面積約100公頃的鯉魚潭為花蓮縣最大湖泊，經複合參數指標判定水質長期屬於輕度優養狀態。鯉魚潭於1993、1996及1999年秋冬季皆曾發生嚴重的葡萄藻華，期間並伴隨吳郭魚大量死亡的現象。為評估葡萄藻華對該水域生態系統的影響，故對葡萄藻的生物毒性進行研究。以葡萄藻粗萃取物對分離自鯉魚潭的十五株藻種進行毒性測試，結果顯示除了Microcystis flos-aquae及Oocystis parva，其餘十三株藻種皆表現程度不一的敏感度。對鯉魚潭浮游藻群落的分析結果顯示，藻種對葡萄藻的敏感度與其在葡萄藻華期間所佔有的相對優勢度，呈現密切相關。本研究亦使用葡萄藻粗萃取物對鯉魚潭的優勢浮游動物進行毒性測試，結果顯示佔有浮游動物群落60 %以上組成的橈足類較枝角類敏感得多。前述測試結果顯示，葡萄藻可能藉由相剋物質的作用，對鯉魚潭的浮游生物群落造成影響。
批次培養結果顯示，在磷或氮源成為限制因子而細胞進入指數生長末期時，葡萄藻能產生並釋出大量的毒性物質。此時期的培養液之營養鹽濃度與葡萄藻華時期的鯉魚潭水質條件最為接近，顯示該水域提供適當的化學環境，使葡萄藻能生長良好並藉由相剋作用與其它藻類競爭。
葡萄藻毒性物質溶於有機溶劑中，經薄層層析、高效液相層析等技術進一步分離及純化，並以核磁共振及質譜等技術鑑定結構後，確認為包括棕櫚酸 (16:0)、油酸 (18:1ω9)、亞麻油酸 (18:2ω6)及α-亞麻脂酸 (18:3ω3)在內的數種自由態脂肪酸。其中，三種18碳不飽和脂肪酸對多數測試藻株及甲殼綱浮游動物具有明顯毒性，半抑制濃度EC50 (或LC50)約在10-20μg‧mL-1之間。葡萄藻的總自由態脂肪酸含量佔細胞乾重的2.25%左右，遠高於其它七個分屬藍綠菌、綠藻及渦鞭毛藻門的測試藻種。
葡萄藻粗萃取物及其主要脂肪酸經由阻斷電子傳遞鏈，而能抑制藻細胞的光合作用，對呼吸作用則皆無明顯影響。三種18碳不飽和脂肪酸在低於明顯抑制光合作用的施加劑量下，即可造成藻細胞的死亡；而DCMU在完全抑制光合作用的高劑量下，卻不會在24小時內造成藻細胞死亡。顯示對光合作用的抑制並非葡萄藻對其它藻類的關鍵致毒機制。
三種18碳不飽和脂肪酸在施加三十分鐘內會造成藻細胞5-10%鉀離子滲漏，並在測試的四小時內持續增加。進一步的研究顯示，此些18碳不飽和脂肪酸會對藻細胞膜上的P型ATPase造成抑制，對V型及F型ATPase則無明顯影響。由於P型ATPase (尤其是H+-ATPase)為藻細胞維持膜電位的最重要酵素，不飽和脂肪酸造成藻細胞之鉀離子滲漏可能與P型ATPase受抑制有關。另一方面，葡萄藻粗萃取物及其主要脂肪酸亦會對枝角類及橈足類的Na+/K+-ATPase產生專一性抑制；而對F型及V型ATPase，甚至其它P型ATPase皆無任何抑制效果。顯示葡萄藻毒性物質可能經由抑制Na+/K+-ATPase活性，導致浮游動物因滲透壓失衡而死亡。

With a surface area of ca. 100 ha, the Lake Li-Yu is the largest lake in Hwa-Lian county, Taiwan. It is an eutrophic lake with Botryococcus braunii blooms occurred in autumn and winter in the years of 1993, 1996 and 1999. Death of fish, mainly Tilapia, were observed in the lake during B. braunii blooming. In order to elucidate the effects of algal blooms in this aquatic ecosystem, the toxicity of B. braunii was studied. Fifteen strains of phytoplankton were isolated from the lake and the cultures were tested for the toxicity of B. braunii extract on them. Results showed that except Microcystis flos-aquae and Oocystis parva, all other species exhibited various degrees of sensitivity to B. braunii. Population analysis of the tested species in the lake showed that the sensitivity to B. braunii was closely correlated with the abundance of these phytoplankton species in the lake during Botryococcus blooming. Toxic effects of B. braunii extract were also observed to the zooplankton collected from the Lake. In them, copepods showed higher sensitivity to crude extracts from B. braunii than cladocerans. All of these results suggest that B. braunii might play an allelopathic effect on other plankton.
In batch culture, the toxicities of B. braunii were enhanced when P- or N-nutrient became a limitation. The maximum toxicity was measured when the growth of cells was near the end of exponential growth phase. Under this condition, the nutrient concentrations in culture medium were close to those measured during B. braunii bloom, suggesting that the chemical environment in the Lake Li-Yu might create a favorable condition for B. braunii to survive and to compete with other plankton species.
The fractions containing active substances of B. braunii were soluble in organic solvents. Further isolation, purification, and identification of active substances, by employing TLC, HPLC, NMR, and GC-MS techniques, indicated that they were fatty acids, including palmitic acid (16:0), oleic acid (18:1ω9), linoleic acid (18:2ω6) and α-linolenic acid (18:3ω3). The highest toxicity was revealed in C18-unsaturated fatty acids whose EC50 (or LC50) were found between 10-20μg‧mL-1 against phytoplankton as well as against zooplankton. Total free fatty acid content might be as high as 2.25% of dry weight, which was much higher than other 7 tested species of cyanophytes, chlorophytes and dinophytes.
Addition of B. braunii extract inhibited the photosynthetic electron flow of tested algal cells, while the respiration rate was affected little. Similar effects were also observed when C18-unsaturated fatty acids were added to the algal cultures. It is noteworthy that the tested cells would die even when the added concentrations were lower than IC50 value of inhibition of photosynthesis. However, the photosynthetic electron flow inhibitor, DCMU, did not cause the death of tested algal cells within 24 hrs. This implicates that the inhibition of electron flow was not the only cause of death B. braunii exerted to other algal species.
It was observed that addition of C18-unsaturated fatty acids to algal cultures caused a remarkable K+-leakage from tested cells within 30 minutes. The amount of released K+ might as high as 5-10% of total K+ in the cells. Enzymatic assays of ATPase showed that C18-unsaturated fatty acids did inhibit P-type, but not V- or F-type ATPase of the tested algal cells. Because P-type ATPase (especially H+-ATPase) is related to the maintenance of membrane potential, inhibition of this type of membrane enzyme might be attributed to K+-leakage from the cells. C18-unsaturated fatty acids did not exert inhibitory effect on V-, F-, or other P-type ATPase, but on Na+/K+-ATPase of copepods and cladocerans. It is assumed that toxic effects of B. braunii on crustacean zooplankton might be related to the interference of osmoregulation through inhibiting the Na+/K+-ATPase activity.

中 文 摘 要………………………………………………………..…….1
英 文 摘 要…………………………………….....……………………3
縮 寫 表……………………………………………………………………5
一、 前 言……………………………………………………………....6
二、材料與方法………………………………………………………….13
重要藥品與儀器列表……………………………………….......13
2.1 葡萄藻毒性物質的純化……………….………………………....15
2.2 葡萄藻毒性物質之化學結構鑑定……….............…………17
2.3 藻細胞自由態脂肪酸的定性與定量………………………….....18
2.4 藻株對葡萄藻毒性物質的敏感度測試……………….........…19
2.5 浮游動物對葡萄藻毒性物質的敏感度測試………………...22
2.6 葡萄藻於不同氮、磷濃度下的批次培養實驗……...........…24
2.7 葡萄藻抑制物質對藻類光合系統的抑制測試………….……....25
2.8 自由態脂肪酸對藻細胞之鉀離子滲漏的影響………….........28
2.9 自由態脂肪酸對藻細胞之P型ATPase活性的抑制測試.……...…29
2.10自由態脂肪酸對浮游動物之Na+/K+-ATPase活性的抑制測試……31
2.11 鯉魚潭水質及浮游生物資料的月調查………………………....32
三、結 果…………………………………………………………………34
3.1 葡萄藻毒性物質之定性與定量…………………………….......34
3.2 葡萄藻於不同營養鹽濃度下的毒性物質產生量之變化……...…37
3.3 毒性物質對於葡萄藻在鯉魚潭形成優勢所扮演的角色………….41
3.4 葡萄藻毒性物質對其它藻類的致毒機制………………….……..44
3.5 葡萄藻毒性物質對浮游動物的致毒機制……………….……....48
四、討 論…………………………………………………………………50
五、結 論………………………………………………………......…60
六、參 考 文 獻……………………………………..…………………62
表……………………………………………………...…………………71
圖…………………………………………………...………………....81

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