖及特性分析
論文名稱(外文):Molecular cloning and characterization of ghrelin gene in orange-spotted grouper ( Epinephelus coioides )
指導教授:陸振岡
指導教授(外文):Jenn-Kan Lu
學位類別:碩士
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:2007
畢業學年度:95
語文別:中文
論文頁數:137
中文關鍵詞:生長素釋質促生長素分泌素攝食分子選殖
外文關鍵詞:ghrelinGHSfeedingmolecular cloning
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生長素釋質 (Ghrelin) 為一種內生型促生長荷爾蒙分泌素 (growth hormone secretagogue, GHS),能促進GH 由腦下垂體分泌,近來年證實ghrelin 也與攝食行為有關。本論文目標是研究點帶石斑 (Epinephelus coioides) 生長、攝食調控因子 ─ ghrelin 基因之分子選殖及其特性分析。根據已知脊椎動物ghrelin 之胺基酸及核酸序列設計退化性引子,以點帶石斑胃組織之cDNA 使用RT-PCR及RACE等方法選殖出ghrelin基因cDNA 序列,其全長為1485 個核苷酸,可轉譯出102 個胺基酸序列。本研究亦選殖出Ghrelin 基因體DNA 為3257 個核苷酸,ghrelin基因結構由四個編碼子及三個插入子所構成。經由Real-time PCR 分析結果顯示點帶石斑ghrelin 基因廣泛表現於各組織中,且以胃表現量最高。在攝食前胃中ghrelin基因及腦中NPY及GH基因表現量逐漸上升,並於攝食後下降。飢餓使ghrelin 基因表現量上升,此與GH 及NPY 基因表現情形相類似。人工合成的人類ghrelin及八碳、十碳修飾的石斑ghrelin皆能促進體外培養腦組織中之GH 及NPY 基因表現。內腦腔 (I.C.V.) 注射八碳修飾之石班ghrelin 可促進腦中GH 及NPY 基因表現。於石斑飢餓時注射葡萄糖,造成胃中ghrelin 基因表現量下降。高脂肪飼料能顯著的降低ghrelin 基因表現量。使用飼料及棉花進行胃部填充能使ghrelin 基因表現量下降。具厭食症的石斑魚其胃中ghrelin 基因表現量顯著的高於正常攝食之石斑。魚類運輸撈捕時所造成的緊迫使ghrelin 基因表現量降低,低溫處理則不會影響ghrelin 表現量。誘引劑可增加ghrelin 基因表現量,推測ghrelin 基因表現量可能受到嗅覺的影響。
Ghrelin an endogeneous growth hormone secretagogues (GHSs), that stimulating growth hormone secretion from pituitary. In recent study show that ghrelin is related in appetite. In this study, we have cloned and characterization of the full-length cDNA of ghrelin gene from orange-spotted grouper (Epinephelus coioides). We base on the ghrelin sequence of vertebrate animals to design degenerate primers. Use the cDNA of stomach tissue to clone the ghrelin cDNA sequence by RT-PCR and RACE methods. Grouper full-length ghrelin cDNA was 1485 bp in length and deduced 102 amino acids. The length of ghrelin genomic DNA was 3257 bp, and the genomic DNA structure was composed of four exons and three introns. Real-time quantitative PCR analysis showed ghrelin gene distribution in every tissues, and dominant expressed in stomach. Grouper maintained on a scheduled feeding regimen show that ghrelin, GH and NPY mRNA level increase before the feeding time, and the level rapid decrease after the feeding time. Food deprivation cause the ghrelin, GH and NPY mRNA level increase. The synthetic humen ghrelin and the grouper ghrelin that was modified by octanoic acid and decanoic acid all cause the GH and NPY mRNA level increase in vitro. Brain intracerebroventricular (i.c.v) injection of synthetic grouper ghrelin showed the stimulation of GH and NPY mRNA level. Glucose intraperitoneally (IP) injection decreased the ghrelin mRNA level. Use feed and cotton to fill the stomach showed that ghrelin gene expression was significantly decreased. Higher ghrelin mRNA was found in anorexia grouper. To handle fish would cause stress that decreased the ghrelin mRNA level in grouper. Attractant increased ghrelin mRNA expression, showed that ghrelin might related to olfaction.
謝辭 ……………………………………………………………… i
中文摘要 ………………………………………………………… ii
英文摘要 ………………………………………………………… iii
目錄 ………………………………………………………… iv
表目錄 ………………………………………………………… vii
圖目錄 ………………………………………………………… viii
壹、前言 ………………………………………………………… 1
一、石斑魚之生物學與養殖概況…………………… 1
二、動物的生長……………………………………… 3
三、飽足、攝食與能量平衡………………………… 7
四、促生長素分泌素 (GHS)………………………… 11
五、生長素釋質受體 (GHS-R)……………………… 13
六、生長素釋質 (ghrelin)………………………… 15
七、本研究目標……………………………………… 20
貳、實驗材料……………………………………………………… 24
參、實驗方法……………………………………………………… 25
1、點帶石斑 ghrelin 基因cDNA 序列選殖………… 25
2、點帶石斑ghrelin 基因全長cDNA 之選殖……… 33
3、點帶石斑ghrelin基因之基因體DNA序列分子選殖 38
4、點帶石斑ghrelin 基因在各組織間之表現模式… 40
5、點帶石斑ghrelin基因在胃部組織不同區域之表現模式 ……………………………………………………………41
6、固定攝食時間前後,點帶石斑胃組織ghrelin基因及腦組織NPY、GH 基因之表現模式…………………………………… 42
7、在連續饑餓的情況下,點帶石斑胃組織ghrelin 及腦組織NPY、GH基因之表現模式…………………………………… 43
8、三種不同的人工合成ghrelin 胜肽對體外培養之點帶石斑全腦組織NPY 及GH 基因表現之影響……………………… 43
9、點帶石斑腦室內注射石斑魚ghrelin 胜肽 (grouper ghrelin peptide) 對石斑魚NPY 及GH 基因表現量之影響…… 44
10、點帶石斑腦室內注射人類ghrelin 胜肽 (humen ghrelin peptide) 對石斑魚NPY 及GH 基因表現量之影響…… 45
11、腹腔內注射glucose 對點帶石斑於胃組織ghrelin基因表現量之影響…………………………………………………… 46
12、投餵高脂肪飼料對點帶石斑魚胃組織ghrelin 基因表現量之影響………………………………………………………… 47
13、空腹時進行胃部填充對點帶石斑ghrelin 基因表現量之影響……………………………………………………………… 47
14、探討正常攝食與產生厭食 (anorexia) 現象的石斑魚胃組織ghrelin基因表現量的差異……………………………… 48
15、探討與魚類運輸時捕撈及低溫對胃組織中ghrelin 基因表現量之影響…………………………………………………… 49
16、誘引劑對點帶石斑ghrelin 基因表現量之影響 50
肆、結果 ………………………………………………………… 51
一、分子選殖點帶石斑ghrelin cDNA 全長及與其他物種之比對……………………………………………………………… 51
二、點帶石斑ghrelin 基因體DNA (Genomic DNA) 之分子選殖………………………………………………………………… 53
三、ghrelin基因在點帶石斑魚之各組織之表現情形 54
四、ghrelin 基因於點帶石斑胃組織不同部位之表現情形…………………………………………………………………… 54
五、固定攝食週期下,點帶石斑腦組織之GH、NPY 基因及胃組織之ghrelin 基因在攝食前後之表現情形………………… 54
六、在連續饑餓 (food deprivation) 下,點帶石斑腦組織之GH、NPY基因及胃組織之ghrelin 基因表現情形………… 55
七、三種不同的ghrelin 蛋白質對石斑體外培養的腦組織NPY 及GH 基因表現量之影響…………………………………… 55
八、腦腔內 (I.C.V.) 注射石斑ghrelin 蛋白質對石斑NPY 及GH 基因表現量之影響…………………………………… 56
九、腦腔內 (I.C.V.) 注射人類ghrelin 蛋白質對石斑NPY基因表現量之影響…………………………………………… 56
十、空腹時以腹腔內 (I.P.) 注射葡萄糖溶液對石斑ghrelin 基因表現量之影響……………………………………… 57
十一、以高脂肪飼料進行投餵對ghrelin 基因表現量之影響…………………………………………………………………… 57
十二、空腹時進行胃部填充對點帶石斑ghrelin 基因表現量之影響…………………………………………………………… 58
十三、厭食症 (anorexia) 對石斑ghrelin 基因表現量之影響………………………………………………………………… 58
十四、魚類運輸過程之低溫及捕撈所造成的緊迫 (stress) 對胃組織ghrelin 基因表現量之影響……………… 58
十五、誘引劑對點帶石斑ghrelin 基因表現量之影響59
伍、討論 ………………………………………………………… 60
一、分子選殖點帶石斑ghrelin cDNA 全長及與其他物種之比對……………………………………………………………… 60
二、點帶石斑ghrelin 基因體DNA (Genomic DNA) 之分子選殖………………………………………………………………… 62
三、ghrelin 基因在點帶石斑魚之各組織之表現情 63
四、ghrelin 基因於點帶石斑胃組織不同部位之表現情形…………………………………………………………………… 64
五、固定攝食週期下,點帶石斑腦組織之GH、NPY 基因及胃組織之ghrelin 基因在攝食前後之表現情形………………… 65
六、在連續饑餓 (food deprivation) 下,點帶石斑腦組織之GH、NPY基因及胃組織之ghrelin 基因表現情形………… 66
七、三種不同的ghrelin 蛋白質對石斑體外培養的腦組織NPY 及GH 基因表現量之影響………………………………………68
八、腦腔內 (I.C.V.) 注射石斑ghrelin 蛋白質對石斑NPY 及GH 基因表現量之影響…………………………………… 70
九、腦腔內 (I.C.V.) 注射人類ghrelin 蛋白質對石斑NPY基因表現量之影響…………………………………………… 72
十、空腹時以腹腔內 (I.P.) 注射葡萄糖溶液對石斑ghrelin 基因表現量之影響……………………………………… 72
十一、以高脂肪飼料對ghrelin 基因表現量之影響 74
十二、空腹時進行胃部填充對點帶石斑ghrelin 基因表現量之影響…………………………………………………………… 75
十三、厭食症 (anorexia) 對石斑ghrelin 基因表現量之影響………………………………………………………………… 76
十四、魚類運輸過程之低溫及捕撈所造成的緊迫 (stress) 對胃組織ghrelin 基因表現量之影響……………… 77
十五、誘引劑對點帶石斑ghrelin 基因表現量之影響78
陸、參考文獻 ……………………………………………… 80
圖表 ………………………………………………………… 99
附錄 ………………………………………………………… 131


表目錄

表一、硬骨魚類促進及抑制攝食之胜肽………………………… 99
表二、各物種ghrelin cDNA 轉譯區域 (coding region) 核酸序列之同一性 (identitiy) 比較……………………………………… 100
表三、各物種ghrelin pre-pro peptide 胺基酸序列之相似性 (similarity) 比較……………………………………………… 101

圖目錄

圖一、GH-IGF 軸線 (axis) …………………………………… 102
圖二、典型的Growth hormone secretagogues (GHSs)…………103
圖三、GHRH 與GHS 促進腦下垂體分泌GH 之模式圖…………… 104
圖四、GHS receptor 之構形…………………………………… 105
圖五、Ghrelin 序列及醯化修飾 (acylation) ……………… 106
圖六、人工合成ghrelin 的醯化修飾 (Acylation) 過程…… 107
圖七、Ghrelin 的生物功能……………………………………… 108
圖八、點帶石斑ghrelin 基因cDNA 及演譯胺基酸序列……… 109
圖九、各物種ghrelin cDNA 轉譯區域序列比對 (I) ………… 110
圖十、各物種ghrelin cDNA 轉譯區域序列比對 (II) ……… 111
圖十一、各物種ghrelin cDNA 轉譯區域序列比對 (III) …… 112
圖十二、各物種ghrelin 基因演譯之胺基酸序列比對………… 113
圖十三、點帶石斑pre-pro ghrelin 胺基酸序列之演化樹型圖 114
圖十四、點帶石斑ghrelin基因體DNA (genomic DNA) 序列… 115
圖十五、點帶石斑gherlin 基因結構與轉錄及後轉錄作用意示圖…………………………………………………………………… 116
圖十六、不同脊椎動物ghrelin 基因結構之比較……………… 117
圖十七、點帶石斑組織中ghrelin 基因之表現情形…………… 118
圖十八、點帶石斑胃各區域之ghrelin 基因表現情形………… 119
圖十九、點帶石斑ghrelin、GH 及NPY 基因在固定攝食週期前
後之表現模式……………………………………………………… 120
圖二十、點帶石斑ghrelin、GH 及NPY 基因在連續饑餓情況下之表現模式………………………………………………………………… 121
圖二十一、三種合成的 ghrelin 蛋白質對體外培養之點帶石斑腦組織NPY 及GH 基因表現量之影響……………………………………122
圖二十二、腦腔內 (I.C.V.) 注射化學合成之點帶石斑ghrelin (octanoyl) 對腦內GH 及NPY 基因表現量之影響……………… 123
圖二十三、腦腔內 (I.C.V.) 注射不同濃度人類ghrelin蛋白質後,在不同時間腦中NPY 基因表現之的變化………………………… 124
圖二十四、腹腔 (I.P.) 注射不同濃度之葡萄糖溶液,對石斑魚胃組織ghrelin 基因表現之影響……………………………………… 125
圖二十五、高脂肪飼料對石斑魚胃組織ghrelin 基因表現之影響…………………………………………………………………… 126
圖二十六、攝食時間前進行胃部填充對石斑魚胃組織ghrelin 基
因表現之影響……………………………………………………… 127
圖二十七、正常攝食與患厭食症 (anorexia) 之點帶石斑魚胃組織中ghrelin 基因表現之情形………………………………………… 128
圖二十八、捕撈及低溫處理對點帶石斑胃組織ghrelin 基因表現
之影響……………………………………………………………… 129
圖二十九、誘引劑對點帶石斑ghrelin 基因表現量之影響…… 130
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