本論文為探討維生素C (L-ascorbic acid, AA)及其四種衍生物﹐包括L-ascorbyl-2-sulfate (C2S)﹑L-ascorbyl-2-polyphosphate (C2PP)﹑L-ascorbyl-2- monophosphate-Na (C2MP-Na)與L-ascorbyl-2-monophosphate-Mg (C2MP-Mg)﹐及與他種營養素包含銅和維生素E間之交互作用﹐對草蝦稚蝦成長﹑免疫反應﹑緊迫耐受性以及疾病抵抗力之影響。實驗結果顯示﹐適量及高量 (適量之五倍)之維生素Ｃ及其衍生物均可促進草蝦之免疫反應﹐包括血球細胞計數 (total haemocyte count, THC)﹑血球細胞超氧陰離子 (superoxide anion, O2-)產率及酚氧化酵素 (phenoloxidase, PO)活性﹐提高草蝦之硝酸-氮 (3,800 mg/L)及低鹽度 (0 ppt)緊迫試驗的耐受性﹐以及增加草蝦對於病原性弧菌 (Vibrio damsela與V. vulnificus)感染之抵抗力。提高飼料中維生素C及其衍生物之含量(適量之五倍)﹐可提高草蝦組織 (肝胰臟與肌肉)中維生素C之濃度及其對於病原性弧菌感染之抵抗力。在不同維生素Ｃ衍生物之間﹐草蝦之THC以餵予C2MP-Mg及C2PP組顯著優於C2MP-Na﹑AA及C2S各組﹐而草蝦組織中對病原性弧菌感染之抵抗力則依序為AA=C2MP-Mg=C2PP>C2MP-Na>C2S。基礎飼料中分別添加銅 (0, 10, 20, 30, 40, 80, 160 mg /kg)及維生素E (0, 25, 50, 75, 100, 150, 200, 400 mg/kg)﹐探討草蝦銅及維生素E之需求及其與免疫反應之相關性。實驗結果顯示﹐以草蝦增重率 (weight gain)﹑飼料效率 (feed efficiency)及蝦體銅蓄積量以多元迴歸分析及其免疫反應(THC及O2-產率)之表現﹐顯示草蝦銅之最適需求量為10-30 mg Cu/kg；以增重率及THC以多元迴歸分析求得草蝦維生素E之最適需求量為85-89 mg vitamin E/kg。在維生素C (C2MP-Mg)與銅之交互作用方面﹐提高飼料中維生素C含量 (適量之五倍)﹐減少草蝦對於飼料中高量銅 (適量之八倍)的吸收﹐降低蝦體內銅之蓄積量﹐改善飼料中高量銅對於草蝦成長及免疫反應(THC及O2-產率)造成的抑制作用。在維生素C (C2MP-Mg)與維生素E之交互作用方面﹐提高飼料中維生素C含量(適量之五倍)﹐可促進草蝦之維生素E再生﹐提高肝胰臟與肌肉中維生素E濃度﹐降低組織中代表脂肪氧化程度的TBA (thiobabituric acid)反應物濃度﹐改善低量 (適量之1/4)或高量 (適量之4倍)維生素E對草蝦成長及免疫反應 (THC﹑O2-產率及superoxidase dismutase)的抑制作用﹐以及增加草蝦對於V. damsela弧菌感染的抵抗力。綜合本研究之結果﹐維生素C﹑銅及維生素E皆為草蝦維持成長與免疫反應之必需營養素。維生素Ｃ及其衍生物可促進草蝦之免疫反應﹑緊迫耐受性以及疾病抵抗力﹐而且提高飼料中維生素C及其衍生物的含量(適量之五倍)﹐可提高草蝦之疾病抵抗力﹑降低飼料中高量銅對於草蝦的毒性以及促進維生素C對於維生素E之節約作用。此外﹐不同維生素C衍生物之間對於草蝦稚蝦免疫反應及疾病抵抗力具有不同之效應﹐整體而言﹐以C2MP-Mg及C2PP效應較其他維生素Ｃ衍生物為高。草蝦肝胰臟與肌肉中的維生素C濃度﹐也許可作為草蝦之抗病指標。

The effects of L-ascorbic acid (AA) and its four derivatives namely, L-ascorbyl-2-sulfate (C2S), L-ascorbyl-2-polyphosphate (C2PP), L-ascorbyl-2- monophosphate-Na (C2MP-Na) and L-ascorbyl-2-monophosphate-Mg (C2MP- Mg), on the growth, immune responses, stress tolerances and disease resistances for juvenile grass shrimp, Penaeus monodon, were investigated. The interaction between ascorbate and dietary vitamin E and copper (Cu) were also studied. Significantly (P<0.01) higher immune responses including total haemocyte count (THC), superoxide anion (O2-) production ratio and phenoloxidase (PO) activity, stress tolerances with nitrate (3,800 mg/L) and low salitity (0 ppt) exposed, and resistances against vibriosis (Vibrio damsela and V. vulnificus) infection were observed in shrimp fed diets supplemented with either adequate or high level (5X adequate) of ascorbate than shrimp fed the vitamin C deprived control diet regardless of the ascorbate source. Shrimp fed high ascorbate containing diets had higher vitamin C concentration in tissue (heapatopancreas and muscle) and disease resistances than shrimp fed diets with adequate level of ascorbate. Among the ascorbate sources, shrimp fed C2MP-Mg and C2PP containing diets had higher THC than shrimp fed AA, C2S and C2MP-Na containing diets. The vitamin C concentration in tissue and the effects of ascorbate derivatives on resistance against vibriosis infection are in the order of : AA = C2PP = C2MP-Mg > C2MP-Na > C2S, regardless of the supplementation level. Graded levels of dietary copper (Cu) (0, 10, 20, 30, 40, 80, 160 mg Cu/kg) and dietary vitamin E (0, 25, 50, 75, 100, 150, 200, 400 mg vitamin E/kg) were each fed to the shrimp to quantify their requirements. Analysis by polynomial regression of WG (weight gain), feed efficiency and by linear regression of the whole-body Cu retention of shrimp indicated that the adequate dietary Cu concentration in growing P. monodon is about 15-21 mg Cu/kg. The immune indicators (THC and O2- production ratio) suggest that an adequate dietary Cu concentration for immune responses is about 10-30 mg Cu/kg. Polynomial regression analysis of WG, THC and hepatopancreatic vitamin E concentration of the shrimp indicated that a level of 85-89 mg vitamin E/kg is required for maximal growth and immune responses of P. monodon. The interaction of vitamin C (C2MP-Mg) with either adequate or high (8X adequate) dietary Cu indicated that increase of dietary vitamin C improved growth and immune responses and prevented tissue Cu accumulation in P. monodon fed with high dietary Cu. A 3 x 3 factorial study in which shrimp were fed either low (1/2 adequate, 20mg/kg), adequate (40 mg/kg) or high (5X adequate, 200 mg/kg) vitamin C (C2MP-Mg) diets and each with low (1/4 adequate, 25), adequate (100 mg/kg) and high (4X adequate, 400 mg/kg) vitamin E (DL-α-TOA) revealed that hepatopancreatic and muscle vitamin E concentrations were increased as dietary vitamin C increased. Tissue thiobabituric acid value decreased as dietary vitamin C increased. Increasing of dietary vitamin C improved growth, immune responses (THC, O2- and superoxide dismutase) and vibriosis resistance (V. damsela) for shrimp fed with low and high vitamin E diets. In summary, this study demonstrates that vitamin C and its derivatives, copper and vitamin E are immunostimulant for P. monodon. Increase of dietary ascorbate derivatives enhances resistance of P. monodon against vibriosis infection. The immune stimulating effects and disease resistance of P. monodon by ascorbate derivatives are in the order of C2MP-Mg=C2PP>C2MP-Na=C2S. High C2MP-Mg level increased copper antagonism effect and improves copper-related immunotoxicity. Increase of C2MP-Mg show a vitamin E sparing effect on growth, immune responses and disease resistance of P. monodon. The hepatopancreatic and muscle vitamin C concentration may used as disease resistance indicator for P. monodon.

目錄
總摘要 I
Abstract III
第一章 文獻整理 1
1-1 前言 2
1-2 文獻回顧 4
1-3 研究方向 18
第二章 維生素C及其衍生物對草蝦稚蝦免疫反應相關性之研究 21
2-1 摘要 22
2-2 前言 23
2-3 材料與方法 24
2-4 結果 34
2-5 討論 37
2-6 結論 44
第三章 維生素C及其衍生物對草蝦稚蝦緊迫耐受性與疾病抵抗之研究 45
3-1 摘要 46
3-2 前言 47
3-3 材料與方法 48
3-4 結果 51
3-5 討論 58
3-6 結論 63
第四章 草蝦稚蝦銅之需求及其對免疫反應相關性之研究 65
4-1 摘要 66
4-2 前言 67
4-3 材料與方法 68
4-4 結果 73
4-5 討論 79
4-6 結論 81
第五章 維生素C與銅之交互作用對草蝦稚蝦免疫反應之影響 83
5-1 摘要 84
5-2 前言 85
5-3 材料與方法 86
5-4 結果 92
5-5 討論 97
5-6 結論 99
第六章 草蝦稚蝦維生素E之需求及其對免疫反應相關性之研究 101
6-1 摘要 102
6-2 前言 103
6-3 材料與方法 105
6-4 結果 110
6-5 討論 114
6-6 結論 121
第七章 維生素C與維生素E之交互作用對草蝦稚蝦疾病抵抗之探討 123
7-1 摘要 124
7-2 前言 126
7-3 材料與方法 127
7-4 結果 132
7-5 討論 139
7-6 結論 141
總結論 143
參考文獻 147
<附錄> 本論文已發表部份之全文 169
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附錄2. Lee, M.H., Shiau, S.Y., 2002. Dietary copper requirement of juvenile grass shrimp, Penaeus monodon, and effects on non-specific immune responses. Fish & Shellfish Immunology 13, 259-270. 183
附錄3. Lee, M.H., Shiau, S.Y., 2003. Increase of dietary vitamin C improves haemocyt e respiratory burst response and growth of juvenile grass shrimp, Penaeus monodon, fed with high dietary copper. Fish & Shellfish Immunology 14, 305-315. 197

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