CN114774477B - 使用了重组酵母的乙醇的制造方法 - Google Patents
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Abstract
在具有木糖代谢能力的酵母的木糖同化和乙醇发酵时代谢培养基中的乙酸,降低乙酸浓度。乙醇的制造方法,具有以下工序:将导入了木糖异构酶基因和乙醛脱氢酶基因的重组酵母,用含有木糖的培养基进行培养,从而进行乙醇发酵木。
Description
本申请是申请日为2014年2月27日、申请号为201480010549.7、发明名称为“使用了重组酵母的乙醇的制造方法”的专利申请的分案申请。
技术领域
本发明涉及使用了具有木糖代谢能力的重组酵母的乙醇的制造方法。
背景技术
纤维素系生物质作为乙醇等有用醇和/或有机酸的原料而被有效利用。为了在利用了纤维素系生物质的乙醇制造中提高乙醇生产量,开发了作为底物能够利用5碳单糖木糖的酵母。例如,专利文献1公开了染色体中插入了来源于树干毕赤酵母(Pichiastipitis)的木糖还原酶基因和木糖醇脱氢酶基因以及来源于酿酒酵母(S.cerevisiae)的木酮糖激酶基因的酵母。
另外,已知乙酸在纤维素系生物质的水解物中大量含有,抑制酵母的乙醇发酵。特别地,已知在导入了木糖同化基因的酵母中,显著地抑制以木糖为糖源的乙醇发酵(非专利文献1和2)
另一方面,将用纤维素酶糖化纤维素系生物质而得的物质进行发酵后的醪中,主要包含未发酵残渣、难发酵残渣、酶和发酵用微生物。通过将包含醪的反应液在接着进行的发酵中利用,可以实现发酵用微生物的再利用,降低发酵用微生物的新加入量,实现低成本化。但是,此时,醪所含的乙酸也同时被带入,其结果是有时发酵培养基所含的乙酸的浓度上升,抑制乙醇发酵。另外,即使在将发酵槽内的醪送入保持减压的闪蒸罐,在闪蒸罐内除去乙醇后,将醪送回发酵槽这样的连续发酵法中,也难以从醪中去除乙酸,因而由乙酸产生的发酵抑制成为重要的问题。
为了避免由乙酸产生的发酵抑制,有通过乙醇发酵中一般使用的菌株酿酒酵母的LPP1基因和/或ENA1基因的过表达(非专利文献3)、FPS1基因的破坏(非专利文献4)来改善在乙酸存在下乙醇的发酵能力的报告。然而,这些报告是以葡萄糖作为底物时的乙醇发酵的结果,对由于乙酸而发酵被显著抑制的以木糖作为底物时的乙醇发酵的效果不明。另外,即使使用这些报告的突变酵母,也不能实现减少在如上所述的发酵菌体再利用和/或连续发酵时成为问题的乙酸的带入量。
作为避免由乙酸产生的发酵抑制的其他方法,考虑使培养基中的乙酸在乙醇发酵的同时代谢。但是,乙酸的代谢是需氧的反应,与乙醇的代谢途径重复。因此,虽然如果在需氧条件下进行发酵则可能使乙酸代谢,但同时目的物质乙醇也被代谢。
为了在乙醇不被代谢的厌氧条件下使乙酸代谢,有通过在破坏了酿酒酵母的甘油生产途径的GPD1·GPD2基因的菌株中导入编码乙醛脱氢酶(EC 1.2.1.10)的mhpF基因,从而同化乙酸的报告(非专利文献5、专利文献2)。此外,乙醛脱氢酶催化以下的可逆反应。
此外,利用GPD1·GPD2基因的甘油生产途径如下述式所示,是将代谢中产生的剩余的辅酶NADH氧化成NAD+的途径。
0.5葡萄糖+NADH+H++ATP→甘油+NAD++ADP+Pi
于是通过破坏GPD1·GPD2基因来破坏该反应途径,通过导入mhpF来供给剩余的辅酶NADH,进行下述的反应。
乙酰辅酶A+NADH+H+→乙醛+NAD++辅酶A
另外,乙酰辅酶A由乙酸通过乙酰辅酶A合成酶而合成,乙醛被转换成乙醇,因而最终形成下述反应式,在剩余的辅酶NADH被氧化的同时,乙酸也被代谢。
乙酸+2NADH+2H++ATP→乙醇+NAD++AMP+Pi
如上所述,对酵母赋予乙酸代谢能力的机理,需要破坏甘油途径。然而,已知GPD1基因和GPD2基因的重复破坏株发酵能力大幅降低,产业水平上的实用性低。另外,非专利文献5和专利文献2不是关于木糖同化酵母的报告,在木糖同化时是否有效果是不明的。
此外,还有对GPD1基因和GPD2基因的非破坏株导入mhpF基因的报告(非专利文献6)。然而,非专利文献6中虽然通过mhpF基因的导入而乙酸的生产量减少,但没有培养基中的乙酸减少这样的报告。进而,该非专利文献6也还不是关于木糖同化酵母的报告。
此外,有关于导入了木糖异构酶(XI)基因(来源于白蚁的肠内原生生物)的木糖同化酵母的报告(专利文献3)、以及对导入有XI基因(来源于Piromyces sp.E2)的木糖同化酵母进一步导入了乙醛脱氢酶基因(来源于青春双歧杆菌(Bifidobacteriumadolescentis0)的报告(专利文献4)。但是,这些文献中关于木糖同化时的乙酸同化几乎没有报告。
如上,现有技术中并未报告在同化木糖的同时进行乙醇发酵的条件下,使乙酸高效地代谢·分解的技术。
现有技术文献
专利文献
专利文献1:日本特开2009-195220号公报
专利文献2:WO 2011/010923
专利文献3:日本特开2011-147445号公报
专利文献4:日本特开2010-239925号公报
非专利文献
非专利文献1:FEMS Yeast Research,vol.9,2009,358-364
非专利文献2:Enzyme and Microbial Technology 33,2003,786-792
非专利文献3:Biotechnol.Bioeng.2009 103(3):500-512
非专利文献4:Biotechnol.Lett.2011 33:277-284
非专利文献5:Appl.Environ.Microbiol.2010 76:190-195
非专利文献6:Biotechnol.Lett.2011 33:1375-1380
发明内容
发明要解决的课题
因此,本发明鉴于上述事实,目的是特别地提供乙醇的制造方法,其中,使用了具有木糖代谢能力的酵母中的在木糖同化和乙醇发酵时能够代谢培养基中的乙酸而降低乙酸浓度的重组酵母。
用于解决课题的方法
为了实现上述目的,本发明者们进行了深入研究,结果发现,对具有木糖代谢能力的酵母导入特定的乙醛脱氢酶基因而得的重组酵母,在包含木糖的培养基中进行乙醇发酵时,能够代谢培养基中的乙酸,从而完成了本发明。
本发明包含以下内容。
(1)乙醇的制造方法,具有以下工序:将导入了木糖异构酶基因和乙醛脱氢酶基因的重组酵母,用含有木糖的培养基进行培养,从而进行乙醇发酵。
(2)根据(1)所述的乙醇的制造方法,其特征在于,所述木糖异构酶基因是编码以下的(a)或(b)的蛋白质的基因,
(a)具有序列号4所示的氨基酸序列的蛋白质,
(b)具有相对于序列号4所示的氨基酸序列有70%以上的同一性的氨基酸序列、且具有将木糖变为木酮糖的酶活性的蛋白质。
(3)根据(1)所述的乙醇的制造方法,其特征在于,所述乙醛脱氢酶基因编码来源于大肠杆菌的乙醛脱氢酶。
(4)根据(3)所述的乙醇的制造方法,其特征在于,所述来源于大肠杆菌的乙醛脱氢酶是以下的(a)或(b)的蛋白质,
(a)具有序列号2或20所示的氨基酸序列的蛋白质,
(b)具有相对于序列号2或20所示的氨基酸序列有70%以上的同一性的氨基酸序列、且具有乙醛脱氢酶活性的蛋白质。
(5)根据(1)所述的乙醇的制造方法,其特征在于,所述乙醛脱氢酶基因编码来源于拜氏梭菌(Clostridium beijerinckii)的乙醛脱氢酶。
(6)根据(5)所述的乙醇的制造方法,其特征在于,所述来源于拜氏梭菌(Clostridium beijerinckii)的乙醛脱氢酶是以下的(a)或(b)的蛋白质,
(a)具有序列号22所示的氨基酸序列的蛋白质,
(b)具有相对于序列号22所示的氨基酸序列有70%以上的同一性的氨基酸序列、且具有乙醛脱氢酶活性的蛋白质。
(7)根据(1)所述的乙醇的制造方法,其特征在于,所述乙醛脱氢酶基因编码来源于莱茵衣藻(Chlamydomonas reinhardtii)的乙醛脱氢酶。
(8)根据(5)所述的乙醇的制造方法,其特征在于,所述来源于莱茵衣藻(Chlamydomonas reinhardtii)的乙醛脱氢酶是以下的(a)或(b)的蛋白质,
(a)具有序列号24所示的氨基酸序列的蛋白质,
(b)具有相对于序列号24所示的氨基酸序列有70%以上的同一性的氨基酸序列、且具有乙醛脱氢酶活性的蛋白质。
(9)根据(1)所述的乙醇的制造方法,其特征在于,所述重组酵母是进一步导入了木酮糖激酶基因的重组酵母。
(10)根据(1)所述的乙醇的制造方法,其特征在于,所述重组酵母是导入了下述基因的重组酵母,所述基因编码选自构成戊糖磷酸途径中的非氧化过程的途径的酶群中的酶。
(11)根据(10)所述的乙醇的制造方法,其特征在于,构成戊糖磷酸途径中的非氧化过程的途径的酶群是核糖-5-磷酸异构酶、核酮糖-5-磷酸-3-差向异构酶、转酮醇酶和转醛醇酶。
(12)根据(1)所述的乙醇的制造方法,其特征在于,所述培养基含有纤维素,在所述乙醇发酵中,至少同时进行所述纤维素的糖化。
(13)根据(1)所述的乙醇的制造方法,其特征在于,所述重组酵母高表达具有将乙醛转换为乙醇的活性的醇脱氢酶基因。
(14)根据(1)所述的乙醇的制造方法,其特征在于,所述重组酵母是具有将乙醇转换为乙醛的活性的醇脱氢酶基因的表达量降低了的重组酵母。
本说明书包含作为本申请的优先权基础的日本专利申请2013-037501号和日本特愿2014-36652号的说明书和/或附图所记载的内容。。
发明的效果
在本发明的乙醇的制造方法中,可以降低培养基中的乙酸浓度,从而可以有效地避免由乙酸引起的发酵抑制。其结果是,本发明的乙醇的制造方法可以较高地维持以木糖作为糖源的乙醇发酵的效率,可以实现优异的乙醇收率。因此,本发明的乙醇的制造方法,例如,在再利用重组酵母时和/或在连续培养中使用时可以降低乙酸的带入量,维持优异的乙醇收率。
附图说明
图1是显示pUC-HIS3U-P_HOR7-XKS1-T_TDH3-P_TDH2-hph-T_CYC1-HIS3D的构成示意图。
图2是显示pUC-R67-HOR7p-RsXI-T_TDH3-TRP1d-R45的构成示意图。
图3是显示pUC-LEU2U-P_HOR7-TAL1-T_TDH3-P_HOR7-TKL1-T_TDH3-HIS3-LEU2D的构成示意图。
图4是显示pUC-GRE3U-P_HOR7-RPE1-T_TDH3-P_HOR7-RKI1-T_TDH3-LEU2-GRE3D的构成示意图。
图5是显示pCR-ADH2U-URA3-ADH2D的构成示意图。
图6是显示pCR-ADH2part-T_CYC1-P_TDH3-ADH1-T_ADH1-URA3-ADH2D的构成示意图。
图7pCR-ADH2part-T_CYC1-ERO1_T-mhpF-HOR7_P-URA3-ADH2D的构成示意图。
图8是显示pCR-ADH2part-T_CYC1-P_TDH3-ADH1-T_ADH1-ERO1_T-mhpF-HOR7_P-URA3-ADH2D的构成示意图。
图9是显示pCR-ADH2U-ERO1_T-mhpF-HOR7_P-URA3-ADH2D的构成示意图。
图10是显示pCR-ADH2U-P_TDH3-ADH1-T_ADH1-ERO1_T-mhpF-HOR7_P-URA3-ADH2D的构成示意图。
图11是显示pCR-ADH2part-T_CYC1-URA3-ADH2D的构成示意图。
具体实施方式
以下,使用附图和实施例更详细地说明本发明。
本发明的乙醇的制造方法是,使用具有木糖代谢能力、且导入了乙醛脱氢酶基因的重组酵母,由培养基所含的糖源合成乙醇的方法。在本发明的乙醇的制造方法中,具有能够通过上述重组酵母来代谢培养基所含的乙酸,随着乙醇发酵而培养基中的乙酸浓度降低这样的特征。
<重组酵母>
本发明的乙醇的制造方法中使用的重组酵母,是导入了木糖异构酶基因和乙醛脱氢酶基因的酵母,并且是具有木糖代谢能力的酵母。这里,具有木糖代谢能力的酵母,包含通过对本来不具有木糖代谢能力的酵母导入木糖异构酶基因从而被赋予了木糖代谢能力的酵母、通过对本来不具有木糖代谢能力的酵母导入木糖异构酶基因和其他木糖代谢相关基因从而被赋予了木糖代谢能力的酵母酵母、本来具有木糖代谢能力的酵母的任一者的含义。
具有木糖代谢能力的酵母可以同化培养基中所含的木糖而生产乙醇。此外,培养基中所含的木糖,既可以是通过将以木糖作为构成糖的木聚糖和/或半纤维素等进行糖化的工艺而得的,也可以是培养基所含的木聚糖和/或半纤维素等被糖化酶糖化而供给至培养基中的。后者的情况下,是指所谓同时糖化发酵的体系。
作为木糖异构酶基因(XI基因),不特别限定,可以使用来源于任何生物种的基因。例如,可以不特别限制地使用日本特开2011-147445号公报所公开的来源于白蚁的肠内原生生物的多种木糖异构酶基因。另外,作为木糖异构酶基因,还可以利用来源于作为厌氧性的霉的Piromyces sp.E2种(特表2005-514951号公报)、来源于作为厌氧性的霉的Cyllamyces aberensi、来源于作为细菌的多形拟杆菌(Bacteroides thetaiotaomicron)、来源于作为细菌的Clostridium phytofermentans、来源于鼠灰链霉菌(Streptomycesmurinus)簇的基因。
具体地,作为木糖异构酶基因,优选使用来源于黄胸散白蚁(Reticulitermessperatus)肠内原生生物的木糖异构酶基因。该来源于黄胸散白蚁(Reticulitermessperatus)肠内原生生物的木糖异构酶基因的编码区的碱基序列和该基因所编码的蛋白质的氨基酸序列分别示于序列号3和4。
但是,作为木糖异构酶基因,不限于序列号3和4所特定的,也可以是碱基序列和/或氨基酸序列不同但处于旁系同源物(paralog)的关系或狭义的同源物的关系的基因。
另外,木糖异构酶基因不限于这些序列号3和4所特定的,例如,还可以是编码具有相对于序列号4的氨基酸序列有70%以上、优选80%以上、更优选90%以上、最优选95%以上的序列类似性或同一性的氨基酸序列,且具有木糖异构酶活性的蛋白质的基因。序列类似性和同一性的值,可以通过安装了BLAST算法的BLASTN和/或BLASTX程序来算出(默认的设定)。此外,序列类似性的值,可以算出在将一对氨基酸序列两两比对分析时完全一致的氨基酸残基、和物理化学上功能类似的氨基酸残基的合计,作为比较的全氨基酸残基中的上述合计数的比例而算出。此外,同一性的值,可以算出在将一对氨基酸序列两两比对分析时完全一致的氨基酸残基,作为比较的全氨基酸残基中的上述氨基酸残基数的比例算出。
进而,木糖异构酶基因不限于这些序列号3和4所特定的,还可以是例如,编码具有相对于序列号4的氨基酸序列置换、缺失、插入或添加了1或数个氨基酸的氨基酸序列、且具有木糖异构酶活性的蛋白质的基因。这里,数个例如为2~30个,优选为2~20个,更优选为2~10个,最优选为2~5个。
进而另外,木糖异构酶基因不限于这些序列号3和4所特定的,也可以是例如,与包含序列号3的碱基序列的DNA的互补链的全部或一部分在严格条件下杂交、且编码具有木糖异构酶活性的蛋白质的基因。这里所说的“严格条件”是指形成所谓特异性杂种、不形成非特异性杂种的条件,可以参照例如Molecular Cloning:A Laboratory Manual(ThirdEdition)来适宜确定。具体地,可以根据DNA杂交(southern hybridization)时的温度和/或溶液所含的盐浓度、和DNA杂交(southern hybridization)的洗涤工序时的温度和/或溶液所含的盐浓度来设定严格度。更详细地,作为严格条件,例如钠浓度为25~500mM、优选为25~300mM,温度为42~68℃、优选为42~65℃。更具体地为5×SSC(83mM NaCl、83mM柠檬酸钠)、温度42℃。
如上所述,关于包含与序列号3不同的碱基序列的基因、或编码与序列号4不同的氨基酸序列的基因是否作为木糖异构酶基因发挥功能,只要制作将该基因插入适当的启动子和终止子等之间而得的表达载体,使用该表达载体转化例如大肠杆菌等宿主,测定表达的蛋白质的木糖异构酶活性即可。木糖异构酶活性是指将木糖异构化为木酮糖的活性。因此,木糖异构酶活性可以如下评价:作为底物准备包含木糖的溶液,使检查对象蛋白质在适当温度下作用,测定木糖的减少量和/或木酮糖的生成量。
特别是,作为木糖异构酶基因,优选使用编码包含对序列号4所示的氨基酸序列中的特定的氨基酸残基导入了特定的突变的氨基酸序列、且木糖异构酶活性提高了的突变型木糖异构酶的基因。具体地,作为编码突变型木糖异构酶的基因,可以列举编码序列号4所示的氨基酸序列中的第337位的天冬酰胺被替换为半胱氨酸的氨基酸序列的基因。包含序列号4所示的氨基酸序列中的第337位的天冬酰胺被替换为半胱氨酸的氨基酸序列的木糖异构酶,与野生型的木糖异构酶相比具有优异的木糖异构酶活性。此外,突变型木糖异构酶不限于将上述第337位的天冬酰胺替换为半胱氨酸的突变体,既可以是将上述第337位的天冬酰胺替换成了除了半胱氨酸以外的氨基酸的突变体,也可以是除了上述第337位的天冬酰胺之外进而将不同的氨基酸残基替换成了其他氨基酸的突变体,还可以是替换了除了所述第337位的天冬酰胺以外的其他的氨基酸残基的突变体。
另一方面,除了木糖异构酶基因以外的木糖代谢相关基因,是包含编码将木糖转换为木糖醇的木糖还原酶的木糖还原酶基因、编码将木糖醇转换为木酮糖的木糖醇脱氢酶的木糖醇脱氢酶基因和编码将木酮糖磷酸化而生成木酮糖5-磷酸的木酮糖激酶的木酮糖激酶基因的含义。此外,通过木酮糖激酶而生成的木酮糖5-磷酸进入戊糖磷酸途径而被代谢。
作为木糖代谢相关基因,不特别限定,可列举来源于树干毕赤酵母(Pichiastipitis)的木糖还原酶基因和木糖醇脱氢酶基因、来源于酿酒酵母(Saccharomycescerevisiae)的木酮糖激酶基因(参照Eliasson A.et al.,Appl.Environ.Microbiol,66:3381-3386和Toivari MN et al.,Metab.Eng.3:236-249)。此外,作为木糖还原酶基因,可利用来源于热带假丝酵母(Candida tropicalis)和/或近平滑假丝酵母(Candidaprapsilosis)的木糖还原酶基因。作为木糖醇脱氢酶基因,可利用来源于热带假丝酵母(Candida tropicalis)和/或近平滑假丝酵母(Candida prapsilosis)的木糖醇脱氢酶基因。作为木酮糖激酶基因,可利用来源于树干毕赤酵母(Pichia stipitis)的木酮糖激酶基因。
另外,作为本来具有木糖代谢能力的酵母,不特别限定,可列举树干毕赤酵母(Pichia stipitis)、热带假丝酵母(Candida tropicalis)和近平滑假丝酵母(Candidaprapsilosis)等。
另一方面,作为导入具有木糖代谢能力的酵母中的乙醛脱氢酶基因,不特别限定,可以使用来源于任何生物的基因。另外,在乙醛脱氢酶基因使用来源于除了酵母等真菌以外的生物,例如,细菌和/或动物、植物、昆虫、藻类的基因的情况下,优选使用根据导入的酵母中的密码子使用频率而改变了碱基序列的基因。
更具体地,作为乙醛脱氢酶基因,可以使用大肠杆菌中的mhpF基因,和/或如Applied and Environmental Microbiology,May 2004,p.2892-2897,Vol.70,No.5中所公开的那样使用溶组织内阿米巴(Entamoeba histolytica)中的ALDH1基因。这里,大肠杆菌中的mhpF基因的碱基序列和由mhpF基因所编码的蛋白质的氨基酸序列分别示于序列号1和2。
但是,作为乙醛脱氢酶基因,不限于序列号1和2所特定的,只要是EC编号1.2.1.10所定义的酶,就也可以是碱基序列和/或氨基酸序列不同但处于旁系同源物(paralog)的关系或狭义的同源物的关系的基因。作为乙醛脱氢酶基因,可列举例如,大肠杆菌中的adhE基因、来源于拜氏梭菌(Clostridium beijerinckii)的乙醛脱氢酶基因和来源于莱茵衣藻(Chlamydomonas reinhardtii)的乙醛脱氢酶基因。这里,大肠杆菌中的adhE基因的碱基序列和由adhE基因所编码的蛋白质的氨基酸序列分别示于序列号19和20。另外,来源于拜氏梭菌(Clostridium beijerinckii)的乙醛脱氢酶基因的碱基序列和该基因所编码的蛋白质的氨基酸序列分别示于序列号21和22。进而,来源于莱茵衣藻(Chlamydomonasreinhardtii)的乙醛脱氢酶基因乙醛脱氢酶基因的碱基序列和该基因所编码的蛋白质的氨基酸序列分别示于序列号23和24。
另外,乙醛脱氢酶基因不限于这些序列号1和2所特定的,只要是由EC编号1.2.1.10所定义的酶,也可以是碱基序列、氨基酸序列不同但处于旁系同源物(paralog)的关系或狭义的同源物的关系的基因。作为乙醛脱氢酶基因,可列举例如,大肠杆菌中的adhE基因、来源于拜氏梭菌(Clostridium beijerinckii)的乙醛脱氢酶基因和来源于莱茵衣藻(Chlamydomonas reinhardtii)的乙醛脱氢酶基因。这里,大肠杆菌中的adhE基因的碱基序列和adhE基因所编码的蛋白质的氨基酸序列分别示于序列号19和20。另外,来源于拜氏梭菌(Clostridium beijerinckii)的乙醛脱氢酶基因的碱基序列和该基因所编码的蛋白质的氨基酸序列分别示于序列号21和22。进而,来源于莱茵衣藻(Chlamydomonasreinhardtii)的乙醛脱氢酶基因的碱基序列和该基因所编码的蛋白质的氨基酸序列分别示于序列号23和24。
另外,乙醛脱氢酶基因不限于这些序列号1和2、19和20、21和22以及23和24所特定的,也可以是例如,编码具有相对于序列号2、20、22或24的氨基酸序列有70%以上、优选为80%以上、更优选为90%以上、最优选为95%以上的序列类似性或同一性的氨基酸序列、且具有乙醛脱氢酶活性的蛋白质的基因。序列类似性和同一性的值,可以通过安装了BLAST算法的BLASTN和/或BLASTX程序来算出(默认的设定)。此外,序列类似性的值,可以算出在将一对氨基酸序列两两比对分析时完全一致的氨基酸残基、和物理化学上功能类似的氨基酸残基的合计,作为比较的全氨基酸残基中的上述合计数的比例而算出。此外,同一性的值,可以算出在将一对氨基酸序列两两比对分析时完全一致的氨基酸残基,作为比较的全氨基酸残基中的上述氨基酸残基数的比例算出。
进而,乙醛脱氢酶基因不限于这些序列号1和2、19和20、21和22以及23和24所特定的,还可以是例如,编码具有相对于序列号2、20、22或24的氨基酸序列置换、缺失、插入或添加了1或数个氨基酸的氨基酸序列、且具有乙醛脱氢酶活性的蛋白质的基因。这里,数个例如为2~30个,优选为2~20个,更优选为2~10个,最优选为2~5个。
进而另外,乙醛脱氢酶基因不限于这些序列号1和2、19和20、21和22以及23和24所特定的,也可以是例如,与包含序列号1、19、21或23的碱基序列的DNA的互补链的全部或一部分在严格条件下杂交、且编码具有乙醛脱氢酶活性的蛋白质的基因。这里所说的“严格条件”是指形成所谓特异性杂种、不形成非特异性杂种的条件,可以参照例如MolecularCloning:A Laboratory Manual(Third Edition)来适宜确定。具体地,可以根据DNA杂交(southern hybridization)时的温度和/或溶液所含的盐浓度、和DNA杂交(southernhybridization)的洗涤工序时的温度和/或溶液所含的盐浓度来设定严格度。更详细地,作为严格条件,例如钠浓度为25~500mM、优选为25~300mM,温度为42~68℃、优选为42~65℃。更具体地为5×SSC(83mM NaCl、83mM柠檬酸钠)、温度42℃。
如上所述,关于包含与序列号1、19、21或23不同的碱基序列的基因、或编码与序列号2、20、22或24不同的氨基酸序列的基因是否作为乙醛脱氢酶基因发挥功能,只要制作将该基因插入适当的启动子和终止子等之间而得的表达载体,使用该表达载体转化例如大肠杆菌等宿主,测定表达的蛋白质的乙醛脱氢酶活性即可。乙醛脱氢酶活性可以如下测定:作为底物准备包含乙醛、辅酶A和NAD+的溶液,在适当的温度下与检查对象蛋白质作用,将生成的乙酰磷酸通过磷酸乙酰转移酶的作用而转换为乙酰磷酸,从而测定,或者将生成的NADH进行分光学测量而测定。
此外,本发明的乙醇的制造方法中使用的重组酵母也可以是具有木糖代谢能力、且至少导入了乙醛脱氢酶基因、进而导入了其他基因的酵母。作为其他基因不特别限定,例如,可以是导入了参与葡萄糖等的糖代谢的基因的酵母。作为一例,重组酵母可以通过导入β-葡糖苷酶基因而制成具有β-葡糖苷酶活性的酵母。
这里β-葡糖苷酶活性是指催化糖的β-糖苷键水解的反应的活性。即,β-葡糖苷酶可以将纤维二糖等纤维寡糖分解成葡萄糖。β-葡糖苷酶基因也可以作为细胞表层呈递型基因而导入。这里,细胞表层呈递型基因,是指进行了改变而使得由该基因所编码的蛋白质以在细胞的表层展示的方式表达的基因。例如,细胞表层呈递型β葡糖苷酶基因,是将β葡糖苷酶基因与细胞表层定位蛋白质基因融合而得的基因。细胞表层定位蛋白质是指固定于酵母的细胞表层、存在于细胞表层的蛋白质。可列举例如作为凝集性蛋白质的α-或a-凝集素、FLO蛋白质等。通常细胞表层定位蛋白质在N-末端侧具有分泌信序列号并在C-末端侧具有GPI锚附着识别信号。虽然在具有分泌信号方面与分泌性蛋白质相同,但细胞表层定位蛋白质在介由GPI锚而被固定输送到细胞膜上这方面与分泌性蛋白质不同。细胞表层定位蛋白质在通过细胞膜时,GPI锚附着识别序列信号被选择性地切断,在新突出的C末端部分与GPI锚结合从而被固定在细胞膜上。之后GPI锚的根部被磷脂酰肌醇依赖性磷脂酶C(PI-PLC)切断。接着,从细胞膜被切下的蛋白质插入细胞壁并被固定在细胞表层,从而定位于细胞表层(例如,参照日本特开2006-174767号公报)。
作为β-葡糖苷酶基因,不特别限定,可列举例如来自棘孢曲霉(Aspergillusaculeatus)的β-葡糖苷酶基因(Murai et al.,Appl.Environ.Microbiol.64:4857-4861)。此外,作为β-葡糖苷酶基因,还可利用来自米曲霉(Aspergillus oryzae)的β-葡糖苷酶基因、来自噬纤维梭菌(Clostridiumcellulovorans)的β-葡糖苷酶基因和来自扣囊复膜酵母(Saccharomycopsis fibligera)的β-葡糖苷酶基因等。
另外,本发明的乙醇的制造方法中使用的重组酵母,还可以是在β葡糖苷酶基因的基础之上,或者除了β葡糖苷酶基因以外,还导入了编码构成纤维素酶的其他酶的基因的重组酵母。作为除了β葡糖苷酶以外的构成纤维素酶的酶,可以列举从结晶纤维素的末端使纤维二糖游离的外切型的纤维二糖水解酶(CBH1和CBH2)、不能分解结晶纤维素但随机切断非结晶纤维素(无定形纤维素)链的内切型的内切葡聚糖酶(EG)。
另外,作为导入重组酵母的其他基因,可列举具有将乙醛转换为乙醇的活性的醇脱氢酶基因(ADH1基因)、具有将乙酸转换为乙酰辅酶A的活性的乙酰辅酶A合成酶基因(ACS1基因)、具有将乙醛转换为乙酸的活性的基因(ALD4基因、ALD5基因和ALD6基因)。此外,还可以破坏具有将乙醇转换为乙醛的活性醇脱氢酶基因(ADH2基因)。
进而,在本发明的乙醇的制造方法中使用的重组酵母,优选为具有以下特征的重组酵母,所述特征是高表达具有将乙醛转换为乙醇的活性的醇脱氢酶基因(ADH1基因)。为了高表达该基因,可列举将内在的该基因的启动子置换为高表达用启动子、将能表达地具有该基因的表达载体导入酵母这样的方法。
酿酒酵母(Saccharomyces cerevisiae)的ADH1基因的碱基序列和由该基因所编码的蛋白质的氨基酸序列分别示于序列号5和6。但是,作为高表达对象的醇脱氢酶基因,不限于由序列号5和6所特定的基因,也可以是碱基序列和/或氨基酸序列不同但处于旁系同源物(paralog)的关系或狭义的同源物的关系的基因。
另外,醇脱氢酶基因不限于由这些序列号5和6所特定的,也可以是例如,编码具有相对于序列号6的氨基酸序列有70%以上、优选为80%以上、更优选为90%以上、最优选为95%以上的序列类似性或同一性的氨基酸序列、且具有醇脱氢酶活性的蛋白质的基因。序列类似性和同一性的值,可以通过安装了BLAST算法的BLASTN和/或BLASTX程序来算出(默认的设定)。此外,序列类似性的值,可以算出在将一对氨基酸序列两两比对分析时完全一致的氨基酸残基、和物理化学上功能类似的氨基酸残基的合计,作为比较的全氨基酸残基中的上述合计数的比例而算出。此外,同一性的值,可以算出在将一对氨基酸序列两两比对分析时完全一致的氨基酸残基,作为比较的全氨基酸残基中的上述氨基酸残基数的比例算出。
进而,醇脱氢酶基因不限于由这些序列号5和6所特定的,也还可以是例如,编码具有相对于序列号6的氨基酸序列的氨基酸序列置换、缺失、插入或添加了1或数个氨基酸的氨基酸序列、且具有醇脱氢酶的蛋白质的基因。这里,数个例如为2~30个,优选为2~20个,更优选为2~10个,最优选为2~5个。
进而另外,醇脱氢酶基因不限于由这些序列号5和6所特定的,也可以是例如,与包含序列号5的碱基序列的DNA的互补链的全部或一部分在严格条件下杂交、且编码具有醇脱氢酶活性的蛋白质的基因。这里所说的“严格条件”是指形成所谓特异性杂种、不形成非特异性杂种的条件,可以参照例如Molecular Cloning:A Laboratory Manual(ThirdEdition)来适宜确定。具体地,可以根据DNA杂交(southern hybridization)时的温度和/或溶液所含的盐浓度、和DNA杂交(southern hybridization)的洗涤工序时的温度和/或溶液所含的盐浓度来设定严格度。更详细地,作为严格条件,例如钠浓度为25~500mM、优选为25~300mM,温度为42~68℃、优选为42~65℃。更具体地为5×SSC(83mM NaCl、83mM柠檬酸钠)、温度42℃。
如上所述,关于包含与序列号5不同的碱基序列的基因、或编码与序列号6不同的氨基酸序列的基因是否作为具有将乙醛转换为乙醇的活性的醇脱氢酶基因发挥功能,只要制作将该基因插入适当的启动子和终止子等之间而得的表达载体,使用该表达载体转化例如酵母等宿主,测定表达的蛋白质的醇脱氢酶活性即可。将乙醛转换为乙醇的醇脱氢酶活性可以如下测定:作为底物准备包含醛和NADH或NADPH的溶液,在适当的温度下与检查对象蛋白质作用,测量生成的醇,或者分光测量NAD+或NADP+,从而测定。
进而,本发明的乙醇的制造方法中使用的重组酵母优选具有如下特征,所述特征是具有将乙醇转换为醛的活性的醇脱氢酶基因(ADH2基因)的表达量下降了。为了使该基因的表达量下降,可列举改变内在的该基因的启动子、使该基因缺失这样的方法。使该基因缺失时,既可以使二倍体的重组酵母中存在的一对ADH2基因中的一方缺失,也可以使两方缺失。作为抑制基因表达的方法,可列举所谓转座子法、转基因法、转录后基因沉默法、RNAi法、无义介导的衰减(Nonsense mediated decay,NMD)法、核酶法、反义法、miRNA(micro-RNA,微RNA)法、siRNA(small interfering RNA,小干扰RNA)法等。
酿酒酵母(Saccharomyces cerevisiae)的ADH2基因的碱基序列和由该基因所编码的蛋白质的氨基酸序列分别示于序列号7和8。但是,作为对象的醇脱氢酶基因,不限于序列号7和8所特定的,也可以是碱基序列和/或氨基酸序列不同但处于旁系同源物(paralog)的关系或狭义的同源物的关系的基因。
另外,醇脱氢酶基因不限于由这些序列号7和8所特定的,也可以是例如,编码具有相对于序列号8的氨基酸序列有70%以上、优选为80%以上、更优选为90%以上、最优选为95%以上的序列类似性或同一性的氨基酸序列、且具有醇脱氢酶活性的蛋白质的基因。序列类似性和同一性的值,可以通过安装了BLAST算法的BLASTN和/或BLASTX程序来算出(默认的设定)。此外,序列类似性的值,可以算出在将一对氨基酸序列两两比对分析时完全一致的氨基酸残基、和物理化学上功能类似的氨基酸残基的合计,作为比较的全氨基酸残基中的上述合计数的比例而算出。此外,同一性的值,可以算出在将一对氨基酸序列两两比对分析时完全一致的氨基酸残基,作为比较的全氨基酸残基中的上述氨基酸残基数的比例算出。
进而,醇脱氢酶基因不限于由这些序列号7和8所特定的,也还可以是例如,编码具有相对于序列号8的氨基酸序列置换、缺失、插入或添加了1或数个氨基酸的氨基酸序列、且具有醇脱氢酶活性的蛋白质的基因。这里,数个例如为2~30个,优选为2~20个,更优选为2~10个,最优选为2~5个。
进而另外,醇脱氢酶基因不限于这些序列号7和8所特定的,也可以是例如,与包含序列号7的碱基序列的DNA的互补链的全部或一部分在严格条件下杂交、且编码具有醇脱氢酶活性的蛋白质的基因。这里所说的“严格条件”是指形成所谓特异性杂种、不形成非特异性杂种的条件,可以参照例如Molecular Cloning:A Laboratory Manual(Third Edition)来适宜确定。具体地,可以根据DNA杂交(southern hybridization)时的温度和/或溶液所含的盐浓度、和DNA杂交(southern hybridization)的洗涤工序时的温度和/或溶液所含的盐浓度来设定严格度。更详细地,作为严格条件,例如钠浓度为25~500mM、优选为25~300mM,温度为42~68℃、优选为42~65℃。更具体地为5×SSC(83mM NaCl、83mM柠檬酸钠)、温度42℃。
如上所述,包含与序列号7不同的碱基序列的基因、或编码与序列号8不同的氨基酸序列的基因是否作为具有将乙醇转换为醛的活性的醇脱氢酶基因发挥功能,只要制作将该基因插入适当的启动子和终止子等之间而得的表达载体,使用该表达载体转化例如酵母等宿主,测定表达的蛋白质的醇脱氢酶活性即可。将乙醇转换为醛的醇脱氢酶活性可以如下测定:作为底物准备包含醇和NAD+或NADP+的溶液,在适当的温度下与检查对象蛋白质作用,测量生成的醛,或者分光测量NADH或NADPH,从而测定。
进而,作为导入重组酵母中的其他基因,可列举参与构成生物质的半纤维素所含的5碳糖L-阿拉伯糖的代谢途径的基因。作为这样的基因,可列举例如,来源于原核生物的L-阿拉伯糖异构酶基因、L-核酮糖激酶基因、L-核酮糖-5-磷酸-4-差向异构酶基因、和/或来源于真核生物的L-阿糖醇-4-脱氢酶基因、L-木糖还原酶基因。
特别是作为导入重组酵母中的其他基因,可列举能够促进培养基中的木糖的利用的基因。具体地,可列举编码具有以木酮糖作为底物生成木酮糖-5-磷酸的活性的木酮糖激酶的基因。通过导入木酮糖激酶基因,能够提高戊糖磷酸途径的代谢通量。
进而重组酵母中可以导入编码选自构成戊糖磷酸途径中的非氧化过程的途径的酶群中的酶的基因。作为构成戊糖磷酸途径中的非氧化过程的途径的酶,可列举核糖-5-磷酸异构酶、核酮糖-5-磷酸-3-差向异构酶、转酮醇酶和转醛醇酶。优选导入1种以上编码这些酶的基因。另外,更优选将这些基因中的2种以上组合导入,进一步优选3种以上组合导入,最优选导入全种类的基因。
更具体地作为木酮糖激酶(XK)基因,可以不特别限制来源生物地使用。此外XK基因由同化木酮糖的细菌、酵母等大量微生物保持。关于XK基因的信息可以通过NCBI的HP等的检索来适宜获得。优选列举来源于酵母、乳酸菌、大肠杆菌、植物等的XK基因。作为XK基因,可列举例如,来源于酿酒酵母(S.cerevisiae)S288C株的XK基因XKS1(GenBank:Z72979)(CDS的编码区的碱基序列和氨基酸序列)。
另外,更具体地,转醛醇酶(TAL)基因、转酮醇酶(TKL)基因、核酮糖-5-磷酸差向异构酶(RPE)基因、核糖-5-磷酸酮异构酶(RKI)基因可以不特别限定来源生物地使用。这些基因在具有戊糖磷酸途径的大量生物中保持。例如,酿酒酵母(S.cerevisiae)等通用酵母也保持这些基因。关于这些基因的信息可以通过登录NCBI等的HP来适宜获得。优选列举来源于真核细胞或酵母等与宿主真核细胞同一属、更优选与宿主真核细胞同一种的各基因。作为TAL基因优选使用TAL1基因,作为TKL基因优选使用TKL1基因和TKL2基因,作为RPE基因优选使用RPE1基因,作为RKI基因优选使用RKI1基因。例如,作为这些基因,可列举作为来源于酿酒酵母(S.cerevisiae)S288株的TAL1基因的TAL1基因(GenBank:U19102)、来源于酿酒酵母(S.cerevisiae)S288株的TKL1基因(GenBank:X73224)、来源于酿酒酵母(S.cerevisiae)S288株的RPE1基因(Genbank:X83571)、来源于酿酒酵母(S.cerevisiae)S288株的RKI1基因(GenBank:Z75003)。
<重组酵母的制作>
可以通过将上述的木糖异构酶基因和乙醛脱氢酶基因导入成为宿主的酵母基因组中,来制作能够在本发明中使用的重组酵母。这里,木糖异构酶基因和乙醛脱氢酶基因既可以导入不具有木糖代谢能力的酵母中,也可以导入本来具有木糖代谢能力的酵母中,还可以与木糖代谢相关基因一起导入不具有木糖代谢能力的酵母中。另外,将木糖异构酶基因、乙醛脱氢酶基因和上述的基因导入酵母时,既可以将全部基因同时导入,也可以利用不同的表达载体逐次导入。
作为能用作宿主的酵母,不特别限定,可列举休哈塔假丝酵母(CandidaShehatae)、树干毕赤酵母(Pichia stipitis)、嗜鞣管囊酵母(Pachysolen tannophilus)、酿酒酵母(Saccharomyces cerevisiae)和粟酒裂殖酵母(Schizosaccaromyces pombe)等酵母,特别优选为酿酒酵母(Saccharomyces cerevisiae)。此外,作为酵母,既可以是为了实验方面的方便性而使用的实验株,也可以是为了实用方面的有用性而使用的工业株(实用株)。作为工业株,可列举例如用于葡萄酒、清酒和/或烧酎制作的酵母株。
此外,作为成为宿主的酵母,优选使用具有同宗配合性的酵母。根据日本特开2009-34036号公报所公开的方法,通过利用具有同宗配合性的酵母,可以简便地向基因组中导入多拷贝基因。具有同宗配合性的酵母与同宗配合酵母为同义。作为具有同宗配合性的酵母,不特别限定,也可以使用任何酵母。作为具有同宗配合性的酵母,可列举酿酒酵母OC-2株(NBRC2260),但不限于此。此外,作为具有同宗配合性的酵母,可列举醇酵母(台研396号、NBRC0216)(出处:“アルコール酵母的諸特性(醇酵母的诸特性)”酒研会报、No37、p18-22(1998.8))、在巴西和冲绳分离出的乙醇生产酵母(出处:“ブラジルと沖縄で分離したSaccharomyces cerevisiae野生株的遺伝学的性質(在巴西和冲绳分离的酿酒酵母野生株的遗传学性质)”日本农艺化学会志、Vol.65、No.4、p759-762(1991.4))和180(出处:“アルコール発酵力の強い酵母的スクリーニング(醇发酵能力强的酵母的筛选)”日本酿造协会志、Vol.82、No.6、p439-443(1987.6))等。另外,即使是显示异宗配合的表型的酵母,也可通过以可表达的方式导入HO基因而制成具有同宗配合性的酵母来使用。因此,在本发明中,具有同宗配合性的酵母是也包含以可表达的方式导入了HO基因的酵母的含义。
其中,酿酒酵母(Saccharomyces cerevisiae)OC-2株是一直以来在葡萄酒酿酒的现场被利用的被确认了安全性的菌株,因而优选。此外酿酒酵母(Saccharomycescerevisiae)OC-2株如后述实施例所述的那样,是在高糖浓度的条件下启动子活性优异的菌株,因而优选。特别是酿酒酵母(Saccharomyces cerevisiae)OC-2株在高糖浓度条件下丙酮酸脱羧酶基因(PDC1)的启动子活性优异,因而优选。
另外,作为导入的基因的启动子,不特别限定,可利用例如,甘油醛-3-磷酸脱氢酶基因(TDH3)的启动子、3-磷酸甘油酸激酶基因(PGK1)的启动子、高渗透压应答7基因(HOR7)的启动子等。其中,丙酮酸脱羧酶基因(PDC1)的启动子高表达下游的目的基因的能力高,因而优选。
即,上述基因可以与调控表达的启动子和/或其他表达调控区一起导入酵母的基因组中。或者,上述基因也可以以通过成为宿主的酵母的基因组中本来存在基因的启动子和/或其他表达调控区而进行表达调控的方式导入。
作为导入上述基因的方法,可以应用已知作为酵母的转化方法的现有公知的任何方法。具体来说,例如,可以以例如电穿孔法“Meth.Enzym.,194,p182(1990)”、原生质球法“Proc.Natl.Acad.Sci.USA,75p1929(1978)”、乙酸锂法“J.Bacteriology,153,p163(1983)”、Proc.Natl.Acad.Sci.USA,75p1929(1978)、Methods in yeast genetics,2000Edition:A Cold Spring Harbor Laboratory Course Manual等记载的方法实施,但不限于此。
<乙醇制造>
在使用以上所说明的重组酵母制造乙醇时,用至少含有木糖的培养基进行乙醇发酵培养。即,进行乙醇发酵的培养基,作为碳源至少含有木糖。此外,培养基中也可以预先含有葡萄糖等其他碳源。
另外,在乙醇发酵中利用的培养基所含的木糖可以来源于生物质。换句话说,在乙醇发酵中利用的培养基也可以是包含纤维素系生物质、糖化纤维素系生物质所含的半纤维素而生成木糖的半纤维素酶的组成。这里,作为纤维素系生物质,可以是实施了现有公知的前处理的物质。作为前处理,不特别限定,可列举例如,将木质素用微生物分解的处理、和/或纤维素系生物质的粉碎处理等。另外,作为前处理,也可以应用例如,将粉碎了的纤维素系生物质浸渍于稀硫酸溶液和/或碱溶液、离子液体中的处理、水热处理、微粉碎处理这样的处理。通过这些前处理,可以提高生物质的糖化率。
此外,使用以上所说明的重组酵母制造乙醇时,上述培养基还可以是进一步包含纤维素和纤维素酶的组成。此时,上述培养基中含有纤维素酶作用于纤维素而生成的葡萄糖。在乙醇发酵中利用的培养基含有纤维素的情况下,该纤维素可以来源于生物质。换句话说,在乙醇发酵中利用的培养基还可以是包含能够糖化纤维素系生物质所含的纤维素的纤维素酶的组成。
另外,在乙醇发酵中利用的培养基,还可以添加将纤维素系生物质糖化处理后的糖化液。此时,糖化液包含残存的纤维素和/或纤维素酶、以及来源于纤维素系生物质所含的半纤维素的木糖。
如上,本发明的乙醇的制造方法包含至少以木糖作为糖源的乙醇发酵的工序。本发明的乙醇的制造方法能够通过以木糖作为糖源的乙醇发酵来制造乙醇。在本发明的利用了重组酵母的乙醇的制造方法中,在乙醇发酵之后从培养基中回收乙醇。对乙醇的回收方法没有特别限制,可采用现有公知的任何方法。例如,在上述乙醇发酵结束后,通过固液分离操作将含有乙醇的液层、与含有重组酵母和/或固体成分的固层分离。然后将液层中所含的乙醇通过蒸馏法进行分离·纯化,从而可以回收纯度高的乙醇。此外,乙醇的纯化度可以根据乙醇的使用目的而适宜调整。
一般地,在利用来源于生物质的糖来制造乙醇的情况下,在上述的前处理和/或糖化处理中有时产生乙酸和/或糠醛这样的发酵抑制物质。特别地关于乙酸,已知其抑制酵母的生长·增殖,使以木糖作为糖源的乙醇发酵的效率下降。
然而,在本发明中,由于使用导入了木糖异构酶基因和乙醛脱氢酶基因的重组酵母,因而可以代谢培养基所含的乙酸,从而可以将培养基所含的乙酸浓度抑制得较低。因此,本发明的乙醇的制造方法,与使用了未导入木糖异构酶基因和乙醛脱氢酶基因的酵母时相比,可以实现优异的乙醇收率。
另外,根据本发明的乙醇的制造方法,由于即使在将重组酵母培养规定时间之后培养基中的乙酸浓度也低,因而,即使在利用培养规定时间之后的培养基的一部分重新开始培养的连续培养体系中使用,也能够降低乙酸的带入量。另外,根据本发明的乙醇的制造方法,即使在乙醇发酵工序结束之后回收菌体而再利用的情况下,也由于同样的理由而能够降低乙酸的带入量。
另外,本发明的乙醇的制造方法,可以成为将培养基所含的纤维素用纤维素酶进行糖化的工序、与以木糖和糖化所生成的葡萄糖作为糖源的乙醇发酵的工序同时进行的所谓同时糖化发酵处理。这里,同时糖化发酵处理是指将糖化纤维素系生物质的工序与乙醇发酵工序不区分地同时实施的处理。
此外,作为糖化方法没有特别限制,可列举利用纤维素酶和/或半纤维素酶等纤维素酶制剂的酶法等。纤维素酶制剂含有参与纤维素链和半纤维素链的分解的多种酶,显示内切葡聚糖酶活性、内切木聚糖酶活性、纤维二糖水解酶活性、葡糖苷酶活性和木糖苷酶活性等多种活性。作为纤维素酶制剂,没有特别限制,可列举例如,由里氏木霉(Trichodermareesei)、和/或解纤维顶孢霉(Acremonium cellulolyticus)等所生产的纤维素酶。作为纤维素酶制剂,也可以使用市售品。
在同时糖化发酵处理中,向包含纤维素系生物质(也可以在前处理后)的培养基中加入纤维素酶制剂和上述重组微生物,在规定的温度范围内培养该重组酵母。作为培养温度没有特别限制,考虑乙醇发酵的效率可以为25~45℃,优选为30~40℃。此外培养液的pH值优选为4~6。此外,培养时也可以进行搅拌和/或振荡。进而,也可以是先在酶的最适温度(40~70℃)下进行糖化,然后,使温度下降至规定的温度(30~40℃)再添加酵母这样的不规则的同时糖化发酵。
实施例
以下使用实施例更详细地说明本发明,但本发明的技术范围并不限制于以下实施例。
〔实施例1〕
在本实施例中,制作导入了木糖异构酶基因和大肠杆菌的乙醛脱氢酶基因(mhpF基因)的重组酵母,评价该重组酵母的乙酸代谢能力。
<导入载体的制作>
(1)XKS1基因导入用载体
作为来源于酿酒酵母(S.cerevisiae)的木酮糖激酶(XK)基因的酵母导入用载体,制作图1所示的pUC-HIS3U-P_HOR7-XKS1-T_TDH3-P_TDH2-hph-T_CYC1-HIS3D。该载体包含:5’侧添加了HOR7启动子和3’侧添加了TDH3终止子的作为来源于酿酒酵母(S.cerevisiae)NBRC304株的XK基因的XKS1基因(genebank:X61377)、成为向酵母基因组上的同源重组区域的组氨酸合成酶(HIS3)基因的上游约500bp的区域(HIS3U)以及该基因内的约500bp的区域(HIS3D)、以及5’侧添加了TDH2启动子和3’侧添加了CYC1终止子的潮霉素磷酸转移酶(hph)基因(标志物基因)。此外,同源重组区域的外侧导入了限制性酶Sse8387I的位点。另外,来源于酿酒酵母(S.cerevisiae)NBRC304株的XKS1基因的编码区的碱基序列和该基因所编码的木酮糖激酶的氨基酸序列分别示于序列号9和10。
(2)XI基因导入用载体
作为来源于黄胸散白蚁(Reticulitermes speratus)肠内原生生物的木糖异构酶(RsXI-C1、参照日本特开2011-147445)基因的酵母导入用载体,制作图2所示的pUC-R67-HOR7p-RsXI-T_TDH3-TRP1d-R45。该载体包含:5’侧添加了HOR7启动子和3’侧添加了TDH3终止子的RsXI-C1基因、成为向酵母基因组上的同源重组区域的rRNA基因(rDNA)的同源序列R45和R67、以及缺失启动子部分而降低了表达量的TRP1d标志物基因。此外,同源重组区域的外侧导入了限制性酶Sse8387I的位点。通过R45和R67,包含RsXI-C1的基因被多拷贝导入到第12号染色体上的rDNA座。进而TRP1d标志物在以多拷贝被导入染色体上的情况下才作为标志物发挥功能。因此,通过利用本载体,多拷贝导入成为可能。此外,在本实施例中RsXI-C1基因,使用将全区域设计成根据酵母的密码子使用频率而转换了使用的密码子的碱基序列,基于该碱基序列全合成而得的基因。本实施例中设计的RsXI-C1基因的碱基序列和该基因所编码的木糖异构酶的氨基酸序列分别示于序列号3和4。
(3)TAL1·TKL1基因导入用载体
作为来源于酿酒酵母(S.cerevisiae)的转醛醇酶1(TAL1)基因和转酮醇酶1(TKL1)基因的酵母导入用载体,制作图3所示的pUC-LEU2U-P_HOR7-TAL1-T_TDH3-P_HOR7-TKL1-T_TDH3-HIS3-LEU2D。该载体包含:5’侧添加了HOR7启动子和3’侧添加了TDH3终止子的作为来源于酿酒酵母(S.cerevisiae)S288株的TAL1基因的TAL1基因(genebank:U19102);5’侧添加了HOR7启动子和3’侧添加了TDH3终止子的作为来源于酿酒酵母(S.cerevisiae)S288株的TKL1基因的TKL1基因(genebank:X73224);成为向酵母基因组上的同源重组区域的亮氨酸合成酶(LEU2)基因从3’侧末端到上游约500bp的区域(LEU2U)以及该基因5’侧末端上游的约450bp的区域(LEU2D);以及组氨酸合成酶(HIS3)基因(标志物基因)。此外,同源重组区域的外侧导入了限制性酶Sse8387I的位点。另外,来源于酿酒酵母(S.cerevisiae)S288株的TAL1基因的编码区的碱基序列和该基因所编码的转醛醇酶1的氨基酸序列分别示于序列号11和12。进而来源于酿酒酵母(S.cerevisiae)S288株的TKL1基因的编码区的碱基序列和该基因所编码的转酮醇酶1的氨基酸序列分别示于序列号13和14。
(4)RPE1·RKI1基因导入、GRE3基因破坏用载体
作为来源于酿酒酵母(S.cerevisiae)的核酮糖磷酸差向异构酶1(RPE1)基因和核糖磷酸酮异构酶(RKI1)基因的酵母导入用载体,制作了图4所示的pUC-GRE3U-P_HOR7-RPE1-T_TDH3-P_HOR7-RKI1-T_TDH3-LEU2-GRE3D。该载体包含:5’侧添加了HOR7启动子和3’侧添加了TDH3终止子的作为来源于酿酒酵母(S.cerevisiae)S288株的RPE1基因的RPE1基因(genebank:X83571);5’侧添加了HOR7启动子和3’侧添加了TDH3终止子的来源于酿酒酵母(S.cerevisiae)S288株的RKI1基因(genebank:Z75003);成为向酵母基因组上的同源重组和用于破坏醛糖还原酶3(GRE3)基因的区域的包含GRE3基因的3’末端区域约500bp的约800bp的区域(GRE3U)和GRE3基因的上游约1000bp的区域(GRE3D);以及亮氨酸合成酶(LEU2)基因(标志物基因)。此外,同源重组区域的外侧导入了限制性酶Sse8387I的位点。另外,来源于酿酒酵母(S.cerevisiae)S288株的RPE1基因的编码区的碱基序列和该基因所编码的核酮糖磷酸差向异构酶1的氨基酸序列分别示于序列号15和16。进而,来源于酿酒酵母(S.cerevisiae)S288株的RKI1基因的编码区的碱基序列和该基因所编码的核糖磷酸酮异构酶的氨基酸序列分别示于序列号17和18。
(5)ADH2基因破坏用载体
作为宿主中内在的ADH2基因的破坏用载体,制作了图5所示的pCR-ADH2U-URA3-ADH2D。该载体中,作为向酵母基因组上的同源重组和用于破坏醇脱氢酶2(ADH2)基因的区域,包含ADH2基因的上游约700bp的区域(ADH2U)、ADH2基因的下流约800bp的区域(ADH2D)、以及乳清苷-5’-磷酸脱羧酶(URA3)基因(标志物基因)。
(6)ADH1基因导入用载体
作为醇脱氢酶1(ADH1)基因的酵母导入用载体,制作了图6所示的pCR-ADH2part-T_CYC1-P_TDH3-ADH1-T_ADH1-URA3-ADH2D。该载体包含:5’侧添加了TDH3启动子和3’侧添加了ADH1终止子的来源于酿酒酵母(S.cerevisiae)S288株的ADH1基因(genebank:Z74828.1)、成为向酵母基因组上的同源重组区域的ADH2基因的从3’侧末端到上游约450bp的区域(ADH2part)和从3’侧末端到下流的约700bp的区域(ADH2D)、作为ADH2的终止子的CYC1终止子、以及URA3基因(标志物基因)。
(7)mhpF基因导入用载体
作为来源于大肠杆菌(E.coli)的乙醛脱氢酶(mhpF)基因的酵母导入用载体,制作了图7所示的pCR-ADH2part-T_CYC1-ERO1_T-mhpF-HOR7_P-URA3-ADH2D。该载体包含:5’侧添加了HOR7启动子和3’侧添加了ERO1终止子的来源于大肠杆菌(E.coli)的乙醛脱氢酶基因(mhpF基因)、成为向酵母基因组上的同源重组区域的ADH2基因的从3’侧末端到上游约450bp的区域(ADH2part)和从3’侧末端到下流的约700bp的区域(ADH2D)、作为ADH2的终止子的CYC1终止子、以及包含URA3基因的基因(标志物基因)。此外,本实施例中,mhpF基因使用将全区域设计成根据酵母的密码子使用频率而转换了使用的密码子的碱基序列,基于该碱基序列全合成而得的基因。本实施例中设计的mhpF基因的碱基序列和该基因所编码的乙醛脱氢酶的氨基酸序列分别示于序列号1和2。
(8)mhpF·ADH1基因导入用载体
作为mhpF基因和ADH1基因的酵母导入用载体,制作了图8所示的pCR-ADH2part-T_CYC1-P_TDH3-ADH1-T_ADH1-ERO1_T-mhpF-HOR7_P-URA3-ADH2D。该载体包含:5’侧添加了HOR7启动子和3’侧添加了ERO1终止子的mhpF基因(与上述(7)相同)、5’侧添加了TDH3启动子和3’侧添加了ADH1终止子的来源于酿酒酵母(S.cerevisiae)S288株的ADH1基因(与上述(6)相同)、成为向酵母基因组上的同源重组区域的ADH2基因的从3’侧末端到上游约450bp的区域(ADH2part)、和从3’侧末端到下流的约700bp的区域(ADH2D)、作为ADH2的终止子的CYC1终止子、以及URA3基因(标志物基因)。
(9)mhpF基因导入、ADH2破坏用载体
作为mhpF基因的酵母导入和ADH2基因破坏用载体,制作了图9所示的pCR-ADH2U-ERO1_T-mhpF-HOR7_P-URA3-ADH2D。该载体包含:5’侧添加了HOR7启动子和3’侧添加了ERO1终止子的mhpF基因(与上述(7)相同)、成为向酵母基因组上的同源重组和用于破坏ADH2基因的区域的ADH2基因的上游约700bp的区域(ADH2U)和ADH2基因的上游约800bp的区域(ADH2D)、以及URA3基因(标志物基因)。
(10)mhpF基因和ADH1基因导入、ADH2破坏用载体
作为mhpF基因和ADH1基因的酵母导入以及ADH2基因破坏用载体,制作了图10所示的pCR-ADH2U-P_TDH3-ADH1-T_ADH1-ERO1_T-mhpF-HOR7_P-URA3-ADH2D。该载体包含:5’侧添加了HOR7启动子和3’侧添加了ERO1终止子的mhpF基因(与上述(7)相同)、5’侧添加了TDH3启动子和3’侧添加了ADH1终止子的来源于酿酒酵母(S.cerevisiae)S288株的ADH1基因(与上述(6)相同)、成为向酵母基因组上的同源重组和用于破坏ADH2基因的区域的ADH2基因的上游约700bp的区域(ADH2U)和ADH2基因的上游约800bp的区域(ADH2D)、以及URA3基因(标志物基因)。
(11)对照载体(仅标志物基因)
作为仅导入标志物基因的对照载体,制作了图11所示的pCR-ADH2part-T_CYC1-URA3-ADH2D。该载体包含:成为向酵母基因组上的同源重组区域的ADH2基因的从3’侧末端到上游约450bp的区域(ADH2part)和从3’侧末端到下流的约700bp的区域(ADH2D)、作为ADH2的终止子的CYC1终止子、以及URA3基因(标志物基因)。
<载体导入酵母株的制作>
用添加了5-氟乳清酸的培养基筛选作为2倍体酵母的酿酒酵母(Saccharomycescerevisiae)OC2-T株(Saitoh,S.等,J.Ferment.Bioeng.1996年、81卷98-103)(Boeke,J.D.,et al.1987Methods Enzymol.;154:164-75.),以成为尿嘧啶营养缺陷型的菌株作为宿主。
酵母的转化使用Frozen-EZ Yeast Transformation II(ZYMO RESEARCH),按照附带的实验步骤进行。首先,使用将pUC-HIS3U-P_HOR7-XKS1-T_TDH3-P_TDH2-hph-T_CYC1-HIS3D载体用限制性酶Sse8387I消化而得的片段,进行OC2-T株的转化,涂布于YPD+HYG琼脂培养基,将生长的集落纯化。将纯化后的选择株命名为OC100株。接下来,使用将pUC-LEU2U-P_HOR7-TAL1-T_TDH3-P_HOR7-TKL1-T_TDH3-HIS3-LEU2D载体用限制性酶Sse8387I消化而得的片段,进行OC100株的转化,涂布于不含组氨酸的SD琼脂培养基(Mthods in YeastGenetics,Cold Spring Harbor Laboratory Press),将生长的集落纯化。将纯化后的选择株命名为OC300株。接下来,使用将pUC-GRE3U-P_HOR7-RPE1-T_TDH3-P_HOR7-RKI1-T_TDH3-LEU2-GRE3D载体用限制性酶Sse8387I消化而得的片段,进行OC300株的转化,涂布于不含亮氨酸的SD琼脂培养基,将生长的集落纯化。将纯化后的选择株命名为OC600株。接下来,使用将pUC-R67-HOR7p-RsXI-T_TDH3-TRP1d-R45载体用限制性酶Sse8387I消化而得的片段,进行OC600株的转化,涂布于不含色氨酸的SD琼脂培养基,将生长的集落纯化。将纯化后的选择株命名为OC700株。如上制作的OC700株中导入有RsXI-C1基因、XK基因、TAL1基因、TKL1基因、RPE1基因和RKI1基因。
接下来,使用将pCR-ADH2U-URA3-ADH2D、pCR-ADH2part-T_CYC1-P_TDH3-ADH1-T_ADH1-URA3-ADH2D、pCR-ADH2part-T_CYC1-ERO1_T-mhpF-HOR7_P-URA3-ADH2D、pCR-ADH2part-T_CYC1-P_TDH3-ADH1-T_ADH1-ERO1_T-mhpF-HOR7_P-URA3-ADH2D、pCR-ADH2U-ERO1_T-mhpF-HOR7_P-URA3-ADH2D、pCR-ADH2U-P_TDH3-ADH1-T_ADH1-ERO1_T-mhpF-HOR7_P-URA3-ADH2D、pCR-ADH2part-T_CYC1-URA3-ADH2D的各载体的同源重组部位之间用PCR扩增而得的片段,进行OC700株的转化,涂布于不含尿嘧啶的SD琼脂培养基,将生长的集落纯化。将纯化后的选择株分别命名为Uz1048、Uz1047、Uz928、Uz1012、Uz926、Uz736和Uz1049株。
<发酵试验>
从如上所得的Uz1048、Uz1047、Uz928、Uz1012、Uz926、Uz736和Uz1049系统的菌株中分别挑选发酵能力高的菌株,如下实施烧瓶发酵试验。首先,在分装有20ml葡萄糖浓度20g/L的YPD液体培养基(酵母提取物10g/L、胨20g/L、葡萄糖20g/L)的100ml容量的带挡板的烧瓶中接种供试株,以30℃120rpm进行24小时培养。集菌后,接种于分装有10mlD20X60YAc6培养基(葡萄糖20g/L、木糖60g/L、酵母提取物10g/L、乙酸6g/L)的20ml容量的烧瓶中(菌浓度0.3g干燥菌体/L),以振荡培养(80rpm、振幅35mm、30℃)进行发酵试验。此外,盖在烧瓶上的塞子是通有内径1.5mm的针的橡胶制,通过在针的最前部安装止回阀而使得烧瓶保持厌氧环境。
发酵开始65小时后进行取样,对于发酵液中的葡萄糖、木糖、乙酸、乙醇,使用HPLC(LC-10A;岛津制作所),在下述条件测定。
柱:AminexHPX-87H
移动相:0.01N H2SO4
流量:0.6ml/min
温度:30℃
检测器:差示折射率检测器RID-10A
<发酵试验结果>
上述发酵试验的结果示于表1。
表1
如由表1判断的那样,与mhpF过表达株相比,在过表达mhpF基因和ADH1基因的同时破坏了ADH2基因的Uz736株中,木糖的同化速度大幅提高,其结果乙醇的生产性提高。由于仅ADH2破坏的菌株、仅ADH1过表达的菌株的木糖的同化速度不改善,因而考虑存在协同效果。另外,在Uz736株中,培养基中的乙酸的浓度有意义地减少,明确了乙酸同化能力也提高。
〔实施例2〕
本实施例中,制作导入了木糖异构酶基因和大肠杆菌的mhpF基因、adhE基因、来源于拜氏梭菌(Clostridium beijerinckii)的乙醛脱氢酶基因或来源于莱茵衣藻(Chlamydomonas reinhardtii)的乙醛脱氢酶基因的重组酵母。本实施例中制作的重组酵母中,破坏了内在的一对ADH2基因中的一方或两方。
<导入载体的制作>
(1)XI·XKS1·TKL1·TAL1·RKI1·RPE1基因导入和GRE3基因破坏用质粒
制作在GRE3基因座破坏GRE3基因、同时包含将来源于黄胸散白蚁(Reticulitermes speratus)肠内原生生物的木糖异构酶基因的第377位的氨基酸从天冬酰胺替换为半胱氨酸、木糖的同化速度提高了的突变基因(XI_N337C)、来源于酵母的木酮糖激酶(XKS1)基因、戊糖磷酸途径的转酮醇酶1(TKL1)基因、转醛醇酶1(TAL1)基因、核酮糖磷酸差向异构酶1(RPE1)基因和核糖磷酸酮异构酶(RKI1)基因导入酵母所需的序列的质粒pUC-5U_GRE3-P_HOR7-TKL1-TAL1-FBA1_P-P_ADH1-RPE1-RKI1-TE F1_P-P_TDH1-XI_N337C-T_DIT1-P_TDH3-XKS1-T_HIS3-LoxP-G418-LoxP-3U_GRE3。
该质粒以包含5’侧添加了HOR7启动子的来源于酿酒酵母(Saccharomycescerevisiae)BY4742株的TKL1基因、添加了FBA1启动子的TAL1基因、添加了ADH1启动子的RKI1基因、添加了TEF1启动子的RPE1基因、添加了TDH1启动子和DIT1终止子的XI_N337C(全合成将全长根据酵母的密码子使用频率而转换了密码子的序列而得的基因)、添加了TDH3启动子和HIS3终止子的XKS1基因、作为向酵母基因组上的同源重组区域的GRE3基因的从5’侧末端到上游约700bp的区域的基因序列(GRE3U)和GRE3基因从3’侧末端到下流的约800bp的区域的DNA序列(GRE3D)、以及作为标志物的包含G418基因的基因序列(G418标志物)的方式构建。此外,标志物基因是能够通过在两侧导入LoxP序列而除去标志物的序列。
此外,本质粒所含的各DNA序列可以使用表2的引物来扩增。为了使各DNA片段结合,使用对表2的引物以与邻接DNA序列重复约15bp的方式添加了DNA序列的引物,以酿酒酵母(Saccharomyces cerevisiae)BY4742基因组、XI_N337C合成基因DNA、LoxP序列的合成DNA为模板扩增目的DNA片段,使用In-Fusion HD Cloning Kit(タカラバイオ)等依次使DNA片段结合,克隆到质粒pUC19中而制成最终目的质粒。
表2
(2)mhpF·ADH1基因导入和ADH2基因破坏用质粒
制作在ADH2基因座破坏ADH2基因、同时包含将来源于大肠杆菌(E.coli)的乙醛脱氢酶基因(mhpF)和来源于酵母的醇脱氢酶1(ADH1)基因导入酵母所需的序列的质粒pUC-5U_ADH2-P_TDH3-ADH1-T_ADH1-DIT1_T-mhpF-HOR7_P-URA3-3U_ADH2。
该质粒以包含5’侧添加了TDH3启动子的来源于酿酒酵母(Saccharomycescerevisiae)BY4742株的ADH1基因、添加了HOR7启动子和DIT1终止子的mhpF基因(全合成将全长根据酵母的密码子使用频率而转换了密码子的序列而得的基因)、作为向酵母基因组上的同源重组区域的ADH2基因的从5’侧末端到上游约700bp的区域的基因序列(ADH2U)和ADH2基因从3’侧末端到下流的约800bp的区域的DNA序列(ADH2D)、以及作为标志物的包含URA3基因的基因序列(URA3标志物)的方式构建。
此外,本质粒所含的各DNA序列可以使用表3的引物来扩增。为了使各DNA片段结合,使用对表3的引物以与邻接DNA序列重复约15bp的方式添加了DNA序列的引物,以酿酒酵母(Saccharomyces cerevisiae)BY4742基因组或mhpF合成基因DNA为模板扩增目的DNA片段,使用In-Fusion HD Cloning Kit等依次使DNA片段结合,克隆到质粒pUC19中而制成最终目的质粒。
表3
(3)adhE·ADH1基因导入和ADH2基因破坏用质粒
制作在ADH2基因座破坏ADH2基因、同时包含将来源于大肠杆菌(E.coli)的乙醛脱氢酶基因(adhE)和来源于酵母的醇脱氢酶1(ADH1)基因导入酵母所需的序列的质粒pUC-5U_ADH2-P_TDH3-ADH1-T_ADH1-DIT1_T-adhE-HOR7_P-URA3-3U_ADH2。
该质粒以包含5’侧添加了TDH3启动子的来源于酿酒酵母(Saccharomycescerevisiae)BY4742株的ADH1基因、添加了HOR7启动子和DIT1终止子的adhE基因(NCBI登录号NP_415757.1、全合成将全长根据酵母的密码子使用频率而转换了密码子的序列而得的基因)、作为向酵母基因组上的同源重组区域的ADH2基因的从5’侧末端到上游约700bp的区域的基因序列(ADH2U)和ADH2基因从3’侧末端到下流的约800bp的区域的DNA序列(ADH2D)、以及作为标志物的包含URA3基因的基因序列(URA3标志物)的方式构建。
此外,本质粒所含的各DNA序列可以使用表4的引物来扩增。为了使各DNA片段结合,使用对表4的引物以与邻接DNA序列重复约15bp的方式添加了DNA序列的引物,以质粒pUC-5U_ADH2-P_TDH3-ADH1-T_ADH1-DIT1_T-mhpF-HOR7_P-URA3-3U_ADH2或adhE合成基因DNA为模板扩增目的DNA片段,使用In-Fusion HD Cloning Kit等依次使DNA片段结合,克隆到质粒pUC19中而制成最终目的质粒。
表4
(4)来源于拜氏梭菌(Clostridium beijerinckii)的乙醛脱氢酶基因·ADH1基因导入和ADH2基因破坏用质粒
制作在ADH2基因座破坏ADH2基因、同时包含将来源于拜氏梭菌(Clostridiumbeijerinckii)的乙醛脱氢酶基因和来源于酵母的醇脱氢酶1(ADH1)基因导入酵母所需的序列的质粒pUC-5U_ADH2-P_TDH3-ADH1-T_ADH1-DIT1_T-CloADH-HOR7_P-URA3-3U_ADH2。
该质粒以包含5’侧添加了TDH3启动子的来源于酿酒酵母(Saccharomycescerevisiae)BY4742株的ADH1基因、添加了HOR7启动子和DIT1终止子的来源于拜氏梭菌(Clostridium beijerinckii)的乙醛脱氢酶基因(NCBI登录号YP_001310903.1、全合成将全长根据酵母的密码子使用频率而转换了密码子的序列而得的基因)、作为向酵母基因组上的同源重组区域的ADH2基因的从5’侧末端到上游约700bp的区域的基因序列(ADH2U)和ADH2基因从3’侧末端到下流的约800bp的区域的DNA序列(ADH2D)、以及作为标志物的包含URA3基因的基因序列(URA3标志物)的方式构建。
此外,本质粒所含的各DNA序列可以使用表5的引物来扩增。为了使各DNA片段结合,使用对表5的引物以与邻接DNA序列重复约15bp的方式添加了DNA序列的引物,以质粒pUC-5U_ADH2-P_TDH3-ADH1-T_ADH1-DIT1_T-mhpF-HOR7_P-URA3-3U_ADH2或来源于拜氏梭菌(Clostridium beijerinckii)的乙醛脱氢的合成基因DNA为模板扩增目的DNA片段,使用In-Fusion HD Cloning Kit等依次使DNA片段结合,克隆到质粒pUC19中而制成最终目的质粒。
表5
(5)来源于莱茵衣藻(Chlamydomonas reinhardtii)的乙醛脱氢酶基因·ADH1基因导入和ADH2基因破坏用质粒
制作在ADH2基因座破坏ADH2基因、同时包含将来源于莱茵衣藻(Chlamydomonasreinhardtii)的乙醛脱氢酶基因和来源于酵母的醇脱氢酶1(ADH1)基因导入酵母所需的序列的质粒pUC-5U_ADH2-P_TDH3-ADH1-T_ADH1-DIT1_T-ChlaADH1-HOR7_P-URA3-3U_ADH2。
该质粒以包含5’侧添加了TDH3启动子的来源于酿酒酵母(Saccharomycescerevisiae)BY4742株的ADH1基因、添加了HOR7启动子并添加了DIT1终止子的来源于莱茵衣藻(Chlamydomonas reinhardtii)的乙醛脱氢酶基因(NCBI登录号5729132,全合成将全长根据酵母的密码子使用频率而转换了密码子的序列而得的基因)、作为向酵母基因组上的同源重组区域的ADH2基因的从5’侧末端到上游约700bp的区域的基因序列(ADH2U)和ADH2基因从3’侧末端到下流的约800bp的区域的DNA序列(ADH2D)、以及作为标志物的包含URA3基因的基因序列(URA3标志物)的方式构建。
此外,本质粒所含的各DNA序列可以使用表6的引物来扩增。为了使各DNA片段结合,使用对表6的引物以与邻接DNA序列重复约15bp的方式添加了DNA序列的引物,以质粒pUC-5U_ADH2-P_TDH3-ADH1-T_ADH1-DIT1_T-mhpF-HOR7_P-URA3-3U_ADH2或来源于莱茵衣藻(Chlamydomonas reinhardtii)的乙醛脱氢的合成基因DNA为模板扩增目的DNA片段,使用In-Fusion HD Cloning Kit等依次使DNA片段结合,克隆到质粒pUC19中而制成最终目的质粒。
表6
(6)mhpF基因导入用质粒
制作不在ADH2基因座破坏ADH2基因、并包含在ADH2基因座附近将来源于大肠杆菌(E.coli)的乙醛脱氢酶基因(mhpF)导入酵母所需的序列的质粒pUC-ADH2-T_CYC1-DIT1_T-mhpF-HOR7_P-URA3-3U_ADH2。
该质粒以包含5’侧添加了HOR7启动子并添加了DIT1终止子的来源于酿酒酵母(Saccharomyces cerevisiae)BY4742株的mhpF基因(全合成将全长根据酵母的密码子使用频率而转换了密码子的序列而得的基因)、作为向酵母基因组上的同源重组区域的ADH2基因和ADH2基因从3’侧末端到下流的约800bp的区域的DNA序列(ADH2D)、以及作为标志物的包含URA3基因的基因序列(URA3标志物)的方式构建。
此外,本质粒所含的各DNA序列可以使用表7的引物来扩增。为了使各DNA片段结合,使用对表7的引物以与邻接DNA序列重复约15bp的方式添加了DNA序列的引物,以质粒pUC-5U_ADH2-P_TDH3-ADH1-T_ADH1-DIT1_T-mhpF-HOR7_P-URA3-3U_ADH2或酿酒酵母(Saccharomyces cerevisiae)BY4742基因组为模板扩增目的DNA片段,使用In-Fusion HDCloning Kit等依次使DNA片段结合,克隆到质粒pUC19中而制成最终目的质粒。
表7
<载体导入酵母株的制作>
用添加了5-氟乳清酸的培养基筛选作为2倍体酵母的酿酒酵母(Saccharomycescerevisiae)OC2株(NBRC2260)(Boeke,J.D.,et al.1987Methods Enzymol.;154:164-75.),以成为尿嘧啶营养缺陷型的菌株(OC2U)作为宿主。酵母的转化使用Frozen-EZ YeastTransformation II(ZYMO RESEARCH),按照附带的实验步骤进行。
使用将上述(1)中制作的质粒:pUC-5U_GRE3-P_HOR7-TKL1-TAL1-FBA1_P-P_ADH1-RPE1-RKI1-TEF1_P-P_TDH1-XI_N337C-T_DIT1-P_TDH3-XKS1-T_HIS3-LoxP-G418-LoxP-3U_GRE3的同源重组部位用PCR扩增而得的片段,进行OC2U株的转化,涂布于包含G418的YPD琼脂培养基,将生长的集落纯化。将纯化后的选择株命名为Uz1252株。使本菌株在胞子形成培养基(1%磷酸钾、0.1%酵母提取物、0.05%葡萄糖、2%琼脂)中形成胞子,利用同宗配合性进行2倍化。获得在作为2倍体的染色体的GRE3基因座区域插入了突变型XI基因、TKL1基因、TAL1基因、RPE1基因、RKI1基因和XKS1基因、且GRE3基因被破坏的菌株。将此作为Uz1252-3株。
并且,使用将上述(2)中制作的质粒:pUC-5U_ADH2-P_TDH3-ADH1-T_ADH1-DIT1_T-mhpF-HOR7_P-URA3-3U_ADH2、上述(3)中制作的质粒:pUC-5U_ADH2-P_TDH3-ADH1-T_ADH1-DIT1_T-adhE-HOR7_P-URA3-3U_ADH2、上述(4)中制作的质粒:pUC-5U_ADH2-P_TDH3-ADH1-T_ADH1-DIT1_T-CloADH-HOR7_P-URA3-3U_ADH2、上述(5)中制作的质粒:pUC-5U_ADH2-P_TDH3-ADH1-T_ADH1-DIT1_T-ChlaADH1-HOR7_P-URA3-3U_ADH2、和上述(6)中制作的质粒:pUC-ADH2-T_CYC1-DIT1_T-mhpF-HOR7_P-URA3-3U_ADH2的各质粒的同源重组部位之间用PCR扩增而得的片段,进行上述Uz1252-3株的转化,涂布于不含尿嘧啶的SD琼脂培养基,将生长的集落纯化。将纯化后的选择株分别命名为Uz1317株、Uz1298株、Uz1296株、Uz1330株、和Uz1320株。
此外,确认了这些全部菌株发生了杂合(1拷贝)重组。对于所得的Uz1317株、Uz1298株和Uz1296株,使其在胞子形成培养基中形成胞子,利用同宗配合性进行2倍化,将所得的菌株分别命名为Uz1319株、Uz1318株和Uz1311株。
另外,作为对照,使用以OC2基因组为模板用PCR扩增而得的尿嘧啶基因,进行OC2U株的转化,涂布于不含尿嘧啶的SD琼脂培养基,将生长的集落纯化,命名为Uz1313株。使Uz1313株在胞子形成培养基中形成胞子,利用同宗配合性进行2倍化,命名为Uz1323株。
此外,本实施例中制作的株的基因型归纳于表8。
表8
<发酵试验>
从如上制成的菌株中分别选择发酵能力高的2株,如下实施烧瓶发酵试验。首先,在分装有20ml葡萄糖浓度20g/L的YPD液体培养基(酵母提取物10g/L、胨20g/L、葡萄糖20g/L)的100ml容量的带挡板的烧瓶中接种供试株,以30℃120rpm进行24小时培养。集菌后,接种于分装有8ml D60X80YPAc4培养基(葡萄糖60g/L、木糖80g/L、酵母提取物10g/L、胨20g/L、乙酸4g/L)或D40X80YPAc2培养基(葡萄糖40g/L、木糖80g/L、酵母提取物10g/L、胨20g/L、乙酸2g/L)的10ml容量的烧瓶中,通过振荡培养(80rpm、振幅35mm、30℃)进行发酵试验。此外,盖在烧瓶上的塞子是通有内径1.5mm针的橡胶制,通过在针的最前部安装止回阀来使烧瓶保持厌氧环境。
对于发酵液中的葡萄糖、木糖、乙醇,使用HPLC(LC-10A;岛津制作所),在下述条件测定。
柱:AminexHPX-87H
移动相:0.01N H2SO4
流量:0.6ml/min
温度:30℃
检测器:差示折射率检测器RID-10A
<发酵试验结果>
将使用D60X80YPAc4培养基、发酵时间为66小时的发酵试验(加入菌浓度0.3g干燥菌体/L)的结果示于表9和10。此外,表9和10所示的数据是独立获得的重组株3株的数据平均值。
表9
表10
将使用D40X80YPAc2培养基、发酵时间为42小时的发酵试验(加入菌浓度0.24g干燥菌体/L)的结果示于表11和12,另外将对于杂合导入株使用D40X80YPAc2培养基、发酵时间为42小时的发酵试验(加入菌浓度0.3g干燥菌体/L)的结果示于表13。此外,表11~13所示的数据是独立获得的重组株3株的数据平均值。
表11
表12
表13
如由表9~13可知的那样,与对照相比,杂合或纯合地破坏了ADH2、同时过表达ADH1和3种乙醛脱氢酶的任一种的菌株,木糖的同化速度大幅提高,乙酸的减少量也多,其结果乙醇的生产性提高。此外,关于乙酸的减少量,与ADH2的杂合导入株相比,ADH2的纯合导入株更多。另一方面,仅表达乙醛脱氢酶mhpF的菌株,木糖的同化速度降低,乙酸基本不减少,乙醇的生产性不提高。
本说明书中引用的全部出版物、专利和专利申请均直接作为参考而纳入本说明书中。
序列表
<110> 丰田自动车株式会社
<120> 使用了重组酵母的乙醇的制造方法
<130> PH-5535-PCT
<150> JP 2013-037501
<151> 2013-2-27
<150> JP 2014-036652
<151> 2014-2-27
<160> 100
<170> PatentIn 版本 3.5
<210> 1
<211> 951
<212> DNA
<213> 大肠杆菌(Escherichia coli)
<220>
<221> CDS
<222> (1)..(951)
<400> 1
atg tca aag aga aaa gtt gct att atc ggt agt gga aac atc ggt act 48
Met Ser Lys Arg Lys Val Ala Ile Ile Gly Ser Gly Asn Ile Gly Thr
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Asp Leu Met Ile Lys Ile Leu Arg His Gly Gln His Leu Glu Met Ala
20 25 30
gtt atg gtc ggt atc gat cct cag tca gac gga ctt gct aga gcc aga 144
Val Met Val Gly Ile Asp Pro Gln Ser Asp Gly Leu Ala Arg Ala Arg
35 40 45
aga atg ggt gtt gct act aca cat gaa ggt gtt att gga ttg atg aac 192
Arg Met Gly Val Ala Thr Thr His Glu Gly Val Ile Gly Leu Met Asn
50 55 60
atg cca gag ttt gca gat att gac atc gtt ttc gat gct aca agt gca 240
Met Pro Glu Phe Ala Asp Ile Asp Ile Val Phe Asp Ala Thr Ser Ala
65 70 75 80
ggt gct cac gtt aag aat gac gct gcc ttg aga gaa gct aaa cct gat 288
Gly Ala His Val Lys Asn Asp Ala Ala Leu Arg Glu Ala Lys Pro Asp
85 90 95
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Ile Arg Leu Ile Asp Leu Thr Pro Ala Ala Ile Gly Pro Tyr Cys Val
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Pro Val Val Asn Leu Glu Ala Asn Val Asp Gln Leu Asn Val Asn Met
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Val Thr Cys Gly Gly Gln Ala Thr Ile Pro Met Val Ala Ala Val Ser
130 135 140
aga gtt gct aga gtc cat tat gcc gaa att atc gca tcc atc gct tca 480
Arg Val Ala Arg Val His Tyr Ala Glu Ile Ile Ala Ser Ile Ala Ser
145 150 155 160
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Lys Ser Ala Gly Pro Gly Thr Arg Ala Asn Ile Asp Glu Phe Thr Glu
165 170 175
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Thr Thr Ser Arg Ala Ile Glu Val Val Gly Gly Ala Ala Lys Gly Lys
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Ala Ile Ile Val Leu Asn Pro Ala Glu Pro Pro Leu Met Met Arg Asp
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Thr Val Tyr Val Leu Ser Asp Glu Ala Ser Gln Asp Asp Ile Glu Ala
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Ser Ile Asn Glu Met Ala Glu Ala Val Gln Ala Tyr Val Pro Gly Tyr
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Arg Leu Lys Gln Arg Val Gln Phe Glu Val Ile Pro Gln Asp Lys Pro
245 250 255
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Val Asn Leu Pro Gly Val Gly Gln Phe Ser Gly Leu Lys Thr Ala Val
260 265 270
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Trp Leu Glu Val Glu Gly Ala Ala His Tyr Leu Pro Ala Tyr Ala Gly
275 280 285
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Asn Leu Asp Ile Met Thr Ser Ser Ala Leu Ala Thr Ala Glu Lys Met
290 295 300
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Ala Gln Ser Leu Ala Arg Lys Ala Gly Glu Ala Ala
305 310 315
<210> 2
<211> 316
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 2
Met Ser Lys Arg Lys Val Ala Ile Ile Gly Ser Gly Asn Ile Gly Thr
1 5 10 15
Asp Leu Met Ile Lys Ile Leu Arg His Gly Gln His Leu Glu Met Ala
20 25 30
Val Met Val Gly Ile Asp Pro Gln Ser Asp Gly Leu Ala Arg Ala Arg
35 40 45
Arg Met Gly Val Ala Thr Thr His Glu Gly Val Ile Gly Leu Met Asn
50 55 60
Met Pro Glu Phe Ala Asp Ile Asp Ile Val Phe Asp Ala Thr Ser Ala
65 70 75 80
Gly Ala His Val Lys Asn Asp Ala Ala Leu Arg Glu Ala Lys Pro Asp
85 90 95
Ile Arg Leu Ile Asp Leu Thr Pro Ala Ala Ile Gly Pro Tyr Cys Val
100 105 110
Pro Val Val Asn Leu Glu Ala Asn Val Asp Gln Leu Asn Val Asn Met
115 120 125
Val Thr Cys Gly Gly Gln Ala Thr Ile Pro Met Val Ala Ala Val Ser
130 135 140
Arg Val Ala Arg Val His Tyr Ala Glu Ile Ile Ala Ser Ile Ala Ser
145 150 155 160
Lys Ser Ala Gly Pro Gly Thr Arg Ala Asn Ile Asp Glu Phe Thr Glu
165 170 175
Thr Thr Ser Arg Ala Ile Glu Val Val Gly Gly Ala Ala Lys Gly Lys
180 185 190
Ala Ile Ile Val Leu Asn Pro Ala Glu Pro Pro Leu Met Met Arg Asp
195 200 205
Thr Val Tyr Val Leu Ser Asp Glu Ala Ser Gln Asp Asp Ile Glu Ala
210 215 220
Ser Ile Asn Glu Met Ala Glu Ala Val Gln Ala Tyr Val Pro Gly Tyr
225 230 235 240
Arg Leu Lys Gln Arg Val Gln Phe Glu Val Ile Pro Gln Asp Lys Pro
245 250 255
Val Asn Leu Pro Gly Val Gly Gln Phe Ser Gly Leu Lys Thr Ala Val
260 265 270
Trp Leu Glu Val Glu Gly Ala Ala His Tyr Leu Pro Ala Tyr Ala Gly
275 280 285
Asn Leu Asp Ile Met Thr Ser Ser Ala Leu Ala Thr Ala Glu Lys Met
290 295 300
Ala Gln Ser Leu Ala Arg Lys Ala Gly Glu Ala Ala
305 310 315
<210> 3
<211> 1320
<212> DNA
<213> 黄胸散白蚁(Reticulitermes speratus)肠内原生生物
<220>
<221> CDS
<222> (1)..(1320)
<400> 3
atg tct caa att ttt aag gat atc cca gtt att aaa tat gaa ggt cca 48
Met Ser Gln Ile Phe Lys Asp Ile Pro Val Ile Lys Tyr Glu Gly Pro
1 5 10 15
gct tcc aag aat cct ttg agt ttc aaa tac tac gat gca aac aag gtt 96
Ala Ser Lys Asn Pro Leu Ser Phe Lys Tyr Tyr Asp Ala Asn Lys Val
20 25 30
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Ile Asp Gly Lys Pro Met Lys Glu His Leu Arg Tyr Ala Met Ala Trp
35 40 45
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Trp His Asn Leu Cys Ala Thr Gly Gln Asp Met Phe Gly Pro Gly Thr
50 55 60
gca gat aaa tcc ttc ggt agt aag aca gtt ggt acc atg gaa cat gca 240
Ala Asp Lys Ser Phe Gly Ser Lys Thr Val Gly Thr Met Glu His Ala
65 70 75 80
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His Ala Lys Val Asp Ala Gly Phe Glu Phe Met Ser Lys Leu Gly Val
85 90 95
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Glu Tyr Phe Cys Phe His Asp Ala Asp Leu Val Pro Glu Ala Asp Thr
100 105 110
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Leu Ser Glu Thr Asn Lys Arg Leu Asp Glu Ile Ala Glu His Ile Val
115 120 125
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Ala Lys Gln Lys Ala Thr Gly Ile Lys Cys Leu Trp Gly Thr Ala Asn
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Leu Phe Ser Asn Pro Arg Phe Leu Asn Gly Ser Gly Ser Ser Asn Ser
145 150 155 160
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Ala Asp Val Tyr Ala Tyr Ala Ala Ala Gln Ile Lys Lys Ala Leu Asp
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Leu Thr Val Lys Phe Gly Gly Val Gly Tyr Val Phe Trp Gly Gly Arg
180 185 190
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Glu Gly Tyr Glu Thr Leu Leu Asn Thr Asp Val Lys Phe Glu Gln Glu
195 200 205
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Asn Ile Ala Asn Leu Met His Leu Ala Val Thr Tyr Gly Arg Ser Ile
210 215 220
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Gly Phe Lys Gly Asp Phe Tyr Ile Glu Pro Lys Pro Lys Glu Pro Thr
225 230 235 240
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Lys His Gln Tyr Asp Phe Asp Ala Ala Thr Thr Ile Gly Phe Ile Arg
245 250 255
caa tac ggt ttg gaa aag gat ttc aag ttg aac atc gaa gca aac cat 816
Gln Tyr Gly Leu Glu Lys Asp Phe Lys Leu Asn Ile Glu Ala Asn His
260 265 270
gct aca tta gca ggt cat acc ttc caa cat gat ttg aga atc tct gct 864
Ala Thr Leu Ala Gly His Thr Phe Gln His Asp Leu Arg Ile Ser Ala
275 280 285
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Ile Asn Gly Met Leu Gly Ser Val Asp Ala Asn Thr Gly Asp Pro Leu
290 295 300
tta ggt tgg gat acc gat gaa ttt cct tat tcc gtt tac gat acc act 960
Leu Gly Trp Asp Thr Asp Glu Phe Pro Tyr Ser Val Tyr Asp Thr Thr
305 310 315 320
ttg gct atg tac gaa att att aag gca ggt ggt ttg acc ggt ggt ttg 1008
Leu Ala Met Tyr Glu Ile Ile Lys Ala Gly Gly Leu Thr Gly Gly Leu
325 330 335
aat ttt gat tcc aag gtt aga aga cca agt tac aca cat gaa gat ttg 1056
Asn Phe Asp Ser Lys Val Arg Arg Pro Ser Tyr Thr His Glu Asp Leu
340 345 350
ttt tac ggt ttc att ttg ggt atg gat tct ttc gct ttg ggt ttg att 1104
Phe Tyr Gly Phe Ile Leu Gly Met Asp Ser Phe Ala Leu Gly Leu Ile
355 360 365
aaa gca aag gct ttg att gca gat ggt aga ttg gat tca ttc gtt aag 1152
Lys Ala Lys Ala Leu Ile Ala Asp Gly Arg Leu Asp Ser Phe Val Lys
370 375 380
gat aga tac gct tct tac ggt tca ggt att ggt gct aag att aga gat 1200
Asp Arg Tyr Ala Ser Tyr Gly Ser Gly Ile Gly Ala Lys Ile Arg Asp
385 390 395 400
cat tct gca act ttg gaa gaa tta gca gct tat gca tta gct aaa gat 1248
His Ser Ala Thr Leu Glu Glu Leu Ala Ala Tyr Ala Leu Ala Lys Asp
405 410 415
aca gtt gct ttg cct ggt tcc ggt aga caa gaa tac tta gaa agt att 1296
Thr Val Ala Leu Pro Gly Ser Gly Arg Gln Glu Tyr Leu Glu Ser Ile
420 425 430
att aac caa att ttg ttt caa taa 1320
Ile Asn Gln Ile Leu Phe Gln
435
<210> 4
<211> 439
<212> PRT
<213> 黄胸散白蚁(Reticulitermes speratus)肠内原生生物
<400> 4
Met Ser Gln Ile Phe Lys Asp Ile Pro Val Ile Lys Tyr Glu Gly Pro
1 5 10 15
Ala Ser Lys Asn Pro Leu Ser Phe Lys Tyr Tyr Asp Ala Asn Lys Val
20 25 30
Ile Asp Gly Lys Pro Met Lys Glu His Leu Arg Tyr Ala Met Ala Trp
35 40 45
Trp His Asn Leu Cys Ala Thr Gly Gln Asp Met Phe Gly Pro Gly Thr
50 55 60
Ala Asp Lys Ser Phe Gly Ser Lys Thr Val Gly Thr Met Glu His Ala
65 70 75 80
His Ala Lys Val Asp Ala Gly Phe Glu Phe Met Ser Lys Leu Gly Val
85 90 95
Glu Tyr Phe Cys Phe His Asp Ala Asp Leu Val Pro Glu Ala Asp Thr
100 105 110
Leu Ser Glu Thr Asn Lys Arg Leu Asp Glu Ile Ala Glu His Ile Val
115 120 125
Ala Lys Gln Lys Ala Thr Gly Ile Lys Cys Leu Trp Gly Thr Ala Asn
130 135 140
Leu Phe Ser Asn Pro Arg Phe Leu Asn Gly Ser Gly Ser Ser Asn Ser
145 150 155 160
Ala Asp Val Tyr Ala Tyr Ala Ala Ala Gln Ile Lys Lys Ala Leu Asp
165 170 175
Leu Thr Val Lys Phe Gly Gly Val Gly Tyr Val Phe Trp Gly Gly Arg
180 185 190
Glu Gly Tyr Glu Thr Leu Leu Asn Thr Asp Val Lys Phe Glu Gln Glu
195 200 205
Asn Ile Ala Asn Leu Met His Leu Ala Val Thr Tyr Gly Arg Ser Ile
210 215 220
Gly Phe Lys Gly Asp Phe Tyr Ile Glu Pro Lys Pro Lys Glu Pro Thr
225 230 235 240
Lys His Gln Tyr Asp Phe Asp Ala Ala Thr Thr Ile Gly Phe Ile Arg
245 250 255
Gln Tyr Gly Leu Glu Lys Asp Phe Lys Leu Asn Ile Glu Ala Asn His
260 265 270
Ala Thr Leu Ala Gly His Thr Phe Gln His Asp Leu Arg Ile Ser Ala
275 280 285
Ile Asn Gly Met Leu Gly Ser Val Asp Ala Asn Thr Gly Asp Pro Leu
290 295 300
Leu Gly Trp Asp Thr Asp Glu Phe Pro Tyr Ser Val Tyr Asp Thr Thr
305 310 315 320
Leu Ala Met Tyr Glu Ile Ile Lys Ala Gly Gly Leu Thr Gly Gly Leu
325 330 335
Asn Phe Asp Ser Lys Val Arg Arg Pro Ser Tyr Thr His Glu Asp Leu
340 345 350
Phe Tyr Gly Phe Ile Leu Gly Met Asp Ser Phe Ala Leu Gly Leu Ile
355 360 365
Lys Ala Lys Ala Leu Ile Ala Asp Gly Arg Leu Asp Ser Phe Val Lys
370 375 380
Asp Arg Tyr Ala Ser Tyr Gly Ser Gly Ile Gly Ala Lys Ile Arg Asp
385 390 395 400
His Ser Ala Thr Leu Glu Glu Leu Ala Ala Tyr Ala Leu Ala Lys Asp
405 410 415
Thr Val Ala Leu Pro Gly Ser Gly Arg Gln Glu Tyr Leu Glu Ser Ile
420 425 430
Ile Asn Gln Ile Leu Phe Gln
435
<210> 5
<211> 1047
<212> DNA
<213> 酿酒酵母(Saccharomyces cerevisiae)
<220>
<221> CDS
<222> (1)..(1047)
<400> 5
atg tct atc cca gaa act caa aaa ggt gtt atc ttc tac gaa tcc cac 48
Met Ser Ile Pro Glu Thr Gln Lys Gly Val Ile Phe Tyr Glu Ser His
1 5 10 15
ggt aag ttg gaa tac aaa gat att cca gtt cca aag cca aag gcc aac 96
Gly Lys Leu Glu Tyr Lys Asp Ile Pro Val Pro Lys Pro Lys Ala Asn
20 25 30
gaa ttg ttg atc aac gtt aaa tac tct ggt gtc tgt cac act gac ttg 144
Glu Leu Leu Ile Asn Val Lys Tyr Ser Gly Val Cys His Thr Asp Leu
35 40 45
cac gct tgg cac ggt gac tgg cca ttg cca gtt aag cta cca tta gtc 192
His Ala Trp His Gly Asp Trp Pro Leu Pro Val Lys Leu Pro Leu Val
50 55 60
ggt ggt cac gaa ggt gcc ggt gtc gtt gtc ggc atg ggt gaa aac gtt 240
Gly Gly His Glu Gly Ala Gly Val Val Val Gly Met Gly Glu Asn Val
65 70 75 80
aag ggc tgg aag atc ggt gac tac gcc ggt atc aaa tgg ttg aac ggt 288
Lys Gly Trp Lys Ile Gly Asp Tyr Ala Gly Ile Lys Trp Leu Asn Gly
85 90 95
tct tgt atg gcc tgt gaa tac tgt gaa ttg ggt aac gaa tcc aac tgt 336
Ser Cys Met Ala Cys Glu Tyr Cys Glu Leu Gly Asn Glu Ser Asn Cys
100 105 110
cct cac gct gac ttg tct ggt tac acc cac gac ggt tct ttc caa caa 384
Pro His Ala Asp Leu Ser Gly Tyr Thr His Asp Gly Ser Phe Gln Gln
115 120 125
tac gct acc gct gac gct gtt caa gcc gct cac att cct caa ggt acc 432
Tyr Ala Thr Ala Asp Ala Val Gln Ala Ala His Ile Pro Gln Gly Thr
130 135 140
gac ttg gcc caa gtc gcc ccc atc ttg tgt gct ggt atc acc gtc tac 480
Asp Leu Ala Gln Val Ala Pro Ile Leu Cys Ala Gly Ile Thr Val Tyr
145 150 155 160
aag gct ttg aag tct gct aac ttg atg gcc ggt cac tgg gtt gct atc 528
Lys Ala Leu Lys Ser Ala Asn Leu Met Ala Gly His Trp Val Ala Ile
165 170 175
tcc ggt gct gct ggt ggt cta ggt tct ttg gct gtt caa tac gcc aag 576
Ser Gly Ala Ala Gly Gly Leu Gly Ser Leu Ala Val Gln Tyr Ala Lys
180 185 190
gct atg ggt tac aga gtc ttg ggt att gac ggt ggt gaa ggt aag gaa 624
Ala Met Gly Tyr Arg Val Leu Gly Ile Asp Gly Gly Glu Gly Lys Glu
195 200 205
gaa tta ttc aga tcc atc ggt ggt gaa gtc ttc att gac ttc act aag 672
Glu Leu Phe Arg Ser Ile Gly Gly Glu Val Phe Ile Asp Phe Thr Lys
210 215 220
gaa aag gac att gtc ggt gct gtt cta aag gcc act gac ggt ggt gct 720
Glu Lys Asp Ile Val Gly Ala Val Leu Lys Ala Thr Asp Gly Gly Ala
225 230 235 240
cac ggt gtc atc aac gtt tcc gtt tcc gaa gcc gct att gaa gct tct 768
His Gly Val Ile Asn Val Ser Val Ser Glu Ala Ala Ile Glu Ala Ser
245 250 255
acc aga tac gtt aga gct aac ggt acc acc gtt ttg gtc ggt atg cca 816
Thr Arg Tyr Val Arg Ala Asn Gly Thr Thr Val Leu Val Gly Met Pro
260 265 270
gct ggt gcc aag tgt tgt tct gat gtc ttc aac caa gtc gtc aag tcc 864
Ala Gly Ala Lys Cys Cys Ser Asp Val Phe Asn Gln Val Val Lys Ser
275 280 285
atc tct att gtt ggt tct tac gtc ggt aac aga gct gac acc aga gaa 912
Ile Ser Ile Val Gly Ser Tyr Val Gly Asn Arg Ala Asp Thr Arg Glu
290 295 300
gct ttg gac ttc ttc gcc aga ggt ttg gtc aag tct cca atc aag gtt 960
Ala Leu Asp Phe Phe Ala Arg Gly Leu Val Lys Ser Pro Ile Lys Val
305 310 315 320
gtc ggc ttg tct acc ttg cca gaa att tac gaa aag atg gaa aag ggt 1008
Val Gly Leu Ser Thr Leu Pro Glu Ile Tyr Glu Lys Met Glu Lys Gly
325 330 335
caa atc gtt ggt aga tac gtt gtt gac act tct aaa taa 1047
Gln Ile Val Gly Arg Tyr Val Val Asp Thr Ser Lys
340 345
<210> 6
<211> 348
<212> PRT
<213> 酿酒酵母(Saccharomyces cerevisiae)
<400> 6
Met Ser Ile Pro Glu Thr Gln Lys Gly Val Ile Phe Tyr Glu Ser His
1 5 10 15
Gly Lys Leu Glu Tyr Lys Asp Ile Pro Val Pro Lys Pro Lys Ala Asn
20 25 30
Glu Leu Leu Ile Asn Val Lys Tyr Ser Gly Val Cys His Thr Asp Leu
35 40 45
His Ala Trp His Gly Asp Trp Pro Leu Pro Val Lys Leu Pro Leu Val
50 55 60
Gly Gly His Glu Gly Ala Gly Val Val Val Gly Met Gly Glu Asn Val
65 70 75 80
Lys Gly Trp Lys Ile Gly Asp Tyr Ala Gly Ile Lys Trp Leu Asn Gly
85 90 95
Ser Cys Met Ala Cys Glu Tyr Cys Glu Leu Gly Asn Glu Ser Asn Cys
100 105 110
Pro His Ala Asp Leu Ser Gly Tyr Thr His Asp Gly Ser Phe Gln Gln
115 120 125
Tyr Ala Thr Ala Asp Ala Val Gln Ala Ala His Ile Pro Gln Gly Thr
130 135 140
Asp Leu Ala Gln Val Ala Pro Ile Leu Cys Ala Gly Ile Thr Val Tyr
145 150 155 160
Lys Ala Leu Lys Ser Ala Asn Leu Met Ala Gly His Trp Val Ala Ile
165 170 175
Ser Gly Ala Ala Gly Gly Leu Gly Ser Leu Ala Val Gln Tyr Ala Lys
180 185 190
Ala Met Gly Tyr Arg Val Leu Gly Ile Asp Gly Gly Glu Gly Lys Glu
195 200 205
Glu Leu Phe Arg Ser Ile Gly Gly Glu Val Phe Ile Asp Phe Thr Lys
210 215 220
Glu Lys Asp Ile Val Gly Ala Val Leu Lys Ala Thr Asp Gly Gly Ala
225 230 235 240
His Gly Val Ile Asn Val Ser Val Ser Glu Ala Ala Ile Glu Ala Ser
245 250 255
Thr Arg Tyr Val Arg Ala Asn Gly Thr Thr Val Leu Val Gly Met Pro
260 265 270
Ala Gly Ala Lys Cys Cys Ser Asp Val Phe Asn Gln Val Val Lys Ser
275 280 285
Ile Ser Ile Val Gly Ser Tyr Val Gly Asn Arg Ala Asp Thr Arg Glu
290 295 300
Ala Leu Asp Phe Phe Ala Arg Gly Leu Val Lys Ser Pro Ile Lys Val
305 310 315 320
Val Gly Leu Ser Thr Leu Pro Glu Ile Tyr Glu Lys Met Glu Lys Gly
325 330 335
Gln Ile Val Gly Arg Tyr Val Val Asp Thr Ser Lys
340 345
<210> 7
<211> 1047
<212> DNA
<213> 酿酒酵母(Saccharomyces cerevisiae)
<220>
<221> CDS
<222> (1)..(1047)
<400> 7
atg tct att cca gaa act caa aaa gcc att atc ttc tac gaa tcc aac 48
Met Ser Ile Pro Glu Thr Gln Lys Ala Ile Ile Phe Tyr Glu Ser Asn
1 5 10 15
ggc aag ttg gag cat aag gat atc cca gtt cca aag cca aag ccc aac 96
Gly Lys Leu Glu His Lys Asp Ile Pro Val Pro Lys Pro Lys Pro Asn
20 25 30
gaa ttg tta atc aac gtc aag tac tct ggt gtc tgc cac acc gat ttg 144
Glu Leu Leu Ile Asn Val Lys Tyr Ser Gly Val Cys His Thr Asp Leu
35 40 45
cac gct tgg cat ggt gac tgg cca ttg cca act aag tta cca tta gtt 192
His Ala Trp His Gly Asp Trp Pro Leu Pro Thr Lys Leu Pro Leu Val
50 55 60
ggt ggt cac gaa ggt gcc ggt gtc gtt gtc ggc atg ggt gaa aac gtt 240
Gly Gly His Glu Gly Ala Gly Val Val Val Gly Met Gly Glu Asn Val
65 70 75 80
aag ggc tgg aag atc ggt gac tac gcc ggt atc aaa tgg ttg aac ggt 288
Lys Gly Trp Lys Ile Gly Asp Tyr Ala Gly Ile Lys Trp Leu Asn Gly
85 90 95
tct tgt atg gcc tgt gaa tac tgt gaa ttg ggt aac gaa tcc aac tgt 336
Ser Cys Met Ala Cys Glu Tyr Cys Glu Leu Gly Asn Glu Ser Asn Cys
100 105 110
cct cac gct gac ttg tct ggt tac acc cac gac ggt tct ttc caa gaa 384
Pro His Ala Asp Leu Ser Gly Tyr Thr His Asp Gly Ser Phe Gln Glu
115 120 125
tac gct acc gct gac gct gtt caa gcc gct cac att cct caa ggt act 432
Tyr Ala Thr Ala Asp Ala Val Gln Ala Ala His Ile Pro Gln Gly Thr
130 135 140
gac ttg gct gaa gtc gcg cca atc ttg tgt gct ggt atc acc gta tac 480
Asp Leu Ala Glu Val Ala Pro Ile Leu Cys Ala Gly Ile Thr Val Tyr
145 150 155 160
aag gct ttg aag tct gcc aac ttg aga gca ggc cac tgg gcg gcc att 528
Lys Ala Leu Lys Ser Ala Asn Leu Arg Ala Gly His Trp Ala Ala Ile
165 170 175
tct ggt gct gct ggt ggt cta ggt tct ttg gct gtt caa tat gct aag 576
Ser Gly Ala Ala Gly Gly Leu Gly Ser Leu Ala Val Gln Tyr Ala Lys
180 185 190
gcg atg ggt tac aga gtc tta ggt att gat ggt ggt cca gga aag gaa 624
Ala Met Gly Tyr Arg Val Leu Gly Ile Asp Gly Gly Pro Gly Lys Glu
195 200 205
gaa ttg ttt acc tcg ctc ggt ggt gaa gta ttc atc gac ttc acc aaa 672
Glu Leu Phe Thr Ser Leu Gly Gly Glu Val Phe Ile Asp Phe Thr Lys
210 215 220
gag aag gac att gtt agc gca gtc gtt aag gct acc aac ggc ggt gcc 720
Glu Lys Asp Ile Val Ser Ala Val Val Lys Ala Thr Asn Gly Gly Ala
225 230 235 240
cac ggt atc atc aat gtt tcc gtt tcc gaa gcc gct atc gaa gct tct 768
His Gly Ile Ile Asn Val Ser Val Ser Glu Ala Ala Ile Glu Ala Ser
245 250 255
acc aga tac tgt agg gcg aac ggt act gtt gtc ttg gtt ggt ttg cca 816
Thr Arg Tyr Cys Arg Ala Asn Gly Thr Val Val Leu Val Gly Leu Pro
260 265 270
gcc ggt gca aag tgc tcc tct gat gtc ttc aac cac gtt gtc aag tct 864
Ala Gly Ala Lys Cys Ser Ser Asp Val Phe Asn His Val Val Lys Ser
275 280 285
atc tcc att gtc ggc tct tac gtg ggg aac aga gct gat acc aga gaa 912
Ile Ser Ile Val Gly Ser Tyr Val Gly Asn Arg Ala Asp Thr Arg Glu
290 295 300
gcc tta gat ttc ttt gcc aga ggt cta gtc aag tct cca ata aag gta 960
Ala Leu Asp Phe Phe Ala Arg Gly Leu Val Lys Ser Pro Ile Lys Val
305 310 315 320
gtt ggc tta tcc agt tta cca gaa att tac gaa aag atg gag aag ggc 1008
Val Gly Leu Ser Ser Leu Pro Glu Ile Tyr Glu Lys Met Glu Lys Gly
325 330 335
caa att gct ggt aga tac gtt gtt gac act tct aaa taa 1047
Gln Ile Ala Gly Arg Tyr Val Val Asp Thr Ser Lys
340 345
<210> 8
<211> 348
<212> PRT
<213> 酿酒酵母(Saccharomyces cerevisiae)
<400> 8
Met Ser Ile Pro Glu Thr Gln Lys Ala Ile Ile Phe Tyr Glu Ser Asn
1 5 10 15
Gly Lys Leu Glu His Lys Asp Ile Pro Val Pro Lys Pro Lys Pro Asn
20 25 30
Glu Leu Leu Ile Asn Val Lys Tyr Ser Gly Val Cys His Thr Asp Leu
35 40 45
His Ala Trp His Gly Asp Trp Pro Leu Pro Thr Lys Leu Pro Leu Val
50 55 60
Gly Gly His Glu Gly Ala Gly Val Val Val Gly Met Gly Glu Asn Val
65 70 75 80
Lys Gly Trp Lys Ile Gly Asp Tyr Ala Gly Ile Lys Trp Leu Asn Gly
85 90 95
Ser Cys Met Ala Cys Glu Tyr Cys Glu Leu Gly Asn Glu Ser Asn Cys
100 105 110
Pro His Ala Asp Leu Ser Gly Tyr Thr His Asp Gly Ser Phe Gln Glu
115 120 125
Tyr Ala Thr Ala Asp Ala Val Gln Ala Ala His Ile Pro Gln Gly Thr
130 135 140
Asp Leu Ala Glu Val Ala Pro Ile Leu Cys Ala Gly Ile Thr Val Tyr
145 150 155 160
Lys Ala Leu Lys Ser Ala Asn Leu Arg Ala Gly His Trp Ala Ala Ile
165 170 175
Ser Gly Ala Ala Gly Gly Leu Gly Ser Leu Ala Val Gln Tyr Ala Lys
180 185 190
Ala Met Gly Tyr Arg Val Leu Gly Ile Asp Gly Gly Pro Gly Lys Glu
195 200 205
Glu Leu Phe Thr Ser Leu Gly Gly Glu Val Phe Ile Asp Phe Thr Lys
210 215 220
Glu Lys Asp Ile Val Ser Ala Val Val Lys Ala Thr Asn Gly Gly Ala
225 230 235 240
His Gly Ile Ile Asn Val Ser Val Ser Glu Ala Ala Ile Glu Ala Ser
245 250 255
Thr Arg Tyr Cys Arg Ala Asn Gly Thr Val Val Leu Val Gly Leu Pro
260 265 270
Ala Gly Ala Lys Cys Ser Ser Asp Val Phe Asn His Val Val Lys Ser
275 280 285
Ile Ser Ile Val Gly Ser Tyr Val Gly Asn Arg Ala Asp Thr Arg Glu
290 295 300
Ala Leu Asp Phe Phe Ala Arg Gly Leu Val Lys Ser Pro Ile Lys Val
305 310 315 320
Val Gly Leu Ser Ser Leu Pro Glu Ile Tyr Glu Lys Met Glu Lys Gly
325 330 335
Gln Ile Ala Gly Arg Tyr Val Val Asp Thr Ser Lys
340 345
<210> 9
<211> 1803
<212> DNA
<213> 酿酒酵母(Saccharomyces cerevisiae)
<220>
<221> CDS
<222> (1)..(1803)
<400> 9
atg ttg tgt tca gta att cag aga cag aca aga gag gtt tcc aac aca 48
Met Leu Cys Ser Val Ile Gln Arg Gln Thr Arg Glu Val Ser Asn Thr
1 5 10 15
atg tct tta gac tca tac tat ctt ggg ttt gat ctt tcg acc caa caa 96
Met Ser Leu Asp Ser Tyr Tyr Leu Gly Phe Asp Leu Ser Thr Gln Gln
20 25 30
ctg aaa tgt ctc gcc att aac cag gac cta aaa att gtc cat tca gaa 144
Leu Lys Cys Leu Ala Ile Asn Gln Asp Leu Lys Ile Val His Ser Glu
35 40 45
aca gtg gaa ttt gaa aag gat ctt ccg cat tat cac aca aag aag ggt 192
Thr Val Glu Phe Glu Lys Asp Leu Pro His Tyr His Thr Lys Lys Gly
50 55 60
gtc tat ata cac ggc gac act atc gaa tgt ccc gta gcc atg tgg tta 240
Val Tyr Ile His Gly Asp Thr Ile Glu Cys Pro Val Ala Met Trp Leu
65 70 75 80
gag gct cta gat ctg gtt ctc tcg aaa tat cgc gag gct aaa ttt cca 288
Glu Ala Leu Asp Leu Val Leu Ser Lys Tyr Arg Glu Ala Lys Phe Pro
85 90 95
ttg aac aaa gtt atg gcc gtc tca ggg tcc tgc cag cag cac ggg tct 336
Leu Asn Lys Val Met Ala Val Ser Gly Ser Cys Gln Gln His Gly Ser
100 105 110
gtc tac tgg tcc tcc caa gcc gaa tct ctg tta gag caa ttg aat aag 384
Val Tyr Trp Ser Ser Gln Ala Glu Ser Leu Leu Glu Gln Leu Asn Lys
115 120 125
aaa ccg gaa aaa gat tta ttg cac tac gtg agc tct gta gca ttt gca 432
Lys Pro Glu Lys Asp Leu Leu His Tyr Val Ser Ser Val Ala Phe Ala
130 135 140
agg caa acc gcc ccc aat tgg caa gac cac agt act gca aag caa tgt 480
Arg Gln Thr Ala Pro Asn Trp Gln Asp His Ser Thr Ala Lys Gln Cys
145 150 155 160
caa gag ttt gaa gag tgc ata ggt ggg cct gaa aaa atg gct caa tta 528
Gln Glu Phe Glu Glu Cys Ile Gly Gly Pro Glu Lys Met Ala Gln Leu
165 170 175
aca ggg tcc aga gcc cat ttt aga ttt act ggt cct caa att ctg aaa 576
Thr Gly Ser Arg Ala His Phe Arg Phe Thr Gly Pro Gln Ile Leu Lys
180 185 190
att gca caa tta gaa cca gaa gct tac gaa aaa aca aag acc att tct 624
Ile Ala Gln Leu Glu Pro Glu Ala Tyr Glu Lys Thr Lys Thr Ile Ser
195 200 205
tta gtg tct aat ttt ttg act tct atc tta gtg ggc cat ctt gtt gaa 672
Leu Val Ser Asn Phe Leu Thr Ser Ile Leu Val Gly His Leu Val Glu
210 215 220
tta gag gag gca gat gcc tgt ggt atg aac ctt tat gat ata cgt gaa 720
Leu Glu Glu Ala Asp Ala Cys Gly Met Asn Leu Tyr Asp Ile Arg Glu
225 230 235 240
aga aaa ttc agt gat gag cta cta cat cta att gat agt tct tct aag 768
Arg Lys Phe Ser Asp Glu Leu Leu His Leu Ile Asp Ser Ser Ser Lys
245 250 255
gat aaa act atc aga caa aaa tta atg aga gca ccc atg aaa aat ttg 816
Asp Lys Thr Ile Arg Gln Lys Leu Met Arg Ala Pro Met Lys Asn Leu
260 265 270
ata gcg ggt acc atc tgt aaa tat ttt att gag aag tac ggt ttc aat 864
Ile Ala Gly Thr Ile Cys Lys Tyr Phe Ile Glu Lys Tyr Gly Phe Asn
275 280 285
aca aac tgc aag gtc tct ccc atg act ggg gat aat tta gcc act ata 912
Thr Asn Cys Lys Val Ser Pro Met Thr Gly Asp Asn Leu Ala Thr Ile
290 295 300
tgt tct tta ccc ctg cgg aag aat gac gtt ctc gtt tcc cta gga aca 960
Cys Ser Leu Pro Leu Arg Lys Asn Asp Val Leu Val Ser Leu Gly Thr
305 310 315 320
agt act aca gtt ctt ctg gtc acc gat aag tat cac ccc tct ccg aac 1008
Ser Thr Thr Val Leu Leu Val Thr Asp Lys Tyr His Pro Ser Pro Asn
325 330 335
tat cat ctt ttc att cat cca act ctg cca aac cat tat atg ggt atg 1056
Tyr His Leu Phe Ile His Pro Thr Leu Pro Asn His Tyr Met Gly Met
340 345 350
att tgt tat tgt aat ggt tct ttg gca agg gag agg ata aga gac gag 1104
Ile Cys Tyr Cys Asn Gly Ser Leu Ala Arg Glu Arg Ile Arg Asp Glu
355 360 365
tta aac aaa gaa cgg gaa aat aat tat gag aag act aac gat tgg act 1152
Leu Asn Lys Glu Arg Glu Asn Asn Tyr Glu Lys Thr Asn Asp Trp Thr
370 375 380
ctt ttt aat caa gct gtg cta gat gac tca gaa agt agt gaa aat gaa 1200
Leu Phe Asn Gln Ala Val Leu Asp Asp Ser Glu Ser Ser Glu Asn Glu
385 390 395 400
tta ggt gta tat ttt cct ctg ggg gag atc gtt cct agc gta aaa gcc 1248
Leu Gly Val Tyr Phe Pro Leu Gly Glu Ile Val Pro Ser Val Lys Ala
405 410 415
ata aac aaa agg gtt atc ttc aat cca aaa acg ggt atg att gaa aga 1296
Ile Asn Lys Arg Val Ile Phe Asn Pro Lys Thr Gly Met Ile Glu Arg
420 425 430
gag gtg gcc aag ttc aaa gac aag agg cac gat gcc aaa aat att gta 1344
Glu Val Ala Lys Phe Lys Asp Lys Arg His Asp Ala Lys Asn Ile Val
435 440 445
gaa tca cag gct tta agt tgc agg gta aga ata tct ccc ctg ctt tcg 1392
Glu Ser Gln Ala Leu Ser Cys Arg Val Arg Ile Ser Pro Leu Leu Ser
450 455 460
gat tca aac gca agc tca caa cag aga ctg aac gaa gat aca atc gtg 1440
Asp Ser Asn Ala Ser Ser Gln Gln Arg Leu Asn Glu Asp Thr Ile Val
465 470 475 480
aag ttt gat tac gat gaa tct ccg ctg cgg gac tac cta aat aaa agg 1488
Lys Phe Asp Tyr Asp Glu Ser Pro Leu Arg Asp Tyr Leu Asn Lys Arg
485 490 495
cca gaa agg act ttt ttt gta ggt ggg gct tct aaa aac gat gct att 1536
Pro Glu Arg Thr Phe Phe Val Gly Gly Ala Ser Lys Asn Asp Ala Ile
500 505 510
gtg aag aag ttt gct caa gtc att ggt gct aca aag ggt aat ttt agg 1584
Val Lys Lys Phe Ala Gln Val Ile Gly Ala Thr Lys Gly Asn Phe Arg
515 520 525
cta gaa aca cca aac tca tgt gcc ctt ggt ggt tgt tat aag gcc atg 1632
Leu Glu Thr Pro Asn Ser Cys Ala Leu Gly Gly Cys Tyr Lys Ala Met
530 535 540
tgg tca ttg tta tat gac tct aat aaa att gca gtt cct ttt gat aaa 1680
Trp Ser Leu Leu Tyr Asp Ser Asn Lys Ile Ala Val Pro Phe Asp Lys
545 550 555 560
ttt ctg aat gac aat ttt cca tgg cat gta atg gaa agc ata tcc gat 1728
Phe Leu Asn Asp Asn Phe Pro Trp His Val Met Glu Ser Ile Ser Asp
565 570 575
gtg gat aat gaa aat tgg gat cgc tat aat tcc aag att gtc ccc tta 1776
Val Asp Asn Glu Asn Trp Asp Arg Tyr Asn Ser Lys Ile Val Pro Leu
580 585 590
agc gaa ctg gaa aag act ctc atc taa 1803
Ser Glu Leu Glu Lys Thr Leu Ile
595 600
<210> 10
<211> 600
<212> PRT
<213> 酿酒酵母(Saccharomyces cerevisiae)
<400> 10
Met Leu Cys Ser Val Ile Gln Arg Gln Thr Arg Glu Val Ser Asn Thr
1 5 10 15
Met Ser Leu Asp Ser Tyr Tyr Leu Gly Phe Asp Leu Ser Thr Gln Gln
20 25 30
Leu Lys Cys Leu Ala Ile Asn Gln Asp Leu Lys Ile Val His Ser Glu
35 40 45
Thr Val Glu Phe Glu Lys Asp Leu Pro His Tyr His Thr Lys Lys Gly
50 55 60
Val Tyr Ile His Gly Asp Thr Ile Glu Cys Pro Val Ala Met Trp Leu
65 70 75 80
Glu Ala Leu Asp Leu Val Leu Ser Lys Tyr Arg Glu Ala Lys Phe Pro
85 90 95
Leu Asn Lys Val Met Ala Val Ser Gly Ser Cys Gln Gln His Gly Ser
100 105 110
Val Tyr Trp Ser Ser Gln Ala Glu Ser Leu Leu Glu Gln Leu Asn Lys
115 120 125
Lys Pro Glu Lys Asp Leu Leu His Tyr Val Ser Ser Val Ala Phe Ala
130 135 140
Arg Gln Thr Ala Pro Asn Trp Gln Asp His Ser Thr Ala Lys Gln Cys
145 150 155 160
Gln Glu Phe Glu Glu Cys Ile Gly Gly Pro Glu Lys Met Ala Gln Leu
165 170 175
Thr Gly Ser Arg Ala His Phe Arg Phe Thr Gly Pro Gln Ile Leu Lys
180 185 190
Ile Ala Gln Leu Glu Pro Glu Ala Tyr Glu Lys Thr Lys Thr Ile Ser
195 200 205
Leu Val Ser Asn Phe Leu Thr Ser Ile Leu Val Gly His Leu Val Glu
210 215 220
Leu Glu Glu Ala Asp Ala Cys Gly Met Asn Leu Tyr Asp Ile Arg Glu
225 230 235 240
Arg Lys Phe Ser Asp Glu Leu Leu His Leu Ile Asp Ser Ser Ser Lys
245 250 255
Asp Lys Thr Ile Arg Gln Lys Leu Met Arg Ala Pro Met Lys Asn Leu
260 265 270
Ile Ala Gly Thr Ile Cys Lys Tyr Phe Ile Glu Lys Tyr Gly Phe Asn
275 280 285
Thr Asn Cys Lys Val Ser Pro Met Thr Gly Asp Asn Leu Ala Thr Ile
290 295 300
Cys Ser Leu Pro Leu Arg Lys Asn Asp Val Leu Val Ser Leu Gly Thr
305 310 315 320
Ser Thr Thr Val Leu Leu Val Thr Asp Lys Tyr His Pro Ser Pro Asn
325 330 335
Tyr His Leu Phe Ile His Pro Thr Leu Pro Asn His Tyr Met Gly Met
340 345 350
Ile Cys Tyr Cys Asn Gly Ser Leu Ala Arg Glu Arg Ile Arg Asp Glu
355 360 365
Leu Asn Lys Glu Arg Glu Asn Asn Tyr Glu Lys Thr Asn Asp Trp Thr
370 375 380
Leu Phe Asn Gln Ala Val Leu Asp Asp Ser Glu Ser Ser Glu Asn Glu
385 390 395 400
Leu Gly Val Tyr Phe Pro Leu Gly Glu Ile Val Pro Ser Val Lys Ala
405 410 415
Ile Asn Lys Arg Val Ile Phe Asn Pro Lys Thr Gly Met Ile Glu Arg
420 425 430
Glu Val Ala Lys Phe Lys Asp Lys Arg His Asp Ala Lys Asn Ile Val
435 440 445
Glu Ser Gln Ala Leu Ser Cys Arg Val Arg Ile Ser Pro Leu Leu Ser
450 455 460
Asp Ser Asn Ala Ser Ser Gln Gln Arg Leu Asn Glu Asp Thr Ile Val
465 470 475 480
Lys Phe Asp Tyr Asp Glu Ser Pro Leu Arg Asp Tyr Leu Asn Lys Arg
485 490 495
Pro Glu Arg Thr Phe Phe Val Gly Gly Ala Ser Lys Asn Asp Ala Ile
500 505 510
Val Lys Lys Phe Ala Gln Val Ile Gly Ala Thr Lys Gly Asn Phe Arg
515 520 525
Leu Glu Thr Pro Asn Ser Cys Ala Leu Gly Gly Cys Tyr Lys Ala Met
530 535 540
Trp Ser Leu Leu Tyr Asp Ser Asn Lys Ile Ala Val Pro Phe Asp Lys
545 550 555 560
Phe Leu Asn Asp Asn Phe Pro Trp His Val Met Glu Ser Ile Ser Asp
565 570 575
Val Asp Asn Glu Asn Trp Asp Arg Tyr Asn Ser Lys Ile Val Pro Leu
580 585 590
Ser Glu Leu Glu Lys Thr Leu Ile
595 600
<210> 11
<211> 1008
<212> DNA
<213> 酿酒酵母(Saccharomyces cerevisiae)
<220>
<221> CDS
<222> (1)..(1008)
<400> 11
atg tct gaa cca gct caa aag aaa caa aag gtt gct aac aac tct cta 48
Met Ser Glu Pro Ala Gln Lys Lys Gln Lys Val Ala Asn Asn Ser Leu
1 5 10 15
gaa caa ttg aaa gcc tcc ggc act gtc gtt gtt gcc gac act ggt gat 96
Glu Gln Leu Lys Ala Ser Gly Thr Val Val Val Ala Asp Thr Gly Asp
20 25 30
ttc ggc tct att gcc aag ttt caa cct caa gac tcc aca act aac cca 144
Phe Gly Ser Ile Ala Lys Phe Gln Pro Gln Asp Ser Thr Thr Asn Pro
35 40 45
tca ttg atc ttg gct gct gcc aag caa cca act tac gcc aag ttg atc 192
Ser Leu Ile Leu Ala Ala Ala Lys Gln Pro Thr Tyr Ala Lys Leu Ile
50 55 60
gat gtt gcc gtg gaa tac ggt aag aag cat ggt aag acc acc gaa gaa 240
Asp Val Ala Val Glu Tyr Gly Lys Lys His Gly Lys Thr Thr Glu Glu
65 70 75 80
caa gtc gaa aat gct gtg gac aga ttg tta gtc gaa ttc ggt aag gag 288
Gln Val Glu Asn Ala Val Asp Arg Leu Leu Val Glu Phe Gly Lys Glu
85 90 95
atc tta aag att gtt cca ggc aga gtc tcc acc gaa gtt gat gct aga 336
Ile Leu Lys Ile Val Pro Gly Arg Val Ser Thr Glu Val Asp Ala Arg
100 105 110
ttg tct ttt gac act caa gct acc att gaa aag gct aga cat atc att 384
Leu Ser Phe Asp Thr Gln Ala Thr Ile Glu Lys Ala Arg His Ile Ile
115 120 125
aaa ttg ttt gaa caa gaa ggt gtc tcc aag gaa aga gtc ctt att aaa 432
Lys Leu Phe Glu Gln Glu Gly Val Ser Lys Glu Arg Val Leu Ile Lys
130 135 140
att gct tcc act tgg gaa ggt att caa gct gcc aaa gaa ttg gaa gaa 480
Ile Ala Ser Thr Trp Glu Gly Ile Gln Ala Ala Lys Glu Leu Glu Glu
145 150 155 160
aag gac ggt atc cac tgt aat ttg act cta tta ttc tcc ttc gtt caa 528
Lys Asp Gly Ile His Cys Asn Leu Thr Leu Leu Phe Ser Phe Val Gln
165 170 175
gca gtt gcc tgt gcc gag gcc caa gtt act ttg att tcc cca ttt gtt 576
Ala Val Ala Cys Ala Glu Ala Gln Val Thr Leu Ile Ser Pro Phe Val
180 185 190
ggt aga att cta gac tgg tac aaa tcc agc act ggt aaa gat tac aag 624
Gly Arg Ile Leu Asp Trp Tyr Lys Ser Ser Thr Gly Lys Asp Tyr Lys
195 200 205
ggt gaa gcc gac cca ggt gtt att tcc gtc aag aaa atc tac aac tac 672
Gly Glu Ala Asp Pro Gly Val Ile Ser Val Lys Lys Ile Tyr Asn Tyr
210 215 220
tac aag aag tac ggt tac aag act att gtt atg ggt gct tct ttc aga 720
Tyr Lys Lys Tyr Gly Tyr Lys Thr Ile Val Met Gly Ala Ser Phe Arg
225 230 235 240
agc act gac gaa atc aaa aac ttg gct ggt gtt gac tat cta aca att 768
Ser Thr Asp Glu Ile Lys Asn Leu Ala Gly Val Asp Tyr Leu Thr Ile
245 250 255
tct cca gct tta ttg gac aag ttg atg aac agt act gaa cct ttc cca 816
Ser Pro Ala Leu Leu Asp Lys Leu Met Asn Ser Thr Glu Pro Phe Pro
260 265 270
aga gtt ttg gac cct gtc tcc gct aag aag gaa gcc ggc gac aag att 864
Arg Val Leu Asp Pro Val Ser Ala Lys Lys Glu Ala Gly Asp Lys Ile
275 280 285
tct tac atc agc gac gaa tct aaa ttc aga ttc gac ttg aat gaa gac 912
Ser Tyr Ile Ser Asp Glu Ser Lys Phe Arg Phe Asp Leu Asn Glu Asp
290 295 300
gct atg gcc act gaa aaa ttg tcc gaa ggt atc aga aaa ttc tct gcc 960
Ala Met Ala Thr Glu Lys Leu Ser Glu Gly Ile Arg Lys Phe Ser Ala
305 310 315 320
gat att gtt act cta ttc gac ttg att gaa aag aaa gtt acc gct taa 1008
Asp Ile Val Thr Leu Phe Asp Leu Ile Glu Lys Lys Val Thr Ala
325 330 335
<210> 12
<211> 335
<212> PRT
<213> 酿酒酵母(Saccharomyces cerevisiae)
<400> 12
Met Ser Glu Pro Ala Gln Lys Lys Gln Lys Val Ala Asn Asn Ser Leu
1 5 10 15
Glu Gln Leu Lys Ala Ser Gly Thr Val Val Val Ala Asp Thr Gly Asp
20 25 30
Phe Gly Ser Ile Ala Lys Phe Gln Pro Gln Asp Ser Thr Thr Asn Pro
35 40 45
Ser Leu Ile Leu Ala Ala Ala Lys Gln Pro Thr Tyr Ala Lys Leu Ile
50 55 60
Asp Val Ala Val Glu Tyr Gly Lys Lys His Gly Lys Thr Thr Glu Glu
65 70 75 80
Gln Val Glu Asn Ala Val Asp Arg Leu Leu Val Glu Phe Gly Lys Glu
85 90 95
Ile Leu Lys Ile Val Pro Gly Arg Val Ser Thr Glu Val Asp Ala Arg
100 105 110
Leu Ser Phe Asp Thr Gln Ala Thr Ile Glu Lys Ala Arg His Ile Ile
115 120 125
Lys Leu Phe Glu Gln Glu Gly Val Ser Lys Glu Arg Val Leu Ile Lys
130 135 140
Ile Ala Ser Thr Trp Glu Gly Ile Gln Ala Ala Lys Glu Leu Glu Glu
145 150 155 160
Lys Asp Gly Ile His Cys Asn Leu Thr Leu Leu Phe Ser Phe Val Gln
165 170 175
Ala Val Ala Cys Ala Glu Ala Gln Val Thr Leu Ile Ser Pro Phe Val
180 185 190
Gly Arg Ile Leu Asp Trp Tyr Lys Ser Ser Thr Gly Lys Asp Tyr Lys
195 200 205
Gly Glu Ala Asp Pro Gly Val Ile Ser Val Lys Lys Ile Tyr Asn Tyr
210 215 220
Tyr Lys Lys Tyr Gly Tyr Lys Thr Ile Val Met Gly Ala Ser Phe Arg
225 230 235 240
Ser Thr Asp Glu Ile Lys Asn Leu Ala Gly Val Asp Tyr Leu Thr Ile
245 250 255
Ser Pro Ala Leu Leu Asp Lys Leu Met Asn Ser Thr Glu Pro Phe Pro
260 265 270
Arg Val Leu Asp Pro Val Ser Ala Lys Lys Glu Ala Gly Asp Lys Ile
275 280 285
Ser Tyr Ile Ser Asp Glu Ser Lys Phe Arg Phe Asp Leu Asn Glu Asp
290 295 300
Ala Met Ala Thr Glu Lys Leu Ser Glu Gly Ile Arg Lys Phe Ser Ala
305 310 315 320
Asp Ile Val Thr Leu Phe Asp Leu Ile Glu Lys Lys Val Thr Ala
325 330 335
<210> 13
<211> 2043
<212> DNA
<213> 酿酒酵母(Saccharomyces cerevisiae)
<220>
<221> CDS
<222> (1)..(2043)
<400> 13
atg act caa ttc act gac att gat aag cta gcc gtc tcc acc ata aga 48
Met Thr Gln Phe Thr Asp Ile Asp Lys Leu Ala Val Ser Thr Ile Arg
1 5 10 15
att ttg gct gtg gac acc gta tcc aag gcc aac tca ggt cac cca ggt 96
Ile Leu Ala Val Asp Thr Val Ser Lys Ala Asn Ser Gly His Pro Gly
20 25 30
gct cca ttg ggt atg gca cca gct gca cac gtt cta tgg agt caa atg 144
Ala Pro Leu Gly Met Ala Pro Ala Ala His Val Leu Trp Ser Gln Met
35 40 45
cgc atg aac cca acc aac cca gac tgg atc aac aga gat aga ttt gtc 192
Arg Met Asn Pro Thr Asn Pro Asp Trp Ile Asn Arg Asp Arg Phe Val
50 55 60
ttg tct aac ggt cac gcg gtc gct ttg ttg tat tct atg cta cat ttg 240
Leu Ser Asn Gly His Ala Val Ala Leu Leu Tyr Ser Met Leu His Leu
65 70 75 80
act ggt tac gat ctg tct att gaa gac ttg aaa cag ttc aga cag ttg 288
Thr Gly Tyr Asp Leu Ser Ile Glu Asp Leu Lys Gln Phe Arg Gln Leu
85 90 95
ggt tcc aga aca cca ggt cat cct gaa ttt gag ttg cca ggt gtt gaa 336
Gly Ser Arg Thr Pro Gly His Pro Glu Phe Glu Leu Pro Gly Val Glu
100 105 110
gtt act acc ggt cca tta ggt caa ggt atc tcc aac gct gtt ggt atg 384
Val Thr Thr Gly Pro Leu Gly Gln Gly Ile Ser Asn Ala Val Gly Met
115 120 125
gcc atg gct caa gct aac ctg gct gcc act tac aac aag ccg ggc ttt 432
Ala Met Ala Gln Ala Asn Leu Ala Ala Thr Tyr Asn Lys Pro Gly Phe
130 135 140
acc ttg tct gac aac tac acc tat gtt ttc ttg ggt gac ggt tgt ttg 480
Thr Leu Ser Asp Asn Tyr Thr Tyr Val Phe Leu Gly Asp Gly Cys Leu
145 150 155 160
caa gaa ggt att tct tca gaa gct tcc tcc ttg gct ggt cat ttg aaa 528
Gln Glu Gly Ile Ser Ser Glu Ala Ser Ser Leu Ala Gly His Leu Lys
165 170 175
ttg ggt aac ttg att gcc atc tac gat gac aac aag atc act atc gat 576
Leu Gly Asn Leu Ile Ala Ile Tyr Asp Asp Asn Lys Ile Thr Ile Asp
180 185 190
ggt gct acc agt atc tca ttc gat gaa gat gtt gct aag aga tac gaa 624
Gly Ala Thr Ser Ile Ser Phe Asp Glu Asp Val Ala Lys Arg Tyr Glu
195 200 205
gcc tac ggt tgg gaa gtt ttg tac gta gaa aat ggt aac gaa gat cta 672
Ala Tyr Gly Trp Glu Val Leu Tyr Val Glu Asn Gly Asn Glu Asp Leu
210 215 220
gcc ggt att gcc aag gct att gct caa gct aag tta tcc aag gac aaa 720
Ala Gly Ile Ala Lys Ala Ile Ala Gln Ala Lys Leu Ser Lys Asp Lys
225 230 235 240
cca act ttg atc aaa atg acc aca acc att ggt tac ggt tcc ttg cat 768
Pro Thr Leu Ile Lys Met Thr Thr Thr Ile Gly Tyr Gly Ser Leu His
245 250 255
gcc ggc tct cac tct gtg cac ggt gcc cca ttg aaa gca gat gat gtt 816
Ala Gly Ser His Ser Val His Gly Ala Pro Leu Lys Ala Asp Asp Val
260 265 270
aaa caa cta aag agc aaa ttc ggt ttc aac cca gac aag tcc ttt gtt 864
Lys Gln Leu Lys Ser Lys Phe Gly Phe Asn Pro Asp Lys Ser Phe Val
275 280 285
gtt cca caa gaa gtt tac gac cac tac caa aag aca att tta aag cca 912
Val Pro Gln Glu Val Tyr Asp His Tyr Gln Lys Thr Ile Leu Lys Pro
290 295 300
ggt gtc gaa gcc aac aac aag tgg aac aag ttg ttc agc gaa tac caa 960
Gly Val Glu Ala Asn Asn Lys Trp Asn Lys Leu Phe Ser Glu Tyr Gln
305 310 315 320
aag aaa ttc cca gaa tta ggt gct gaa ttg gct aga aga ttg agc ggc 1008
Lys Lys Phe Pro Glu Leu Gly Ala Glu Leu Ala Arg Arg Leu Ser Gly
325 330 335
caa cta ccc gca aat tgg gaa tct aag ttg cca act tac acc gcc aag 1056
Gln Leu Pro Ala Asn Trp Glu Ser Lys Leu Pro Thr Tyr Thr Ala Lys
340 345 350
gac tct gcc gtg gcc act aga aaa tta tca gaa act gtt ctt gag gat 1104
Asp Ser Ala Val Ala Thr Arg Lys Leu Ser Glu Thr Val Leu Glu Asp
355 360 365
gtt tac aat caa ttg cca gag ttg att ggt ggt tct gcc gat tta aca 1152
Val Tyr Asn Gln Leu Pro Glu Leu Ile Gly Gly Ser Ala Asp Leu Thr
370 375 380
cct tct aac ttg acc aga tgg aag gaa gcc ctt gac ttc caa cct cct 1200
Pro Ser Asn Leu Thr Arg Trp Lys Glu Ala Leu Asp Phe Gln Pro Pro
385 390 395 400
tct tcc ggt tca ggt aac tac tct ggt aga tac att agg tac ggt att 1248
Ser Ser Gly Ser Gly Asn Tyr Ser Gly Arg Tyr Ile Arg Tyr Gly Ile
405 410 415
aga gaa cac gct atg ggt gcc ata atg aac ggt att tca gct ttc ggt 1296
Arg Glu His Ala Met Gly Ala Ile Met Asn Gly Ile Ser Ala Phe Gly
420 425 430
gcc aac tac aaa cca tac ggt ggt act ttc ttg aac ttc gtt tct tat 1344
Ala Asn Tyr Lys Pro Tyr Gly Gly Thr Phe Leu Asn Phe Val Ser Tyr
435 440 445
gct gct ggt gcc gtt aga ttg tcc gct ttg tct ggc cac cca gtt att 1392
Ala Ala Gly Ala Val Arg Leu Ser Ala Leu Ser Gly His Pro Val Ile
450 455 460
tgg gtt gct aca cat gac tct atc ggt gtc ggt gaa gat ggt cca aca 1440
Trp Val Ala Thr His Asp Ser Ile Gly Val Gly Glu Asp Gly Pro Thr
465 470 475 480
cat caa cct att gaa act tta gca cac ttc aga tcc cta cca aac att 1488
His Gln Pro Ile Glu Thr Leu Ala His Phe Arg Ser Leu Pro Asn Ile
485 490 495
caa gtt tgg aga cca gct gat ggt aac gaa gtt tct gcc gcc tac aag 1536
Gln Val Trp Arg Pro Ala Asp Gly Asn Glu Val Ser Ala Ala Tyr Lys
500 505 510
aac tct tta gaa tcc aag cat act cca agt atc att gct ttg tcc aga 1584
Asn Ser Leu Glu Ser Lys His Thr Pro Ser Ile Ile Ala Leu Ser Arg
515 520 525
caa aac ttg cca caa ttg gaa ggt agc tct att gaa agc gct tct aag 1632
Gln Asn Leu Pro Gln Leu Glu Gly Ser Ser Ile Glu Ser Ala Ser Lys
530 535 540
ggt ggt tac gta cta caa gat gtt gct aac cca gat att att tta gtg 1680
Gly Gly Tyr Val Leu Gln Asp Val Ala Asn Pro Asp Ile Ile Leu Val
545 550 555 560
gct act ggt tcc gaa gtg tct ttg agt gtt gaa gct gct aag act ttg 1728
Ala Thr Gly Ser Glu Val Ser Leu Ser Val Glu Ala Ala Lys Thr Leu
565 570 575
gcc gca aag aac atc aag gct cgt gtt gtt tct cta cca gat ttc ttc 1776
Ala Ala Lys Asn Ile Lys Ala Arg Val Val Ser Leu Pro Asp Phe Phe
580 585 590
act ttt gac aaa caa ccc cta gaa tac aga cta tca gtc tta cca gac 1824
Thr Phe Asp Lys Gln Pro Leu Glu Tyr Arg Leu Ser Val Leu Pro Asp
595 600 605
aac gtt cca atc atg tct gtt gaa gtt ttg gct acc aca tgt tgg ggc 1872
Asn Val Pro Ile Met Ser Val Glu Val Leu Ala Thr Thr Cys Trp Gly
610 615 620
aaa tac gct cat caa tcc ttc ggt att gac aga ttt ggt gcc tcc ggt 1920
Lys Tyr Ala His Gln Ser Phe Gly Ile Asp Arg Phe Gly Ala Ser Gly
625 630 635 640
aag gca cca gaa gtc ttc aag ttc ttc ggt ttc acc cca gaa ggt gtt 1968
Lys Ala Pro Glu Val Phe Lys Phe Phe Gly Phe Thr Pro Glu Gly Val
645 650 655
gct gaa aga gct caa aag acc att gca ttc tat aag ggt gac aag cta 2016
Ala Glu Arg Ala Gln Lys Thr Ile Ala Phe Tyr Lys Gly Asp Lys Leu
660 665 670
att tct cct ttg aaa aaa gct ttc taa 2043
Ile Ser Pro Leu Lys Lys Ala Phe
675 680
<210> 14
<211> 680
<212> PRT
<213> 酿酒酵母(Saccharomyces cerevisiae)
<400> 14
Met Thr Gln Phe Thr Asp Ile Asp Lys Leu Ala Val Ser Thr Ile Arg
1 5 10 15
Ile Leu Ala Val Asp Thr Val Ser Lys Ala Asn Ser Gly His Pro Gly
20 25 30
Ala Pro Leu Gly Met Ala Pro Ala Ala His Val Leu Trp Ser Gln Met
35 40 45
Arg Met Asn Pro Thr Asn Pro Asp Trp Ile Asn Arg Asp Arg Phe Val
50 55 60
Leu Ser Asn Gly His Ala Val Ala Leu Leu Tyr Ser Met Leu His Leu
65 70 75 80
Thr Gly Tyr Asp Leu Ser Ile Glu Asp Leu Lys Gln Phe Arg Gln Leu
85 90 95
Gly Ser Arg Thr Pro Gly His Pro Glu Phe Glu Leu Pro Gly Val Glu
100 105 110
Val Thr Thr Gly Pro Leu Gly Gln Gly Ile Ser Asn Ala Val Gly Met
115 120 125
Ala Met Ala Gln Ala Asn Leu Ala Ala Thr Tyr Asn Lys Pro Gly Phe
130 135 140
Thr Leu Ser Asp Asn Tyr Thr Tyr Val Phe Leu Gly Asp Gly Cys Leu
145 150 155 160
Gln Glu Gly Ile Ser Ser Glu Ala Ser Ser Leu Ala Gly His Leu Lys
165 170 175
Leu Gly Asn Leu Ile Ala Ile Tyr Asp Asp Asn Lys Ile Thr Ile Asp
180 185 190
Gly Ala Thr Ser Ile Ser Phe Asp Glu Asp Val Ala Lys Arg Tyr Glu
195 200 205
Ala Tyr Gly Trp Glu Val Leu Tyr Val Glu Asn Gly Asn Glu Asp Leu
210 215 220
Ala Gly Ile Ala Lys Ala Ile Ala Gln Ala Lys Leu Ser Lys Asp Lys
225 230 235 240
Pro Thr Leu Ile Lys Met Thr Thr Thr Ile Gly Tyr Gly Ser Leu His
245 250 255
Ala Gly Ser His Ser Val His Gly Ala Pro Leu Lys Ala Asp Asp Val
260 265 270
Lys Gln Leu Lys Ser Lys Phe Gly Phe Asn Pro Asp Lys Ser Phe Val
275 280 285
Val Pro Gln Glu Val Tyr Asp His Tyr Gln Lys Thr Ile Leu Lys Pro
290 295 300
Gly Val Glu Ala Asn Asn Lys Trp Asn Lys Leu Phe Ser Glu Tyr Gln
305 310 315 320
Lys Lys Phe Pro Glu Leu Gly Ala Glu Leu Ala Arg Arg Leu Ser Gly
325 330 335
Gln Leu Pro Ala Asn Trp Glu Ser Lys Leu Pro Thr Tyr Thr Ala Lys
340 345 350
Asp Ser Ala Val Ala Thr Arg Lys Leu Ser Glu Thr Val Leu Glu Asp
355 360 365
Val Tyr Asn Gln Leu Pro Glu Leu Ile Gly Gly Ser Ala Asp Leu Thr
370 375 380
Pro Ser Asn Leu Thr Arg Trp Lys Glu Ala Leu Asp Phe Gln Pro Pro
385 390 395 400
Ser Ser Gly Ser Gly Asn Tyr Ser Gly Arg Tyr Ile Arg Tyr Gly Ile
405 410 415
Arg Glu His Ala Met Gly Ala Ile Met Asn Gly Ile Ser Ala Phe Gly
420 425 430
Ala Asn Tyr Lys Pro Tyr Gly Gly Thr Phe Leu Asn Phe Val Ser Tyr
435 440 445
Ala Ala Gly Ala Val Arg Leu Ser Ala Leu Ser Gly His Pro Val Ile
450 455 460
Trp Val Ala Thr His Asp Ser Ile Gly Val Gly Glu Asp Gly Pro Thr
465 470 475 480
His Gln Pro Ile Glu Thr Leu Ala His Phe Arg Ser Leu Pro Asn Ile
485 490 495
Gln Val Trp Arg Pro Ala Asp Gly Asn Glu Val Ser Ala Ala Tyr Lys
500 505 510
Asn Ser Leu Glu Ser Lys His Thr Pro Ser Ile Ile Ala Leu Ser Arg
515 520 525
Gln Asn Leu Pro Gln Leu Glu Gly Ser Ser Ile Glu Ser Ala Ser Lys
530 535 540
Gly Gly Tyr Val Leu Gln Asp Val Ala Asn Pro Asp Ile Ile Leu Val
545 550 555 560
Ala Thr Gly Ser Glu Val Ser Leu Ser Val Glu Ala Ala Lys Thr Leu
565 570 575
Ala Ala Lys Asn Ile Lys Ala Arg Val Val Ser Leu Pro Asp Phe Phe
580 585 590
Thr Phe Asp Lys Gln Pro Leu Glu Tyr Arg Leu Ser Val Leu Pro Asp
595 600 605
Asn Val Pro Ile Met Ser Val Glu Val Leu Ala Thr Thr Cys Trp Gly
610 615 620
Lys Tyr Ala His Gln Ser Phe Gly Ile Asp Arg Phe Gly Ala Ser Gly
625 630 635 640
Lys Ala Pro Glu Val Phe Lys Phe Phe Gly Phe Thr Pro Glu Gly Val
645 650 655
Ala Glu Arg Ala Gln Lys Thr Ile Ala Phe Tyr Lys Gly Asp Lys Leu
660 665 670
Ile Ser Pro Leu Lys Lys Ala Phe
675 680
<210> 15
<211> 717
<212> DNA
<213> 酿酒酵母(Saccharomyces cerevisiae)
<220>
<221> CDS
<222> (1)..(717)
<400> 15
atg gtc aaa cca att ata gct ccc agt atc ctt gct tct gac ttc gcc 48
Met Val Lys Pro Ile Ile Ala Pro Ser Ile Leu Ala Ser Asp Phe Ala
1 5 10 15
aac ttg ggt tgc gaa tgt cat aag gtc atc aac gcc ggc gca gat tgg 96
Asn Leu Gly Cys Glu Cys His Lys Val Ile Asn Ala Gly Ala Asp Trp
20 25 30
tta cat atc gat gtc atg gac ggc cat ttt gtt cca aac att act ctg 144
Leu His Ile Asp Val Met Asp Gly His Phe Val Pro Asn Ile Thr Leu
35 40 45
ggc caa cca att gtt acc tcc cta cgt cgt tct gtg cca cgc cct ggc 192
Gly Gln Pro Ile Val Thr Ser Leu Arg Arg Ser Val Pro Arg Pro Gly
50 55 60
gat gct agc aac aca gaa aag aag ccc act gcg ttc ttc gat tgt cac 240
Asp Ala Ser Asn Thr Glu Lys Lys Pro Thr Ala Phe Phe Asp Cys His
65 70 75 80
atg atg gtt gaa aat cct gaa aaa tgg gtc gac gat ttt gct aaa tgt 288
Met Met Val Glu Asn Pro Glu Lys Trp Val Asp Asp Phe Ala Lys Cys
85 90 95
ggt gct gac caa ttt acg ttc cac tac gag gcc aca caa gac cct ttg 336
Gly Ala Asp Gln Phe Thr Phe His Tyr Glu Ala Thr Gln Asp Pro Leu
100 105 110
cat tta gtt aag ttg att aag tct aag ggc atc aaa gct gca tgc gcc 384
His Leu Val Lys Leu Ile Lys Ser Lys Gly Ile Lys Ala Ala Cys Ala
115 120 125
atc aaa cct ggt act tct gtt gac gtt tta ttt gaa cta gct cct cat 432
Ile Lys Pro Gly Thr Ser Val Asp Val Leu Phe Glu Leu Ala Pro His
130 135 140
ttg gat atg gct ctt gtt atg act gtg gaa cct ggg ttt gga ggc caa 480
Leu Asp Met Ala Leu Val Met Thr Val Glu Pro Gly Phe Gly Gly Gln
145 150 155 160
aaa ttc atg gaa gac atg atg cca aaa gtg gaa act ttg aga gcc aag 528
Lys Phe Met Glu Asp Met Met Pro Lys Val Glu Thr Leu Arg Ala Lys
165 170 175
ttc ccc cat ttg aat atc caa gtc gat ggt ggt ttg ggc aag gag acc 576
Phe Pro His Leu Asn Ile Gln Val Asp Gly Gly Leu Gly Lys Glu Thr
180 185 190
atc ccg aaa gcc gcc aaa gcc ggt gcc aac gtt att gtc gct ggt acc 624
Ile Pro Lys Ala Ala Lys Ala Gly Ala Asn Val Ile Val Ala Gly Thr
195 200 205
agt gtt ttc act gca gct gac ccg cac gat gtt atc tcc ttc atg aaa 672
Ser Val Phe Thr Ala Ala Asp Pro His Asp Val Ile Ser Phe Met Lys
210 215 220
gaa gaa gtc tcg aag gaa ttg cgt tct aga gat ttg cta gat tag 717
Glu Glu Val Ser Lys Glu Leu Arg Ser Arg Asp Leu Leu Asp
225 230 235
<210> 16
<211> 238
<212> PRT
<213> 酿酒酵母(Saccharomyces cerevisiae)
<400> 16
Met Val Lys Pro Ile Ile Ala Pro Ser Ile Leu Ala Ser Asp Phe Ala
1 5 10 15
Asn Leu Gly Cys Glu Cys His Lys Val Ile Asn Ala Gly Ala Asp Trp
20 25 30
Leu His Ile Asp Val Met Asp Gly His Phe Val Pro Asn Ile Thr Leu
35 40 45
Gly Gln Pro Ile Val Thr Ser Leu Arg Arg Ser Val Pro Arg Pro Gly
50 55 60
Asp Ala Ser Asn Thr Glu Lys Lys Pro Thr Ala Phe Phe Asp Cys His
65 70 75 80
Met Met Val Glu Asn Pro Glu Lys Trp Val Asp Asp Phe Ala Lys Cys
85 90 95
Gly Ala Asp Gln Phe Thr Phe His Tyr Glu Ala Thr Gln Asp Pro Leu
100 105 110
His Leu Val Lys Leu Ile Lys Ser Lys Gly Ile Lys Ala Ala Cys Ala
115 120 125
Ile Lys Pro Gly Thr Ser Val Asp Val Leu Phe Glu Leu Ala Pro His
130 135 140
Leu Asp Met Ala Leu Val Met Thr Val Glu Pro Gly Phe Gly Gly Gln
145 150 155 160
Lys Phe Met Glu Asp Met Met Pro Lys Val Glu Thr Leu Arg Ala Lys
165 170 175
Phe Pro His Leu Asn Ile Gln Val Asp Gly Gly Leu Gly Lys Glu Thr
180 185 190
Ile Pro Lys Ala Ala Lys Ala Gly Ala Asn Val Ile Val Ala Gly Thr
195 200 205
Ser Val Phe Thr Ala Ala Asp Pro His Asp Val Ile Ser Phe Met Lys
210 215 220
Glu Glu Val Ser Lys Glu Leu Arg Ser Arg Asp Leu Leu Asp
225 230 235
<210> 17
<211> 777
<212> DNA
<213> 酿酒酵母(Saccharomyces cerevisiae)
<220>
<221> CDS
<222> (1)..(777)
<400> 17
atg gct gcc ggt gtc cca aaa att gat gcg tta gaa tct ttg ggc aat 48
Met Ala Ala Gly Val Pro Lys Ile Asp Ala Leu Glu Ser Leu Gly Asn
1 5 10 15
cct ttg gag gat gcc aag aga gct gca gca tac aga gca gtt gat gaa 96
Pro Leu Glu Asp Ala Lys Arg Ala Ala Ala Tyr Arg Ala Val Asp Glu
20 25 30
aat tta aaa ttt gat gat cac aaa att att gga att ggt agt ggt agc 144
Asn Leu Lys Phe Asp Asp His Lys Ile Ile Gly Ile Gly Ser Gly Ser
35 40 45
aca gtg gtt tat gtt gcc gaa aga att gga caa tat ttg cat gac cct 192
Thr Val Val Tyr Val Ala Glu Arg Ile Gly Gln Tyr Leu His Asp Pro
50 55 60
aaa ttt tat gaa gta gcg tct aaa ttc att tgc att cca aca gga ttc 240
Lys Phe Tyr Glu Val Ala Ser Lys Phe Ile Cys Ile Pro Thr Gly Phe
65 70 75 80
caa tca aga aac ttg att ttg gat aac aag ttg caa tta ggc tcc att 288
Gln Ser Arg Asn Leu Ile Leu Asp Asn Lys Leu Gln Leu Gly Ser Ile
85 90 95
gaa cag tat cct cgc att gat ata gcg ttt gac ggt gct gat gaa gtg 336
Glu Gln Tyr Pro Arg Ile Asp Ile Ala Phe Asp Gly Ala Asp Glu Val
100 105 110
gat gag aat tta caa tta att aaa ggt ggt ggt gct tgt cta ttt caa 384
Asp Glu Asn Leu Gln Leu Ile Lys Gly Gly Gly Ala Cys Leu Phe Gln
115 120 125
gaa aaa ttg gtt agt act agt gct aaa acc ttc att gtc gtt gct gat 432
Glu Lys Leu Val Ser Thr Ser Ala Lys Thr Phe Ile Val Val Ala Asp
130 135 140
tca aga aaa aag tca cca aaa cat tta ggt aag aac tgg agg caa ggt 480
Ser Arg Lys Lys Ser Pro Lys His Leu Gly Lys Asn Trp Arg Gln Gly
145 150 155 160
gtt ccc att gaa att gta cct tcc tca tac gtg agg gtc aag aat gat 528
Val Pro Ile Glu Ile Val Pro Ser Ser Tyr Val Arg Val Lys Asn Asp
165 170 175
cta tta gaa caa ttg cat gct gaa aaa gtt gac atc aga caa gga ggt 576
Leu Leu Glu Gln Leu His Ala Glu Lys Val Asp Ile Arg Gln Gly Gly
180 185 190
tct gct aaa gca ggt cct gtt gta act gac aat aat aac ttc att atc 624
Ser Ala Lys Ala Gly Pro Val Val Thr Asp Asn Asn Asn Phe Ile Ile
195 200 205
gat gcg gat ttc ggt gaa att tcc gat cca aga aaa ttg cat aga gaa 672
Asp Ala Asp Phe Gly Glu Ile Ser Asp Pro Arg Lys Leu His Arg Glu
210 215 220
atc aaa ctg tta gtg ggc gtg gtg gaa aca ggt tta ttc atc gac aac 720
Ile Lys Leu Leu Val Gly Val Val Glu Thr Gly Leu Phe Ile Asp Asn
225 230 235 240
gct tca aaa gcc tac ttc ggt aat tct gac ggt agt gtt gaa gtt acc 768
Ala Ser Lys Ala Tyr Phe Gly Asn Ser Asp Gly Ser Val Glu Val Thr
245 250 255
gaa aag tga 777
Glu Lys
<210> 18
<211> 258
<212> PRT
<213> 酿酒酵母(Saccharomyces cerevisiae)
<400> 18
Met Ala Ala Gly Val Pro Lys Ile Asp Ala Leu Glu Ser Leu Gly Asn
1 5 10 15
Pro Leu Glu Asp Ala Lys Arg Ala Ala Ala Tyr Arg Ala Val Asp Glu
20 25 30
Asn Leu Lys Phe Asp Asp His Lys Ile Ile Gly Ile Gly Ser Gly Ser
35 40 45
Thr Val Val Tyr Val Ala Glu Arg Ile Gly Gln Tyr Leu His Asp Pro
50 55 60
Lys Phe Tyr Glu Val Ala Ser Lys Phe Ile Cys Ile Pro Thr Gly Phe
65 70 75 80
Gln Ser Arg Asn Leu Ile Leu Asp Asn Lys Leu Gln Leu Gly Ser Ile
85 90 95
Glu Gln Tyr Pro Arg Ile Asp Ile Ala Phe Asp Gly Ala Asp Glu Val
100 105 110
Asp Glu Asn Leu Gln Leu Ile Lys Gly Gly Gly Ala Cys Leu Phe Gln
115 120 125
Glu Lys Leu Val Ser Thr Ser Ala Lys Thr Phe Ile Val Val Ala Asp
130 135 140
Ser Arg Lys Lys Ser Pro Lys His Leu Gly Lys Asn Trp Arg Gln Gly
145 150 155 160
Val Pro Ile Glu Ile Val Pro Ser Ser Tyr Val Arg Val Lys Asn Asp
165 170 175
Leu Leu Glu Gln Leu His Ala Glu Lys Val Asp Ile Arg Gln Gly Gly
180 185 190
Ser Ala Lys Ala Gly Pro Val Val Thr Asp Asn Asn Asn Phe Ile Ile
195 200 205
Asp Ala Asp Phe Gly Glu Ile Ser Asp Pro Arg Lys Leu His Arg Glu
210 215 220
Ile Lys Leu Leu Val Gly Val Val Glu Thr Gly Leu Phe Ile Asp Asn
225 230 235 240
Ala Ser Lys Ala Tyr Phe Gly Asn Ser Asp Gly Ser Val Glu Val Thr
245 250 255
Glu Lys
<210> 19
<211> 2676
<212> DNA
<213> 大肠杆菌(Escherichia coli)
<220>
<221> CDS
<222> (1)..(2676)
<400> 19
atg gca gtt acg aac gtt gca gag ctt aat gcc ctt gtg gag agg gtt 48
Met Ala Val Thr Asn Val Ala Glu Leu Asn Ala Leu Val Glu Arg Val
1 5 10 15
aag aaa gca caa agg gaa tac gcc agc ttc acg caa gaa caa gtt gac 96
Lys Lys Ala Gln Arg Glu Tyr Ala Ser Phe Thr Gln Glu Gln Val Asp
20 25 30
aag att ttc agg gca gcc gcg cta gcg gca gct gat gcg aga ata ccc 144
Lys Ile Phe Arg Ala Ala Ala Leu Ala Ala Ala Asp Ala Arg Ile Pro
35 40 45
ctt gct aag atg gca gtc gca gaa agc gga atg gga att gtg gaa gat 192
Leu Ala Lys Met Ala Val Ala Glu Ser Gly Met Gly Ile Val Glu Asp
50 55 60
aag gtc atc aag aat cat ttc gct tct gag tat atc tac aac gcg tat 240
Lys Val Ile Lys Asn His Phe Ala Ser Glu Tyr Ile Tyr Asn Ala Tyr
65 70 75 80
aag gat gag aag act tgc ggc gtc tta tct gag gac gat aca ttc ggc 288
Lys Asp Glu Lys Thr Cys Gly Val Leu Ser Glu Asp Asp Thr Phe Gly
85 90 95
aca ata act atc gct gaa cct ata ggg atc atc tgt ggt ata gtt cca 336
Thr Ile Thr Ile Ala Glu Pro Ile Gly Ile Ile Cys Gly Ile Val Pro
100 105 110
act acg aac cct acc tca act gct ata ttc aag agt ctt att tct ttg 384
Thr Thr Asn Pro Thr Ser Thr Ala Ile Phe Lys Ser Leu Ile Ser Leu
115 120 125
aaa acc cgt aac gct atc atc ttt tct cca cat cca aga gcc aag gac 432
Lys Thr Arg Asn Ala Ile Ile Phe Ser Pro His Pro Arg Ala Lys Asp
130 135 140
gcc acc aac aag gcg gcg gac atc gtg ctg cag gcg gcc atc gcg gcg 480
Ala Thr Asn Lys Ala Ala Asp Ile Val Leu Gln Ala Ala Ile Ala Ala
145 150 155 160
ggc gcg cct aag gat cta att ggc tgg atc gac caa cct agc gtt gaa 528
Gly Ala Pro Lys Asp Leu Ile Gly Trp Ile Asp Gln Pro Ser Val Glu
165 170 175
ctt tcc aat gct ctt atg cac cat cct gat att aat ttg ata ctt gcg 576
Leu Ser Asn Ala Leu Met His His Pro Asp Ile Asn Leu Ile Leu Ala
180 185 190
act gga ggc cct ggg atg gta aag gcc gct tat agc agc ggt aag ccc 624
Thr Gly Gly Pro Gly Met Val Lys Ala Ala Tyr Ser Ser Gly Lys Pro
195 200 205
gcc atc ggt gtt ggc gct gga aat acg ccg gta gtc att gat gag acc 672
Ala Ile Gly Val Gly Ala Gly Asn Thr Pro Val Val Ile Asp Glu Thr
210 215 220
gcc gac att aag agg gca gtc gcg tca gtt ttg atg tct aag act ttt 720
Ala Asp Ile Lys Arg Ala Val Ala Ser Val Leu Met Ser Lys Thr Phe
225 230 235 240
gac aat gga gta atc tgc gcg tca gag cag tcc gtt gtc gtt gtg gat 768
Asp Asn Gly Val Ile Cys Ala Ser Glu Gln Ser Val Val Val Val Asp
245 250 255
tca gta tac gac gcg gtc aga gaa agg ttc gcg act cac gga ggg tac 816
Ser Val Tyr Asp Ala Val Arg Glu Arg Phe Ala Thr His Gly Gly Tyr
260 265 270
cta ctt caa ggg aag gag ttg aaa gcc gtc cag gac gtt atc ctt aag 864
Leu Leu Gln Gly Lys Glu Leu Lys Ala Val Gln Asp Val Ile Leu Lys
275 280 285
aat ggt gca ttg aac gcc gcc att gta ggc caa cca gcg tat aaa ata 912
Asn Gly Ala Leu Asn Ala Ala Ile Val Gly Gln Pro Ala Tyr Lys Ile
290 295 300
gcc gaa ctt gcg ggc ttt agc gta cca gaa aac act aaa ata ctt att 960
Ala Glu Leu Ala Gly Phe Ser Val Pro Glu Asn Thr Lys Ile Leu Ile
305 310 315 320
ggt gaa gtc acc gtg gtt gat gag tca gaa ccc ttc gcc cat gag aaa 1008
Gly Glu Val Thr Val Val Asp Glu Ser Glu Pro Phe Ala His Glu Lys
325 330 335
ctt tcc cca aca ctg gcg atg tac aga gcg aag gac ttt gaa gat gca 1056
Leu Ser Pro Thr Leu Ala Met Tyr Arg Ala Lys Asp Phe Glu Asp Ala
340 345 350
gtg gaa aaa gcc gaa aag ctt gtt gca atg ggc ggt atc ggc cac aca 1104
Val Glu Lys Ala Glu Lys Leu Val Ala Met Gly Gly Ile Gly His Thr
355 360 365
agc tgc ctt tac aca gat cag gat aac caa cca gca aga gtt agt tat 1152
Ser Cys Leu Tyr Thr Asp Gln Asp Asn Gln Pro Ala Arg Val Ser Tyr
370 375 380
ttc ggc cag aaa atg aag acg gct agg ata cta atc aac acc ccg gcc 1200
Phe Gly Gln Lys Met Lys Thr Ala Arg Ile Leu Ile Asn Thr Pro Ala
385 390 395 400
agc caa gga ggc att ggt gac ctg tat aac ttc aag ttg gct cct tct 1248
Ser Gln Gly Gly Ile Gly Asp Leu Tyr Asn Phe Lys Leu Ala Pro Ser
405 410 415
ctt aca ttg ggc tgt ggt tcc tgg ggc gga aac tct atc tct gag aac 1296
Leu Thr Leu Gly Cys Gly Ser Trp Gly Gly Asn Ser Ile Ser Glu Asn
420 425 430
gtt gga cct aag cat ctt atc aat aag aaa acc gtc gct aag cgt gct 1344
Val Gly Pro Lys His Leu Ile Asn Lys Lys Thr Val Ala Lys Arg Ala
435 440 445
gag aac atg ctt tgg cac aaa ctg ccg aag tca atc tat ttc cgt agg 1392
Glu Asn Met Leu Trp His Lys Leu Pro Lys Ser Ile Tyr Phe Arg Arg
450 455 460
ggc tcc ctg cct atc gcg tta gat gaa gtc atc acc gat ggg cat aag 1440
Gly Ser Leu Pro Ile Ala Leu Asp Glu Val Ile Thr Asp Gly His Lys
465 470 475 480
aga gca cta atc gtc aca gat agg ttc cta ttc aac aac ggt tac gca 1488
Arg Ala Leu Ile Val Thr Asp Arg Phe Leu Phe Asn Asn Gly Tyr Ala
485 490 495
gac caa atc acg tct gtt ctg aaa gcg gcc ggt gtc gaa aca gag gtg 1536
Asp Gln Ile Thr Ser Val Leu Lys Ala Ala Gly Val Glu Thr Glu Val
500 505 510
ttc ttc gaa gtg gaa gcc gac ccg aca tta agt att gtc agg aaa gga 1584
Phe Phe Glu Val Glu Ala Asp Pro Thr Leu Ser Ile Val Arg Lys Gly
515 520 525
gct gag ctt gcg aat agt ttt aag ccg gat gtc atc atc gct ttg gga 1632
Ala Glu Leu Ala Asn Ser Phe Lys Pro Asp Val Ile Ile Ala Leu Gly
530 535 540
gga gga tcc ccg atg gat gct gct aag atc atg tgg gtc atg tat gaa 1680
Gly Gly Ser Pro Met Asp Ala Ala Lys Ile Met Trp Val Met Tyr Glu
545 550 555 560
cac ccg gag aca cac ttt gaa gag ttg gcg ttg aga ttc atg gat att 1728
His Pro Glu Thr His Phe Glu Glu Leu Ala Leu Arg Phe Met Asp Ile
565 570 575
agg aaa agg atc tat aaa ttc cct aag atg gga gtg aaa gcc aag atg 1776
Arg Lys Arg Ile Tyr Lys Phe Pro Lys Met Gly Val Lys Ala Lys Met
580 585 590
ata gca gtg acg acc acc agt gga acc ggg agt gaa gtg act ccg ttt 1824
Ile Ala Val Thr Thr Thr Ser Gly Thr Gly Ser Glu Val Thr Pro Phe
595 600 605
gcg gtt gtt act gac gac gct acg ggc cag aag tat ccc cta gcc gac 1872
Ala Val Val Thr Asp Asp Ala Thr Gly Gln Lys Tyr Pro Leu Ala Asp
610 615 620
tat gca ttg acg ccg gat atg gcg att gtg gac gcg aac ctg gtt atg 1920
Tyr Ala Leu Thr Pro Asp Met Ala Ile Val Asp Ala Asn Leu Val Met
625 630 635 640
gat atg cca aag tca ctt tgc gca ttc ggt gga ctt gac gca gtt aca 1968
Asp Met Pro Lys Ser Leu Cys Ala Phe Gly Gly Leu Asp Ala Val Thr
645 650 655
cat gca atg gag gca tat gtg tcc gtg ctt gct tca gag ttt agc gat 2016
His Ala Met Glu Ala Tyr Val Ser Val Leu Ala Ser Glu Phe Ser Asp
660 665 670
gga cag gca ctt caa gcc ttg aag tta cta aag gaa tac ctt ccc gca 2064
Gly Gln Ala Leu Gln Ala Leu Lys Leu Leu Lys Glu Tyr Leu Pro Ala
675 680 685
tct tac cat gag ggc agc aag aac cct gtt gcc cgt gag aga gta cat 2112
Ser Tyr His Glu Gly Ser Lys Asn Pro Val Ala Arg Glu Arg Val His
690 695 700
agt gcg gct acc att gcc ggt atc gca ttc gct aat gct ttc ctg ggt 2160
Ser Ala Ala Thr Ile Ala Gly Ile Ala Phe Ala Asn Ala Phe Leu Gly
705 710 715 720
gtg tgt cat tca atg gcc cat aag ctt gga tcc cag ttt cac atc cca 2208
Val Cys His Ser Met Ala His Lys Leu Gly Ser Gln Phe His Ile Pro
725 730 735
cac gga tta gcc aac gct ctt ctt ata tgt aac gtc atc cgt tat aat 2256
His Gly Leu Ala Asn Ala Leu Leu Ile Cys Asn Val Ile Arg Tyr Asn
740 745 750
gcc aac gac aac cct aca aag caa acc gca ttc tca caa tat gac agg 2304
Ala Asn Asp Asn Pro Thr Lys Gln Thr Ala Phe Ser Gln Tyr Asp Arg
755 760 765
ccg cag gcg agg cgt cgt tat gca gaa ata gcc gat cac cta ggc tta 2352
Pro Gln Ala Arg Arg Arg Tyr Ala Glu Ile Ala Asp His Leu Gly Leu
770 775 780
tcc gcg cca gga gat cgt aca gct gct aag att gaa aag ttg ctt gcg 2400
Ser Ala Pro Gly Asp Arg Thr Ala Ala Lys Ile Glu Lys Leu Leu Ala
785 790 795 800
tgg tta gaa acg ctt aag gcc gaa cta ggc att cca aag tct att aga 2448
Trp Leu Glu Thr Leu Lys Ala Glu Leu Gly Ile Pro Lys Ser Ile Arg
805 810 815
gag gcg gga gtt caa gag gcg gat ttc ctt gct aac gtg gac aag ctt 2496
Glu Ala Gly Val Gln Glu Ala Asp Phe Leu Ala Asn Val Asp Lys Leu
820 825 830
agc gaa gat gca ttc gat gat cag tgc act gga gcc aac ccg aga tat 2544
Ser Glu Asp Ala Phe Asp Asp Gln Cys Thr Gly Ala Asn Pro Arg Tyr
835 840 845
cct ctg att tca gaa ttg aag caa att tta ctt gat aca tac tat ggg 2592
Pro Leu Ile Ser Glu Leu Lys Gln Ile Leu Leu Asp Thr Tyr Tyr Gly
850 855 860
aga gac tac gtt gag gga gag acc gca gcg aag aag gag gcg gcg ccg 2640
Arg Asp Tyr Val Glu Gly Glu Thr Ala Ala Lys Lys Glu Ala Ala Pro
865 870 875 880
gct aag gcc gag aag aaa gca aag aaa tca gct taa 2676
Ala Lys Ala Glu Lys Lys Ala Lys Lys Ser Ala
885 890
<210> 20
<211> 891
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 20
Met Ala Val Thr Asn Val Ala Glu Leu Asn Ala Leu Val Glu Arg Val
1 5 10 15
Lys Lys Ala Gln Arg Glu Tyr Ala Ser Phe Thr Gln Glu Gln Val Asp
20 25 30
Lys Ile Phe Arg Ala Ala Ala Leu Ala Ala Ala Asp Ala Arg Ile Pro
35 40 45
Leu Ala Lys Met Ala Val Ala Glu Ser Gly Met Gly Ile Val Glu Asp
50 55 60
Lys Val Ile Lys Asn His Phe Ala Ser Glu Tyr Ile Tyr Asn Ala Tyr
65 70 75 80
Lys Asp Glu Lys Thr Cys Gly Val Leu Ser Glu Asp Asp Thr Phe Gly
85 90 95
Thr Ile Thr Ile Ala Glu Pro Ile Gly Ile Ile Cys Gly Ile Val Pro
100 105 110
Thr Thr Asn Pro Thr Ser Thr Ala Ile Phe Lys Ser Leu Ile Ser Leu
115 120 125
Lys Thr Arg Asn Ala Ile Ile Phe Ser Pro His Pro Arg Ala Lys Asp
130 135 140
Ala Thr Asn Lys Ala Ala Asp Ile Val Leu Gln Ala Ala Ile Ala Ala
145 150 155 160
Gly Ala Pro Lys Asp Leu Ile Gly Trp Ile Asp Gln Pro Ser Val Glu
165 170 175
Leu Ser Asn Ala Leu Met His His Pro Asp Ile Asn Leu Ile Leu Ala
180 185 190
Thr Gly Gly Pro Gly Met Val Lys Ala Ala Tyr Ser Ser Gly Lys Pro
195 200 205
Ala Ile Gly Val Gly Ala Gly Asn Thr Pro Val Val Ile Asp Glu Thr
210 215 220
Ala Asp Ile Lys Arg Ala Val Ala Ser Val Leu Met Ser Lys Thr Phe
225 230 235 240
Asp Asn Gly Val Ile Cys Ala Ser Glu Gln Ser Val Val Val Val Asp
245 250 255
Ser Val Tyr Asp Ala Val Arg Glu Arg Phe Ala Thr His Gly Gly Tyr
260 265 270
Leu Leu Gln Gly Lys Glu Leu Lys Ala Val Gln Asp Val Ile Leu Lys
275 280 285
Asn Gly Ala Leu Asn Ala Ala Ile Val Gly Gln Pro Ala Tyr Lys Ile
290 295 300
Ala Glu Leu Ala Gly Phe Ser Val Pro Glu Asn Thr Lys Ile Leu Ile
305 310 315 320
Gly Glu Val Thr Val Val Asp Glu Ser Glu Pro Phe Ala His Glu Lys
325 330 335
Leu Ser Pro Thr Leu Ala Met Tyr Arg Ala Lys Asp Phe Glu Asp Ala
340 345 350
Val Glu Lys Ala Glu Lys Leu Val Ala Met Gly Gly Ile Gly His Thr
355 360 365
Ser Cys Leu Tyr Thr Asp Gln Asp Asn Gln Pro Ala Arg Val Ser Tyr
370 375 380
Phe Gly Gln Lys Met Lys Thr Ala Arg Ile Leu Ile Asn Thr Pro Ala
385 390 395 400
Ser Gln Gly Gly Ile Gly Asp Leu Tyr Asn Phe Lys Leu Ala Pro Ser
405 410 415
Leu Thr Leu Gly Cys Gly Ser Trp Gly Gly Asn Ser Ile Ser Glu Asn
420 425 430
Val Gly Pro Lys His Leu Ile Asn Lys Lys Thr Val Ala Lys Arg Ala
435 440 445
Glu Asn Met Leu Trp His Lys Leu Pro Lys Ser Ile Tyr Phe Arg Arg
450 455 460
Gly Ser Leu Pro Ile Ala Leu Asp Glu Val Ile Thr Asp Gly His Lys
465 470 475 480
Arg Ala Leu Ile Val Thr Asp Arg Phe Leu Phe Asn Asn Gly Tyr Ala
485 490 495
Asp Gln Ile Thr Ser Val Leu Lys Ala Ala Gly Val Glu Thr Glu Val
500 505 510
Phe Phe Glu Val Glu Ala Asp Pro Thr Leu Ser Ile Val Arg Lys Gly
515 520 525
Ala Glu Leu Ala Asn Ser Phe Lys Pro Asp Val Ile Ile Ala Leu Gly
530 535 540
Gly Gly Ser Pro Met Asp Ala Ala Lys Ile Met Trp Val Met Tyr Glu
545 550 555 560
His Pro Glu Thr His Phe Glu Glu Leu Ala Leu Arg Phe Met Asp Ile
565 570 575
Arg Lys Arg Ile Tyr Lys Phe Pro Lys Met Gly Val Lys Ala Lys Met
580 585 590
Ile Ala Val Thr Thr Thr Ser Gly Thr Gly Ser Glu Val Thr Pro Phe
595 600 605
Ala Val Val Thr Asp Asp Ala Thr Gly Gln Lys Tyr Pro Leu Ala Asp
610 615 620
Tyr Ala Leu Thr Pro Asp Met Ala Ile Val Asp Ala Asn Leu Val Met
625 630 635 640
Asp Met Pro Lys Ser Leu Cys Ala Phe Gly Gly Leu Asp Ala Val Thr
645 650 655
His Ala Met Glu Ala Tyr Val Ser Val Leu Ala Ser Glu Phe Ser Asp
660 665 670
Gly Gln Ala Leu Gln Ala Leu Lys Leu Leu Lys Glu Tyr Leu Pro Ala
675 680 685
Ser Tyr His Glu Gly Ser Lys Asn Pro Val Ala Arg Glu Arg Val His
690 695 700
Ser Ala Ala Thr Ile Ala Gly Ile Ala Phe Ala Asn Ala Phe Leu Gly
705 710 715 720
Val Cys His Ser Met Ala His Lys Leu Gly Ser Gln Phe His Ile Pro
725 730 735
His Gly Leu Ala Asn Ala Leu Leu Ile Cys Asn Val Ile Arg Tyr Asn
740 745 750
Ala Asn Asp Asn Pro Thr Lys Gln Thr Ala Phe Ser Gln Tyr Asp Arg
755 760 765
Pro Gln Ala Arg Arg Arg Tyr Ala Glu Ile Ala Asp His Leu Gly Leu
770 775 780
Ser Ala Pro Gly Asp Arg Thr Ala Ala Lys Ile Glu Lys Leu Leu Ala
785 790 795 800
Trp Leu Glu Thr Leu Lys Ala Glu Leu Gly Ile Pro Lys Ser Ile Arg
805 810 815
Glu Ala Gly Val Gln Glu Ala Asp Phe Leu Ala Asn Val Asp Lys Leu
820 825 830
Ser Glu Asp Ala Phe Asp Asp Gln Cys Thr Gly Ala Asn Pro Arg Tyr
835 840 845
Pro Leu Ile Ser Glu Leu Lys Gln Ile Leu Leu Asp Thr Tyr Tyr Gly
850 855 860
Arg Asp Tyr Val Glu Gly Glu Thr Ala Ala Lys Lys Glu Ala Ala Pro
865 870 875 880
Ala Lys Ala Glu Lys Lys Ala Lys Lys Ser Ala
885 890
<210> 21
<211> 1407
<212> DNA
<213> 拜氏梭菌(Clostridium beijerinckii)
<220>
<221> CDS
<222> (1)..(1407)
<400> 21
atg aat aag gat acc ttg att cca act aca aag gat ttg aag gtt aag 48
Met Asn Lys Asp Thr Leu Ile Pro Thr Thr Lys Asp Leu Lys Val Lys
1 5 10 15
aca aac ggt gaa aac atc aac ttg aaa aat tac aag gat aac tct tca 96
Thr Asn Gly Glu Asn Ile Asn Leu Lys Asn Tyr Lys Asp Asn Ser Ser
20 25 30
tgt ttc ggt gtt ttc gaa aac gtt gaa aac gct att tcc agt gct gtt 144
Cys Phe Gly Val Phe Glu Asn Val Glu Asn Ala Ile Ser Ser Ala Val
35 40 45
cat gca caa aag att ttg tca ttg cat tac act aaa gaa caa aga gaa 192
His Ala Gln Lys Ile Leu Ser Leu His Tyr Thr Lys Glu Gln Arg Glu
50 55 60
aag att atc aca gaa atc aga aag gct gca ttg caa aat aag gaa gtt 240
Lys Ile Ile Thr Glu Ile Arg Lys Ala Ala Leu Gln Asn Lys Glu Val
65 70 75 80
ttg gca acc atg atc ttg gaa gaa act cat atg ggt aga tac gaa gat 288
Leu Ala Thr Met Ile Leu Glu Glu Thr His Met Gly Arg Tyr Glu Asp
85 90 95
aag atc ttg aag cat gaa tta gtt gct aaa tac aca cca ggt act gaa 336
Lys Ile Leu Lys His Glu Leu Val Ala Lys Tyr Thr Pro Gly Thr Glu
100 105 110
gat ttg acc act aca gca tgg tcc ggt gac aat ggt tta act gtt gtt 384
Asp Leu Thr Thr Thr Ala Trp Ser Gly Asp Asn Gly Leu Thr Val Val
115 120 125
gaa atg agt cct tat ggt gtt att ggt gct att act cca tct aca aat 432
Glu Met Ser Pro Tyr Gly Val Ile Gly Ala Ile Thr Pro Ser Thr Asn
130 135 140
cct acc gaa act gtt att tgt aac tca atc ggt atg att gct gct ggt 480
Pro Thr Glu Thr Val Ile Cys Asn Ser Ile Gly Met Ile Ala Ala Gly
145 150 155 160
aat gct gtt gtt ttt aac ggt cat cca tgt gct aag aaa tgt gtt gct 528
Asn Ala Val Val Phe Asn Gly His Pro Cys Ala Lys Lys Cys Val Ala
165 170 175
ttc gca gtt gaa atg atc aat aag gct atc atc tct tgt ggt ggt cca 576
Phe Ala Val Glu Met Ile Asn Lys Ala Ile Ile Ser Cys Gly Gly Pro
180 185 190
gaa aat ttg gtt acc act att aag aac cct act atg gaa tct ttg gat 624
Glu Asn Leu Val Thr Thr Ile Lys Asn Pro Thr Met Glu Ser Leu Asp
195 200 205
gca att att aag cat cca tca att aag ttg tta tgt ggt aca ggt ggt 672
Ala Ile Ile Lys His Pro Ser Ile Lys Leu Leu Cys Gly Thr Gly Gly
210 215 220
cct ggt atg gtt aag acc ttg ttg aac tct ggt aaa aag gct att ggt 720
Pro Gly Met Val Lys Thr Leu Leu Asn Ser Gly Lys Lys Ala Ile Gly
225 230 235 240
gct ggt gca ggt aac cca cct gtt att gtt gat gat aca gct gat atc 768
Ala Gly Ala Gly Asn Pro Pro Val Ile Val Asp Asp Thr Ala Asp Ile
245 250 255
gaa aag gca ggt aga tct atc atc gaa ggt tgt tca ttc gat aac aac 816
Glu Lys Ala Gly Arg Ser Ile Ile Glu Gly Cys Ser Phe Asp Asn Asn
260 265 270
ttg cct tgt atc gct gaa aag gaa gtt ttc gtt ttc gaa aac gtt gca 864
Leu Pro Cys Ile Ala Glu Lys Glu Val Phe Val Phe Glu Asn Val Ala
275 280 285
gat gat ttg atc tcc aac atg tta aag aat aac gct gtt att att aat 912
Asp Asp Leu Ile Ser Asn Met Leu Lys Asn Asn Ala Val Ile Ile Asn
290 295 300
gaa gat caa gtt tca aag tta att gat ttg gtt tta caa aag aat aac 960
Glu Asp Gln Val Ser Lys Leu Ile Asp Leu Val Leu Gln Lys Asn Asn
305 310 315 320
gaa act caa gaa tac ttc atc aat aag aaa tgg gtt ggt aaa gat gct 1008
Glu Thr Gln Glu Tyr Phe Ile Asn Lys Lys Trp Val Gly Lys Asp Ala
325 330 335
aag ttg ttc tta gat gaa atc gat gtt gaa tct cca tca aac gtt aag 1056
Lys Leu Phe Leu Asp Glu Ile Asp Val Glu Ser Pro Ser Asn Val Lys
340 345 350
tgt atc atc tgt gaa gtt aat gca aac cat cct ttt gtt atg aca gaa 1104
Cys Ile Ile Cys Glu Val Asn Ala Asn His Pro Phe Val Met Thr Glu
355 360 365
ttg atg atg cca atc ttg cct atc gtt aga gtt aag gat att gat gaa 1152
Leu Met Met Pro Ile Leu Pro Ile Val Arg Val Lys Asp Ile Asp Glu
370 375 380
gct att aaa tac gct aag atc gca gaa caa aac aga aag cat tcc gca 1200
Ala Ile Lys Tyr Ala Lys Ile Ala Glu Gln Asn Arg Lys His Ser Ala
385 390 395 400
tac atc tat agt aag aac atc gat aat ttg aac aga ttt gaa aga gaa 1248
Tyr Ile Tyr Ser Lys Asn Ile Asp Asn Leu Asn Arg Phe Glu Arg Glu
405 410 415
atc gat aca acc atc ttc gtt aaa aat gct aag tct ttc gca ggt gtt 1296
Ile Asp Thr Thr Ile Phe Val Lys Asn Ala Lys Ser Phe Ala Gly Val
420 425 430
ggt tac gaa gct gaa ggt ttt act aca ttc acc att gca ggt tcc act 1344
Gly Tyr Glu Ala Glu Gly Phe Thr Thr Phe Thr Ile Ala Gly Ser Thr
435 440 445
ggt gaa ggt att aca agt gct aga aac ttc act aga caa aga aga tgt 1392
Gly Glu Gly Ile Thr Ser Ala Arg Asn Phe Thr Arg Gln Arg Arg Cys
450 455 460
gtt tta gca ggt taa 1407
Val Leu Ala Gly
465
<210> 22
<211> 468
<212> PRT
<213> 拜氏梭菌(Clostridium beijerinckii)
<400> 22
Met Asn Lys Asp Thr Leu Ile Pro Thr Thr Lys Asp Leu Lys Val Lys
1 5 10 15
Thr Asn Gly Glu Asn Ile Asn Leu Lys Asn Tyr Lys Asp Asn Ser Ser
20 25 30
Cys Phe Gly Val Phe Glu Asn Val Glu Asn Ala Ile Ser Ser Ala Val
35 40 45
His Ala Gln Lys Ile Leu Ser Leu His Tyr Thr Lys Glu Gln Arg Glu
50 55 60
Lys Ile Ile Thr Glu Ile Arg Lys Ala Ala Leu Gln Asn Lys Glu Val
65 70 75 80
Leu Ala Thr Met Ile Leu Glu Glu Thr His Met Gly Arg Tyr Glu Asp
85 90 95
Lys Ile Leu Lys His Glu Leu Val Ala Lys Tyr Thr Pro Gly Thr Glu
100 105 110
Asp Leu Thr Thr Thr Ala Trp Ser Gly Asp Asn Gly Leu Thr Val Val
115 120 125
Glu Met Ser Pro Tyr Gly Val Ile Gly Ala Ile Thr Pro Ser Thr Asn
130 135 140
Pro Thr Glu Thr Val Ile Cys Asn Ser Ile Gly Met Ile Ala Ala Gly
145 150 155 160
Asn Ala Val Val Phe Asn Gly His Pro Cys Ala Lys Lys Cys Val Ala
165 170 175
Phe Ala Val Glu Met Ile Asn Lys Ala Ile Ile Ser Cys Gly Gly Pro
180 185 190
Glu Asn Leu Val Thr Thr Ile Lys Asn Pro Thr Met Glu Ser Leu Asp
195 200 205
Ala Ile Ile Lys His Pro Ser Ile Lys Leu Leu Cys Gly Thr Gly Gly
210 215 220
Pro Gly Met Val Lys Thr Leu Leu Asn Ser Gly Lys Lys Ala Ile Gly
225 230 235 240
Ala Gly Ala Gly Asn Pro Pro Val Ile Val Asp Asp Thr Ala Asp Ile
245 250 255
Glu Lys Ala Gly Arg Ser Ile Ile Glu Gly Cys Ser Phe Asp Asn Asn
260 265 270
Leu Pro Cys Ile Ala Glu Lys Glu Val Phe Val Phe Glu Asn Val Ala
275 280 285
Asp Asp Leu Ile Ser Asn Met Leu Lys Asn Asn Ala Val Ile Ile Asn
290 295 300
Glu Asp Gln Val Ser Lys Leu Ile Asp Leu Val Leu Gln Lys Asn Asn
305 310 315 320
Glu Thr Gln Glu Tyr Phe Ile Asn Lys Lys Trp Val Gly Lys Asp Ala
325 330 335
Lys Leu Phe Leu Asp Glu Ile Asp Val Glu Ser Pro Ser Asn Val Lys
340 345 350
Cys Ile Ile Cys Glu Val Asn Ala Asn His Pro Phe Val Met Thr Glu
355 360 365
Leu Met Met Pro Ile Leu Pro Ile Val Arg Val Lys Asp Ile Asp Glu
370 375 380
Ala Ile Lys Tyr Ala Lys Ile Ala Glu Gln Asn Arg Lys His Ser Ala
385 390 395 400
Tyr Ile Tyr Ser Lys Asn Ile Asp Asn Leu Asn Arg Phe Glu Arg Glu
405 410 415
Ile Asp Thr Thr Ile Phe Val Lys Asn Ala Lys Ser Phe Ala Gly Val
420 425 430
Gly Tyr Glu Ala Glu Gly Phe Thr Thr Phe Thr Ile Ala Gly Ser Thr
435 440 445
Gly Glu Gly Ile Thr Ser Ala Arg Asn Phe Thr Arg Gln Arg Arg Cys
450 455 460
Val Leu Ala Gly
465
<210> 23
<211> 2865
<212> DNA
<213> 莱茵衣藻(Chlamydomonas reinhardtii)
<220>
<221> CDS
<222> (1)..(2865)
<400> 23
atg atg agt tcc tct ctg gtt agt ggt aag aga gtt gcc gta cct tct 48
Met Met Ser Ser Ser Leu Val Ser Gly Lys Arg Val Ala Val Pro Ser
1 5 10 15
gcc gca aaa ccg tgc gct gca gtt cct cta cct agg gtt gct ggc aga 96
Ala Ala Lys Pro Cys Ala Ala Val Pro Leu Pro Arg Val Ala Gly Arg
20 25 30
aga acg gct gca aga gtt gtg tgc gaa gca gcg cct tca ggt gct gcc 144
Arg Thr Ala Ala Arg Val Val Cys Glu Ala Ala Pro Ser Gly Ala Ala
35 40 45
cct gcc tct cca aaa gca gag gca gct gct cct gtt gca gct gct ccc 192
Pro Ala Ser Pro Lys Ala Glu Ala Ala Ala Pro Val Ala Ala Ala Pro
50 55 60
gct aca cct cat gca gaa gtt aag aaa gaa aga gcg cca gca act gac 240
Ala Thr Pro His Ala Glu Val Lys Lys Glu Arg Ala Pro Ala Thr Asp
65 70 75 80
gaa gcg ttg aca gag tta aag gct ttg cta aag aga gca caa act gca 288
Glu Ala Leu Thr Glu Leu Lys Ala Leu Leu Lys Arg Ala Gln Thr Ala
85 90 95
caa gct cag tat tct aca tac act caa gaa cag gtc gac gaa atc ttt 336
Gln Ala Gln Tyr Ser Thr Tyr Thr Gln Glu Gln Val Asp Glu Ile Phe
100 105 110
cgt gca gcc gct gaa gca gct aac gca gcc aga ata cct ttg gcc aaa 384
Arg Ala Ala Ala Glu Ala Ala Asn Ala Ala Arg Ile Pro Leu Ala Lys
115 120 125
atg gca gtg gaa gaa act cgt atg gga gtg gct gaa gat aaa gtt gtt 432
Met Ala Val Glu Glu Thr Arg Met Gly Val Ala Glu Asp Lys Val Val
130 135 140
aag aat cac ttt gcg tca gaa ttc att tat aat aag tat aaa cac acg 480
Lys Asn His Phe Ala Ser Glu Phe Ile Tyr Asn Lys Tyr Lys His Thr
145 150 155 160
aaa act tgt ggt gtg atc gag cat gac cca gca gga ggg att cag aaa 528
Lys Thr Cys Gly Val Ile Glu His Asp Pro Ala Gly Gly Ile Gln Lys
165 170 175
gtg gct gag cct gta ggt gta ata gct ggt atc gtt cca act act aat 576
Val Ala Glu Pro Val Gly Val Ile Ala Gly Ile Val Pro Thr Thr Asn
180 185 190
ccc act tct acg gca ata ttt aag tcc cta cta agc tta aag acc aga 624
Pro Thr Ser Thr Ala Ile Phe Lys Ser Leu Leu Ser Leu Lys Thr Arg
195 200 205
aac gcc ttg gtt ttg tgt ccc cac cct agg gca gct aaa agt aca atc 672
Asn Ala Leu Val Leu Cys Pro His Pro Arg Ala Ala Lys Ser Thr Ile
210 215 220
gca gct gcc aga ata gtg aga gat gcg gct gtc gca gct gga gct cct 720
Ala Ala Ala Arg Ile Val Arg Asp Ala Ala Val Ala Ala Gly Ala Pro
225 230 235 240
ccc aat ata att tct tgg gtg gaa aca ccg tcg ttg cca gta agt caa 768
Pro Asn Ile Ile Ser Trp Val Glu Thr Pro Ser Leu Pro Val Ser Gln
245 250 255
gcc ttg atg caa gct act gaa att aac ttg att tta gct act ggt ggt 816
Ala Leu Met Gln Ala Thr Glu Ile Asn Leu Ile Leu Ala Thr Gly Gly
260 265 270
ccc gcc atg gta aga gct gca tac agt tca ggg aat cct tct ttg gga 864
Pro Ala Met Val Arg Ala Ala Tyr Ser Ser Gly Asn Pro Ser Leu Gly
275 280 285
gtt ggc gct ggg aat aca ccg gct ttg ata gat gaa acc gca gat gta 912
Val Gly Ala Gly Asn Thr Pro Ala Leu Ile Asp Glu Thr Ala Asp Val
290 295 300
gcc atg gca gtc tct tcc att cta ctg agc aaa acc ttc gac aat ggt 960
Ala Met Ala Val Ser Ser Ile Leu Leu Ser Lys Thr Phe Asp Asn Gly
305 310 315 320
gtg att tgt gct tca gaa caa agc gtt gtg gtc gtt gcc aag gct tat 1008
Val Ile Cys Ala Ser Glu Gln Ser Val Val Val Val Ala Lys Ala Tyr
325 330 335
gac gct gtc agg aca gaa ttc gtt agg aga ggt gcc tac ttt cta acg 1056
Asp Ala Val Arg Thr Glu Phe Val Arg Arg Gly Ala Tyr Phe Leu Thr
340 345 350
gaa gat gat aag gta aag gtt aga gcg gga gtc gta gta gat ggc aaa 1104
Glu Asp Asp Lys Val Lys Val Arg Ala Gly Val Val Val Asp Gly Lys
355 360 365
ttg aat cca aat ata gtt ggc cag tca att cca aaa ctt gcc gcc cta 1152
Leu Asn Pro Asn Ile Val Gly Gln Ser Ile Pro Lys Leu Ala Ala Leu
370 375 380
ttt ggc ata aag gtt cct caa ggc aca aaa gtt ttg att ggt gag gta 1200
Phe Gly Ile Lys Val Pro Gln Gly Thr Lys Val Leu Ile Gly Glu Val
385 390 395 400
gag aaa atc ggt cca gaa gaa gct tta tct caa gag aaa ctt tgc ccc 1248
Glu Lys Ile Gly Pro Glu Glu Ala Leu Ser Gln Glu Lys Leu Cys Pro
405 410 415
ata ctg gca atg tat aga gct cca gat tac gac cat ggt gtc aaa atg 1296
Ile Leu Ala Met Tyr Arg Ala Pro Asp Tyr Asp His Gly Val Lys Met
420 425 430
gct tgt gag tta ata atg tac ggt ggt gcc gga cat acg agc gtt ttg 1344
Ala Cys Glu Leu Ile Met Tyr Gly Gly Ala Gly His Thr Ser Val Leu
435 440 445
tac acc aat ccc ctt aat aat gcg cac att cag cag tat caa tca gct 1392
Tyr Thr Asn Pro Leu Asn Asn Ala His Ile Gln Gln Tyr Gln Ser Ala
450 455 460
gtc aaa aca gtg aga atc ttg att aac act cca gct agt caa ggt gca 1440
Val Lys Thr Val Arg Ile Leu Ile Asn Thr Pro Ala Ser Gln Gly Ala
465 470 475 480
atc ggt gat ctt tac aat ttt cat ctt gat ccc tct ctt aca ttg ggc 1488
Ile Gly Asp Leu Tyr Asn Phe His Leu Asp Pro Ser Leu Thr Leu Gly
485 490 495
tgt ggt aca tgg ggc tca act tct gta tca acg aat gtt ggt ccc caa 1536
Cys Gly Thr Trp Gly Ser Thr Ser Val Ser Thr Asn Val Gly Pro Gln
500 505 510
cat ctt ttg aac atc aaa acc gta aca gcc agg agg gaa aac atg tta 1584
His Leu Leu Asn Ile Lys Thr Val Thr Ala Arg Arg Glu Asn Met Leu
515 520 525
tgg ttt cgt gtt cca cct aaa atc tac ttt aaa ggt gga tgt tta gaa 1632
Trp Phe Arg Val Pro Pro Lys Ile Tyr Phe Lys Gly Gly Cys Leu Glu
530 535 540
gtt gcg tta acc gac ctg aga ggg aaa agc aga gct ttc att gta act 1680
Val Ala Leu Thr Asp Leu Arg Gly Lys Ser Arg Ala Phe Ile Val Thr
545 550 555 560
gat aag cca tta ttt gac atg ggt tat gct gat aaa gtc acg cac att 1728
Asp Lys Pro Leu Phe Asp Met Gly Tyr Ala Asp Lys Val Thr His Ile
565 570 575
tta gac tcc att aac gtg cat cat caa gtt ttc tac cac gtt aca ccg 1776
Leu Asp Ser Ile Asn Val His His Gln Val Phe Tyr His Val Thr Pro
580 585 590
gac cct aca ttg gct tgc att gaa gcg ggg tta aaa gag atc ttg gaa 1824
Asp Pro Thr Leu Ala Cys Ile Glu Ala Gly Leu Lys Glu Ile Leu Glu
595 600 605
ttt aaa cca gac gtt ata atc gct tta ggc gga ggg agt cca atg gat 1872
Phe Lys Pro Asp Val Ile Ile Ala Leu Gly Gly Gly Ser Pro Met Asp
610 615 620
gcc gct aag atc atg tgg cta atg tac gaa tgc cca gac acc aga ttt 1920
Ala Ala Lys Ile Met Trp Leu Met Tyr Glu Cys Pro Asp Thr Arg Phe
625 630 635 640
gat gga cta gcc atg agg ttc atg gat att aga aag aga gtt tat gag 1968
Asp Gly Leu Ala Met Arg Phe Met Asp Ile Arg Lys Arg Val Tyr Glu
645 650 655
gtt ccg gaa ttg ggc aag aaa gca acc atg gtg tgc att cct aca acc 2016
Val Pro Glu Leu Gly Lys Lys Ala Thr Met Val Cys Ile Pro Thr Thr
660 665 670
tct ggt aca ggt tcc gaa gtc act cca ttt tct gta gtt aca gat gaa 2064
Ser Gly Thr Gly Ser Glu Val Thr Pro Phe Ser Val Val Thr Asp Glu
675 680 685
cgt ttg ggc gca aaa tat cca ctg gca gat tat gca ttg act ccg tcg 2112
Arg Leu Gly Ala Lys Tyr Pro Leu Ala Asp Tyr Ala Leu Thr Pro Ser
690 695 700
atg gcc att gtc gat cca caa ttg gtg tta aac atg cct aag aaa tta 2160
Met Ala Ile Val Asp Pro Gln Leu Val Leu Asn Met Pro Lys Lys Leu
705 710 715 720
acc gcc tgg ggt ggt att gac gca ttg act cat gcg ctt gag tcg tat 2208
Thr Ala Trp Gly Gly Ile Asp Ala Leu Thr His Ala Leu Glu Ser Tyr
725 730 735
gtc tcc att tgt gcg act gat tat acg aag gga tta tca aga gaa gcc 2256
Val Ser Ile Cys Ala Thr Asp Tyr Thr Lys Gly Leu Ser Arg Glu Ala
740 745 750
ata tca cta ctg ttc aag tac ttg ccc agg gca tat gca aac ggg tcg 2304
Ile Ser Leu Leu Phe Lys Tyr Leu Pro Arg Ala Tyr Ala Asn Gly Ser
755 760 765
aat gat tac tta gct aga gaa aag gtt cac tat gcc gcg act att gct 2352
Asn Asp Tyr Leu Ala Arg Glu Lys Val His Tyr Ala Ala Thr Ile Ala
770 775 780
ggt atg gcc ttt gca aac gca ttc tta gga ata tgt cac tcc atg gcg 2400
Gly Met Ala Phe Ala Asn Ala Phe Leu Gly Ile Cys His Ser Met Ala
785 790 795 800
cat aaa tta ggc gct gct tat cat gtg cct cat ggt cta gct aat gca 2448
His Lys Leu Gly Ala Ala Tyr His Val Pro His Gly Leu Ala Asn Ala
805 810 815
gcc tta att tcg cat gtc ata cgt tat aac gct acc gat atg cca gcg 2496
Ala Leu Ile Ser His Val Ile Arg Tyr Asn Ala Thr Asp Met Pro Ala
820 825 830
aaa caa gcc gct ttc cca caa tac gag tat cct act gca aag caa gac 2544
Lys Gln Ala Ala Phe Pro Gln Tyr Glu Tyr Pro Thr Ala Lys Gln Asp
835 840 845
tat gct gat ttg gcc aac atg ctg ggt ctt gga ggt aat acc gta gat 2592
Tyr Ala Asp Leu Ala Asn Met Leu Gly Leu Gly Gly Asn Thr Val Asp
850 855 860
gag aag gtc atc aag ttg att gaa gca gta gaa gaa ctg aaa gct aaa 2640
Glu Lys Val Ile Lys Leu Ile Glu Ala Val Glu Glu Leu Lys Ala Lys
865 870 875 880
gtt gac ata cca ccg act att aag gaa atc ttt aac gat cca aag gtg 2688
Val Asp Ile Pro Pro Thr Ile Lys Glu Ile Phe Asn Asp Pro Lys Val
885 890 895
gat gcg gat ttt ctg gct aat gtc gac gct ttg gcc gag gat gca ttt 2736
Asp Ala Asp Phe Leu Ala Asn Val Asp Ala Leu Ala Glu Asp Ala Phe
900 905 910
gat gat caa tgt aca gga gca aat cca aga tac cca ttg atg gct gat 2784
Asp Asp Gln Cys Thr Gly Ala Asn Pro Arg Tyr Pro Leu Met Ala Asp
915 920 925
tta aaa cag tta tac ctt gat gcc cat gct gcc cca att tta cca gtc 2832
Leu Lys Gln Leu Tyr Leu Asp Ala His Ala Ala Pro Ile Leu Pro Val
930 935 940
aaa acc ctt gaa ttc ttc tcc aaa atc aac taa 2865
Lys Thr Leu Glu Phe Phe Ser Lys Ile Asn
945 950
<210> 24
<211> 954
<212> PRT
<213> 莱茵衣藻(Chlamydomonas reinhardtii)
<400> 24
Met Met Ser Ser Ser Leu Val Ser Gly Lys Arg Val Ala Val Pro Ser
1 5 10 15
Ala Ala Lys Pro Cys Ala Ala Val Pro Leu Pro Arg Val Ala Gly Arg
20 25 30
Arg Thr Ala Ala Arg Val Val Cys Glu Ala Ala Pro Ser Gly Ala Ala
35 40 45
Pro Ala Ser Pro Lys Ala Glu Ala Ala Ala Pro Val Ala Ala Ala Pro
50 55 60
Ala Thr Pro His Ala Glu Val Lys Lys Glu Arg Ala Pro Ala Thr Asp
65 70 75 80
Glu Ala Leu Thr Glu Leu Lys Ala Leu Leu Lys Arg Ala Gln Thr Ala
85 90 95
Gln Ala Gln Tyr Ser Thr Tyr Thr Gln Glu Gln Val Asp Glu Ile Phe
100 105 110
Arg Ala Ala Ala Glu Ala Ala Asn Ala Ala Arg Ile Pro Leu Ala Lys
115 120 125
Met Ala Val Glu Glu Thr Arg Met Gly Val Ala Glu Asp Lys Val Val
130 135 140
Lys Asn His Phe Ala Ser Glu Phe Ile Tyr Asn Lys Tyr Lys His Thr
145 150 155 160
Lys Thr Cys Gly Val Ile Glu His Asp Pro Ala Gly Gly Ile Gln Lys
165 170 175
Val Ala Glu Pro Val Gly Val Ile Ala Gly Ile Val Pro Thr Thr Asn
180 185 190
Pro Thr Ser Thr Ala Ile Phe Lys Ser Leu Leu Ser Leu Lys Thr Arg
195 200 205
Asn Ala Leu Val Leu Cys Pro His Pro Arg Ala Ala Lys Ser Thr Ile
210 215 220
Ala Ala Ala Arg Ile Val Arg Asp Ala Ala Val Ala Ala Gly Ala Pro
225 230 235 240
Pro Asn Ile Ile Ser Trp Val Glu Thr Pro Ser Leu Pro Val Ser Gln
245 250 255
Ala Leu Met Gln Ala Thr Glu Ile Asn Leu Ile Leu Ala Thr Gly Gly
260 265 270
Pro Ala Met Val Arg Ala Ala Tyr Ser Ser Gly Asn Pro Ser Leu Gly
275 280 285
Val Gly Ala Gly Asn Thr Pro Ala Leu Ile Asp Glu Thr Ala Asp Val
290 295 300
Ala Met Ala Val Ser Ser Ile Leu Leu Ser Lys Thr Phe Asp Asn Gly
305 310 315 320
Val Ile Cys Ala Ser Glu Gln Ser Val Val Val Val Ala Lys Ala Tyr
325 330 335
Asp Ala Val Arg Thr Glu Phe Val Arg Arg Gly Ala Tyr Phe Leu Thr
340 345 350
Glu Asp Asp Lys Val Lys Val Arg Ala Gly Val Val Val Asp Gly Lys
355 360 365
Leu Asn Pro Asn Ile Val Gly Gln Ser Ile Pro Lys Leu Ala Ala Leu
370 375 380
Phe Gly Ile Lys Val Pro Gln Gly Thr Lys Val Leu Ile Gly Glu Val
385 390 395 400
Glu Lys Ile Gly Pro Glu Glu Ala Leu Ser Gln Glu Lys Leu Cys Pro
405 410 415
Ile Leu Ala Met Tyr Arg Ala Pro Asp Tyr Asp His Gly Val Lys Met
420 425 430
Ala Cys Glu Leu Ile Met Tyr Gly Gly Ala Gly His Thr Ser Val Leu
435 440 445
Tyr Thr Asn Pro Leu Asn Asn Ala His Ile Gln Gln Tyr Gln Ser Ala
450 455 460
Val Lys Thr Val Arg Ile Leu Ile Asn Thr Pro Ala Ser Gln Gly Ala
465 470 475 480
Ile Gly Asp Leu Tyr Asn Phe His Leu Asp Pro Ser Leu Thr Leu Gly
485 490 495
Cys Gly Thr Trp Gly Ser Thr Ser Val Ser Thr Asn Val Gly Pro Gln
500 505 510
His Leu Leu Asn Ile Lys Thr Val Thr Ala Arg Arg Glu Asn Met Leu
515 520 525
Trp Phe Arg Val Pro Pro Lys Ile Tyr Phe Lys Gly Gly Cys Leu Glu
530 535 540
Val Ala Leu Thr Asp Leu Arg Gly Lys Ser Arg Ala Phe Ile Val Thr
545 550 555 560
Asp Lys Pro Leu Phe Asp Met Gly Tyr Ala Asp Lys Val Thr His Ile
565 570 575
Leu Asp Ser Ile Asn Val His His Gln Val Phe Tyr His Val Thr Pro
580 585 590
Asp Pro Thr Leu Ala Cys Ile Glu Ala Gly Leu Lys Glu Ile Leu Glu
595 600 605
Phe Lys Pro Asp Val Ile Ile Ala Leu Gly Gly Gly Ser Pro Met Asp
610 615 620
Ala Ala Lys Ile Met Trp Leu Met Tyr Glu Cys Pro Asp Thr Arg Phe
625 630 635 640
Asp Gly Leu Ala Met Arg Phe Met Asp Ile Arg Lys Arg Val Tyr Glu
645 650 655
Val Pro Glu Leu Gly Lys Lys Ala Thr Met Val Cys Ile Pro Thr Thr
660 665 670
Ser Gly Thr Gly Ser Glu Val Thr Pro Phe Ser Val Val Thr Asp Glu
675 680 685
Arg Leu Gly Ala Lys Tyr Pro Leu Ala Asp Tyr Ala Leu Thr Pro Ser
690 695 700
Met Ala Ile Val Asp Pro Gln Leu Val Leu Asn Met Pro Lys Lys Leu
705 710 715 720
Thr Ala Trp Gly Gly Ile Asp Ala Leu Thr His Ala Leu Glu Ser Tyr
725 730 735
Val Ser Ile Cys Ala Thr Asp Tyr Thr Lys Gly Leu Ser Arg Glu Ala
740 745 750
Ile Ser Leu Leu Phe Lys Tyr Leu Pro Arg Ala Tyr Ala Asn Gly Ser
755 760 765
Asn Asp Tyr Leu Ala Arg Glu Lys Val His Tyr Ala Ala Thr Ile Ala
770 775 780
Gly Met Ala Phe Ala Asn Ala Phe Leu Gly Ile Cys His Ser Met Ala
785 790 795 800
His Lys Leu Gly Ala Ala Tyr His Val Pro His Gly Leu Ala Asn Ala
805 810 815
Ala Leu Ile Ser His Val Ile Arg Tyr Asn Ala Thr Asp Met Pro Ala
820 825 830
Lys Gln Ala Ala Phe Pro Gln Tyr Glu Tyr Pro Thr Ala Lys Gln Asp
835 840 845
Tyr Ala Asp Leu Ala Asn Met Leu Gly Leu Gly Gly Asn Thr Val Asp
850 855 860
Glu Lys Val Ile Lys Leu Ile Glu Ala Val Glu Glu Leu Lys Ala Lys
865 870 875 880
Val Asp Ile Pro Pro Thr Ile Lys Glu Ile Phe Asn Asp Pro Lys Val
885 890 895
Asp Ala Asp Phe Leu Ala Asn Val Asp Ala Leu Ala Glu Asp Ala Phe
900 905 910
Asp Asp Gln Cys Thr Gly Ala Asn Pro Arg Tyr Pro Leu Met Ala Asp
915 920 925
Leu Lys Gln Leu Tyr Leu Asp Ala His Ala Ala Pro Ile Leu Pro Val
930 935 940
Lys Thr Leu Glu Phe Phe Ser Lys Ile Asn
945 950
<210> 25
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 25
tgggaatatt accgctcgaa g 21
<210> 26
<211> 29
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 26
ctttaaaaaa tttccaattt tcctttacg 29
<210> 27
<211> 18
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 27
tcgcagccac gggtcaac 18
<210> 28
<211> 37
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 28
ttttattatt agtctttttt ttttttgaca atatctg 37
<210> 29
<211> 31
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 29
atgactcaat tcactgacat tgataagcta g 31
<210> 30
<211> 32
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 30
atattcttta ttggctttat acttgaatgg tg 32
<210> 31
<211> 24
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 31
gacgttgatt taaggtggtt ccgg 24
<210> 32
<211> 25
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 32
atgtctgaac cagctcaaaa gaaac 25
<210> 33
<211> 37
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 33
tttgaatatg tattacttgg ttatggttat atatgac 37
<210> 34
<211> 25
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 34
actggtagag agcgactttg tatgc 25
<210> 35
<211> 22
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 35
gctttcaatt catttgggtg tg 22
<210> 36
<211> 30
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 36
tgtatatgag atagttgatt gtatgcttgg 30
<210> 37
<211> 28
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 37
atggtcaaac caattatagc tcccagta 28
<210> 38
<211> 25
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 38
aaatggatat tgatctagat ggcgg 25
<210> 39
<211> 24
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 39
cttggtgtgt catcggtagt aacg 24
<210> 40
<211> 17
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 40
atggctgccg gtgtccc 17
<210> 41
<211> 36
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 41
ttgtaattaa aacttagatt agattgctat gctttc 36
<210> 42
<211> 18
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 42
aggaacagcc gtcaaggg 18
<210> 43
<211> 28
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 43
cttccctttt acagtgcttc ggaaaagc 28
<210> 44
<211> 32
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 44
tttgttttgt gtgtaaattt agtgaagtac tg 32
<210> 45
<211> 28
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 45
atgtctcaaa tttttaagga tatcccag 28
<210> 46
<211> 29
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 46
ttattgaaac aaaatttggt taataatac 29
<210> 47
<211> 28
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 47
taaagtaaga gcgctacatt ggtctacc 28
<210> 48
<211> 24
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 48
ttactccgca acgcttttct gaac 24
<210> 49
<211> 26
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 49
tagcgttgaa tgttagcgtc aacaac 26
<210> 50
<211> 41
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 50
tttgtttgtt tatgtgtgtt tattcgaaac taagttcttg g 41
<210> 51
<211> 27
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 51
atgttgtgtt cagtaattca gagacag 27
<210> 52
<211> 26
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 52
ttagatgaga gtcttttcca gttcgc 26
<210> 53
<211> 25
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 53
tgacaccgat tatttaaagc tgcag 25
<210> 54
<211> 18
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 54
agagcgcgcc tcgttcag 18
<210> 55
<211> 27
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 55
aattccgctg tatagctcat atctttc 27
<210> 56
<211> 39
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 56
aacgaggcgc gctctaattc cgctgtatag ctcatatct 39
<210> 57
<211> 24
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 57
acgacatcgt cgaatatgat tcag 24
<210> 58
<211> 33
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 58
tattaattta gtgtgtgtat ttgtgtttgt gtg 33
<210> 59
<211> 23
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 59
atgagccata ttcaacggga aac 23
<210> 60
<211> 28
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 60
ttacaaccaa ttaaccaatt ctgattag 28
<210> 61
<211> 37
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 61
tgcatgtcta ctaaactcac aaattagagc ttcaatt 37
<210> 62
<211> 39
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 62
gggtaataac tgatataatt aaattgaagc tctaatttg 39
<210> 63
<211> 59
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 63
cccataactt cgtatagcat acattatacg aagttattga caccgattat ttaaagctg 59
<210> 64
<211> 25
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 64
gtatgctgca gctttaaata atcgg 25
<210> 65
<211> 26
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 65
tccagccagt aaaatccata ctcaac 26
<210> 66
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 66
aagggggaag gtgtggaatc 20
<210> 67
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 67
ctatgggact tccgggaaac 20
<210> 68
<211> 38
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 68
tgtgtattac gatatagtta atagttgata gttgattg 38
<210> 69
<211> 26
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 69
tagcgttgaa tgttagcgtc aacaac 26
<210> 70
<211> 41
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 70
tttgtttgtt tatgtgtgtt tattcgaaac taagttcttg g 41
<210> 71
<211> 28
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 71
atgtctatcc cagaaactca aaaaggtg 28
<210> 72
<211> 28
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 72
ttgtcctctg aggacataaa atacacac 28
<210> 73
<211> 24
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 73
ttactccgca acgcttttct gaac 24
<210> 74
<211> 28
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 74
taaagtaaga gcgctacatt ggtctacc 28
<210> 75
<211> 18
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 75
ttatgcggcc tctcctgc 18
<210> 76
<211> 28
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 76
atgtcaaaga gaaaagttgc tattatcg 28
<210> 77
<211> 37
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 77
ttttattatt agtctttttt ttttttgaca atatctg 37
<210> 78
<211> 18
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 78
tcgctcgcag ccacgggt 18
<210> 79
<211> 21
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Claims (5)
1.乙醇的制造方法,具有以下工序:将导入了木糖异构酶基因和乙醛脱氢酶基因、高表达具有将乙醛转换为乙醇的活性的醇脱氢酶基因、且具有将乙醇转换为乙醛的活性的醇脱氢酶基因的表达量降低了的重组酵母,用含有木糖的培养基进行培养,从而进行乙醇发酵,其中所述重组酵母能够代谢所述含有木糖的培养基中的乙酸,
所述木糖异构酶基因编码序列号4所示的氨基酸序列中的第337位的天冬酰胺被替换成了半胱氨酸的氨基酸序列,
所述乙醛脱氢酶基因编码来源于大肠杆菌的乙醛脱氢酶、来源于拜氏梭菌的乙醛脱氢酶、或者来源于莱茵衣藻的乙醛脱氢酶,
所述来源于大肠杆菌的乙醛脱氢酶是由序列号2或20所示的氨基酸序列组成的蛋白质,
所述来源于拜氏梭菌的乙醛脱氢酶是由序列号22所示的氨基酸序列组成的蛋白质,
所述来源于莱茵衣藻的乙醛脱氢酶是由序列号24所示的氨基酸序列组成的蛋白质。
2.根据权利要求1所述的乙醇的制造方法,其特征在于,所述重组酵母是进一步导入了木酮糖激酶基因的重组酵母。
3.根据权利要求1所述的乙醇的制造方法,其特征在于,所述重组酵母是导入了下述基因的重组酵母,所述基因编码选自构成戊糖磷酸途径中的非氧化过程的途径的酶群中的酶。
4.根据权利要求3所述的乙醇的制造方法,其特征在于,构成戊糖磷酸途径中的非氧化过程的途径的酶群是核糖-5-磷酸异构酶、核酮糖-5-磷酸-3-差向异构酶、转酮醇酶和转醛醇酶。
5.根据权利要求1所述的乙醇的制造方法,其特征在于,所述培养基含有纤维素,在所述乙醇发酵中,至少同时进行所述纤维素的糖化。
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CN104024419A (zh) | 2011-10-24 | 2014-09-03 | 丰田自动车株式会社 | 使用了重组酵母的乙醇的制造方法 |
JP6027559B2 (ja) | 2013-03-28 | 2016-11-16 | 株式会社豊田中央研究所 | キシロースイソメラーゼ活性を有するタンパク質及びその利用 |
BR112017005959A2 (pt) * | 2014-10-15 | 2017-12-19 | Jx Nippon Oil & Energy Corp | levedura capaz de produzir etanol a partir de xilose |
JP6303984B2 (ja) | 2014-10-31 | 2018-04-04 | トヨタ自動車株式会社 | 連続培養によるエタノールの製造方法及び連続培養装置 |
JP6697250B2 (ja) * | 2015-12-02 | 2020-05-20 | 株式会社豊田中央研究所 | ゲノム再編方法及びその利用 |
JP6447583B2 (ja) * | 2016-06-24 | 2019-01-09 | トヨタ自動車株式会社 | 組換え酵母、及びそれを用いたエタノールの製造方法 |
JP6097869B1 (ja) * | 2016-08-01 | 2017-03-15 | 新日鉄住金エンジニアリング株式会社 | エタノールの製造方法 |
JP6879111B2 (ja) | 2017-08-02 | 2021-06-02 | トヨタ自動車株式会社 | 組換え酵母及びこれを用いたエタノールの製造方法 |
WO2019058260A1 (en) | 2017-09-19 | 2019-03-28 | Lallemand Hungary Liquidity Management Llc | TOLERANCE TO ACETATE TOXICITY IN RECOMBINANT MICROBIAL HOST CELLS |
JP2019068788A (ja) * | 2017-10-11 | 2019-05-09 | トヨタ自動車株式会社 | 組換え酵母及びこれを用いたエタノールの製造方法 |
JP7298674B2 (ja) * | 2017-11-06 | 2023-06-27 | トヨタ自動車株式会社 | エタノール発酵によるエタノール生産性の向上に関与する変異遺伝子及びこれを用いたエタノールの製造方法 |
JP7078887B2 (ja) | 2017-11-06 | 2022-06-01 | トヨタ自動車株式会社 | エタノール発酵によるエタノール生産性の向上に関与する変異遺伝子及びこれを用いたエタノールの製造方法 |
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EP3775219A1 (en) * | 2018-03-27 | 2021-02-17 | Basf Se | Xylose metabolizing yeast |
WO2019188839A1 (ja) | 2018-03-27 | 2019-10-03 | 積水化学工業株式会社 | エタノールの製造方法及びエタノール組成物 |
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CN113046252B (zh) * | 2021-03-23 | 2022-07-05 | 江南大学 | 一株乙醛脱氢酶高产菌株的分离与鉴定 |
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US20160002674A1 (en) | 2016-01-07 |
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CA2901974C (en) | 2020-01-14 |
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