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研究生：	劉建宏 Liu, Chien-Hung
論文名稱：	脂肪分解酵素菌株之篩選及其應用 Isolation and application of lipase producing bacteria
指導教授：	張嘉修 Chang, Jo-Shu
學位類別：	博士 Doctor
系所名稱：	工學院 - 化學工程學系 Department of Chemical Engineering
論文出版年：	2008
畢業學年度：	96
語文別：	英文
論文頁數：	209
中文關鍵詞：	酵素動力學、醱酵策略、Burkholderia sp. 、酵素固定化、生質柴油、反應曲面法、菌種篩選、脂解酵素、廚餘、矽藻土
外文關鍵詞：	Bacterial strain isolation, biodiesel, Burkholderia sp., celite, enzyme immobilization, enzyme kinetics, fermentation strategy, food wastes, lipase, response surface methodology (RSM)
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本研究自廚餘中篩選能分泌脂解酵素(lipase)的菌種，並嘗試以醱酵方法提升該菌種脂解酵素之產量，並將該酵素進行固定化以利酵素回收利用，最後再將該固定化脂解酵素應用於生質柴油製備。本研究首先參考文獻中較可行的脂解酵素菌種篩選方法，並搭配自行開發的篩選策略來進行菌種篩選，由工研院提供之廚餘樣本中篩選出33株具有脂解酵素的菌種，其中有16株的脂解酵素在pH 6.0之活性優於在pH 9.0時。而在這33株菌中，以利用策略C (screening strategy C)所篩選到的菌種Burkholderia sp. C20具有最大的轉酯化活性。從文獻得知，源於Burkholderia的脂解酵素具有合成生質柴油的能力，因此本研究選擇該菌種為主要研究對象，並嘗試提升其脂解酵素產量，以利於後續之應用。
接著，本研究嘗試以實驗設計方法提高Burkholderia sp. C20菌株的脂解酵素產量，所採用的實驗設計方法為反應曲面法(response surface methodology)。在實驗設計上，先利用二水準變因篩選方法，選取與脂解酵素產量較相關的影響因子，再以反應曲面法求得各重要因子的最佳參數值。本研究以25-1二水準因子法篩選可能影響脂解酵素產量的因子，所檢測之變因包括溫度、pH、olive oil濃度、CASO濃度及NaCl濃度。經過結果分析，CASO濃度與NaCl濃度對脂解酵素產量的影響較大。因此，本研究以反應曲面法求得此兩項因子的最佳參數值，分別為0.12%及0.16%。利用此最佳條件生產脂解酵素，其產量可以提升約5倍(即由0.8 U/ml提升到3.9 U/ml)。為進一步提昇脂解酵素之產量以利於進行生質柴油之製備，本研究進一步以醱酵槽來進行脂解酵素之放大生產，並藉由影響因子(如通氣速率、轉速、pH control與培養時間等)之調整與olive oil添加方式，尋找較佳之醱酵策略，以提升脂解酵素的生產效率。實驗結果發現，當通氣速率控制在0.5到2 vvm之間，脂解酵素的總生產速率可從0.057提升到0.076 U/ml-h。此外，在配合最佳的攪拌速率(100 rpm)操作，可再提升脂解酵素的總生產速率至0.09 U/ml-h。以上實驗結果顯示，對Burkholderia sp. C20而言，脂解酵素屬於混合生長相關型產物，其產率係數(yield coefficient)為524 U/g dry cell weight。本研究亦發現，將pH控制在7.0時有助於細胞生長，而將pH控制在6.0時有助於lipase生產。而以逐步添加olive oil的方式，則可提升該菌株在搖瓶中的脂解酵素產量，且在醱酵槽實驗中得到驗證。
本研究接著將所生產之Burkholderia lipase進行固定化，以利脂解酵素之回收與重複使用。藉由薄膜過濾方式與矽藻土(celite)共價鍵結的方式，將Burkholderia sp. C20生產的脂解酵素進行固定化。結果發現，在70 g/L粗酵素濃度、pH 7.0與反應時間3小時為最佳固定化條件。在此條件之下，矽藻土上有最大的酵素負載量，約每克載體上有62.9毫克粗酵素，此時的固定化酵素活性為每克載體上有273.5 U的活性。由脂肪水解最佳化實驗得知，固定化酵素的最佳水解反應條件分別為CN-lipase：55℃與pH 9.0以及celite-lipase：55℃與pH 10.0。再者，在油脂水解動力學研究中發現，不論是游離態或固定化的脂解酵素都符合Michaelis-Menten模式，並藉由此模式求得其動力學參數。其中，所求得之游離態脂解酵素之動力學參數為vmax = 133.33 U/mg與Km = 0.09 mM；CN-lipase的動力學參數為vmax = 31.25 U/mg與Km = 9.44 mM；celite-lipase的動力學參數為vmax = 11.29 U/mg與Km = 12.06 mM。此結果顯示，該脂解酵素經由固定化程序之後，會降低其最大反應速率(vmax變小)與其對基質的親和性(Km變大)。最後，本研究進一步利用Burkholderia sp. C20的固定化脂解酵素來製備生質柴油。結果發現以celite-lipase製備生質柴油時，其最佳的celite-lipase添加量為0.6 g，甲醇/油的莫爾比為3。在此條件下，生質柴油的轉化率可達43%。

Approaches for rapid screening of lipase-producing bacteria were first developed and the feasibility assessment of the screening methods was performed. From food waste samples, the proposed screening procedures allowed isolation of thirty-three pure bacteria possessing lipase activity. Among them, sixteen strains expressing higher lipase activity at acidic pH (pH 6.0) than at alkaline pH (pH 9.0). Among the 33 strains, the isolate having best transesterification activity was obtained by strategy C (SSC) and was identified as Burkholderia sp. C20. Literature shows that the Burkholderia lipase could be used to produce biodiesel. Thus, in this study, Burkholderia sp. C20 was used as a target lipase producing strain for biodiesel production.
Next, an experimental design tool (i.e., response surface methodology; RSM) was used to improve lipase production from Burkholderia sp. C20. The factors affecting the lipase production of Burkholderia sp. C20 were screened by two-level factorial design and the optimal conditions for lipase production were determined by response surface methodology (RSM). Preliminary batch tests were employed to obtain the favorable conditions for lipase activity analysis and found that the optimal temperature and pH for lipase activity assay was 55oC and 9.0, respectively. Comparison of cell growth and lipase activity profiles shows that the fermentation time of 67 h was suitable for harvesting lipase products in the batch culture of Burkholderia sp. C20. A two-level design of 25-1 experiments was applied to identify the most significant influential factors out of five factors; namely, temperature, pH, and concentrations of olive oil, CASO and NaCl. After analysis with two-level design, concentrations of CASO and NaCl were selected for RSM analysis, predicting an optimal composition of 0.12% and 0.16% for CASO and NaCl, respectively. Using the optimal conditions, lipase production by Burkholderia sp. C20 was enhanced nearly 5 fold (from 0.8 to 3.9 U/ml). However, it is insufficient to provide enough amount of lipase from flask cultivation for biodiesel production. Therefore, a 5-L fermentor was used for scale-up production of lipase using the Burkholderia sp. C20 strain. The target factors affecting fermentative lipase production were aeration rate, agitation rate, and incubation time. By adjusting the aeration rate from 0.5 to 2 vvm, the overall lipase productivity increased from 0.057 to 0.076 U/ml-h. In addition, lipase production by fermentation could be further improved by optimizing agitation speed at 100 rpm, giving an overall lipase productivity of 0.09 U/ml-h. The formation of lipase from Burkholderia sp. C20 was found to be a mixed growth-associated event with a yield coefficient of 524 U/g-dry-cell-weight. It was also found that pH controlled at 7.0 was preferable to the cell growth while pH controlled at 6.0 was adaptable to lipase production. A stepwise addition of olive oil was found to be able to enhance lipase production in flask experiments and the results were also confirmed in fermentor tests.
To enhance the recovery, efficiency and reusability of the lipase for further applications in biodiesel synthesis, the lipase originating from Burkholderia sp. C20 was immobilized onto cellulose nitrate (CN) membrane and celite via filtration and covalent bonding, respectively. The optimal condition for lipase immobilized onto celite matrix was as follows: crude lipase concentration, 70 g/L; pH 7.0; incubation time, 3 h. Under the optimal conditions, the maximum immobilized lipase activity was about 273.5 U/g celite while the lipase loading amount was around 62.9 mg per gram celite. The optimal lipolysis condition of CN-lipase and celite-lipase was pH 9.0, 55℃ and pH 10.0, 55℃, respectively. Kinetic analysis shows that the dependence of lipolytic activity of free and immobilized lipase on oil substrate can be described by Michaelis–Menten model with good agreement. The estimated kinetic constants for free lipase were vmax = 133.33 U/mg protein and Km = 0.09 mM; for CN-lipase were vmax = 31.25 U/mg protein and Km = 9.44 mM; for celite-lipase were vmax = 11.29 U/mg protein and Km = 12.06 mM, respectively. As a result, the employment of lipase immobilization would lead to a decrease in vmax and an increase in Km. Finally, biodiesel was produced by the immobilized lipase from Burkholderia sp. C20. Celite-lipase was used to produce biodiesel. The optimal condition of biodiesel production by celite-lipase was 0.6 g celite-lipase and a methanol-to-olive oil molar ratio of 3. Under that optimal condition, the maximum conversion of olive oil to biodiesel was 43%.

Abstract (Chinese)……………………………………………………………………I
Abstract (English)…………………………………………………………………III
Acknowledgement………………………………………………………………………VI
Contents………………………………………………………………………………VIII
List of tables………………………………………………………………………XV
List of figures……………………………………………………………………XVII
Chapter 1 Introduction………………………………………………………………1
1 Motivation and purpose…………………………………………………………1
2 Content of this dissertation…………………………………………………5
Chapter 2 Literature review…………………………………………………………8
1 Alternative energy………………………………………………………………8
1.1 Bioethanol and biomethanol………………………………………………9
1.2 Hydrogen………………………………………………………………………10
1.3 Biodiesel……………………………………………………………………11
1.4 Other alternative fuels…………………………………………………14
2 Biodiesel production…………………………………………………………15
2.1 Alkaline catalysis…………………………………………………………16
2.2 Acidic catalysis……………………………………………………………19
2.3 Enzymatic catalysis………………………………………………………20
2.4 Supercritical and other methods………………………………………22
3 Lipase……………………………...……………………………………………23
3.1 Introduction to lipase……………………………………………………23
3.2 Characterization of lipase structure…………………………………27
3.3 Application of lipase………………………………………………………28
4 Isolation of microorganisms producing lipase…………………………32
4.1 Screening of strains possessing lipase activity: qualitative method……………32
4.2 Measurement of lipase activity: quantitative method…………………………34
5 Immobilization of enzymes………………………………………………………40
5.1 Classification of immobilization methods……………………………………40
5.2 Immobilization of lipase………………………………………………………44
6 Response surface methodology (RSM)……………………………………………46
6.1 Introduction to RSM…………………………………………………………51
6.2 RSM for lipase production……………………………………………………54
7 Fermentation technology…………………………………………………………54
7.1 Introduction to fermentation technology………………………………………56
7.2 Fermentation technology for lipase production………………………………58
Chapter 3 Materials and methods………………………………………………………62
1 Chemicals and materials……………………………………………………………62
2 Equipment…………………………………………………………………………64
3 Methods for analysis and measurement…………………………………………66
3.1 Preparation of emulsified substrate for pH-stat assay………………………66
3.2 General pH-stat assay for measuring lipase activity…………………………66
3.3 The definition of lipase activity………………………………………………66
3.4 Measurement of cell concentration……………………………………………68
3.5 Emulsification assay…………………………………………………………68
3.6 Total protein concentration analysis…………………………………………68
3.7 Scanning electron microscopy (SEM) analysis………………………………70
3.8 Measurement of biodiesel……………………………………………………70
3.9 Calculation of biodiesel conversion……………………………………………72
3.10 Preparation of crude lipase powder from bacteria……………………………72
3.11 Measurement of transesterification activity…………………………………72
3.12 Definition of transesterification activity……………………………………73
3.13 Stock of bacteria……………………………………………………………73
3.14 Measurement of probe response time (p)……………………………………73
3.15 Estimation of kLa ..…………………………………………………………74
4 Isolation of bacteria producing lipase……………………………………………75
4.1 Procedures for isolating lipase-producing bacterial strains……………………75
4.2 Definition of the percentage of bacteria of real lipase or acidic lipase…79
4.3 Cultivation of pure bacterial isolates for lipase assay…………………………79
4.4 Analysis of lipase activity at acidic pH with microwave deactivation treatment…………………………………………………………………79
4.5 Effect of microwave treatment on the activity of CRL (Candida rugosa lipase)………………………………………………………………………80
4.6 Effect of microwave treatment on olive oil and oleic acid……………………80
5 Optimization of lipase production from Burkholderia sp. C20 in flasks…………81
5.1 The desired strain for lipase production………………………………………81
5.2 Measurement of lipase activity by pH-stat……………………………………81
5.3 Time course profile of cell growth and lipase production……………………81
5.4 Screening of factors affecting lipase production significantly………………82
5.5 The method of path of steepest ascent…………………………………………84
5.6 Response surface methodology………………………………………………84
5.7 Confirmation of the predicted optimal value…………………………………87
6 Lipase production from Bulkholderia sp. C20 by fermentation……………………87
6.1 Strain and general cultivation procedure for fermentor experiments…………87
6.2 Lipase activity assay……………………………………………………………88
6.3 Effect of aeration rate and agitation speed on lipase productivity……………88
6.4 Effect of pH control on lipase productivity……………………………………88
6.5 General procedure for flask experiments………………………………………89
6.6 Effect of gum arabic on lipase production in flask experiments………………89
6.7 Effect of oleic acid on lipase production in flask experiments………………90
6.8 Stepwise addition of olive oil in flask experiments……………………………90
6.9 Lipase production in fermentor with stepwise addition of olive oil…………90
7 Immobilization of Burkholderia lipase……………………………………………90
7.1 Preparation of crude lipase solution via Burkholderia strain…………………90
7.2 Emulsification assay…………………………………………………………91
7.3 Lipase activity assay…………………………………………………………91
7.4 Immobilization of lipase by filtration with cellulose nitrate (CN) membrane………………………………………………………………………91
7.5 Activation of celite……………………………………………………………93
7.6 Effect of crude lipase concentration on celite-lipase activity…………………93
7.7 Effect of pH of reaction solution on celite-lipase activity……………………95
7.8 Effect of incubation time on celite-lipase activity……………………………95
7.9 Definition of lipase loading efficiency (LLE)…………………………………95
8 Characterization of free and immobilized Burkholdeira lipase……………………96
8.1 Preparation of free and immobilized Burkholderia lipase……………………96
8.2 Lipase activity under different pH……………………………………………96
8.3 Lipase activity under different temperature…………………………………98
8.4 Thermal stability of free and immobilized lipase……………………………98
8.5 Kinetics analysis………………………………………………………………98
8.6 Experimental procedure for kinetic analysis…………………………………99
9 Biodiesel production by immobilized lipase………………………………………99
9.1 Preparation of immobilized lipase……………………………………………99
9.2 Biodiesel production in shaker by different immobilized lipase……………100
9.3 Synthesis of biodiesel by CN- and celite-lipase by stirring…………………100
9.4 Effect of celite-lipase concentration on biodiesel production………………101
9.5 Effect of molar ratio of methanol to olive oil on biodiesel production………101
Chapter 4 Isolation of bacteria producing lipase………………………………………………………102
1 Introduction………………………………………………………………………102
2 Determination of acidic lipase activity using microwave deactivation treatment……………………………………………………………………102
3 Developing strategies for isolation of acidic lipase-producing bacterial strains………………………………………………………………………104
4 Transesterification of the isolated strains…………………………………………109
5 Comparison of the four isolation strategies………………………………………109
Chapter 5 Optimization of lipase production from Burkholderia sp. C20 in flasks…………………………………………………………………………117
1 Introduction……………………………………………………………………117
2 Time-course profiles of cell growth and lipase activity………………………117
3 Growth associated lipase production……………………………………………119
4 Two-level factorial design and the method of path of steepest ascent……………120
5 Response surface methodology (RSM)…………………………………………124
Chapter 6 Fermentative production of lipase from indigenous Burkholderia sp. C20…………………………………………………………………………128
1 Introduction...……………………………………………………………………128
2 Effect of aeration rate and agitation speed on lipase productivity………………128
3 Dependence of lipase production and cell growth………………………………136
4 Effect of pH-control on lipase productivity……………………………………138
5 Lipase production by stepwise addition of olive oil in flasks……………………141
6 Stepwise addition of olive oil in fermentor………………………………………143
7 Effect of gum arabic and oleic acid on lipase production………………………145
Chapter 7 Immobilization of Burkholderia lipase……………………………………148
1 Introduction………………………………………………………………………148
2 Immobilization of lipase on cellulose nitrate (CN) membrane…………………148
3 Effect of lipase concentration on immobilization of lipase onto celite…………153
4 Effect of pH on lipase immobilization onto celite………………………………153
5 Time course profile of celite-lipase activity during immobilization……………156
Chapter 8 Characterization of free and immobilized Burkholdeira lipase…………158
1 Introduction………………………………………………………………………158
2 Optimization of lipolytic reaction by free and immobilized lipase………………158
3 Thermal stability of free and immobilized lipase………………………………163
4 Kinetic study of free lipase and immobilized lipase……………………………165
Chapter 9 Biodiesel production by immobilized lipase………………………………170
1 Introduction………………………………………………………………………170
2 Pretests of biodiesel production in a shaker by different immobilized lipase……………………………………………………………………………………170
3 Synthesis of biodiesel by CN- and celite-lipase by stirring………………………172
4 Improvement of biodiesel production by celite-lipase……………………………174
Chapter 10 Conclusions and future work……………………………………………178
1 Conclusions………………………………………………………………………178
2 Future work……………………………………………………………………180
References………………………………………………………………………………181
Appendix curriculum vitae………………………………………………………………207
 A.1 Curriculum vitae…………………………………………………………………207
 A.2 Refereed journal paper…………………………………………………………207
 A.3 Conference paper…………………………………………………………………208
                                    

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