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研究生：	陳姍玗 Chen, Shan-yu
論文名稱：	生物界面活性劑-鼠李糖酯之醱酵策略開發 Development of fermentation strategy for rhamnolipid production
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：	2007
畢業學年度：	95
語文別：	英文
論文頁數：	186
中文關鍵詞：	饋料培養、pH-stat 、16S rDNA 、生物界面活性劑、菌種篩選、fill-and-draw 、鼠李糖酯、綠膿桿菌、培養基最適化、回應曲面實驗設計法
外文關鍵詞：	fed-batch culture, fill-and-draw operation, medium optimization, biosurfactant, 16S rDNA, pH-stat, Pseudomonas aeruginosa, strain isolation, rhamnolipid, response surface methodology
相關次數：	點閱：144 下載：5
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本研究首先由受到柴油污染之土壤所篩選出具有生產具表面活性物質之本土菌株S2，該菌株所生產之生物界面活性劑不僅可大幅降低水的表面張力，並對於煤油及柴油具有優異之乳化能力。利用16S rRNA基因序列比對方式鑑定該菌種屬於Pseudomonas aeruginosa菌種，並將該菌株稱為P. aeruginosa S2。該菌株之16S rDNA部分基因序列已上傳至NCBI基因庫中，其登錄編號(accession number)為EF151192。由本土菌株P. aeruginosa S2生產出來的胞外生物界面活性劑經質譜儀鑑定乃屬於醣脂類生物界面活性劑-鼠李糖酯(rhamnolipid)。將所生產的鼠李糖酯經純化後可使水的表面張力由72降低至29.4 mN/m，對煤油之乳化指數E24(%)也可達73.5%，其臨界微胞濃度(CMC)約為170 mg/l 。
為了增進P. aeruginosa S2之鼠李糖酯(RL)產量，本研究先以不同的碳源及氮源進行探討，尋求較適合該菌株之生物界面活性劑生產的碳源及氮源，由實驗結果發現4%葡萄糖及50 mM之硝酸氨對於鼠李糖酯的產量有較佳的表現，另外，也針對了碳氮源的比例進行研究，其結果顯示最佳的碳氮源比例為11:4 。此外本研究引入反應曲面(response surface)實驗設計法的概念，進行該菌株生產RL培養基之微量金屬濃度最適化，首先以二水準設計法(two-level design) 將培養基中影響較小的生產界面活性劑因子排除，接著再以陡升法(method of path of steepest ascent) 尋找出下一階段的實驗設計之培養基組成範圍中心點，最後以回應曲面法找出最佳的培養基組成。由實驗結果發現，Mg2+及Fe2+對於RL之生產是不可或缺的因子，而利用實驗設計法所得之最適化培養基於37 oC、200 rpm 之條件下培養7天可獲得2.37 g/l之rhamnolipid。
由於生物界面活性劑的產量低與成本高，使得生物界面活性劑無法被廣泛的應用，因此亟需發展一套效率高與成本低之rhamnolipid醱酵生產技術，以提昇生物界面活性劑之產業競爭力。本研究以水溶性基質glucose為主要碳源，並以5 L醱酵槽進行RL之生產。首先，利用批次實驗進行最佳醱酵條件之研究，由實驗結果發現，醱酵之溫度、轉速、pH以及消泡劑種類均會影響RL之產量，其最佳醱酵槽之操作條件為：37oC、250 rpm、pH 6.8，以此條件進行醱酵可獲得RL的最高產量為5.31 g/l，此產量較佳於搖瓶實驗的兩倍。接著本研究以pH-stat的饋料策略進行醱酵，以達促進RL產量之目的。由實驗結果顯示以6% glucose為饋料基質，可使RL之產量提昇至6.06 g/l，其生產速率可達172.5 mg/h/l。此外為了更有效率的進行RL之生產，本研究結合pH-stat及fill-and-draw之饋料策略進行RL之量產測試，其結果顯示，除了可持續醱酵生產RL達500 h外，並可於第二個饋料週期將RL之產量提昇至9.4 g/l。此研究結果證實pH-stat醱酵策略確可提昇RL之濃度，並有助於增加該生物界面活性劑商品化之競爭力。

Pseudomonas aeruginosa S2 was isolated from a diesel-contaminated soil site located in southern Taiwan. The strain was selected for its ability to produce extracellular products able to reduce surface tension and emulsify diesel and kerosene. The S2 strain was identified by comparing its 16S rDNA sequence with those available in NCBI gene bank. The accession number is EF151192. The extracellular surface active agent produced by the indigenous strain P. aeruginosa S2 was identified as rhamnolipid, which is one of the most commonly used biosurfactants with the ability to reduce surface tension of water from 72 to 29.4 mN/m and excellent emulsification index (E24) of 73.5 %. Meanwhile, the critical micelle concentration of the rhamnolipid product was 170 mg/l.
To improve production yield of rhamnolipid with P. aeruginosa S2, various carbon and nitrogen sources were screened to select favorable ones leading to better biosurfactant production yield. It was found that using 4% glucose could attain better rhamnolipid yield, while 50 mM NH4NO3 appeared to be the most preferable nitrogen source. Meanwhile, the effect of carbon to nitrogen ratio (C/N ratio) on rhamnolipid yield was also investigated and the optimal C/N ratio was identified as approximately 11.4. Moreover, response surface methodology (RSM) was applied to optimize the trace element concentration for rhamnolipid production. Results from two-level design indicate that concentrations of MgSO4 and FeSO4 were the most significant factors affecting rhamnolipid production. Using steepest ascent method and RSM analysis, an optimal medium composition was determined, giving a rhamnolipid production yield of 2.37 g/l in 100 h at 37oC and 200 rpm agitation.
Rhamnolipid production was also performed in a well-controlled 5 liter laboratory-scale jar fermentor. The effect of pH, temperature, antifoaming strategy, and agitation rate on rhamnolipid production was investigated. Using the optimal medium and operating condition (at 37 oC, pH 6.8 and 250 rpm agitation) further elevated the biosurfactant production yield to 5.31 g/l (in 97 h), which is over 2 fold higher than the best results obtained from shake-flask tests. To further improve the rhamnolipid yield, a pH-stat fed-batch culture was performed by maintaining a constant pH of 6.8 through manipulating glucose feeding. The effect of influent glucose concentration on rhamnolipid yield and productivity was investigated. Using the pH-stat culture, a maximum rhamnolipid concentration (6.06 g/l) and production rate (172.5 mg/h/l) was obtained with 6% glucose in the feed. Moreover, combining pH-stat culture with fill-and-draw operation allowed a stable repeated fed-batch operation for ca. 500 h. A marked increase in rhamnolipid production was achieved, leading to the highest rhamnolipid concentration of ca. 9.4 g/l during the second repeated run.

Contents 

Abstract (Chinese)	I
Abstract (English)	V
Acknowledgement	VII
Contents	IX
List of tables	XVI
List of figures	XIX
Notations	XXIV
Chapter 1 Introduction	1
1.1 Background	1
1.2 Motivation and purpose	4
1.3 Construction of Dissertation	5
Chapter 2 Literature review	9
2.1 Surfactants	10
2.2 Biosurfactants	14
2.3 Rhamnolipid	23
2.3.1 Chemical structure of rhamnolipid	23
2.3.2 Genetics of rhamnolipid synthesis	25
2.3.3 Biosynthesis of rhamnolipid	27
2.3.4 Physicochemical properties of rhamnolipid	29
2.3.5 Rhamnolipid production	30
2.3.6 Purification of rhamnolipid	37
2.3.7 Applications of rhamnolipid	37
2.4 Fed-batch fermentation strategy	39
2.5 Identification of bacteria	44
2.6 Introduction of statistical experimental design………………...…. 47
2.6.1 The 2k factorial design	48
2.6.2 The method of steepest ascent	50
2.6.3 Response surface methodology	51
Chapter 3 Materials and Methods 	55
3.1 Chemicals and materials	55
3.2 Equipment	58
3.3 Isolation of identification of rhamnolipid producing microorganisms	61
3.3.1 Strain isolation	61
3.3.2 Sodium dodecyl sulfate polyacrylamide gel electrophoresis (SDS-PAGE)	62
3.3.3 Amplified 16S rDNA	65
3.3.4 Analysis of 16S rDNA sequence	65
3.4 Flask experiments	68
3.4.1 Microorganisms and cultivation	68
3.4.2 Culture medium for rhamnolipid production	68
3.4.3 Carbon sources and nitrogen sources for rhamnolipid production	69
3.4.4 Trace element for rhamnolipid production	69
3.5 Statistical experimental design	70
3.5.1 Two-level factorial design	70
3.5.2 Response surface methodology	70
3.6 Fermentor experiments	75
3.6.1 Batch culture	75
3.6.2 pH-stat fed-batch culture	76
3.6.3 Constant feeding fed-batch culture	77
3.6.4 Combined pH-stat and fill-and-draw process	77
3.7 Analytical methods and rhamnolipid purification	78
3.7.1 Emulsification test	78
3.7.2 Measurements of surface tension	78
3.7.3 Determination of cell concentration	79
3.7.4 Determination of dry cell weight (DCW)	79
3.7.5 Measurement of glucose concentration	80
3.7.6 Quantification of rhamnolipid	81
3.7.7 Estimation of NH3-N concentration	83
3.7.8 Isolation and purification of rhamnolipid	84
3.7.9 Characterization of purified rhamnolipid product	84
3.8 Transient data analysis	85
Chapter 4 Isolation and Identification of Rhamnolipid Producing Bacterial Strain	87
4.1 Introduction	87
4.2 Isolation of rhamnolipid-producing bacterial strain	87
4.3 Identification of biosurfactant-producing bacterium	90
4.4 Rhamnolipid production of Pseudomonas sp. isolates	92
4.5 Identification of rhamnolipid-producing Pseudomonas sp. S2	94
Chapter 5 Characterization of Rhamnolipid Product Produced by Pseudomonas aeruginosa S2	97
5.1 Introduction	97
5.2 Purification and identification of rhamnolipid	97
5.3 Surface tension reduction of purified rhamnolipid	101
5.4 Emulsification activity of the rhamnolipid product	103
5.5 Thermal stability of rhamnolipid	105
Chapter 6 Fermentation Medium for Rhamnolipid Production	107
6.1 Introduction	107
6.2 Effect of carbon sources on rhamnolipid production	107
6.3 Effect of nitrogen sources on rhamnolipid production	110
6.4 Effect of carbon to nitrogen ratio on rhamnolipid production	112
6.5 Effect of trace element on rhamnolipid production	114
6.6 Optimization of biosurfactants production by response surface methodology	120
Chapter 7 Exploring Fermentor Operation Conditions for Rhamnolipid Production	123
7.1 Introduction	123
7.2 Effect of antifoam agent on rhamnolipid production	123
7.3 Batch rhamnolipid fermentation without pH control	126
7.4 Effect of pH on rhamnolipid production	128
7.5 Effect of temperature on rhamnolipid production	130
7.6 Effect of agitation rate on rhamnolipid	132
7.7 Rhamnolipid production using predetermined favorable conditions	134
7.8 The effect of alkaline pH on rhamnolipid production	136
7.9 Rhamnolipid production using repeated batch fermentation	138
Chapter 8 Rhamnolipid Production with Fed-Batch Fermentation	141
8.1 Introduction	141
8.2 Rhamnolipipd production by fed-batch operation with the addition of glucose alone	142
8.3 Effect of initial glucose concentration on rhamnolipid production using pH-stat fed-batch fermentation	145
8.4 Effect of feeding glucose concentration on rhamnolipid production using pH-stat fed-batch fermentation	147
8.5 Rhamnolipid production with constant feeding fed-batch fermentation	148
8.6 Rhamnolipid production with combined pH-stat and fill-and-draw operation	153   
Chapter 9 Conclusions	159
Chapter 10 Prospects and future work	163
References	165
Appendix Curriculum vitae	і


 
List of Tables

Table 2.1 	Biosurfactants produced by the microorganisms	21
Table 2.1 	Comparison of the microbial production of rhamnolipids by Pseudomonas sp.	34
Table 2.3	The design matrix	50
Table 3.1	 Soil characteristic and properties	63
Table 3.2  The composition of BH medium	63
Table 3.3  The composition of NB medium	63
Table 3.4 	The composition of LB( Luria-Bertani broth) medium	64
Table 3.5	SDS-PAGE composition	64
Table 3.6 	PCR component	66
Table 3.7	Cycling parametars for PCR	67
Table 3.8	The composition of 5X TBE Buffer	67
Table 3.9	The composition of MSI medium	67
Table 3.10 The composition of modified MSI medium	68
Table 3.11	Two-level factorial design for rhamnolipid production by     P. aeruginosa S2	72
Table 3.12	The method of steepest ascent for rhamnolipid production by  P. aeruginosa S2	73
Table 3.13	Experimental design according to response surface methodology (RSM) for rhamnolipid production by          P. aeruginosa S2	74
Table 4.1	Surface activity and emulsification activity of selection strains	89
Table 5.1	Comparison of surface activity of rhamnolipid with and without autoclave treatment	106
Table 6.1	Two-level factorial design for rhamnolipid production by     P. aeruginosa S2	117
Table 6.2	Experimental design according to response surface methodology (RSM) for rhamnolipid production by          P. aeruginosa S2	121
Table 7.1	Effect of pH on rhamnolipid production with P. aeruginosa S2 in batch ferementor. (37oC, 250 rpm agitation)	129
Table 7.2	Effect of temperature on rhamnolipid production with        P. aeruginosa S2 in batch fermentor. (pH 6.8, 250 rpm agitation)	131
Table 7.3	Effect of agitation rate on rhamnolipid production with       P. aeruginosa S2 in batch fermentor	133
Table 8.1		Performance of rhamnolipid production with fed-batch fermentation using pH-stat and constant feeding strategies (37oC, pH 6.8, 250 rpm agitation)	151
 
List of Figures

Figure 1.1 	The fish bone chart describing the research scope            of this study	6
Figure 2.1 	Relationship between surfactant concentration, critical micelle concentration and surface activity	14
Figure 2.2 	Sophorolipid lipid (lactonic form) from Torulopsis magnoliae.	16
Figure 2.3 	Rhamnolipids from Pseudomonas aeruginosa	17
Figure 2.4  Surfactin from Bacillus subtilis	17
Figure 2.5 	Lichenysin from Bacillus licheniformis	17
Figure 2.6 	Biosurfactant produced by Corynebacterium lepus	18
Figure 2.7 	Emulsan from Acinetobacter calcoaceticus	18
Figure 2.8 	Genes involved in the synthesis and regulation of rhamnolipid in P. aeruginosa.	27
Figure 2.9 	Biosynthesis of rhamnolipids R1 and R2 from P. aeruginosa	28
Figure 2.10 	An example of conventional drawing of a phylogenetic tree       (a phenogram)	47
Figure 2.11 The 23 factorial design	49	
Figure 2.12 The central composite designs for p = 3 factors	53	
Figure 3.1  A 5 L well-controlled fermentor	76
Figure 3.2 	Calibration curve of dry cell weight of P. aeruginosa S2	80
Figure 3.3 	Calibration curve of glucose concentration	81
Figure 3.4 	Calibration curve of L-rhamnose concentration	82
Figure 3.5 	Calibration coefficient between rhamnolipid and rhamnose	83
Figure 4.1	SDS-PAGE analysis of the supernatant after sonication with S-series isolates	91
Figure 4.2	Rhamnolipid production of different Pseudomonas species (i.e., strains S2, F72, and J16)	93
Figure 4.3	The dendrogram phylogenic tree of strain S2	95
Figure 4.4 	Partial 16S rDNA gene sequence (1455 bp) of P. aeruginosa S2………	96
Figure 5.1	Mass spectrometry spectrograms of rhamnolipid purified from batch cultures of P. aeruginosa S2	99
Figure 5.2	HPLC of rhamnolipid purified from batch cultures of        P. aeruginosa S2	100
Figure 5.3 	Surface tension reduction profile with purified rhamnolipid from P. aeruginosa S2	102
Figure 5.4 	Emulsification index profile (for kerosene) with purified rhamnolipid from P. aeruginosa S2	104
Figure 6.1	Effect of carbon source on rhamnolipid production by        P. aeruginosa S2	109
Figure 6.2 	Effect of nitrogen source on rhamnolipid production by       P. aeruginosa S2	111
Figure 6.3		Effect of carbon to nitrogen ratio on rhamnolipid production by P. aeruginosa S2..	113
Figure 6.4		Effect of teace element on MSI medium for rhamnolipid production by P. aeruginosa S2	116
Figure 6.5	Two-level factorial design for rhamnolipid production by     P. aeruginosa S2	118
Figure 6.6	Method of path of steepest ascent analysis for rhamnolipid production by P. aeruginosa S2	119
Figure 6.7	Response surface analysis of Mg2+ and Fe2+ concentrations on rhamnolipid production by P. aeruginosa S2	122
Figure 7.1	Time-course profile of cell growth, pH and rhamnolipid production during batch fermentation with P. aeruginosa S2 with antifoam 204	125
Figure 7.2	Time-course profile of cell growth, pH and rhamnolipid production during batch fermentation with P. aeruginosa S2 on MSI medium containing 4% glucose	127
Figure 7.3	Time-course profile of cell growth, glucose consumption and rhamnolipid production during batch fermentation with       P. aeruginosa S2 on MSI medium containing 4% glucose. (Operation conditions: pH 6.8, 250 rpm and 37oC)	135
Figure 7.4	Time-course profile of cell growth, pH and rhamnolipid production with P. aeruginosa S2 under a pH control strategy, in which culture pH was initially controlled at 6.8 but was not controlled when pH increased to above 6.8	137
Figure 7.5	Repeated batch fermentation for rhamnolipid production with  P. aeruginosa S2.  The dotted line represents the feeding time of second run	139
Figure 8.1	Time-course profiles of cell growth, and rhamnolipid production during rhamnolipipd fermentation by addition of glucose of P. aeruginosa S2. The dotted line represents the onset of fed-batch feeding  	144
Figure 8.2	Effect of initial glucose concentration with pH-stat fed-batch fermentation for rhamnolipid production	146
Figure 8.3	Performance of rhamnolipid production with fed-batch fermentation using pH-stat feeding strategies. (37oC, pH 6.8, 250 rpm agitation)	150
Figure 8.4	Fed-batch fermentation using constant feeding and pH-stat feeding strategies with P. aeruginosa S2. for rhamnolipid production	152
Figure 8.5	Combined pH-stat fed-batch and fill-draw operation for rhamnolipid production with P. aeruginosa S2 using 6% glucose alone in the feed	157
Figure 8.6	Combined pH-stat fed-batch and fill-draw operation for rhamnolipid production with P. aeruginosa S2. (a) Profiles of cell concentration and rhamnolipid production (b) profiles of glucose consumption and NH4+ concentration. (Feeding medium: MSI medium amended with 6% glucose)	158
                                    

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