本研究自南台灣林地中篩選出能夠分泌纖維水解酵素(cellulases)的菌種，並透過微酵素工程與醱酵工程等角度，來了解其特性、生產方式與應用方式，最後利用系統整合程序將此酵素應用至纖維素生質能源的製備，進而朝向商業發展的可行性與綠能產業的永續發展的目標邁進。本研究首先建立適合篩選纖維水解菌株的方法，並利用所設計的菌株篩選策略，成功地至目標樣本中篩選出12株較具潛力的纖維水解生產菌株，其中Pseudomonas sp. CL3具有相當優異的纖維水解能力，且所生產出的水解酵素種類豐富且活性高，因此本研究選擇該菌種作為主要研究對象。
本研究透過酵素反應活性測試中發現，Pseudomonas sp. CL3 不只能生產纖維水解酶(endo-β-1,4-D-glucanase, exo-β-1,4-D-glucanase, β-1,4-D-glucosidase 和xylanase)，也能生產澱粉水解酶(amylase)和果膠水解酶(pectinase)等酵素。由溫度與pH反應性測試中得知，endo-β-1,4-D-glucanase, exo-β-1,4-D-glucanase, β-1,4-D-glucosidase和xylanase 最適反應溫度分別為50, 45, 45 與 55oC，而最適反應的pH值皆為6。由蛋白質活性電泳膠分析技術，確定所使用的酵素分子量大小分別為endo-β-1,4-D-glucanase, 80 KDa and 100 KDa；exo-β-1,4-D-glucanase, 55 KDa； β-1,4-D-glucosidase, 65 KDa 和xylanase, 20 KDa 。本研究也針對酵素的熱穩定度與其他容易影響酵素活性之因子(如金屬離子與界面活性劑)進行相關測試研究。由研究結果可知，所篩選出的纖維水解菌株，其生產的酵素種類為中溫型酵素；適量的鐵與錳離子對其酵素活性具有顯著促進之作用，而添加生物界面活性劑(表面素)也可以提升酵素活性。然而，由於酵素會吸附於醣脂類界面活性劑(如鼠李醣脂)之醣基，而導致水解效能降低。本研究亦建立固定化纖維素水解酶，將該酵素固定在磁性載體上，經測試纖維二醣酶(β-1,4-D-glucosidase)在此固定化系統下，可重複使用5次，其活性仍保有90%以上。
此外，為了嘗試提升Pseudomonas sp. CL3之纖維水解酶產量，以利後續之應用，本研究使用實驗設計法(回應曲面法; response surface methodology)來提高CL3菌株的纖維水解酶產量。先根據二水準變因篩選法，選取與纖維水解酶產量相關性較高的影響因子，再以反應曲面法求得各個重要因子的最佳參數。本研究以24-1二水準因子法篩選可能影響纖維水解酶產量的因子，所檢測的變因包含rice straw, CMC, NH4NO3 and yeast extract等培養基成分的濃度。經過結果分析，rice straw與yeast extract濃度對纖維水解酶產量的影響較大。因此，透過回應曲面法，本研究求得rice straw與yeast extract兩項因子的最佳參數值，分別為1.32% and 0.32%。而利用此最佳生產配方條件，纖維水解酶的產量可以提升至原來的3.7倍。為了進一步提升纖維水解酶的生產效率，本研究以不同醱酵策略來提升纖維水解酶的生產效率。由實驗結果發現，在CSTR系統中，將水力停留時間控制在17小時時，纖維水解酶的濃度可高達3.3 FPU/ml並且達到穩定生產，此時纖維水解酶的總生產速率可提升至194.12 FPU/L/h，此生產速率為原來批次生產的3.4倍。
本研究接著將所生產之纖維水解酶進行對農業纖維素廢棄物之水解測試，結果發現，透過鹼處理程序所得之蔗渣纖維其水解效率為未處理纖維之3.2倍。再者，由纖維水解動力學研究發現，所生產之Pseudomonas sp. CL3纖維水解酶，其纖維水解動力學非常符合Michaelis-Menten模式，並藉由此模式求得其動力學參數為Vmax= 0.133 g/L/h、Km=7.4 g/L，此結果顯示在蔗渣纖維濃度為14.8 g/L時，其最大葡萄糖產生速率為0.133 g/L/h。為了更進一步了解Pseudomonas sp. CL3纖維水解酶與蔗渣纖維處理量對葡萄糖產率的影響，本研究藉由不同濃度的纖維水解酶與蔗渣負載量進行實驗設計。結果發現，在纖維水解酶濃度為9 FPU/ml時，並且所添加之蔗渣纖維濃度介於50至70 g/L時，此時最大葡萄糖產率可高達0.55 g/L/h。而Pseudomonas sp. CL3纖維水解酶不僅對農業廢棄纖維(二代生質料源)能夠有效水解糖化，還能夠應用至微藻料源(三代生質料源)的水解糖化，經研究證實，其葡萄糖產率可高達0.13 g/L/h，且轉化率可高達90% 以上。
本研究亦利用所開發出的纖維水解酵素搭配進行生質氫能之生產，主要透過批次實驗的同步糖化醱酵(SSF)與兩階段分步糖化醱酵(SHF)進行測試。由研究結果可知，在低糖量基質之產氫醱酵系統中，其氫氣生產速率與氫氣濃度都會偏低，故未來纖維氫氣生產系統應以兩階段分步糖化醱酵(SHF)較為可行。本研究接著利用稻稈水解液進行連續醱酵產氫，在水利停留時間為12小時的條件下，其氫氣生產速率326.7 ml/L/h，氫氣濃度為56.5%，由此可知，本研究所開發出的纖維水解酵素能夠有效結合暗醱酵產氫系統，進行纖維氫氣的生產。
最後，本研究進一步利用所開發最佳的酵素生產與水解糖化程序，結合暗醱酵產氫與微藻固碳系統，進一步針對纖維氫能最佳生產程序及整合低碳排放系統之評估，以達成高效綠色能源生產與永續經營之目標。

In this study, an effective cellulase-producing strain was isolated from forest soil in southern Taiwan using the approaches associated with microbiology, enzyme engineering and fermentation engineering. The enzymes produced were used for the development of sustainable and commercially feasible bioenergy producing processes. Twelve strains were first isolated from the environmental samples. Among them, Pseudomonas sp. CL3 was found to secrete a number of highly active hydrolytic enzymes the best activity, so was selected as the target strain for further studies.
The isolated indigenous bacterium Pseudomonas sp. CL3 was able to produce novel cellulases consisting of endo-β-1,4-D-glucanase (80 and 100 KDa), exo-β-1,4-D-glucanase (55 KDa), β-1,4-D-glucosidase (65 KDa) and xylanase (hemicellulase) with a molecular weight of 20 KDa characterized by enzyme assay and zymography analysis. In addition, the CL3 strain also produced amylase and pectinase. The optimal temperature for enzyme activity was 50, 45, 45 and 55oC for endo-β-1,4-D-glucanase, exo-β-1,4-D-glucanase, β-1,4-D-glucosidase and xylanase, respectively. All the enzymes displayed optimal activity at pH 6.0.
In addition, the effect of reaction temperature and environmental factors (metal ions and surfactant) on the activity of the cellulolytic enzymes originating from Pseudomonas sp. CL3 was intensively studied. It was found that the cellulases produced by Pseudomonas sp. CL3 belong to mesophilc enzymes. The metal ions (Mn2+ and Fe2+) were also significant factors affecting the cellulase activity. Addition of biosurfactant (e.g., surfactin) can enhance the cellulase activity. However, for glycolipid-type biosurfactant (e.g., rhamnolipid), since the cellulases will bind to the sugar structure of the biosurfactant, the addition of rhamnolipid led to a decrease in the hydrolytic activity. The cellulases produced by strain CL3 were also immobilized on the magnetic particles. These results indicate that the β-glucosidase activity of the immobilized enzyme had extremely high stability, and could maintain 90% of its original activity even when the immobilized enzyme particles were reused for 5 cycles.
To further enhance the production of cellulases from the CL3 strain, statistical experimental design methodology was applied to optimize the culture medium composition favoring enzyme synthesis. Four key parameters (CMC, rice straw, NH4NO3, yeast extract) were selected by two-level factorial design. Response surface methodology was then used to identify the optimal composition of the selected parameters, giving an optimal concentration of 1.32% and 0.32% for rice straw and yeast extract, respectively. With this optimal medium, the cellulolytic enzymes production could be markedly elevated to a maximum concentration of 3.8 FPU/ml, which is 3.7 times compared with original medium.
In order to develop efficient and cost-effective process for producing cellulases from CL3 strain, different fermentation strategies were examined. In the CSTR fermentation process, the cellulase titer in the fermentor could be maintained at around 3.3 FPU/ml at a HRT of 17 h with a cellulase productivity of 194.12 FPU/L/h, which is 3.4 times higher than that obtained from the batch system.
Nest, the cellulases produced from CL3 strain was used to perform hydrolysis process for the saccharification of lignocellulosic materials (such as sugarcane bagasse and rice straw). The effects of pretreatment process (chemical and physical method) on the enzymatic hydrolysis were investigated. The alkaline pretreatment seems to be the most effective method able to achieve the highest monosaccharide yield from bagasse through enzymatic saccharification, which is 3.2 times compared with non- pretreated bagasse. Kinetic analysis shows that the dependence of cellulase activity on cellulose substrate can be described by Michaelis-Menten model with good agreement. The estimated kinetic constants for cellulases obtained from CL3 strain were Vmax= 0.133 g/L/h and Km=7.4 g/L, respectively (with an enzyme loading of 4.5 FPU/ml). That is, the maximum glucose productivity from the enzymatic hydrolysis was 0.133 g/L/h at a bagasse loading of 14.8 g/L. The cellulose hydrolysis experiments were also conducted under different concentrations of alkaline-pretreated bagasse. Under different bagasse loadings (50 to 70 g/L), a cellulase dosage of higher than 9 FPU/ml can obtain a great glucose productivity of around 0.55 g/L/h. Moreover, our study also shows that the enzyme from Pseudomonas sp. CL3 can also hydrolyze microalgal biomass (e.g., Chlorella vulgaris), giving a glucose productivity of 0.13 g/L/h, and the yield of around 90%.
The cellulases were also applied in producing biohydrogen from cellulosic feedstock. In the cellulosic hydrogen production process, the experiments were divided into two groups, namely separate hydrolysis and fermentation (SHF) process and simultaneous saccharification and fermentation (SSF) process using alkaline pretreated sugarcane bagasse. It was found that lower H2 content and productivity were obtained when the glucose concentration in the fermentation medium was at a lower level (under 4 g/L). Therefore, the SHF process seems to be more suitable for the biohydrogen production from the pretreated bagasse due to the requirement of high glucose concentration in the medium. Therefore, we used the bagasse hydrolysate to produce hydrogen with Clostridium pasteurianum CH4 using CSTR operations at a HRT of 12 h. Under this condition, the H2 content in the biogas was around 56.5% and the maximum H2 production rate was 326.7 ml/L/h. These results show that the cellulases produced form a newly isolated indigenous bacterium Pseudomonas sp. CL3 were able to combine with dark H2 fermentation system to produce cellulosic hydrogen. Finally, we combine the cellulases production strategy, optimal hydrolysis process, dark H2 fermentation, and microalgae-CO2 capture system to develop a sustainable and low-carbon-emission biohydrogen producing system.

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