樟芝屬於無褶菌目、多孔菌科，據文獻指出具有抗癌、抗腫瘤及保肝之效果。樟芝的有效活性物質包含固醇類、三帖類、多醣體等。由於野生樟芝只寄生在台灣特有的牛樟樹上，不但數量少之又少，而且品質不穩定，所以近年來樟芝已被利用液態醱酵培養以達大量生產的目的。本研究探討脯氨酸和通氣量對於液態醱酵培養樟芝多醣體之影響，進而利用二階段批次醱酵之調控以達增加多醣體產量之目的，此外並利用巨噬細胞之體外測試(in vitro)，分析樟芝多醣體的抗腫瘤生物活性，評估其提升免疫能力的效果。本實驗所使用的樟芝菌株購自新竹食品工業研究所，編號為BCRC 35396。
搖瓶培養：不同氨基酸(精氨酸、麩氨醯酸、異白氨酸、離氨酸、脯氨酸、酥氨酸)在0.3%濃度之下對樟芝液態培養生物質量(biomass)及胞外多醣體(EPS)之影響。實驗結果顯示，離氨酸和脯氨酸分別有利於樟芝菌絲體生物質量及胞外多醣體之生合成，而使生物質量和胞外多醣體產量增加，分別為2.97g L-1 和 218.64mg L-1。進ㄧ步實驗，利用不同初始濃度脯氨酸(0.3%、0.6%、0.9%、1.2%、1.5%)之條件研究對樟芝液態培養生物質量和胞外多醣體之影響，控制組為0% 濃度脯氨酸。實驗結果顯示，樟芝生物質量和胞外多醣體之生合成隨著脯氨酸濃度增加而增加，在1.5%脯氨酸時，其生物質量之最大值為3.85g L-1；胞外多醣體產量則隨脯氨酸濃度上升而增加，直到脯氨酸濃度到達1.2%為止，其最大胞外多醣體產量為247.12mg L-1。
攪拌式醱酵槽培養：不同初始濃度脯氨酸對樟芝液態培養生物質量和多醣體產量之影響。批次醱酵實驗結果顯示在0.3%脯氨酸下有一比生長速率最大值(0.29d-1)；而在 0.9%脯氨酸時，可得生物質量、胞外多醣體、胞內多醣體(IPS)及胞內多醣體內含量(IPS content；mg IPS / g Biomass)最大值分別為 6.57g L-1、204.32g L-1、308.33g L-1和47.15mg g-1。此外，為考慮氧氣對樟芝液態醱酵培養的影響，於是利用不同通氣量(0.2vvm、0.4vvm、0.6vvm、0.8vvm、1.0vvm、1.2vvm)來探討對樟芝液態培養生物質量和多醣體產量的影響。批次醱酵實驗結果顯示在 通氣量0.2vvm之下，有最大胞外多醣體產量為188.65mg L-1；而在通氣量在1.2vvm 時，有最大生物質量產量、最大胞內多醣體產量及最大胞內多醣體內含量分別為5.89g L-1、289.98mg L-1和50.00mg g-1 。為了增加樟芝生物質量和多醣體產量，本研究亦進行二階段批次醱酵培養(添加0.9%脯氨酸培養基於醱酵槽中培養，伴隨前期操作通氣量為1.2vvm以利幫助菌體成長，待菌體成長至一定階段後，於醱酵中期第9天降低操作通氣量至0.2vvm以促進產物多醣體的生成增加)，實驗結果顯示生物質量、胞外多醣體、胞內多醣體及胞內多醣體內含量為 6.98g L-1、255.19g L-1、381.23g L-1 和55.65mg g-1，其中二階段批次醱酵之胞外多醣體產量(255.19g L-1)較0.9%脯氨酸醱酵之胞外多醣體產量(204.32g L-1)增加了24%；較通氣量為0.2vvm 醱酵之胞外多醣體產量(188.65mg L-1)增加了35%。
利用凝膠過濾管柱層析法分離純化樟芝粗多醣體，可得純化的二區分(fraction-1和fraction-2)。接著進行樟芝多醣體與蛋白質含量比分析，實驗結果顯示，二階段批次醱酵培養之樟芝多醣體與蛋白質含量比(胞外多醣體fraction-1：20.78% 和 胞內多醣體：38.69%)比各批次醱酵之含量比相對較高。在樟芝胞外多醣體分子量分析中，實驗結果顯示二階段批次醱酵培養之分子量 (fraction-1：1.24×106 Da)較各批次醱酵培養之分子量高；對於在胞內多醣體分子量分析，二階段批次醱酵之胞內多醣體分子量 (3.30×105 Da)也是比各批次醱酵相對稍高。在樟芝多醣體之中性單醣成分及組成分析顯示，無論是二階段批次醱酵或是各批次醱酵之單糖組成都是以葡萄糖、木糖、半乳糖單醣為主；但是組成比例卻幾乎不變，可以推論得知脯氨酸效應、氧氣質傳和二階段醱酵培養雖然會改變多醣體代謝機制，然而對多醣體本身的組成而言並不會造成影響。
對於樟芝液態醱酵培養之多醣體的生物活性評估(巨噬細胞激素測定)，從實驗結果發現生物活性(murine tumor necrosis factor-alpha concentration, mTNF-α conc.)與分子量成正比關係，多醣體分子量越大則生物活性效果越好。綜觀來說，由於二階段批次醱酵所得之樟芝胞外多醣體的分子量較大，因此其胞外多醣體之生物活性(fraction-1：1779.91 ± 46.52pg/mL和fraction-2：544.93 ± 44.39pg/mL)比各批次醱酵胞外多醣體之生物活性相對較好；同時二階段批次醱酵胞內多醣體之生物活性(567.42 ± 50.83pg/mL)也是比各批次醱酵胞內多醣體之生物活性相對稍好。此外，樟芝多醣體與蛋白質含量比結果以多醣體的生物活性成正比，多醣體與蛋白質含量比愈高則生物活性效果愈好。

Antrodia camphorata (Zang-Zhi) belongs to the aphyllophorales and the poly -poracease family with the effect of remedy for anticancer, antitumor and protection of the liver. Some bioactive compounds of A. camphorata including sesquiterpene lactone, steroids, triterpenoids have been isolated and characterized. Wild A. camphorata parasitizes specifically on Cinnamomum kanehirai hays in a small quantity and its quality is not stable. In recent years, A. camphorata was attempted to be produced on large scale by submerged culture. The main purpose of this research was to investigate the influence of L-Proline and aeration rate on polysaccharide fermentation by submerged culture of A. camphorata, and to increase production of exopolysaccharide (EPS) by regulating two-stage culture strategy. Moreover, the biological activity of polysaccharides from A. camphorate was examined by using TNF-α release capability of polysaccharides on a murine macrophage cell line, RAW264.7. A. camphorata BCRC 35396 was obtained from the Bioresources Collection and Research Center (BCRC) in Food Industry Research and Development Institute (Hsin chu, ROC).
The effects of the different amino acids at 3g/L of (i.e., L-Arginine, L-Glutamine, L-Isoleucine, L-Lysine, L-Proline, and L-Threonine) on production of biomass and EPS were conducted in this study. The result exhibited that L-Lysine and L-Proline stimulated biomass (2.97g L-1) and EPS (218.64mg L-1) production, respectively. In addition, the effects of various concentrations of L-Proline (i.e., 3, 6, 9, 12, 15 g/L) on biomass and EPS production of A. camphorata were conducted in shake-flask cultures. Comparing with controlled 0 g/L of L-Proline, the biomass production increased with L-Proline concentration ranging from 3 to 15 g/L. The optimal biomass concentration was reached to 3.85 g/L as L-Proline concentration was at 15 g/L, and the optimal EPS concentration was reached to 247.12 mg L-1 as L-Proline concentration was at 12 g/L.
The effects of different L-Proline concentration (i.e., 3, 6, 9, 12, 15 g/L) on biomass and EPS production of A. camphorata by submerged culture were conducted in stirred-tank fermentor. The maximal value of the specific growth rate (μ) was 0.29 d-1 at 0.3% L-Proline. At 9g/L of L-Proline, the maximum values were 6.57 g L-1, 204.32 g L-1, 308.33 g L-1, and 47 mg g-1 for biomass, EPS, endopolysaccharide (IPS), and IPS content production, respectively. In addition, the effects of the aeration rates (i.e., 0.2, 0.4, 0.6, 0.8, 1.0, 1.2 vvm) on production of biomass and EPS of A. camphorata by submerged culture were also conducted in stirred-tank fermentor in this study. Controlled with aeration of 0.2 vvm, the maximum value was 188.65mg L-1 for EPS production ; controlled with aeration of 1.2 vvm, the maximum values were 5.89g L-1, 289.98mg L-1, and 50.00mg L-1 for biomass, IPS, and IPS content production, respectively. In order to improve the A. camphorata production, the two-stage culture strategy was used. The productions were reached to 6.98g L-1, 255.19g L-1, 381.23g L-1, and 55.65mg g-1 for biomass, EPS, IPS, and IPS content, respectively.
The crude EPS from submerged culture of A. camphorata was purified by using Silica gel 60 column chromatography, where two EPS peaks were eluted. The protein to polysaccharide ratios from A. camphorata by regulating two-stage culture strategy (fraction-1 of EPS:20.78% ; IPS:38.69%) were superior those of batch cultures in 9g/L L-Proline, aeration rates of 0.2vvm and 1.2vvm. The molecular weight of EPS by regulating two-stage culture strategy (1.24×106 Da of EPS in fraction-1) was higher than those by two the other cultures (i.e., culture in 9g/L L-Proline, aeration rates of 0.2vvm and 1.2vvm), and the similar result was observed in the molecular weight of IPS (3.30×105Da) from regulating two-stage culture strategy. The carbohydrate compositions of polysaccharide produced from A. camphrata consisted of glucose, xylose, and galactose ; the percentages of monosaccharide compositions from polysaccharides in each fraction were almost constant. The results indicate that the monosaccharide compositions were not changed with culture conditions including of L-Proline effect, oxygen transfer, and two-stage culture strategy used in this study.
The results of biological activities (cytokine assay) of polysaccharide from A. camphorata were directly proportion to molecular weight of polysaccharide from A. camphorata. The higher molecular weight of polysaccharide was, the more murine tumor necrosis factor-alpha (mTNF-α) concentration was achieved. The maximal mTNF-α concentration values for EPS in fraction-1, EPS in fraction-2, and IPS were 1779.91 ± 46.52 pg/mL, 544.93 ± 44.39 pg/mL, 567.42 ± 50.83 pg/mL, respectively. Besides, the protein to polysaccharide ratio was closely correlated with the biological activity of polysaccharide from A. camphorata. The higher protein to polysaccharide ratio was, the more mTNF-α concentration was obtained.

中文摘要 ---------------------------------------------------------------------------------- I
英文摘要 -------------------------------------------------------------------------------- IV
誌謝 ------------------------------------------------------------------------------------- VII
目錄 ------------------------------------------------------------------------------------ VIII
圖次索引 ------------------------------------------------------------------------------ XIV
表次索引 ------------------------------------------------------------------------------ XXI
第一章 緒論 ----------------------------------------------------------------------------- 1
1.1 研究動機與背景 --------------------------------------------------------------- 1
1.2 研究目的 ------------------------------------------------------------------------ 4
第二章 文獻回顧 ----------------------------------------------------------------------- 5
2.1真菌菇類介紹 ------------------------------------------------------------------- 5
2.1.1 真菌菇類的抗腫瘤與免疫調節活性 -------------------------------- 6
2.1.2 真菌菇類的生理活性成分 -------------------------------------------- 6
2.2 真菌多醣體介紹 --------------------------------------------------------------- 8
2.2.1 真菌多醣體之特性 ----------------------------------------------------- 8
2.2.2 真菌多醣體之生合成 ------------------------------------------------- 10
2.2.3 真菌多醣體之機能性 ------------------------------------------------- 12
2.3 巨噬細胞與真菌多醣體 ---------------------------------------------------- 15
2.3.1 巨噬細胞與其功能 --------------------------------------------------- 15
2.3.2 巨噬細胞吞噬能力 --------------------------------------------------- 17
2.3.3 多醣體抗腫瘤生物活性之巨噬細胞實驗 ------------------------ 17
2.4 樟芝介紹 ---------------------------------------------------------------------- 18
2.4.1 樟芝多醣體 ------------------------------------------------------------- 20
2.4.2 樟芝化學成分組成 ---------------------------------------------------- 21
2.4.3 樟芝藥理特性 ---------------------------------------------------------- 21
2.5 深層醱酵培養 ---------------------------------------------------------------- 24
2.5.1 影響醱酵的物理因子 ------------------------------------------------- 24
2.5.2 影響醱酵的化學因子 ------------------------------------------------- 29
2.6 氨基酸簡介 ------------------------------------------------------------------- 32
2.6.1 精氨酸；筋氨酸(L-Arginine) ----------------------------------------- 32
2.6.2 麩氨醯酸(L-Glutamine) ----------------------------------------------- 32
2.6.3 異白氨酸(L-Isoleucine) ----------------------------------------------- 33
2.6.4 離氨酸；2,6-二氨己酸(L-Lysine) ------------------------------------ 33
2.6.5 脯氨酸；口比咯口定甲酸(L-Proline) ------------------------------- 34
2.6.6 酥氨酸；羥丁氨酸(L-Threonine) ------------------------------------- 35
　　　 2.7 多醣體之分離與純化 ------------------------------------------------------- 37
2.7.1 透析法 ------------------------------------------------------------------- 37
2.7.2 酒精沉澱法 ------------------------------------------------------------- 37
2.7.3 冷凍離心法 ------------------------------------------------------------- 37
2.7.4 層析法 ------------------------------------------------------------------- 37
2.7.4.1 離子交換管柱層析 ------------------------------------------- 38
2.7.4.2 疏水性作用管柱層析 ---------------------------------------- 40
2.7.4.3 親和性作用管柱層析 ---------------------------------------- 41
2.7.4.4 膠體過濾管柱層析 ------------------------------------------- 41
第三章 材料與方法-------------------------------------------------------------------- 43
3.1 實驗架構 ---------------------------------------------------------------------- 43
3.2 實驗材料 ---------------------------------------------------------------------- 44
3.2.1 樟芝實驗菌株 ---------------------------------------------------------- 44
3.2.2 實驗藥品 ---------------------------------------------------------------- 44
3.2.3 實驗儀器及設備 ------------------------------------------------------- 46
3.2.4 實驗裝置 ---------------------------------------------------------------- 48
3.3 實驗方法 ---------------------------------------------------------------------- 55
3.3.1 菌株保存 ---------------------------------------------------------------- 55
3.3.2 培養基成分 ------------------------------------------------------------- 55
3.3.3 操作條件 ---------------------------------------------------------------- 57
3.4 分析方法 ---------------------------------------------------------------------- 59
3.4.1 醱酵產物分析流程 ---------------------------------------------------- 59
3.4.2 菌體乾重測定(Mycelial growth weight，X) ------------------------ 60
3.4.3 粗多醣體濃度分析(Polysaccharide concentration，P) ------------ 60
3.4.4 葡萄糖濃度分析(Residual glucose concentration，S) ------------- 62
3.4.5 脯氨酸濃度分析 ------------------------------------------------------- 64
3.4.6 菌絲球粒徑測定 ------------------------------------------------------- 65
3.4.7 管柱層析與分離純化 ------------------------------------------------- 66
3.4.8 萃取醱酵產物胞內多醣體之製備 ---------------------------------- 67
3.4.8.1 醱酵產物 --------------------------------------------------------- 67
3.4.8.2 製備流程 --------------------------------------------------------- 67
3.4.9 蛋白質含量分析 ------------------------------------------------------- 68
3.4.10 多醣體分子量分析 -------------------------------------------------- 68
3.4.11 多醣體之中性單醣組成分析 -------------------------------------- 69
3.4.12 多醣體之生物活性測定 -------------------------------------------- 71
3.4.12.1 巨噬細胞株 ---------------------------------------------------- 71
3.4.12.2 巨噬細胞株培養液組成 ------------------------------------- 71
3.4.12.3 巨噬細胞株保存 ---------------------------------------------- 71
3.4.12.4 巨噬細胞株解凍 ---------------------------------------------- 72
3.4.12.5 巨噬細胞株繼代培養 ---------------------------------------- 72
3.4.12.6 巨噬細胞激素測定實驗流程 ------------------------------- 72
第四章 結果與討論 ------------------------------------------------------------------- 75
4.1 樟芝菌絲型態 ---------------------------------------------------------------- 75
4.1.1 固態培養 ---------------------------------------------------------------- 75
4.1.2 搖瓶培養 ---------------------------------------------------------------- 77
4.1.3 攪拌式醱酵槽培養 ---------------------------------------------------- 78
4.2 樟芝搖瓶培養實驗 ---------------------------------------------------------- 79
4.2.1 不同氨基酸培養基對於培養樟芝菌體產量之影響 ------------- 79
4.2.1.1 不同氨基酸對樟芝培養生物質量和胞外多醣體
產量的影響 ---------------------------------------------------- 79
4.2.1.2 不同氨基酸對樟芝轉化率胞外YP/X、YX/S 和YP/S
的影響 ---------------------------------------------------------- 82
4.2.2 不同初始濃度脯氨酸培養基對於培養樟芝菌體產量
之影響 ----------------------------------------------------------------- 86
4.2.2.1 不同初始濃度脯氨酸對樟芝培養生物質量和
胞外多醣體產量的影響 ------------------------------------- 86
4.2.2.3 不同初始濃度脯氨酸對樟芝轉化率胞外YP/X、YX/S
和YP/S 的影響 ------------------------------------------------- 88
4.3 樟芝醱酵槽培養實驗 ------------------------------------------------------- 91
4.3.1 不同初始濃度脯氨酸於攪拌式醱酵槽培養之影響 ------------- 91
4.3.1.1 不同初始濃度脯氨酸對樟芝培養生物質量產量
的影響 ----------------------------------------------------------- 95
4.3.1.2 不同初始濃度脯氨酸對μ、QX和YX/S的影響 --------------- 96

4.3.1.3 不同初始濃度脯氨酸對樟芝培養胞外多醣體產量
的影響 ---------------------------------------------------------- 97
4.3.1.4 不同初始濃度脯氨酸對胞外QP、YP/X和YP/S
的影響 ---------------------------------------------------------- 98
4.3.1.5 不同初始濃度脯氨酸對樟芝培養胞內多醣體產量
和胞內多醣體內含量的影響 ------------------------------- 99
4.3.1.6 不同初始濃度脯氨酸對胞內QP、YP/X和YP/S
的影響 -------------------------------------------------------- 104
4.3.1.7 不同初始濃度脯氨酸對樟芝菌絲球粒徑的影響 ------- 105
4.3.2 不同通氣量於攪拌式醱酵槽培養之影響 ----------------------- 106
4.3.2.1 不同通氣量對樟芝培養生物質量產量的影響 ---------- 111
4.3.2.2 不同通氣量對μ、QX和YX/S的影響 -------------------------- 111
4.3.2.3 不同通氣量對樟芝培養胞外多醣體產量的影響 ------- 113
4.3.2.4 不同通氣量對胞外QP、YP/X和YP/S 的影響 ---------------- 114
4.3.2.5 不同通氣量對樟芝培養胞內多醣體產量和胞內
多醣體內含量的影響 -------------------------------------- 115
4.3.2.6 不同通氣量對胞內QP、YP/X和YP/S 的影響 ---------------- 121
4.3.2.7 不同通氣量對樟芝菌絲球粒徑的影響 ------------------- 122
4.3.3 ¬二階段不同通氣量和脯氨酸操作於攪拌式醱酵槽
培養之影響 -------------------------------------------------------- 123
4.3.3.1 二階段不同通氣量和脯氨酸操作對樟芝培養
胞外多醣體產量、QP、YP/X和YP/S的影響 ---------------- 125
4.3.3.2 二階段不同通氣量和脯氨酸操作對樟芝培養胞內
多醣體產量、多醣體內含量、QP、YP/X和YP/S
的影響 -------------------------------------------------------- 126
4.3.3.3 二階段不同通氣量和脯氨酸操作對樟芝培養
生物質量、μ、QX和YX/S的影響 ---------------------------- 127
4.4 樟芝粗多醣體分離純化結果 --------------------------------------------- 129
4.4.1 不同初始濃度脯氨酸對樟芝胞外粗多醣體純化
之分布 ---------------------------------------------------------------- 129
4.4.2 不同通氣量對樟芝胞外粗多醣體純化之分布 ----------------- 133
4.4.3 二階段不同通氣量和脯氨酸操作對樟芝胞外
粗多醣體純化之分布 ---------------------------------------------- 136
4.5 樟芝多醣體與蛋白質含量影響 ------------------------------------------ 138
4.5.1 樟芝胞外多醣體與蛋白質含量之分析 -------------------------- 138
4.5.2 樟芝胞內多醣體與蛋白質含量之分析 -------------------------- 141
4.6 樟芝多醣體分子量分布影響 --------------------------------------------- 144
4.6.1 樟芝胞外多醣體分子量分布之分析 ----------------------------- 145
4.6.2 樟芝胞內多醣體分子量分布之分析 ----------------------------- 149
4.7 樟芝多醣體之中性單醣成分組成影響 --------------------------------- 152
4.7.1 樟芝胞外多醣體之中性單醣成分之組成分析 ----------------- 153
4.7.2 樟芝胞內多醣體之中性單醣成分之組成分析 ----------------- 156
4.8 樟芝多醣體之生物活性測定影響 --------------------------------------- 158
4.8.1 巨噬細胞之型態 ----------------------------------------------------- 158
4.8.2 樟芝胞外多醣體之生物活性測定之分析 ---------------------- 160
4.8.3 樟芝胞內多醣體之生物活性測定之分析 ---------------------- 165
第五章 結論與建議 ----------------------------------------------------------------- 170
5.1 結論 --------------------------------------------------------------------------- 170
5.2 建議 --------------------------------------------------------------------------- 172
參考文獻 ------------------------------------------------------------------------------- 173
作者簡介 ------------------------------------------------------------------------------- 188

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