本研究選殖海鱺 (Cobia, Rachycentron canadum) 過氧化體增生活化受體 (peroxisome proliferator-activated receptors, PPARs) 基因，並探討養殖期間 PPAR mRNA 之表現及受飼料脂肪酸組成之影響。目的為透過 PPAR 表現和魚體體脂肪蓄積狀況關聯之瞭解，建立以飼料營養組成之操作，調控養殖海鱺體脂肪含量或分佈的技術，以改善肉質品質及市場的競爭力，提升養殖海鱺經濟價值。研究分成四部份，包括 PPARs 基因選殖、養殖過程 PPARs mRNA 表現與體脂肪狀態時間序列表現、成長狀況優劣與 PPARs 表現之關聯和飼料含不同脂肪酸組成對 PPARs 表現之影響。
選殖的海鱺 PPARα、PPARβ 和 PPARγ 基因 cDNA 全長分別為 2046-bp、2707-bp 及 1943-bp，可轉譯 476、510 及 531 個胺基酸。以 PPARs 基因部份片段，進行 RT-PCR 與即時定量 PCR (qPCR) 分析發現，PPARα mRNA 主要表現在血合肉、心臟及肝臟，在頭腎及背肉表現量最低；PPARβ mRNA 表現以心臟、肝臟、腦及幽門垂最多，PPARγ mRNA 主要表現在脂肪組織、肝臟及幽門垂。
養殖海鱺在移入箱網 (9週) 前之中間育成階段，肝臟、腹肉的脂肪細胞的直徑明顯增大呈現hypertrophy；14 週平均體重 330 g，肝臟和腹肉的脂肪積蓄量明顯增加，腹肉和背肉脂肪細胞數目顯著增加，但脂肪細胞明顯變小，呈現 hyperplasia。由 52 週採樣十次綜合結果而言，脂肪的蓄積過程主要先發生在肝臟，接著腹肉，最後在背肉。肝臟 PPARα mRNA表現量與魚體脂肪蓄積、脂肪細胞大小及密度呈顯著負相關 (p ＜ 0.05)。PPARγ mRNA 表現與身體脂肪蓄積、脂肪細胞大小及密度呈顯著正相關。這些結果顯示，PPARα 及PPARγ mRNA 之表現對海鱺肝臟、肌肉及內臟脂肪團中脂質代謝及脂肪蓄積是重要的。
比較蓄養於室外育成池18週，同批而成長呈優劣不同的兩群海鱺，和成長較差之小型魚 (體長約 36.8 公分；體重約 386 公克) 相較，成長較優的大型魚 (體長約54.1公分；體重約 1,287 公克)，肝臟脂肪 (35%) 及中性脂質含量高於小型魚 (26%)，其肝臟、腹肉脂肪細胞體積也有大於小型魚的趨勢。成長優劣狀況顯著影響海鱺肝臟PPARα mRNA 表現量及肝臟脂肪蓄積量。小型魚肝臟 PPARα mRNA 的表現量顯著高於大型魚。
以含15% 油脂飼料餵飼80 g 海鱺十週，探討飼料脂肪酸組成對PPARs mRNA表現的影響。肝臟PPARα mRNA之表現量與飼料C18:3n-3含量呈顯著正相關，而與C18:0含量呈顯著負相關；肝臟PPARβ mRNA表現量與飼料C16:0或C18:0含量呈顯著正相關；PPARγ則與飼料C20:1n-9、C20:5n-3、C20:6n-3含量呈顯著正相關 (p < 0.001)。
本研究結果顯示，海鱺飼料魚油可以部份由植物油取代，除長鏈 (C>20) 不飽和脂肪酸含量外，適當減少C18:3n-3 或增加C18:0含量，也可以增加海鱺體脂肪的蓄積，海鱺體脂肪蓄積之增減明顯與肝臟 PPARs mRNA 的表現具一定相關性，因此可以透過對 PPAR 表現研究，建立飼料配方策略，用來調控養殖海鱺魚體脂肪的含量。

The present study cloned full-length genes of peroxisome proliferators activated receptors (PPARs) of the cobia Rachycentron canadum and investigated their expressions in association with cobias’ body adiposity and lipid-metabolism related physiological parameters. In addition to gene cloning, several studies evaluating the roles of PPARs were carried out, including: a time-series study on cage-farmed cobias from week 5 to week 52 post-hatching, a study comparing fish groups with contrasting growth performance and a study elucidating the effects of dietary fatty acids.
Three isotypes, PPAR α, PPARβ and PPARγ, that were cloned from cobia’s cDNA contained 2046 bp, 2702 bp and 1943 bp, respectively. Their open reading frames encode 476, 510 and 531 amino acids, respectively. The identity in amino acid sequences between the PPARs are 52% (between PPARα and PPARβ), 52% (between PPARα and PPARγ), and 44% (between PPARβ and PPARγ), respectively. RT-PCR and real-time PCR (qPCR) analyses showed that expression of PPARα mRNA predominated in red muscle, heart and liver, and at a lower level in the head kidney and dorsal muscle. PPARβ transcripts were particularly abundant in the heart, liver, brain, and pyloric caeca. In contrast, PPARγ mRNA was detected primarily in the adipose tissues, liver, and pyloric caeca.
In the time-series study, the PPARs expression was related to the body adiposity and lipid-metabolism related physiological parameters of the cobias that were raised for one year to approximately 4.5 Kg in a commercial cage-culture farm. Ten samplings were conducted on weeks 5, 7, 9, 14, 18, 23, 29, 34, 41, and 52 post-hatching. The cobias were raised in an outdoor nursery to 88 g before being transferred to an offshore cage on week 9. The adipocytes in the liver and ventral muscle showed a hypertrophic (increase in cell size) increase towards the end of the nursery phase. Their cell size decreased significantly after the cage transfer and was maintained afterwards a size spectrum dominated by small cells until week 34. The cobias grew rapidly after the offshore transfer and reached 330 g on week 14. They showed a concurrent increase in fat deposition in the liver and ventral muscle and a concurrent hyperplasia increase in density of adipocytes in the ventral and dorsal muscle. Adipocyte hypertrophy was obvious on week 41 and regressed afterwards. As the fish grew, serum phospholipids concentration increased significantly from approximately 380 to 750 mg/dL. Time-series pattern for the specific activity of two NADPH-generating enzymes, malic enzyme and glucose-6- phosphate dehydrogenase, were reciprocal and compensatory. The expression of liver PPARα mRNA was negatively correlated to fat deposition and adiposity. There was a significant increase in body lipid deposition and hepatic PPARγ expression as the fish grew. Hepatic PPARγ expression could be a sufficient parameter describing its expression in whole body. These results showed that PPARγ and PPARα played a pivotal role in the control of lipid metabolic and storage functions in the liver, muscle and visceral fat depot of the cobia.
In the study comparing differential fish growth, two groups of cobias were selected based on their growth performance from a same batch of fish raised in a nursery. The large-size group that was regarded as superior grower was 54.1 cm in total length and 1,287 g in weight; while the small-size fish (inferior grower) was 36.8 cm and 386 g. Compared to large cobias, small cobias showed a similar hepatosomatic index and viscerasomatic index, but a significantly (p ＜ 0.05) smaller mesenteric fat index (MFI).The levels of crude lipid in the liver (35% vs. 26%) and the proportions of neutral lipids in lipid were higher in large cobias than in small cobias. Concentrations of serum phospholipids, free fatty acids and total protein of large cobias were significantly higher than those of small cobias. Adipocyte density of liver and ventral muscle was increased with increasing fish size. The PPARα mRNA expression in the liver of small cobias was significantly higher (p ＜ 0.05) than large cobias, ascribing to possible stress effect from their inferior growth. The growth superiority obviously affected PPARα mRNA expression and fat deposition in the liver. In general, the expression of liver PPARα mRNA was negatively correlated to body weight, body length, MFI, and serum NEFA, as well as lipid concentration, adiposity (adipocyte density and adipocyte size), G6PDH enzyme activity in the liver. The PPARγ mRNA expression in the liver was positively correlated to size of the adipocytes size.
The effects of dietary fatty acids on PPARs expression were evaluated in a 10-week growth trial, in which cobias with an initial weight of 80 g were fed diets containing 15% lipid. Among the lipids, 6% was fish oil and the remaining 9% were fish oil (rich in EPA and DHA), perilla oil (C18:3n-3), safflower oil (C18:2n-6), olive oil (C18:1n-9) or palm oil (C16:0). Significant difference was detected in PPARs mRNA expression among dietary treatments and among tissues. In the liver, among the dietary treatments, significantly higher expression levels of PPARα mRNA were detected in perilla oil and olive oil group, PPARβ mRNA in palm oil group and PPARγ mRNA in fish oil group. Linear regression analysis showed that liver PPARα mRNA expression was positively (p ＜ 0.05) correlated with dietary C18:3n-3 levels and negatively with dietary C18:0 levels. Liver PPARβ mRNA expression was positively correlated to C16:0 or C18:0 levels in diets. The PPARγ expression was positively (p ＜ 0.001) correlated to dietary levels of C20:1n-9, C20:5n-3 and C22:6n-3.
In summary, the mRNA expression pattern of PPARs was tissue or organ-specific with the expression of PPARα occurred predominantly in the liver and PPARγ in the adipose tissues. The expressions of PPARs in the liver were more related to their physiological roles than in other tissues or organs studied in the present study. The expression of PPARα in the liver was shown correlated negatively to body fat deposition; and reciprocally, expression of PPARγ was positively correlated to fat deposition. PPARs mRNA expression was also associated with major dietary fatty acids. Increased dietary C18:0 levels down-regulated PPARα and up-regulated PPARβ. Up-regulation of PPARγ was significantly related to increased levels of highly (C>20) unsaturated fatty acid in diets. Dietary C16–C18 fatty acids on the other hand were more related to expressions of PPARα and PPARβ. These results suggest that fish oil could be partially replaced by plant oils as the lipid source in the diet of the cobia. In addition to highly unsaturated fatty acids, reduction in dietary C18:3n-3 and increase in C18:0 lead to increased fat deposition, implicating a possible strategy to modulate body lipid contents of the cobia through dietary manipulation.

目錄

頁次
中文摘要……………………………………………………………… I
英文摘要……………………………………………………………… III
目錄…………………………………………………………………… VIII
表目錄………………………………………………………………… X
圖目錄………………………………………………………………… XII
一、 前言…………………………………………………………… 1
二、 文獻回顧……………………………………………………… 6
2.1 魚類體脂肪之分佈……………………………… 3
2.2 食物脂肪的運送…………………………………… 9
2.3 魚類脂質生合成…………………………………… 14
2.4 脂肪細胞的分化…………………………………… 17
2.5 脂肪細胞的分化之過程…………………………… 21
2.6 與脂肪細胞分化有關的轉錄因子………………… 23
2.7 PPARs 的生理效應…………………………………… 25
三、 綜合材料與方法……………………………………………… 41
3.1 實驗動物採集………………………………………… 41
3.2 肝臟中 total RNA 之萃取…………………………… 41
3.3 RNA 變性電泳分析…………………………………… 42
3.4 互補去氧核醣核酸（cDNA）之製備…………………… 42
3.5 Primer之設計………………………………………… 43
3.6 即時定量 PCR……………………………………… 43
3.7 一般成分分析………………………………………… 46
3.8 血清脂質含量分析…………………………………… 48
3.9 脂肪細胞直徑分佈分析…………………………… 49
3.10 脂質生合成相關酵素活性分析…………………… 50
四、 過氧化體增生活化受體 (PPARs) 基因選殖 (實驗一) …… 52
4.1 摘要…………………………………………………… 52
4.2 研究緣由……………………………………………… 53
4.3 實驗過程……………………………………………… 54
4.4 結果…………………………………………………… 64
4.5 討論…………………………………………………… 76
五、 海鱺成長各階段 PPARs mRNA 表現與體脂肪之關聯 (實驗二) …………………………………………………………… 82
5.1 摘要…………………………………………………… 82
5.2 研究緣由……………………………………………… 83
5.3 實驗過程……………………………………………… 84
5.4 結果…………………………………………………… 88
5.5 討論…………………………………………………… 107
六、 成長優劣狀況對 PPARs mRNA 表現及其體脂肪之影響 (實驗三) ………………………………………………………… 112
6.1 摘要…………………………………………………… 112
6.2 研究緣由……………………………………………… 113
6.3 實驗過程……………………………………………… 113
6.4 結果………………………………………………… 117
6.5 討論………………………………………………… 130
七、 飼料脂肪酸組成對 PPARs mRNA 表現之影響 (實驗四)…… 138
7.1 摘要…………………………………………………… 138
7.2 研究緣由……………………………………………… 139
7.3 實驗過程……………………………………………… 141
7.4 結果…………………………………………………… 152
7.5 討論…………………………………………………… 158
八、 總結論…………………………………………………………… 162
九、 參考文獻………………………………………………………… 164
附錄一 英文縮寫對照表…………………………………………… 186
附錄二 本論文研究期間發表之相關論文

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