CN113151130A - 一种基因工程菌及其在生物转化甲烷制备异丁醇中的应用 - Google Patents
一种基因工程菌及其在生物转化甲烷制备异丁醇中的应用 Download PDFInfo
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Abstract
本发明公开了一种基因工程菌及其在生物转化甲烷制备异丁醇中的应用。本发明解决的技术问题为:甲烷应用途径单一和沼气、天然气、页岩气的高值利用方式不足等问题。本发明提供了一种重组菌,是在嗜碱甲烷微菌中导入2‑酮异戊酸脱羧酶基因和乙醇脱氢酶基因,得到的重组菌。本发明还提供了一种重组菌,是在嗜碱甲烷微菌中导入2‑酮异戊酸脱羧酶基因,得到的重组菌。本发明还保护所述重组菌在制备异丁醛以及异丁醇中的应用。本发明获得的通过嗜碱甲烷微菌基因工程菌可以实现以甲烷为碳源生产异丁醇。本发明构建得到的嗜碱甲烷微菌基因工程菌对拓展沼气、天然气、页岩气等含有甲烷成分的气体的利用途径具有重大意义。
Description
技术领域
本发明属于生物技术领域,具体涉及一种基因工程菌及其在生物转化甲烷制备异丁醇中的应用,更具体涉及一种嗜碱甲烷微菌基因工程菌及其构建方法及其在生物转化甲烷制备异丁醇中的应用。
背景技术
近年来我国对沼气资源十分重视,随着其生产技术、储藏设备和安全控制系统逐步完善,沼气已成为一种廉价、充足的可再生碳源,但目前我国大量沼气仍以低品位热利用方式为主。随着我国沼气产量不断提高,沼气及其主要成分-甲烷的高端利用途径急需得到扩展和补充。
异丁醇是一种重要的有机化工原料,具有较高的能量密度,较低的挥发性以及较弱的吸水性,在许多领域有着广泛应用,如用于制造石油添加剂、醋酸异丁酯、邻苯二甲酸二异丁酯增塑剂及人造麝香等,是一种潜在的液体燃料。目前报道的利用生物法合成异丁醇都是通过在重组微生物体内构建异源合成路径来实现的,2008年,Atsumi等人证明了在大肠杆菌菌株中利用葡萄糖可以生产异丁醇。
与高成本的葡萄糖相比,沼气是一种相对廉价丰富的碳源,利用微生物作为催化剂转化沼气合成异丁醇不仅缓解了环境污染问题,同时为二代生物燃料的生产提供了一个新的思路。
发明内容
本发明的目的是提供一种基因工程菌及其在生物转化甲烷制备异丁醇中的应用。本发明解决的技术问题为:甲烷应用途径单一和沼气、天然气、页岩气的高值利用方式不足等问题。
本发明提供了一种重组菌,是在嗜碱甲烷微菌中导入2-酮异戊酸脱羧酶基因和乙醇脱氢酶基因,得到的重组菌。
2-酮异戊酸脱羧酶基因和乙醇脱氢酶基因具体可通过重组质粒导入嗜碱甲烷微菌。所述重组质粒为具有2-酮异戊酸脱羧酶基因和乙醇脱氢酶基因的重组质粒。所述重组质粒具体为具有序列表的序列5所示的DNA分子的重组质粒或者具有序列表的序列8所示的DNA分子的重组质粒。所述重组质粒具体可为将载体pAWP89中的“TATTCACACAGGAAACAGCT”和“TAGTTGTCGGGAAGATGCGT”之间的小片段取代为序列表的序列5所示的DNA分子或序列表的序列8所示的DNA分子得到的重组质粒。
所述重组质粒具体可通过三亲结合方法导入嗜碱甲烷微菌。所述三亲结合方法为:将具有所述重组质粒的大肠杆菌、具有辅助质粒PRK600的大肠杆菌和嗜碱甲烷微菌共培养。具有所述重组质粒的大肠杆菌是将所述重组质粒导入大肠杆菌DH5α得到的。具有辅助质粒PRK600的大肠杆菌是将辅助质粒PRK600导入大肠杆菌DH5α得到的。
所述乙醇脱氢酶基因为乙醇脱氢酶A基因或乙醇脱氢酶B基因。
当所述乙醇脱氢酶基因为乙醇脱氢酶A基因时,所述重组菌命名为工程菌5G-IBT-H03。
当所述乙醇脱氢酶基因为乙醇脱氢酶B基因时,所述重组菌命名为工程菌5G-IBT-H04。
本发明还提供了一种重组菌(工程菌5G-IBT-H02),是在嗜碱甲烷微菌中导入2-酮异戊酸脱羧酶基因,得到的重组菌。
2-酮异戊酸脱羧酶基因具体可通过重组质粒导入嗜碱甲烷微菌。所述重组质粒为具有2-酮异戊酸脱羧酶基因的重组质粒。所述重组质粒具体为具有序列表的序列2所示的DNA分子的重组质粒。所述重组质粒具体可为将载体pAWP89中的“TATTCACACAGGAAACAGCT”和“TAGTTGTCGGGAAGATGCGT”之间的小片段取代为序列表的序列2所示的DNA分子得到的重组质粒。
所述重组质粒具体可通过三亲结合方法导入嗜碱甲烷微菌。所述三亲结合方法为:将具有所述重组质粒的大肠杆菌、具有辅助质粒PRK600的大肠杆菌和嗜碱甲烷微菌共培养。具有所述重组质粒的大肠杆菌是将所述重组质粒导入大肠杆菌DH5α得到的。具有辅助质粒PRK600的大肠杆菌是将辅助质粒PRK600导入大肠杆菌DH5α得到的。
本发明还提供了一种重组菌(工程菌5G-IBT-H05),是将具有2-酮异戊酸脱羧酶基因的外源DNA分子整合至嗜碱甲烷微菌的基因组DNA,得到的重组菌。具体的,具有2-酮异戊酸脱羧酶基因的外源DNA分子整合至嗜碱甲烷微菌的基因组DNA中并替换fadE基因。所述替换fadE基因指的是替换基因组DNA中同源臂LF和同源臂RF之间的区段。所述外源DNA分子具体可为序列表的序列10中第1001-4535位核苷酸所示的DNA分子。同源臂LF如序列表的序列10中第1-1000位核苷酸所示。同源臂RF如序列表的序列10中第3536-4535位核苷酸所示。
本发明还保护工程菌5G-IBT-H02、工程菌5G-IBT-H03、工程菌5G-IBT-H04或工程菌5G-IBT-H05在制备异丁醛中的应用。所述应用中,以甲醇为原料。所述应用中,以甲烷和/或沼气和/或天然气和/或页岩气为底物。
本发明还保护工程菌5G-IBT-H03或工程菌5G-IBT-H04在制备异丁醇中的应用。所述应用中,以甲醇为原料。所述应用中,以甲醇和3g/L 3-甲基-2-氧丁酸为原料。所述应用中,以甲烷和/或沼气和/或天然气和/或页岩气为底物。
本发明还提供了一种制备异丁醛的方法,包括如下步骤:发酵工程菌5G-IBT-H02、工程菌5G-IBT-H03、工程菌5G-IBT-H04或工程菌5G-IBT-H05,得到异丁醛。所述发酵具体可为:将工程菌接种至发酵培养基,进行发酵。所述发酵的初始pH为8.0-9.8,具体可为9.0。所述发酵的初始发酵体系的OD600nm值为0.2-0.8,具体可为0.2-0.5或0.5-0.8或0.2或0.5或0.8。所述发酵的温度为24-36℃,具体可为30℃。所述发酵的转速为100~300rpm,具体可为200rpm。所述发酵以甲醇为原料。初始发酵体系中,甲醇的浓度可为1%(体积百分含量)。发酵时,可每12小时加入NADPH或NADPH盐(例如NADPH-Na4)。NADPH或NADPH盐(例如NADPH-Na4)在发酵体系中的浓度可为50mg/L。所述发酵的时间可为12-84小时,具体可为24-48小时。
本发明还提供了一种制备异丁醇的方法,包括如下步骤:发酵工程菌5G-IBT-H03或工程菌5G-IBT-H04,得到异丁醇。所述发酵具体可为:将工程菌接种至发酵培养基,进行发酵。所述发酵的初始pH为8.0-9.8,具体可为9.0。所述发酵的初始发酵体系的OD600nm值为0.2-0.8,具体可为0.2-0.5或0.5-0.8或0.2或0.5或0.8。所述发酵的温度为24-36℃,具体可为30℃。所述发酵的转速为100~300rpm,具体可为200rpm。所述发酵以甲醇为原料。初始发酵体系中,甲醇的浓度可为1%(体积百分含量)。所述发酵以甲醇和3g/L 3-甲基-2-氧丁酸为原料。初始发酵体系中,甲醇的浓度可为1%(体积百分含量),3-甲基-2-氧丁酸的浓度为3g/L。发酵时,可每12小时加入NADPH或NADPH盐(例如NADPH-Na4)。NADPH或NADPH盐(例如NADPH-Na4)在发酵体系中的浓度可为50mg/L。所述发酵的时间可为12-84小时,具体可为24-48小时。
以上任一所述2-酮异戊酸脱羧酶基因为编码2-酮异戊酸脱羧酶的基因。
以上任一所述乙醇脱氢酶基因为编码乙醇脱氢酶的基因。
以上任一所述乙醇脱氢酶A基因为编码乙醇脱氢酶A的基因。
以上任一所述乙醇脱氢酶B基因为编码乙醇脱氢酶B的基因。
以上任一所述嗜碱甲烷微菌具体可为Methylomicrobium buryatense 5GB1S。
所述2-酮异戊酸脱羧酶为如下(a1)或(a2)或(a3):
(a1)序列表的序列1所示的蛋白质;
(a2)来源于乳酸乳球菌且与(a1)具有98%以上同一性且具有2-酮异戊酸脱羧酶功能的蛋白质;
(a3)将(a1)所示的蛋白质经过一个或几个氨基酸残基的取代和/或缺失和/或添加得到的具有2-酮异戊酸脱羧酶功能的蛋白质。
所述2-酮异戊酸脱羧酶基因为如下(b1)或(b2)或(b3):
(b1)编码区如序列表的序列2所示的DNA分子;
(b2)来源于乳酸乳球菌且与(b1)具有98%以上同一性且编码所述蛋白质的DNA分子;
(b3)在严格条件下与(b1)杂交且编码所述蛋白质的DNA分子。
所述乙醇脱氢酶A为如下(c1)或(c2)或(c3):
(c1)序列表的序列3所示的蛋白质;
(c2)来源于运动发酵单胞菌且与(c1)具有98%以上同一性且具有乙醇脱氢酶功能的蛋白质;
(c3)将(c1)所示的蛋白质经过一个或几个氨基酸残基的取代和/或缺失和/或添加得到的具有乙醇脱氢酶功能的蛋白质。
所述乙醇脱氢酶A基因为如下(d1)或(d2)或(d3):
(d1)编码区如序列表的序列4所示的DNA分子;
(d2)来源于运动发酵单胞菌且与(d1)具有98%以上同一性且编码所述蛋白质的DNA分子;
(d3)在严格条件下与(d1)杂交且编码所述蛋白质的DNA分子。
所述乙醇脱氢酶B为如下(e1)或(e2)或(e3):
(e1)序列表的序列6所示的蛋白质;
(e2)来源于运动发酵单胞菌且与(e1)具有98%以上同一性且具有乙醇脱氢酶功能的蛋白质;
(e3)将(e1)所示的蛋白质经过一个或几个氨基酸残基的取代和/或缺失和/或添加得到的具有乙醇脱氢酶功能的蛋白质。
所述乙醇脱氢酶B基因为如下(f1)或(f2)或(f3):
(f1)编码区如序列表的序列7所示的DNA分子;
(f2)来源于运动发酵单胞菌且与(f1)具有98%以上同一性且编码所述蛋白质的DNA分子;
(f3)在严格条件下与(f1)杂交且编码所述蛋白质的DNA分子。
所述严格条件是在2×SSC,0.1%SDS的溶液中,在68℃下杂交并洗膜2次,每次5min,又于0.5×SSC,0.1%SDS的溶液中,在68℃下杂交并洗膜2次,每次15min。
本发明还保护序列表的序列1所示的蛋白质作为2-酮异戊酸脱羧酶的应用。
本发明还保护序列表的序列3所示的蛋白质或序列表的序列6所示的蛋白质作为乙醇脱氢酶的应用。
嗜碱甲烷微菌是一种可以将甲烷作为唯一碳源和能量源的微生物,其特有的甲烷单加氧酶(MMO)可在体内将甲烷氧化为甲醇,然后通过核酮糖单磷酸循环(RuMP)或丝氨酸循环路径,将甲醇/甲醛转化为乙酰辅酶A进入三羧酸循环和其他代谢路径完成理想产物代谢合成过程。嗜碱甲烷微菌具备相对成熟的遗传操作系统,为进行基因工程改造提供了一种简便易行的工具。未发现嗜碱甲烷微菌生产异丁醇的公开报道。
本发明通过表达得2-酮异戊酸脱羧酶基因kivD实现产异丁醛和异丁醇的嗜碱甲烷微菌的构建。2-酮异戊酸在过表达的2-酮异戊酸脱羧酶基因(kivD)作用下脱羧形成异丁醛,随后被过表达的乙醇脱氢酶基因(adhA/B)催化转化为异丁醇。
本发明获得的通过嗜碱甲烷微菌基因工程菌可以实现以甲烷为碳源生产异丁醇的目的。在本发明构建得到的嗜碱甲烷微菌基因工程菌对拓展沼气、天然气、页岩气等含有甲烷成分的气体的利用途径具有重大意义。
附图说明
图1为异丁醛的标准曲线。
图2为异丁醇的标准曲线。
图3为初始OD600nm值为0.2时,不同发酵时间体系的OD600nm值结果。
图4为初始OD600nm值为0.5时,不同发酵时间体系的OD600nm值结果。
图5为初始OD600nm值为0.8时,不同发酵时间体系的OD600nm值结果。
具体实施方式
下面结合具体实施方式对本发明进行进一步的详细描述,给出的实施例仅为了阐明本发明,而不是为了限制本发明的范围。以下提供的实施例可作为本技术领域普通技术人员进行进一步改进的指南,并不以任何方式构成对本发明的限制。
下述实施例中的实验方法,如无特殊说明,均为常规方法,按照本领域内的文献所描述的技术或条件或者按照产品说明书进行。下述实施例中所用的材料、试剂等,如无特殊说明,均可从商业途径得到。实施例中的质粒均已进行测序验证。
如无特殊说明,以下实施例中的定量试验,均设置三次重复实验,结果取平均值。
实施例中所用的嗜碱甲烷微菌为Methylomicrobium buryatense 5GB1S。嗜碱甲烷微菌为Methylomicrobium buryatense 5GB1S又称为原始菌5G-IBT-H01。实施例中的协助菌为E.coli DH5α(pRK600),即含有辅助质粒PRK600的大肠杆菌DH5α。载体pAWP89为环状质粒,如序列表的序列9所示。
pH6.8磷酸盐缓冲液的制备方法:磷酸二氢钾5.44g、磷酸氢二钠10.73g,加水至800ml,溶解后用硫酸调pH至6.8,然后加水定容至1L。
pH9.5碳酸钠缓冲液的制备方法:1M碳酸氢钠水溶液700ml,1M碳酸钠水溶液300ml,混匀,过滤,收集滤液。
500×trace elements:含1.0g/L Na2-EDTA、2.0g/L FeSO4·7H2O、0.8g/LZnSO4·7H2O、0.03g/L MnCl2·4H2O、0.03g/L H3BO3、0.2g/L CoCl2·6H2O、0.6g/L CuCl2·2H2O、0.02g/L NiCl2·6H2O、0.05g/L Na2MoO4·2H2O,余量为水。
固体NMS培养基:由MgSO4·7H2O、CaCl2·6H2O、KNO3、NaCl、500×trace elements、pH6.8磷酸盐缓冲液、pH9.5碳酸钠缓冲液、琼脂粉和水组成。NMS培养基中,含0.2g/LMgSO4·7H2O、0.02g/L CaCl2·6H2O、1g/L KNO3、7.5g/L NaCl、1×trace elements、150g/L琼脂粉,磷酸根离子的浓度为2.3mM,碳酸根离子的浓度为50mM。
液体NMS培养基:与固体NMS培养基相比,差别仅在于不加入琼脂粉。
结合琼脂培养基:由MgSO4·7H2O、CaCl2·6H2O、KNO3、NaCl、500×trace elements、pH6.8磷酸盐缓冲液、pH9.5碳酸钠缓冲液、琼脂粉和水组成。结合琼脂培养基中,含0.2g/LMgSO4·7H2O、0.02g/L CaCl2·6H2O、1g/L KNO3、7.5g/L NaCl、1×trace elements、150g/L琼脂粉,磷酸根离子的浓度为5.8mM,碳酸根离子的浓度为5mM。
实施例1、工程菌5G-IBT-H02的构建
一、制备kivD基因
kivD基因,如序列表的序列2所示,为优化后基因。序列表的序列2所示的DNA分子编码序列表的序列1所示的蛋白质,即2-酮异戊酸脱羧酶(alpha-ketoisovaleratedecarboxylase),其来源于乳酸乳球菌(Lactococcus lactis subsp.lactis)IFPL730菌株。
以人工合成的kivD基因为模板,采用kivD-F和kivD-R组成的引物对进行PCR扩增。
kivD-F:5'-tattcacacaggaaacagctATGTATACCGTTGGCGATTATTTG-3';
kivD-R:5'-acgcatcttcccgacaactaTCACGATTTATTTTGTTCCGCA-3'。
PCR扩增条件:95℃预变性5分钟;92℃变性45秒、63℃退火45秒、72℃延伸3分钟,30个循环;72℃延伸10分钟。
二、构建重组表达载体kivD-pAWP89和重组大肠杆菌
1、取载体pAWP89,采用p89-F和p89-R组成的引物对进行PCR扩增,回收约5kb的线性化片段。
p89-F:5'-TAGTTGTCGGGAAGATGCGT-3';
p89-R:5'-AGCTGTTTCCTGTGTGAATA-3'。
2、采用ClonExpress MultiS One Step Cloning Kit将步骤一得到的PCR扩增产物和步骤1制备的线性化片段连接,得到重组质粒,即为重组表达载体kivD-pAWP89。
3、将重组表达载体kivD-pAWP89导入大肠杆菌DH5α,得到重组大肠杆菌。
三、制备工程菌5G-IBT-H02
将重组表达载体kivD-pAWP89通过三亲结合方法导入嗜碱甲烷微菌,得到工程菌5G-IBT-H02。
具体步骤如下:
1、采用含1%甲醇的固体NMS培养基平板培养嗜碱甲烷微菌3-4天。
2、采用固体LB培养基平板培养协助菌过夜。
3、采用固体LB培养基平板培养步骤二得到的重组大肠杆菌过夜。
4、完成步骤1后,用接种环刮取嗜碱甲烷微菌接种至结合琼脂培养基平板,培养过夜。
5、用接种环分别刮取完成步骤2的协助菌以及完成步骤3的重组大肠杆菌,均涂布接种于完成步骤4的平板,30℃培养48小时。
6、完成步骤5后,从平板上刮取混合菌,涂布于含100μg/ml卡那霉素的固体NMS培养基平板上,30℃培养,可以长出的单菌落即为工程菌5G-IBT-H02。
实施例2、工程菌5G-IBT-H03的构建
一、制备adhA基因
adhA基因,如序列表的序列4所示,为优化后基因。序列表的序列4所示的DNA分子编码序列表的序列3所示的蛋白质,即乙醇脱氢酶A,其来源于运动发酵单胞菌(Zymomonasmobilis)。
以人工合成的adhA基因为模板,采用adhA-F和adhA-R组成的引物对进行PCR扩增。
adhA-F:5'-tattcacacaggaaacagctATGAAAGCGGCGGTTATTACC-3';
adhA-R:5'-acgcatcttcccgacaactaTCAATGATGGGTAAAATCAACAACC-3'。
扩增条件:95℃预变性5分钟;92℃变性45秒、63℃退火45秒、72℃延伸2分钟,30个循环;72℃延伸10分钟。
二、构建重组表达载体adhA-pAWP89和重组大肠杆菌
1、同实施例1的步骤二的1。
2、采用ClonExpress MultiS One Step Cloning Kit将步骤一得到的PCR扩增产物和步骤1制备的线性化片段连接,得到重组质粒,即为重组表达载体adhA-pAWP89。
3、将重组表达载体adhA-pAWP89导入大肠杆菌DH5α,得到重组大肠杆菌。
三、构建重组表达载体kivd-adhA-pAWP89和重组大肠杆菌
1、以人工合成的序列5所示的双链DNA分子为模板,采用kivD-F和adhA-R组成的引物对进行PCR扩增。序列表的序列5中,第1-1647位核苷酸为kivD基因,第1648-1681位核苷酸为RBS序列,第1682-2695位核苷酸为adhA基因。
2、同实施例1的步骤二的1。
3、采用ClonExpress MultiS One Step Cloning Kit将步骤1得到的PCR扩增产物和步骤2制备的线性化片段连接,得到重组质粒,即为重组表达载体kivD-adhA-pAWP89。
4、将重组表达载体kivD-adhA-pAWP89导入大肠杆菌DH5α,得到重组大肠杆菌。
四、制备工程菌5G-IBT-H03
将重组表达载体kivD-adhA-pAWP89通过三亲结合方法导入嗜碱甲烷微菌,得到工程菌5G-IBT-H03。
具体步骤如下:
1、采用含1%甲醇的固体NMS培养基平板培养嗜碱甲烷微菌3-4天。
2、采用固体LB培养基平板培养协助菌过夜。
3、采用固体LB培养基平板培养步骤三得到的重组大肠杆菌过夜。
4、完成步骤1后,用接种环刮取嗜碱甲烷微菌接种至结合琼脂培养基平板,培养过夜。
5、用接种环分别刮取完成步骤2的协助菌以及完成步骤3的重组大肠杆菌,均涂布接种于完成步骤4的平板,30℃培养48小时。
6、完成步骤5后,从平板上刮取混合菌,涂布于含100μg/ml卡那霉素的固体NMS培养基平板上,30℃培养,可以长出的单菌落即为工程菌5G-IBT-H03。
实施例3、工程菌5G-IBT-H04的构建
一、制备adhB基因
adhB基因,如序列表的序列7所示,为优化后基因。序列表的序列7所示的DNA分子编码序列表的序列6所示的蛋白质,即乙醇脱氢酶B,其来源于运动发酵单胞菌(Zymomonasmobilis)。
以人工合成的adhB基因为模板,采用adhB-F和adhB-R组成的引物对进行PCR扩增。
adhB-F:5'-tattcacacaggaaacagctATGGCGTCGTCGACCTTTT-3';
adhB-R:5'-acgcatcttcccgacaactaTCAAAACGCCGACAAAAACA-3'。
扩增条件:95℃预变性5分钟;92℃变性45秒、退火63℃45秒、72℃延伸2分钟,30个循环;72℃延伸10分钟。
二、构建重组表达载体adhB-pAWP89和重组大肠杆菌
1、同实施例1的步骤二的1。
2、采用ClonExpress MultiS One Step Cloning Kit将步骤一得到的PCR扩增产物和步骤1制备的线性化片段连接,得到重组质粒,即为重组表达载体adhB-pAWP89。
3、将重组表达载体adhB-pAWP89导入大肠杆菌DH5α,得到重组大肠杆菌。
三、构建重组表达载体kivd-adhB-pAWP89和重组大肠杆菌
1、以人工合成的序列8所示的双链DNA分子为模板,采用kivD-F和adhB-R组成的引物对进行PCR扩增。序列表的序列8中,第1-1647位核苷酸为kivD基因,第1648-1681位核苷酸为RBS序列,第1682-2833位核苷酸为adhB基因。
2、同实施例1的步骤二的1。
3、采用ClonExpress MultiS One Step Cloning Kit将步骤1得到的PCR扩增产物和步骤2制备的线性化片段连接,得到重组质粒,即为重组表达载体kivD-adhB-pAWP89。
4、将重组表达载体kivD-adhB-pAWP89导入大肠杆菌DH5α,得到重组大肠杆菌。
四、制备工程菌5G-IBT-H04
将重组表达载体kivD-adhB-pAWP89通过三亲结合方法导入嗜碱甲烷微菌,得到工程菌5G-IBT-H04。
具体步骤如下:
1、采用含1%甲醇的固体NMS培养基平板培养嗜碱甲烷微菌3-4天。
2、采用固体LB培养基平板培养协助菌过夜。
3、采用固体LB培养基平板培养步骤三得到的重组大肠杆菌过夜。
4、完成步骤1后,用接种环刮取嗜碱甲烷微菌接种至结合琼脂培养基平板,培养过夜。
5、用接种环分别刮取完成步骤2的协助菌以及完成步骤3的重组大肠杆菌,均涂布接种于完成步骤4的平板,30℃培养48小时。
6、完成步骤5后,从平板上刮取混合菌,涂布于含100μg/ml卡那霉素的固体NMS培养基平板上,30℃培养,可以长出的单菌落即为工程菌5G-IBT-H04。
实施例4、工程菌5G-IBT-H05的构建
一、制备复合片段LF-K+-Kivd-RF
复合片段LF-K+-Kivd-RF为双链DNA分子,如序列表的序列10所示。
序列表的序列10中,第1-1000位核苷酸组成同源臂LF,第1001-1816位核苷酸组成卡那霉素抗性基因,第1889-3535位核苷酸组成kivD基因,第3536-4535位核苷酸组成同源臂RF。同源臂LF和同源臂RF用于同源重组,从而将外源DNA分子整合至嗜碱甲烷微菌的基因组DNA中并替换fadE基因(被替换的是起始密码子开始至终止密码子结束的区段)。
二、制备工程菌5G-IBT-H05
1、采用含1%甲醇的液体NMS培养基培养嗜碱甲烷微菌至OD600nm=2。
2、完成步骤1后,取50ml体系,4℃、5000×g离心10min,收集菌体沉淀,用无菌水洗涤,然后用1000μl无菌水重悬。
3、取50μl步骤2得到的菌体悬液,加入500ng复合片段LF-K+-Kivd-RF,温和混匀,混合液转移至1mm电极杯中,进行电转(电转条件为1.3-2.5kV,25μF,200Ω),然后立即加入1ml液体NMS培养基,然后转移至10ml含0.1%甲醇的NMS液体培养基中,30℃培养过夜。
4、完成步骤3后,室温、5000×g离心10min,收集菌体沉淀,涂布于含100μg/ml卡那霉素的固体NMS培养基平板上,30℃培养,可以长出的单菌落即为工程菌5G-IBT-H05。
实施例5、工程菌转化甲醇生产异丁醛
发酵培养基:含1%(体积百分含量)甲醇的液体NMS培养基。
供试菌分别为:工程菌5G-IBT-H01、工程菌5G-IBT-H02、工程菌5G-IBT-H03、工程菌5G-IBT-H04或工程菌5G-IBT-H05。
将供试菌接种至发酵培养基,进行发酵。初始发酵体系调pH至9.0。设置不同的接种量,使初始发酵体系的OD600nm值分别为0.2、0.5或0.8。发酵过程参数:温度为30℃,转速为200rpm。
发酵过程中,不同时间点检测体系的OD600nm值并取样。
取样样本,4000×g离心2min,收集上清液;采用气相色谱进行检测。气相色谱仪Agilent 7820A,带有DB-WAX柱(30m x 0.32mm x 0.5μm)和火焰离子化检测器。进样量为1μL,分流比为20:1,氢气为载气,流速为30ml/分钟。柱箱温度80℃下保持5分钟,然后以12℃/min的速率加热至230℃。
异丁醛标准品购自Sigma公司。异丁醛标准品的出峰时间为1.667min。异丁醛的标准曲线见图1,函数式:y=0.50483x-1.27826,x为异丁醛浓度mg/L,y为峰面积。
根据目标峰(参照异丁醛标准品的出峰时间)的峰面积和异丁醛的标准曲线函数式,计算取样样本中的异丁醛含量,又称为异丁醛产量,单位为mg/L。
进行三次试验,每次试验至少设置三个重复处理,结果取平均值。
初始OD600nm值为0.2时,不同发酵时间体系的OD600nm值结果见图3。初始OD600nm值为0.5时,不同发酵时间体系的OD600nm值结果见图4。初始OD600nm值为0.8时,不同发酵时间体系的OD600nm值结果见图5。
初始OD600nm值为0.5时,异丁醛产量的结果见表1。初始OD600nm值为0.2时,异丁醛产量的结果见表2。初始OD600nm值为0.8时,异丁醛产量的结果见表3。
表1异丁醛产量(mg/L)
发酵时间 | 5G-IBT-H02 | 5G-IBT-H03 | 5G-IBT-H04 | 5G-IBT-H05 |
24小时 | 3.08±0.18 | 3.33±0.27 | 2.93±0.06 | 2.94±0.10 |
48小时 | 3.53±0.04 | 3.98±0.08 | 11.84±1.12 | 3.29±0.07 |
表2异丁醛产量(mg/L)
发酵时间 | 5G-IBT-H02 | 5G-IBT-H03 | 5G-IBT-H04 | 5G-IBT-H05 |
24小时 | 3.75±0.27 | 3.31±0.41 | 3.17±0.06 | 3.19±0.17 |
48小时 | 3.25±0.20 | 2.69±0.21 | 2.78±0.34 | 2.69±0.22 |
表3异丁醛产量(mg/L)
发酵时间 | 5G-IBT-H02 | 5G-IBT-H03 | 5G-IBT-H04 | 5G-IBT-H05 |
24小时 | 0 | 0 | 0 | 0 |
48小时 | 0 | 0 | 1.4±0.99 | 0 |
实施例6、工程菌转化甲醇生产异丁醛和异丁醇
发酵培养基:含1%(体积百分含量)甲醇的液体NMS培养基。
供试菌分别为:工程菌5G-IBT-H02或工程菌5G-IBT-H04。
将供试菌接种至发酵培养基,进行发酵,分别于发酵12小时后、发酵24小时后、发酵36小时后加入NADPH-Na4并使其在体系中的浓度为50mg/L。初始发酵体系调pH至9.0。初始发酵体系的OD600nm值为0.5。发酵过程参数:温度为30℃,转速为200rpm。
发酵过程中,不同时间点取样。
取样样本,4000×g离心2min,收集上清液;采用气相色谱进行检测。气相色谱仪Agilent 7820A,带有DB-WAX柱(30m x 0.32mm x 0.5μm)和火焰离子化检测器。进样量为1μL,分流比为20:1,氢气为载气,流速为30ml/分钟。柱箱温度80℃下保持5分钟,然后以12℃/min的速率加热至230℃。
异丁醛标准品和异丁醇标准品菌购自Sigma公司。异丁醛标准品的出峰时间为1.667min。异丁醛的标准曲线见图1,函数式:y=0.50483x-1.27826,x为异丁醛浓度(mg/L),y为峰面积。异丁醇标准品的出峰时间为3.169min。异丁醇的标准曲线见图2,函数式:y=0.69032x-2.82354,x为异丁醇浓度(mg/L),y为峰面积。
根据目标峰(参照异丁醛标准品的出峰时间)的峰面积和异丁醛的标准曲线函数式,计算取样样本中的异丁醛含量,又称为异丁醛产量,单位为mg/L。
根据目标峰(参照异丁醇标准品的出峰时间)的峰面积和异丁醇的标准曲线函数式,计算取样样本中的异丁醇含量,又称为异丁醇产量,单位为mg/L。
采用工程菌5G-IBT-H04发酵24小时后,异丁醇产量为4.41mg/L,异丁醛产量为8.92±0.93mg/L。
采用工程菌5G-IBT-H02发酵48小时后,异丁醛产量为13.47±0.62mg/L。
实施例7、工程菌转化甲醇生产异丁醇
发酵培养基:含1%(体积百分含量)甲醇和3g/L 3-甲基-2-氧丁酸的液体NMS培养基。
供试菌为:工程菌5G-IBT-H04。
将供试菌接种至发酵培养基,进行发酵。初始发酵体系调pH至9.0。初始发酵体系的OD600nm值为0.2。发酵过程参数:温度为30℃,转速为200rpm。
发酵过程中,不同时间点取样。
按照实施例6的方法检测异丁醛产量和异丁醇产量。
采用工程菌5G-IBT-H04发酵48小时后,异丁醇产量为2.81mg/L,异丁醛产量为5.78±0.77mg/L。
以上对本发明进行了详述。对于本领域技术人员来说,在不脱离本发明的宗旨和范围,以及无需进行不必要的实验情况下,可在等同参数、浓度和条件下,在较宽范围内实施本发明。虽然本发明给出了特殊的实施例,应该理解为,可以对本发明作进一步的改进。总之,按本发明的原理,本申请欲包括任何变更、用途或对本发明的改进,包括脱离了本申请中已公开范围,而用本领域已知的常规技术进行的改变。按以下附带的权利要求的范围,可以进行一些基本特征的应用。
序列表
<110> 西安交通大学
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ccgccggaaa aaatggattt gtcgattccg cgattggttt tggatggcat tgaagttttg 840
ggctcgttgg ttggcacccg agaagatttg aaagaagcgt ttcaatttgc ggcggaaggc 900
aaagttaaac cgaaagttac caaacgaaaa gttgaagaaa ttaatcaaat ttttgatgaa 960
atggaacatg gcaaatttac cggccgaatg gttgttgatt ttacccatca ttga 1014
<210> 5
<211> 2695
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 5
atgtataccg ttggcgatta tttgttggat cgattgcatg aattgggcat tgaagaaatt 60
tttggcgttc cgggcgatta taatttgcaa tttttggatc aaattatttc gcataaagat 120
atgaaatggg ttggcaatgc gaatgaattg aatgcgtcgt atatggcgga tggctatgcg 180
cgaaccaaaa aagcggcggc gtttttgacc acctttggcg ttggcgaatt gtcggcggtt 240
aatggcttgg cgggctcgta tgcggaaaat ttgccggttg ttgaaattgt tggctcgccg 300
acctcgaaag ttcaaaatga aggcaaattt gttcatcata ccttggcgga tggcgatttt 360
aaacatttta tgaaaatgca tgaaccggtt accgcggcgc gaaccttgtt gaccgcggaa 420
aatgcgaccg ttgaaattga tcgagttttg tcggcgttgt tgaaagaacg aaaaccggtt 480
tatattaatt tgccggttga tgttgcggcg gcgaaagcgg aaaaaccgtc gttgccgttg 540
aaaaaagaaa attcgacctc gaatacctcg gatcaagaaa ttttgaataa aattcaagaa 600
tcgttgaaaa atgcgaaaaa accgattgtt attaccggcc atgaaattat ttcgtttggc 660
ttggaaaaaa ccgttaccca atttatttcg aaaaccaaat tgccgattac caccttgaat 720
tttggcaaat cgtcggttga tgaagcgttg ccgtcgtttt tgggcattta taatggcacc 780
ttgtcggaac cgaatttgaa agaatttgtt gaatcggcgg attttatttt gatgttgggc 840
gttaaattga ccgattcgtc gaccggcgcg tttacccatc atttgaatga aaataaaatg 900
atttcgttga atattgatga aggcaaaatt tttaatgaac gaattcaaaa ttttgatttt 960
gaatcgttga tttcgtcgtt gttggatttg tcggaaattg aatataaagg caaatatatt 1020
gataaaaaac aagaagattt tgttccgtcg aatgcgttgt tgtcgcaaga tcgattgtgg 1080
caagcggttg aaaatttgac ccaatcgaat gaaaccattg ttgcggaaca aggcacctcg 1140
ttttttggcg cgtcgtcgat ttttttgaaa tcgaaatcgc attttattgg ccaaccgttg 1200
tggggctcga ttggctatac ctttccggcg gcgttgggct cgcaaattgc ggataaagaa 1260
tcgcgacatt tgttgtttat tggcgatggc tcgttgcaat tgaccgttca agaattgggc 1320
ttggcgattc gagaaaaaat taatccgatt tgctttatta ttaataatga tggctatacc 1380
gttgaacgag aaattcatgg cccgaatcaa tcgtataatg atattccgat gtggaattat 1440
tcgaaattgc cggaatcgtt tggcgcgacc gaagatcgag ttgtttcgaa aattgttcga 1500
accgaaaatg aatttgtttc ggttatgaaa gaagcgcaag cggatccgaa tcgaatgtat 1560
tggattgaat tgattttggc gaaagaaggc gcgccgaaag ttttgaaaaa aatgggcaaa 1620
ttgtttgcgg aacaaaataa atcgtgaatt ttttcggtaa ctaacacaca ggagaagtca 1680
aatgaaagcg gcggttatta ccaaagatca taccattgaa gttaaagata ccaaattgcg 1740
accgttgaaa tatggcgaag cgttgttgga aatggaatat tgcggcgttt gccataccga 1800
tttgcatgtt aaaaatggcg attttggcga tgaaaccggc cgaattaccg gccatgaagg 1860
cattggcatt gttaaacaag ttggcgaagg cgttacctcg ttgaaagttg gcgatcgagc 1920
gtcggttgcg tggtttttta aaggctgcgg ccattgcgaa tattgcgttt cgggcaatga 1980
aaccttgtgc cgaaatgttg aaaatgcggg ctataccgtt gatggcgcga tggcggaaga 2040
atgcattgtt gttgcggatt attcggttaa agttccggat ggcttggatc cggcggttgc 2100
gtcgtcgatt acctgcgcgg gcgttaccac ctataaagcg gttaaagttt cgcaaattca 2160
accgggccaa tggttggcga tttatggctt gggcggcttg ggcaatttgg cgttgcaata 2220
tgcgaaaaat gtttttaatg cgaaagttat tgcgattgat gttaatgatg aacaattggc 2280
gtttgcgaaa gaattgggcg cggatatggt tattaatccg aaaaatgaag atgcggcgaa 2340
aattattcaa gaaaaagttg gcggcgcgca tgcgaccgtt gttaccgcgg ttgcgaaatc 2400
ggcgtttaat tcggcggttg aagcgattcg agcgggcggc cgagttgttg cggttggctt 2460
gccgccggaa aaaatggatt tgtcgattcc gcgattggtt ttggatggca ttgaagtttt 2520
gggctcgttg gttggcaccc gagaagattt gaaagaagcg tttcaatttg cggcggaagg 2580
caaagttaaa ccgaaagtta ccaaacgaaa agttgaagaa attaatcaaa tttttgatga 2640
aatggaacat ggcaaattta ccggccgaat ggttgttgat tttacccatc attga 2695
<210> 6
<211> 383
<212> PRT
<213> Zymomonas mobilis
<400> 6
Met Ala Ser Ser Thr Phe Tyr Ile Pro Phe Val Asn Glu Met Gly Glu
1 5 10 15
Gly Ser Leu Glu Lys Ala Ile Lys Asp Leu Asn Gly Ser Gly Phe Lys
20 25 30
Asn Ala Leu Ile Val Ser Asp Ala Phe Met Asn Lys Ser Gly Val Val
35 40 45
Lys Gln Val Ala Asp Leu Leu Lys Ala Gln Gly Ile Asn Ser Ala Val
50 55 60
Tyr Asp Gly Val Met Pro Asn Pro Thr Val Thr Ala Val Leu Glu Gly
65 70 75 80
Leu Lys Ile Leu Lys Asp Asn Asn Ser Asp Phe Val Ile Ser Leu Gly
85 90 95
Gly Gly Ser Pro His Asp Cys Ala Lys Ala Ile Ala Leu Val Ala Thr
100 105 110
Asn Gly Gly Glu Val Lys Asp Tyr Glu Gly Ile Asp Lys Ser Lys Lys
115 120 125
Pro Ala Leu Pro Leu Met Ser Ile Asn Thr Thr Ala Gly Thr Ala Ser
130 135 140
Glu Met Thr Arg Phe Cys Ile Ile Thr Asp Glu Val Arg His Val Lys
145 150 155 160
Met Ala Ile Val Asp Arg His Val Thr Pro Met Val Ser Val Asn Asp
165 170 175
Pro Leu Leu Met Val Gly Met Pro Lys Gly Leu Thr Ala Ala Thr Gly
180 185 190
Met Asp Ala Leu Thr His Ala Phe Glu Ala Tyr Ser Ser Thr Ala Ala
195 200 205
Thr Pro Ile Thr Asp Ala Cys Ala Leu Lys Ala Ala Ser Met Ile Ala
210 215 220
Lys Asn Leu Lys Thr Ala Cys Asp Asn Gly Lys Asp Met Pro Ala Arg
225 230 235 240
Glu Ala Met Ala Tyr Ala Gln Phe Leu Ala Gly Met Ala Phe Asn Asn
245 250 255
Ala Ser Leu Gly Tyr Val His Ala Met Ala His Gln Leu Gly Gly Tyr
260 265 270
Tyr Asn Leu Pro His Gly Val Cys Asn Ala Val Leu Leu Pro His Val
275 280 285
Leu Ala Tyr Asn Ala Ser Val Val Ala Gly Arg Leu Lys Asp Val Gly
290 295 300
Val Ala Met Gly Leu Asp Ile Ala Asn Leu Gly Asp Lys Glu Gly Ala
305 310 315 320
Glu Ala Thr Ile Gln Ala Val Arg Asp Leu Ala Ala Ser Ile Gly Ile
325 330 335
Pro Ala Asn Leu Thr Glu Leu Gly Ala Lys Lys Glu Asp Val Pro Leu
340 345 350
Leu Ala Asp His Ala Leu Lys Asp Ala Cys Ala Leu Thr Asn Pro Arg
355 360 365
Gln Gly Asp Gln Lys Glu Val Glu Glu Leu Phe Leu Ser Ala Phe
370 375 380
<210> 7
<211> 1152
<212> DNA
<213> Zymomonas mobilis
<400> 7
atggcgtcgt cgacctttta tattccgttt gttaatgaaa tgggcgaagg ctcgttggaa 60
aaagcgatta aagatttgaa tggctcgggc tttaaaaatg cgttgattgt ttcggatgcg 120
tttatgaata aatcgggcgt tgttaaacaa gttgcggatt tgttgaaagc gcaaggcatt 180
aattcggcgg tttatgatgg cgttatgccg aatccgaccg ttaccgcggt tttggaaggc 240
ttgaaaattt tgaaagataa taattcggat tttgttattt cgttgggcgg cggctcgccg 300
catgattgcg cgaaagcgat tgcgttggtt gcgaccaatg gcggcgaagt taaagattat 360
gaaggcattg ataaatcgaa aaaaccggcg ttgccgttga tgtcgattaa taccaccgcg 420
ggcaccgcgt cggaaatgac ccgattttgc attattaccg atgaagttcg acatgttaaa 480
atggcgattg ttgatcgaca tgttaccccg atggtttcgg ttaatgatcc gttgttgatg 540
gttggcatgc cgaaaggctt gaccgcggcg accggcatgg atgcgttgac ccatgcgttt 600
gaagcgtatt cgtcgaccgc ggcgaccccg attaccgatg cgtgcgcgtt gaaagcggcg 660
tcgatgattg cgaaaaattt gaaaaccgcg tgcgataatg gcaaagatat gccggcgcga 720
gaagcgatgg cgtatgcgca atttttggcg ggcatggcgt ttaataatgc gtcgttgggc 780
tatgttcatg cgatggcgca tcaattgggc ggctattata atttgccgca tggcgtttgc 840
aatgcggttt tgttgccgca tgttttggcg tataatgcgt cggttgttgc gggccgattg 900
aaagatgttg gcgttgcgat gggcttggat attgcgaatt tgggcgataa agaaggcgcg 960
gaagcgacca ttcaagcggt tcgagatttg gcggcgtcga ttggcattcc ggcgaatttg 1020
accgaattgg gcgcgaaaaa agaagatgtt ccgttgttgg cggatcatgc gttgaaagat 1080
gcgtgcgcgt tgaccaatcc gcgacaaggc gatcaaaaag aagttgaaga attgtttttg 1140
tcggcgtttt ga 1152
<210> 8
<211> 2833
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 8
atgtataccg ttggcgatta tttgttggat cgattgcatg aattgggcat tgaagaaatt 60
tttggcgttc cgggcgatta taatttgcaa tttttggatc aaattatttc gcataaagat 120
atgaaatggg ttggcaatgc gaatgaattg aatgcgtcgt atatggcgga tggctatgcg 180
cgaaccaaaa aagcggcggc gtttttgacc acctttggcg ttggcgaatt gtcggcggtt 240
aatggcttgg cgggctcgta tgcggaaaat ttgccggttg ttgaaattgt tggctcgccg 300
acctcgaaag ttcaaaatga aggcaaattt gttcatcata ccttggcgga tggcgatttt 360
aaacatttta tgaaaatgca tgaaccggtt accgcggcgc gaaccttgtt gaccgcggaa 420
aatgcgaccg ttgaaattga tcgagttttg tcggcgttgt tgaaagaacg aaaaccggtt 480
tatattaatt tgccggttga tgttgcggcg gcgaaagcgg aaaaaccgtc gttgccgttg 540
aaaaaagaaa attcgacctc gaatacctcg gatcaagaaa ttttgaataa aattcaagaa 600
tcgttgaaaa atgcgaaaaa accgattgtt attaccggcc atgaaattat ttcgtttggc 660
ttggaaaaaa ccgttaccca atttatttcg aaaaccaaat tgccgattac caccttgaat 720
tttggcaaat cgtcggttga tgaagcgttg ccgtcgtttt tgggcattta taatggcacc 780
ttgtcggaac cgaatttgaa agaatttgtt gaatcggcgg attttatttt gatgttgggc 840
gttaaattga ccgattcgtc gaccggcgcg tttacccatc atttgaatga aaataaaatg 900
atttcgttga atattgatga aggcaaaatt tttaatgaac gaattcaaaa ttttgatttt 960
gaatcgttga tttcgtcgtt gttggatttg tcggaaattg aatataaagg caaatatatt 1020
gataaaaaac aagaagattt tgttccgtcg aatgcgttgt tgtcgcaaga tcgattgtgg 1080
caagcggttg aaaatttgac ccaatcgaat gaaaccattg ttgcggaaca aggcacctcg 1140
ttttttggcg cgtcgtcgat ttttttgaaa tcgaaatcgc attttattgg ccaaccgttg 1200
tggggctcga ttggctatac ctttccggcg gcgttgggct cgcaaattgc ggataaagaa 1260
tcgcgacatt tgttgtttat tggcgatggc tcgttgcaat tgaccgttca agaattgggc 1320
ttggcgattc gagaaaaaat taatccgatt tgctttatta ttaataatga tggctatacc 1380
gttgaacgag aaattcatgg cccgaatcaa tcgtataatg atattccgat gtggaattat 1440
tcgaaattgc cggaatcgtt tggcgcgacc gaagatcgag ttgtttcgaa aattgttcga 1500
accgaaaatg aatttgtttc ggttatgaaa gaagcgcaag cggatccgaa tcgaatgtat 1560
tggattgaat tgattttggc gaaagaaggc gcgccgaaag ttttgaaaaa aatgggcaaa 1620
ttgtttgcgg aacaaaataa atcgtgaatt ttttcggtaa ctaacacaca ggagaagtca 1680
aatggcgtcg tcgacctttt atattccgtt tgttaatgaa atgggcgaag gctcgttgga 1740
aaaagcgatt aaagatttga atggctcggg ctttaaaaat gcgttgattg tttcggatgc 1800
gtttatgaat aaatcgggcg ttgttaaaca agttgcggat ttgttgaaag cgcaaggcat 1860
taattcggcg gtttatgatg gcgttatgcc gaatccgacc gttaccgcgg ttttggaagg 1920
cttgaaaatt ttgaaagata ataattcgga ttttgttatt tcgttgggcg gcggctcgcc 1980
gcatgattgc gcgaaagcga ttgcgttggt tgcgaccaat ggcggcgaag ttaaagatta 2040
tgaaggcatt gataaatcga aaaaaccggc gttgccgttg atgtcgatta ataccaccgc 2100
gggcaccgcg tcggaaatga cccgattttg cattattacc gatgaagttc gacatgttaa 2160
aatggcgatt gttgatcgac atgttacccc gatggtttcg gttaatgatc cgttgttgat 2220
ggttggcatg ccgaaaggct tgaccgcggc gaccggcatg gatgcgttga cccatgcgtt 2280
tgaagcgtat tcgtcgaccg cggcgacccc gattaccgat gcgtgcgcgt tgaaagcggc 2340
gtcgatgatt gcgaaaaatt tgaaaaccgc gtgcgataat ggcaaagata tgccggcgcg 2400
agaagcgatg gcgtatgcgc aatttttggc gggcatggcg tttaataatg cgtcgttggg 2460
ctatgttcat gcgatggcgc atcaattggg cggctattat aatttgccgc atggcgtttg 2520
caatgcggtt ttgttgccgc atgttttggc gtataatgcg tcggttgttg cgggccgatt 2580
gaaagatgtt ggcgttgcga tgggcttgga tattgcgaat ttgggcgata aagaaggcgc 2640
ggaagcgacc attcaagcgg ttcgagattt ggcggcgtcg attggcattc cggcgaattt 2700
gaccgaattg ggcgcgaaaa aagaagatgt tccgttgttg gcggatcatg cgttgaaaga 2760
tgcgtgcgcg ttgaccaatc cgcgacaagg cgatcaaaaa gaagttgaag aattgttttt 2820
gtcggcgttt tga 2833
<210> 9
<211> 5756
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 9
gaccctttcc gacgctcacc gggctggttg ccctcgccgc tgggctggcg gccgtctatg 60
gccctgcaaa cgcgccagaa acgccgtcga agccgtgtgc gagacaccgc ggccgccggc 120
gttgtggata cctcgcggaa aacttggccc tcactgacag atgaggggcg gacgttgaca 180
cttgaggggc cgactcaccc ggcgcggcgt tgacagatga ggggcaggct cgatttcggc 240
cggcgacgtg gagctggcca gcctcgcaaa tcggcgaaaa cgcctgattt tacgcgagtt 300
tcccacagat gatgtggaca agcctgggga taagtgccct gcggtattga cacttgaggg 360
gcgcgactac tgacagatga ggggcgcgat ccttgacact tgaggggcag agtgctgaca 420
gatgaggggc gcacctattg acatttgagg ggctgtccac aggcagaaaa tccagcattt 480
gcaagggttt ccgcccgttt ttcggccacc gctaacctgt cttttaacct gcttttaaac 540
caatatttat aaaccttgtt tttaaccagg gctgcgccct gtgcgcgtga ccgcgcacgc 600
cgaagggggg tgccccccct tctcgaaccc tcccggcccg ctaacgcggg cctcccatcc 660
ccccaggggc tgcgcccctc ggccgcgaac ggcctcaccc caaaaatggc agccaagctg 720
acccgctagg gacgtgaagt cgattccttc gatggttagc aatcaaagac tcagagtgct 780
gtctggagcg tgaatctaac ggtacgtatc tcgattgctc ggtcgctatt cgcactctgc 840
gaaagttcgt accgctcatt cactaggttg cgaatcatga ccaaaatccc ttaacgtgag 900
ttttcgttcc actgagcgtc agaccccgta gaaaagatca aaggatcttc ttgagatcct 960
ttttttctgc gcgtaatctg ctgcttgcaa acaaaaaaac caccgctacc agcggtggtt 1020
tgtttgccgg atcaagagct accaactctt tttccgaagg taactggctt cagcagagcg 1080
cagataccaa atactgtcct tctagtgtag ccgtagttag gccaccactt caagaactct 1140
gtagcaccgc ctacatacct cgctctgcta atcctgttac cagtggctgc tgccagtggc 1200
gataagtcgt gtcttaccgg gttggactca agacgatagt taccggataa ggcgcagcgg 1260
tcgggctgaa cggggggttc gtgcacacag cccagcttgg agcgaacgac ctacaccgaa 1320
ctgagatacc tacagcgtga gctatgagaa agcgccacgc ttcccgaagg gagaaaggcg 1380
gacaggtatc cggtaagcgg cagggtcgga acaggagagc gcacgaggga gcttccaggg 1440
ggaaacgcct ggtatcttta tagtcctgtc gggtttcgcc acctctgact tgagcgtcga 1500
tttttgtgat gctcgtcagg ggggcggagc ctatggaaaa acgccagcaa cgcggccttt 1560
ttacggttcc tggccttttg ctggcctttt gctcacatgt tctttcctgc gttatcccct 1620
gattctgtgg ataaccgtat taccgccttt gagtgagctg ctctgaaatg agctgttgac 1680
aattaatcat cggctcgtat aatgtgtgga ggtattcaca caggaaacag ctatggtgag 1740
caagggcgag gaggtcatca aagagttcat gcgcttcaag gtgcgcatgg agggctccat 1800
gaacggccac gagttcgaga tcgagggcga gggcgagggc cgcccctacg agggcaccca 1860
gaccgccaag ctgaaggtga ccaagggcgg ccccctgccc ttcgcctggg acatcctgtc 1920
cccccagttc atgtacggct ccaaggcgta cgtgaagcac cccgccgaca tccccgatta 1980
caagaagctg tccttccccg agggcttcaa gtgggagcgc gtgatgaact tcgaggacgg 2040
cggtctggtg accgtgaccc aggactcctc cctgcaggac ggcacgctga tctacaaggt 2100
gaagatgcgc ggcaccaact tcccccccga cggccccgta atgcagaaga agaccatggg 2160
ctgggaggcc tccaccgagc gcctgtaccc ccgcgacggc gtgctgaagg gcgagatcca 2220
ccaggccctg aagctgaagg acggcggcca ctacctggtg gagttcaaga ccatctacat 2280
ggccaagaag cccgtgcaac tgcccggcta ctactacgtg gacaccaagc tggacatcac 2340
ctcccacaac gaggactaca ccatcgtgga acagtacgag cgctccgagg gccgccacca 2400
cctgttcctg tacggcatgg acgagctgta caagtagttg tcgggaagat gcgtgatctg 2460
atccttccac tcagcaaaag ttcgatttat tcaacaaagc cgccgtcccg tcaagtcagc 2520
gtaatgctct gccagtgtta caaccaatta accaattctg attagaaaaa ctcatcgagc 2580
atcaaatgaa actgcaattt attcatatca ggattatcaa taccatattt ttgaaaaagc 2640
cgtttctgta atgaaggaga aaactcaccg aggcagttcc ataggatggc aagatcctgg 2700
tatcggtctg cgattccgac tcgtccaaca tcaatacaac ctattaattt cccctcgtca 2760
aaaataaggt tatcaagtga gaaatcacca tgagtgacga ctgaatccgg tgagaatggc 2820
aaaagcttat gcatttcttt ccagacttgt tcaacaggcc agccattacg ctcgtcatca 2880
aaatcactcg catcaaccaa accgttattc attcgtgatt gcgcctgagc gagacgaaat 2940
acgcgatcgc tgttaaaagg acaattacaa acaggaatcg aatgcaaccg gcgcaggaac 3000
actgccagcg catcaacaat attttcacct gaatcaggat attcttctaa tacctggaat 3060
gctgttttcc cggggatcgc agtggtgagt aaccatgcat catcaggagt acggataaaa 3120
tgcttgatgg tcggaagagg cataaattcc gtcagccagt ttagtctgac catctcatct 3180
gtaacatcat tggcaacgct acctttgcca tgtttcagaa acaactctgg cgcatcgggc 3240
ttcccataca atcgatagat tgtcgcacct gattgcccga cattatcgcg agcccattta 3300
tacccatata aatcagcatc catgttggaa tttaatcgcg gcctcgagca agacgtttcc 3360
cgttgaatat ggctcataac accccttgta ttactgttta tgtaagcaga cagttttatt 3420
gttcatgatg atatattttt atcttgtgca atgtaacatc agagattttg agacacaacg 3480
tggctttcga cttccggcaa gctatacgcg ccctagaatt gtcaatttta atcctctgtt 3540
tatcggcagt tcgtagagcg cgccgtgcgt cccgagcgat actgagcgaa gcaagtgcgt 3600
cgagcagtgc ccgcttgttc ctgaaatgcc agtaaagcgc tggctgctga acccccagcc 3660
ggaactgacc ccacaaggcc ctagcgtttg caatgcacca ggtcatcatt gacccaggcg 3720
tgttccacca ggccgctgcc tcgcaactct tcgcaggctt cgccgacctg ctcgcgccac 3780
ttcttcacgc gggtggaatc cgatccgcac atgaggcgga aggtttccag cttgagcggg 3840
tacggctccc ggtgcgagct gaaatagtcg aacatccgtc gggccgtcgg cgacagcttg 3900
cggtacttct cccatatgaa tttcgtgtag tggtcgccag caaacagcac gacgatttcc 3960
tcgtcgatca ggacctggca acgggacgtt ttcttgccac ggtccaggac gcggaagcgg 4020
tgcagcagcg acaccgattc caggtgccca acgcggtcgg acgtgaagcc catcgccgtc 4080
gcctgtaggc gcgacaggca ttcctcggcc ttcgtgtaat accggccatt gatcgaccag 4140
cccaggtcct ggcaaagctc gtagaacgtg aaggtgatcg gctcgccgat aggggtgcgc 4200
ttcgcgtact ccaacacctg ctgccacacc agttcgtcat cgtcggcccg cagctcgacg 4260
ccggtgtagg tgatcttcac gtccttgttg acgtggaaaa tgaccttgtt ttgcagcgcc 4320
tcgcgcggga ttttcttgtt gcgcgtggtg aacagggcag agcgggccgt gtcgtttggc 4380
atcgctcgca tcgtgtccgg ccacggcgca atatcgaaca aggaaagctg catttccttg 4440
atctgctgct tcgtgtgttt cagcaacgcg gcctgcttgg cctcgctgac ctgttttgcc 4500
aggtcctcgc cggcggtttt tcgcttcttg gtcgtcatag ttcctcgcgt gtcgatggtc 4560
atcgacttcg ccaaacctgc cgcctcctgt tcgagacgac gcgaacgctc cacggcggcc 4620
gatggcgcgg gcagggcagg gggagccagt tgcacgctgt cgcgctcgat cttggccgta 4680
gcttgctgga ccatcgagcc gacggactgg aaggtttcgc ggggcgcacg catgacggtg 4740
cggcttgcga tggtttcggc atcctcggcg gaaaaccccg cgtcgatcag ttcttgcctg 4800
tatgccttcc ggtcaaacgt ccgattcatt caccctcctt gcgggattgc cccgactcac 4860
gccggggcaa tgtgccctta ttcctgattt gacccgcctg gtgccttggt gtccagataa 4920
tccaccttat cggcaatgaa gtcggtcccg tagaccgtct ggccgtcctt ctcgtacttg 4980
gtattccgaa tcttgccctg cacgaatacc agctccgcga agtcgctctt cttgatggag 5040
cgcatgggga cgtgcttggc aatcacgcgc accccccggc cgttttagcg gctaaaaaag 5100
tcatggctct gccctcgggc ggaccacgcc catcatgacc ttgccaagct cgtcctgctt 5160
ctcttcgatc ttcgccagca gggcgaggat cgtggcatca ccgaaccgcg ccgtgcgcgg 5220
gtcgtcggtg agccagagtt tcagcaggcc gcccaggcgg cccaggtcgc cattgatgcg 5280
ggccagctcg cggacgtgct catagtccac gacgcccgtg attttgtagc cctggccgac 5340
ggccagcagg taggcctaca ggctcatgcc ggccgccgcc gccttttcct caatcgctct 5400
tcgttcgtct ggaaggcagt acaccttgat aggtgggctg cccttcctgg ttggcttggt 5460
ttcatcagcc atccgcttgc cctcatctgt tacgccggcg gtagccggcc agcctcgcag 5520
agcaggattc ccgttgagca ccgccaggtg cgaataaggg acagtgaaga aggaacaccc 5580
gctcgcgggt gggcctactt cacctatcct gcccggctga cgccgttgga tacaccaagg 5640
aaagtctaca cgaacccttt ggcaaaatcc tgtatatcgt gcgaaaaagg atggatatac 5700
cgaaaaaatc gctataatga ccccgaagca gggttatgca gcggaaaaga tccgtc 5756
<210> 10
<211> 4535
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 10
cggcattgct gcggcaagat tgggacaccg tcgcccgaat cgtcgagctc gggcaacaaa 60
cttatcaagc cttgaaatac gcgcctttcc cggtcgtcgg cgcgccgtcc ggtttggcgc 120
tgggcggcgg ctgcgaaatt ctgctgcatt gcgatgcggt gcaagcccat gccgaactgt 180
ataccggcct cgtcgaaacc ggagtcggtc tggtgccggg ctggggcggc tgtaaggaat 240
tattgcgccg atggcttagc ttgcctaacc gcccgagcgg accgatgccg gcgatttcgc 300
aagccttcga gaccatcgcg ctcgccaaaa cgtccaaatc ggccttgctg gcgaaggaat 360
tactctattt atccgaacac gacggcatca caatgaacaa ggaccgtctg ctggccgacg 420
caaaagcgcg cgtcctgaca atgatcgccg actaccgacc gccggaacct tatgtttatt 480
atttaccggg cgcttcggcc cgcgcggcgc ttgacatagc cgtgcgcaac ttagcgctct 540
cgggcaaagc aaccgcctac gaccgggaaa tcgccggcca actggctttc gtattgagcg 600
gcggagacac cgactccctc gatccgctta gcgaacaaga catattgaac ctggaacgac 660
aagccttttt gcacctggtc aagcaacccg gcactgcagc aaggctcgaa cacatgctaa 720
aaaccggcaa gccgttgaga aactgacggg gcaatcgagc aacgtaacgt tcatgaaggc 780
ctggccatcg ccccggcaaa tgaaagtagt ctggtgacgg agggtgcggt cactcgcccg 840
acaggacgcc gtgaatacgt ccatgtaggc tcgacggcgg ctgtcgatcg agcaaccgca 900
ccctctccgg aaccggcatt gccagaacgg agttttgcat atcgaaaaat tagataaaga 960
acccaaacct acgaactgac acagcaacag gaattaagcc atgagccata ttcaacggga 1020
aacgtcttgc tcgaggccgc gattaaattc caacatggat gctgatttat atgggtataa 1080
atgggctcgc gataatgtcg ggcaatcagg tgcgacaatc tatcgattgt atgggaagcc 1140
cgatgcgcca gagttgtttc tgaaacatgg caaaggtagc gttgccaatg atgttacaga 1200
tgagatggtc agactaaact ggctgacgga atttatgcct cttccgacca tcaagcattt 1260
tatccgtact cctgatgatg catggttact caccactgcg atccccggga aaacagcatt 1320
ccaggtatta gaagaatatc ctgattcagg tgaaaatatt gttgatgcgc tggcagtgtt 1380
cctgcgccgg ttgcattcga ttcctgtttg taattgtcct tttaacagcg atcgcgtatt 1440
tcgtctcgct caggcgcaat cacgaatgaa taacggtttg gttgatgcga gtgattttga 1500
tgacgagcgt aatggctggc ctgttgaaca agtctggaaa gaaatgcata agcttttgcc 1560
attctcaccg gattcagtcg tcactcatgg tgatttctca cttgataacc ttatttttga 1620
cgaggggaaa ttaataggtt gtattgatgt tggacgagtc ggaatcgcag accgatacca 1680
ggatcttgcc atcctatgga actgcctcgg tgagttttct ccttcattac agaaacggct 1740
ttttcaaaaa tatggtattg ataatcctga tatgaataaa ttgcagtttc atttgatgct 1800
cgatgagttt ttctaactct gaaatgagct gttgacaatt aatcatcggc tcgtataatg 1860
tgtggaggta ttcacacagg aaacagctat gtataccgtt ggcgattatt tgttggatcg 1920
attgcatgaa ttgggcattg aagaaatttt tggcgttccg ggcgattata atttgcaatt 1980
tttggatcaa attatttcgc ataaagatat gaaatgggtt ggcaatgcga atgaattgaa 2040
tgcgtcgtat atggcggatg gctatgcgcg aaccaaaaaa gcggcggcgt ttttgaccac 2100
ctttggcgtt ggcgaattgt cggcggttaa tggcttggcg ggctcgtatg cggaaaattt 2160
gccggttgtt gaaattgttg gctcgccgac ctcgaaagtt caaaatgaag gcaaatttgt 2220
tcatcatacc ttggcggatg gcgattttaa acattttatg aaaatgcatg aaccggttac 2280
cgcggcgcga accttgttga ccgcggaaaa tgcgaccgtt gaaattgatc gagttttgtc 2340
ggcgttgttg aaagaacgaa aaccggttta tattaatttg ccggttgatg ttgcggcggc 2400
gaaagcggaa aaaccgtcgt tgccgttgaa aaaagaaaat tcgacctcga atacctcgga 2460
tcaagaaatt ttgaataaaa ttcaagaatc gttgaaaaat gcgaaaaaac cgattgttat 2520
taccggccat gaaattattt cgtttggctt ggaaaaaacc gttacccaat ttatttcgaa 2580
aaccaaattg ccgattacca ccttgaattt tggcaaatcg tcggttgatg aagcgttgcc 2640
gtcgtttttg ggcatttata atggcacctt gtcggaaccg aatttgaaag aatttgttga 2700
atcggcggat tttattttga tgttgggcgt taaattgacc gattcgtcga ccggcgcgtt 2760
tacccatcat ttgaatgaaa ataaaatgat ttcgttgaat attgatgaag gcaaaatttt 2820
taatgaacga attcaaaatt ttgattttga atcgttgatt tcgtcgttgt tggatttgtc 2880
ggaaattgaa tataaaggca aatatattga taaaaaacaa gaagattttg ttccgtcgaa 2940
tgcgttgttg tcgcaagatc gattgtggca agcggttgaa aatttgaccc aatcgaatga 3000
aaccattgtt gcggaacaag gcacctcgtt ttttggcgcg tcgtcgattt ttttgaaatc 3060
gaaatcgcat tttattggcc aaccgttgtg gggctcgatt ggctatacct ttccggcggc 3120
gttgggctcg caaattgcgg ataaagaatc gcgacatttg ttgtttattg gcgatggctc 3180
gttgcaattg accgttcaag aattgggctt ggcgattcga gaaaaaatta atccgatttg 3240
ctttattatt aataatgatg gctataccgt tgaacgagaa attcatggcc cgaatcaatc 3300
gtataatgat attccgatgt ggaattattc gaaattgccg gaatcgtttg gcgcgaccga 3360
agatcgagtt gtttcgaaaa ttgttcgaac cgaaaatgaa tttgtttcgg ttatgaaaga 3420
agcgcaagcg gatccgaatc gaatgtattg gattgaattg attttggcga aagaaggcgc 3480
gccgaaagtt ttgaaaaaaa tgggcaaatt gtttgcggaa caaaataaat cgtgacttca 3540
cctgattagc aagatgcttc agcttgccga agccaagtgt ccaagctggg gttagtcgta 3600
gtcgcaaaaa ctaaaatatg ctgttaccca agctccagct tgggtaacct gttcaggaag 3660
ctctagcttc ccgataatca agaaagattg aacgagcgag cgagtcgggg gtgattgaga 3720
atgcgcggag gttgtgtcgg tctcaagaag ctggagcttc cggtgtgtct ttcccaagct 3780
ggagcttggg aaagagcgtg aaaatgaaag cgatcgcggt cgacgatttc gcgcctgagc 3840
acttgaccgg acgctcttga ctttacgcca ggattttcgt catgtaccgc acgcccaaaa 3900
gcgcgggtgg gtcagtattc atagcgtcct ttgacgggca tttgactagc ccagaagata 3960
tttagagtct taatcaatac ataatgctgc cgggaaaatt actttcccgg ccaatcgttt 4020
tatcaaatcc ctttttgttc cttggtcaat tgcttcaacc agccttccac ttgcggactc 4080
aactcttttt ttccgcttaa tttcaaggtt tgcgccaaat agccgtgttc aaccaacggc 4140
aacggcaaca cgccctgccc gcccatgccg ttatgggctt ttatgctgct atctttgtca 4200
tcggccaata aaatactgcc ggccacctca agcttatcca tatccacaac caacagatta 4260
ttagcgaact tgctggccac ataagcatag tacccgccgc cttttttcat gccgaaatga 4320
acgccatgac atcccgcctc gcaaggcaag cttttgacga ctttattggt tgcggtatcg 4380
acaatcgtga tgctcatgct caaggtattc gctgtcacta catacttacc gtcaggacta 4440
accggcgttt gtatcggcaa caacccgtat gcctcgccgc tgatttttcc cgatatcgga 4500
tcgtaatcgg cggccaggtc gatgtcggac aattt 4535
Claims (10)
1.一种重组菌,是在嗜碱甲烷微菌中导入2-酮异戊酸脱羧酶基因和乙醇脱氢酶基因,得到的重组菌。
2.如权利要求1所述的重组菌,其特征在于:所述乙醇脱氢酶基因为乙醇脱氢酶A基因或乙醇脱氢酶B基因。
3.一种重组菌,是在嗜碱甲烷微菌中导入2-酮异戊酸脱羧酶基因,得到的重组菌。
4.一种重组菌,是将具有2-酮异戊酸脱羧酶基因的外源DNA分子整合至嗜碱甲烷微菌的基因组DNA,得到的重组菌。
5.权利要求1至4中任一所述重组菌在制备异丁醛中的应用。
6.权利要求1或2所述重组菌在制备异丁醇中的应用。
7.一种制备异丁醛的方法,包括如下步骤:发酵权利要求1至4中任一所述重组菌,得到异丁醛。
8.一种制备异丁醇的方法,包括如下步骤:发酵权利要求1或2所述重组菌,得到异丁醇。
9.序列表的序列1所示的蛋白质作为2-酮异戊酸脱羧酶的应用。
10.序列表的序列3所示的蛋白质或序列表的序列6所示的蛋白质作为乙醇脱氢酶的应用。
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