CN109689864B - 苹果酸脱氢酶 - Google Patents
苹果酸脱氢酶 Download PDFInfo
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- CN109689864B CN109689864B CN201780054104.2A CN201780054104A CN109689864B CN 109689864 B CN109689864 B CN 109689864B CN 201780054104 A CN201780054104 A CN 201780054104A CN 109689864 B CN109689864 B CN 109689864B
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Abstract
本发明涉及一种重组宿主细胞，其能够生产二羧酸并且包含突变的苹果酸脱氢酶，从而导致提高的二羧酸生产。本发明还涉及生产二羧酸的方法，所述方法包括在合适的发酵培养基中发酵所述重组宿主细胞并生产所述二羧酸。
Description
技术领域
本发明涉及能够生产二羧酸的重组宿主细胞，以及使用所述重组宿主细胞生产二羧酸的方法。
背景技术
4-碳二羧酸苹果酸、延胡索酸和琥珀酸是大量化学品的潜在前体。例如，琥珀酸可以被转变为1,4-丁二醇(BDO)、四氢呋喃和γ-丁内酯。衍生自琥珀酸的另一产物是通过连接琥珀酸和BDO制造的聚酯聚合物。
用于工业用途的琥珀酸主要是通过马来酸或马来酸酐的催化氢化由丁烷通过石油化学生产的。这些方法被认为是对环境有害且昂贵的。琥珀酸的发酵生产被视为生产琥珀酸的一种吸引人的替代方法，其中可以使用可再生的原料作为碳源。
已经对(重组)酵母中C4-二羧酸的发酵生产进行了若干研究。
例如，EP2495304公开了适合琥珀酸生产的重组酵母，其中编码丙酮酸羧化酶、磷酸烯醇式丙酮酸羧激酶、苹果酸脱氢酶、延胡索酸酶、延胡索酸还原酶和琥珀酸转运体的基因被遗传修饰。
尽管在宿主细胞(例如酵母)中发酵生产二羧酸已经取得了改进，但仍需要进一步改进的用于发酵生产二羧酸的宿主细胞。
发明内容
本发明涉及重组宿主细胞，其能够生产二羧酸并且包含具有苹果酸脱氢酶(MDH)活性的突变多肽。出乎意料地发现：与包含参照MDH多肽的宿主细胞所产生的二羧酸的量相比，根据本发明的宿主细胞产生增加量的二羧酸，所述参照MDH多肽通常是NAD(H)-依赖型苹果酸脱氢酶(EC1.1.1.37)。
本发明因此提供了一种重组宿主细胞，其能够生产二羧酸并且包含编码具有苹果酸脱氢酶活性的突变多肽的核酸序列，其中所述突变多肽包含这样的氨基酸序列，当与包含SEQ ID NO:39所示序列的苹果酸脱氢酶比对时，所述氨基酸序列包含对应于SEQ ID NO:39中的第34位氨基酸的氨基酸残基的一个突变(例如一个替换)。所述具有苹果酸脱氢酶活性的突变多肽可以进一步包含一个或多个另外的突变(例如替换)。特别地，具有苹果酸脱氢酶活性的突变多肽可以进一步包含对应于SEQ ID NO:39中的第35、36、37、38、39和/或40位氨基酸中的任何的一个或多个另外的突变(例如替换)。
本发明还提供了一种重组宿主细胞，其能够生产二羧酸并且包含编码具有苹果酸脱氢酶活性的突变多肽的核酸序列，其中与参照MDH多肽相比，所述突变多肽的NADP(H)-依赖型活性与NAD(H)-依赖型活性之比提高，所述参照MDH多肽通常是NAD(H)-依赖型苹果酸脱氢酶(EC1.1.1.37)。在所述实施方式中，所述突变多肽可以是包含这样的氨基酸序列的突变多肽，当与包含SEQ ID NO:39所示序列的苹果酸脱氢酶比对时，所述氨基酸序列包含对应于SEQ ID NO:39中的第34位氨基酸的氨基酸残基的一个突变(例如一个替换)。
本发明还提供了：
-根据本发明的重组宿主细胞，其中编码具有苹果酸脱氢酶活性的突变多肽的核酸序列在胞质中表达，并且具有苹果酸脱氢酶活性的突变多肽在胞质中有活性。
-根据本发明的重组宿主细胞，其中所述重组宿主细胞具有从磷酸烯醇或丙酮酸盐/酯到琥珀酸盐/酯的活性还原性三羧酸(TCA)通路。
-生产二羧酸的方法，其中所述方法包括在适于生产二羧酸的条件下发酵本发明的重组宿主细胞。二羧酸可以是琥珀酸、苹果酸和/或延胡索酸。所述方法可以进一步包括从发酵培养基中回收二羧酸。
附图说明
图1示出了片段9至12整合的示意图。片段9至12中所示的阴影线部分表示导致(如同源区之间的虚线交叉所示的)重组事件的独特同源重叠区。片段9的5’端和片段12的3’端(由片段9和12中的灰色区域表示)与染色体16上的YPRCtau3基因座同源。同源重组导致片段10和11整合到YPRCtau3基因座中。
图2示出了片段1至8和片段113整合的示意图。片段1至8和113中所示的阴影线部分表示导致(如同源区之间的虚线交叉所示的)重组事件的独特同源重叠区。片段1和片段113与染色体XI上的INT59基因座同源，同源重组导致片段2至8整合到INT59基因座中。
图3示出了片段13、114、115、15和16整合的示意图。片段13、114、115、15和16中所示的阴影线部分表示导致(如同源区之间的虚线交叉所示的)重组事件的独特同源重叠区。片段13和片段16与染色体XV上的INT1基因座同源，同源重组导致片段114、115和15整合到INT1基因座中。
图4示出了片段13、14、16和片段17-110之一整合的示意图。片段13、14、16和片段17-110中所示的阴影线部分表示导致(如同源区之间的虚线交叉所示的)重组事件的独特同源重叠区。片段13和片段16与染色体XV上的INT1基因座同源，同源重组导致片段14和片段17-110之一整合到INT1基因座中。
图5A：培养SUC-1112转化体后在生产培养基的上清液中测量的平均苹果酸滴度，所述SUC-1112转化体表达磷酸烯醇式丙酮酸羧激酶(PCKa)、丙酮酸羧化酶(PYC2)、苹果酸脱氢酶(MDH3)、延胡索酸酶(FUMR和fumB)、二羧酸转运体(DCT_02)并用参照苹果酸脱氢酶(SEQ ID NO:39)或突变的苹果酸脱氢酶转化，与参照序列相比，所述突变的苹果酸脱氢酶在表1中所示的氨基酸位置含有突变。苹果酸滴度如“通用的材料和方法”中所述进行测量，其代表从三个独立克隆获得的平均值。
图5B：在菌株SUC-1112中表达的MDH突变体的NADH特异性苹果酸脱氢酶(MDH)活性(具体突变请参见表1)。显示的是活性，描述为ΔA340/min/mg总蛋白。该值是负的，因为MDH依赖型NADH氧化导致340nm处的吸光度降低。负值越大表示活性越高。如实施例5中所述测量活性。
图5C：在菌株SUC-1112中表达的MDH突变体的NADPH特异性苹果酸脱氢酶(MDH)活性(具体突变请参见表1)。显示的是活性，描述为ΔA340/min/mg总蛋白。该值是负的，因为MDH依赖型NADPH氧化导致340nm处的吸光度降低。负值越大表示活性越高。如实施例5中所述测量活性。
图5D：在菌株SUC-1112中表达的MDH突变体的NADPH依赖型活性:NADH依赖型活性的比值(具体突变请参见表1)。测量活性并如实施例5中所述测定比值。虚线表示NADPH:NADH的比值为1.0。
图6示出了片段116、117、118和片段119整合的示意图。片段116、117、118和片段119中所示的阴影线部分表示导致重组事件的独特同源重叠区，如同源区之间的虚线交叉所示。片段116和片段119与染色体XV上的INT1基因座同源，同源重组导致片段117和片段118整合到INT1基因座中。
图7示出了片段1-5、124和片段120、121、122或123整合的示意图。片段中所示的阴影线部分表示导致(如同源区之间的虚线交叉所示的)重组事件的独特同源重叠区。片段1和片段124与染色体XI上的INT59基因座同源，同源重组导致片段2至5和120、121、122或123整合到INT59基因座中。
序列表说明
SEQ ID NO:1示出了片段2(图2)的核苷酸序列，其包括针对在Saccharomycescerevisiae中表达而经密码子对优化的来自Actinobacillus succinogenes的PEP羧激酶。
SEQ ID NO:2示出了片段3(图2)的核苷酸序列，其包括针对在S.cerevisiae中表达而经密码子对优化的来自S.cerevisiae的丙酮酸羧化酶(PYC2)。
SEQ ID NO:3示出了片段4(图2)的PCR模板的核苷酸序列，其包括在S.cerevisiae中有功能的KanMX选择标记。
SEQ ID NO:4示出了片段5(图2)的核苷酸序列，其包括针对在S.cerevisiae中表达而经密码子对优化的来自Aspergillus niger的推定的二羧酸转运体。
SEQ ID NO:5示出了片段6(图2)的核苷酸序列，其包括针对在S.cerevisiae中表达而经密码子对优化的来自S.cerevisiae的苹果酸脱氢酶(MDH3)。
SEQ ID NO:6示出了片段7(图2)的核苷酸序列，其包括针对在S.cerevisiae中表达而经密码子对优化的来自Escherchia coli的延胡索酸酶(fumB)。
SEQ ID NO:7示出了片段8(图2)的核苷酸序列，其包括针对在S.cerevisiae中表达而经密码子对优化的来自Trypanosoma brucei的延胡索酸还原酶(FRDg)。
SEQ ID NO:8示出了来自Trypanosoma brucei的延胡索酸还原酶(FRDg)的氨基酸序列。
SEQ ID NO:9示出了用于生成片段1(图2)的引物的核苷酸序列。
SEQ ID NO:10示出了用于生成片段1(图2)的引物的核苷酸序列。
SEQ ID NO:11示出了用于生成片段2(图2)的引物的核苷酸序列。
SEQ ID NO:12示出了用于生成片段2(图2)的引物的核苷酸序列。
SEQ ID NO:13示出了用于生成片段3(图2)的引物的核苷酸序列。
SEQ ID NO:14示出了用于生成片段3(图2)的引物的核苷酸序列。
SEQ ID NO:15示出了用于生成片段4(图2)的引物的核苷酸序列。
SEQ ID NO:16示出了用于生成片段4(图2)的引物的核苷酸序列。
SEQ ID NO:17示出了用于生成片段5(图2)的引物的核苷酸序列。
SEQ ID NO:18示出了用于生成片段5(图2)的引物的核苷酸序列。
SEQ ID NO:19示出了用于生成片段6(图2)以及片段120、121、122和123(图7)的引物的核苷酸序列。
SEQ ID NO:20示出了用于生成片段6(图2)以及片段120、121、122和123(图7)的引物的核苷酸序列。
SEQ ID NO:21示出了用于生成片段7(图2)的引物的核苷酸序列。
SEQ ID NO:22示出了用于生成片段7(图2)的引物的核苷酸序列。
SEQ ID NO:23示出了用于生成片段8(图2)的引物的核苷酸序列。
SEQ ID NO:24示出了用于生成片段8(图2)的引物的核苷酸序列。
SEQ ID NO:25示出了用于生成片段13(图3)的引物的核苷酸序列。
SEQ ID NO:26示出了用于生成片段13(图3)的引物的核苷酸序列。
SEQ ID NO:27示出了用于生成片段14(图4)的引物的核苷酸序列。
SEQ ID NO:28示出了用于生成片段115(图3)和片段14(图4)的引物的核苷酸序列。
SEQ ID NO:29示出了用于生成片段15(图3)和片段17至110(图4)的引物的核苷酸序列。
SEQ ID NO:30示出了用于生成片段15(图3)和片段17至110(图4)的引物的核苷酸序列。
SEQ ID NO:31示出了片段15(图3)和片段120(图7)的核苷酸序列，其包括针对在S.cerevisiae中表达而经密码子对优化的编码SEQ ID NO:39的核苷酸序列。
SEQ ID NO:32示出了用于生成片段16(图3)的引物的核苷酸序列。
SEQ ID NO:33示出了用于生成片段16(图3)的引物的核苷酸序列。
SEQ ID NO:34示出了片段9(图1)的核苷酸序列，其包括针对在Saccharomycescerevisiae中表达而经密码子对优化的来自Rhizopus oryzae的延胡索酸酶。
SEQ ID NO:35示出了片段10(图1)的核苷酸序列，其包括Cre重组酶的5’部分。
SEQ ID NO:36示出了片段11(图1)的核苷酸序列，其包括Cre重组酶的3’部分。
SEQ ID NO:37示出了片段12(图1)的核苷酸序列，其包括与YPRCtau3基因座同源的区域。
SEQ ID NO:38示出了片段115(图3)、片段14(图4)和片段117(图6)的PCR模板的核苷酸序列，其包含诺尔丝菌素选择标记。
SEQ ID NO:39示出了缺少3C-末端过氧化物酶体靶向序列的来自S.cerevisiae的苹果酸脱氢酶(MDH3)蛋白质的氨基酸序列。
SEQ ID NO:40示出了用于生成片段113(图2)的引物的核苷酸序列。
SEQ ID NO:41示出了用于生成片段113(图2)的引物的核苷酸序列。
SEQ ID NO:42示出了片段114(图3)的核苷酸序列，其包括针对在Saccharomycescerevisiae中表达而经密码子对优化的来自S.cerevisiae的ZWF1的表达盒。
SEQ ID NO:43示出了用于生成片段114(图3)的引物的核苷酸序列。
SEQ ID NO:44示出了用于生成片段114(图3)的引物的核苷酸序列。
SEQ ID NO:45示出了用于生成片段115(图3)的引物的核苷酸序列。
SEQ ID NO:46示出了来自S.cerevisiae的丙酮酸羧化酶蛋白质的氨基酸序列。
SEQ ID NO:47示出了在第120-122位具有EGY到DAF修饰的来自Actinobacillussuccinogenes的磷酸烯醇式丙酮酸羧激酶的氨基酸序列。
SEQ ID NO:48示出了来自Escherichia coli的延胡索酸酶(fumB)的氨基酸序列。
SEQ ID NO:49示出了缺少前23个N-末端氨基酸的来自Rhizopus oryzae的延胡索酸酶的氨基酸序列。
SEQ ID NO:50示出了来自Aspergillus niger的推定的二羧酸转运体的氨基酸序列。
SEQ ID NO:51示出了来自Kluyveromyces lactis的异柠檬酸裂解酶的氨基酸序列。
SEQ ID NO:52示出了缺少3C-末端过氧化物酶体靶向序列的Saccharomycescerevisiae过氧化物酶体苹果酸合酶(Mls1)氨基酸序列的氨基酸序列。
SEQ ID NO:53示出了包括过氧化物酶体靶向序列SKL的来自S.cerevisiae的苹果酸脱氢酶(MDH3)蛋白质的氨基酸序列。
SEQ ID NO:54示出了用于生成片段116(图6)的引物的核苷酸序列。
SEQ ID NO:55示出了用于生成片段116(图6)的引物的核苷酸序列。
SEQ ID NO:56示出了用于生成片段117(图6)的引物的核苷酸序列。
SEQ ID NO:57示出了用于生成片段117(图6)的引物的核苷酸序列。
SEQ ID NO:58示出了用于生成片段119(图6)的引物的核苷酸序列。
SEQ ID NO:59示出了用于生成片段119(图6)的引物的核苷酸序列。
SEQ ID NO:60示出了用于生成片段118(图6)的引物的核苷酸序列。
SEQ ID NO:61示出了用于生成片段118(图6)的引物的核苷酸序列。
SEQ ID NO:62示出了片段118(图6)的核苷酸序列，其包括针对在S.cerevisiae中表达而经密码子对优化的来自Trypanosoma brucei的延胡索酸还原酶(FRDg)的编码序列。
SEQ ID NO:63示出了用于生成片段124(图7)的引物的核苷酸序列。
SEQ ID NO:64示出了片段121(图7)的核苷酸序列，其包括针对在S.cerevisiae中表达而经密码子对优化的S.cerevisiae MDH3突变体MUT_014的编码序列。
SEQ ID NO:65示出了片段122(图7)的核苷酸序列，其包括针对在S.cerevisiae中表达而经密码子对优化的S.cerevisiae MDH3突变体MUT_015的编码序列。
SEQ ID NO:66示出了片段123(图7)的核苷酸序列，其包括针对在S.cerevisiae中表达而经密码子对优化的S.cerevisiae MDH3突变体MUT_034的编码序列。
SEQ ID NO:67示出了来自Arabidopsis thaliana的延胡索酸酶的氨基酸序列。
具体实施方式
在本说明书和所附的权利要求书中，词语“包括”、“包含”和“具有”及其变形应被理解为“包含在内”。也就是说，在语境允许的情况下，这些词语意图传达的意思是：可以包括其它没有明确列举的要素或整体。
本文中不使用数量词修饰时指的是一个/种或多于一个/种(即一个/种或至少一个/种)对象。例如，“要素”可意味着一个/种要素或多于一个/种要素。
还原性TCA通路是微生物可产生二羧酸的主要通路之一。近年来，其已被证明是微生物生产二羧酸例如琥珀酸的最佳经济选择。还原性TCA通路包括两个需要消耗还原力的反应；即苹果酸脱氢酶反应(草酰乙酸盐/酯还原为苹果酸盐/酯)和延胡索酸还原酶反应(延胡索酸盐/酯还原为琥珀酸盐/酯)。
苹果酸脱氢酶(MDH)使用NAD或NADP作为辅因子(也统称为NAD(P))催化苹果酸盐/酯可逆地转变为草酰乙酸盐/酯。MDH是一种相当普遍存在的酶，在许多代谢通路中起着至关重要的作用，所述代谢通路包括三羧酸循环、氨基酸合成、糖异生、维持氧化/还原平衡和代谢应激。
根据MDH对辅因子的偏好，MDH可被分为NAD(H)-依赖型MDH(NAD-MDH)(EC1.1.1.37)和NADP(H)-依赖型MDH(NADP-MDH)(EC 1.1.1.82)。大多数细菌和古细菌MDH是NAD-MDH。真核MDH同种型均为NAD-MDH，包括线粒体MDH、胞质MDH、乙醛酸循环体MDH和过氧化物酶体MDH，例外是叶绿体NADP-MDH，其是将还原等同物从叶绿体基质转移到胞质所必需的。在酵母Saccharomyces cerevisiae中，已鉴定出了三种内源性苹果酸脱氢酶同工酶，即MDH1、MDH2和MDH3。它们位于线粒体(MDH1)、胞质(MDH2)和过氧化物酶体(MDH3)中，并且均被表征为NAD(H)-依赖型MDH(EC 1.1.1.37)。
对苹果酸脱氢酶家族中的NAD(P)-结合结构域的研究揭示了Rossmann折叠的保守βB-αC基序。脱氢酶区分NADP(H)的能力在于该βB-αC基序的氨基酸序列，其已被预测为辅因子特异性的主要决定簇。例如，在S.cerevisiae过氧化物酶体NAD-MDH(MDH3)中，NAD-结合基序包括第34-40位氨基酸残基，其被发现对于辅因子结合和特异性是重要的。
在本发明的上下文中，出乎意料地发现：当在能够生产二羧酸的重组宿主细胞中(过)表达时，具有MDH活性的多肽的保守NAD结合基序中的一组特定突变赋予提高的二羧酸生产。也就是说，与(过)表达参照MDH多肽的重组宿主细胞相比，在重组宿主细胞中(过)表达具有MDH活性的所述突变多肽通常导致提高的二羧酸生产；“参照MDH多肽”通常是NAD-MDH(EC 1.1.1.37)。同时，已显示：与所述参照MDH多肽相比，在保守NAD-结合基序中具有至少一个突变的所述突变多肽的NADP(H)-依赖型活性与NAD(H)-依赖型活性之比提高。出乎意料地，本发明的发明人进一步显示了：为了获得二羧酸生产的提高，NADP(H)-依赖型活性不必高于NAD(H)-依赖型活性。
因此，本发明的一个目标是提供一种重组宿主细胞，其能够生产二羧酸并且包含具有苹果酸脱氢酶(MDH)活性的突变多肽。
在一个实施方式中，具有苹果酸脱氢酶活性的突变多肽包含这样的氨基酸序列，当与包含SEQ ID NO:39所示序列的苹果酸脱氢酶比对时，所述氨基酸序列包含对应于SEQID NO:39中的第34位氨基酸的氨基酸残基的一个突变。换句话说，所述突变多肽包含在对应于SEQ ID NO:39中的第34位的位置处出现的氨基酸残基的一个突变。
在本发明的一个优选实施方式中，对应于第34位氨基酸(参照SEQ ID NO:39限定)的氨基酸的突变是替换。
更优选地，对应于第34位氨基酸(参照SEQ ID NO:39限定)的氨基酸的替换通常是替换为小氨基酸。合适的小氨基酸包括苏氨酸(T)、丝氨酸(S)、甘氨酸(G)、丙氨酸(A)和脯氨酸(P)。优选的小氨基酸是甘氨酸(G)和丝氨酸(S)。
在本发明的上下文中，“重组宿主细胞”或“经遗传修饰的宿主细胞”是已通过重组DNA技术引入包含编码具有苹果酸脱氢酶活性的突变多肽的核酸序列的核酸、核酸构建体或载体的宿主细胞。
在本文中，“具有苹果酸脱氢酶(MDH)活性的突变多肽”可被称为“突变的/型苹果酸脱氢酶”、“MDH突变体”、“MDH突变多肽”、“突变体”、“突变多肽”等。
在本文中，“苹果酸脱氢酶活性”是将草酰乙酸转变为苹果酸的活性：
苹果酸+受体<＝>草酰乙酸+还原的受体。
术语“多肽”在本文中用于含有多于约7个氨基酸残基的链。本文所有的多肽序列都是从左到右书写，并从氨基末端到羧基末端的方向书写。本文所用的氨基酸单字母代码是本领域公知的，可以在Sambrook等(Molecular Cloning:A Laboratory Manual,第二版Cold Spring Harbor Laboratory,Cold Spring Harbor Laboratory Press,Cold SpringHarbor,NY,1989)中找到。
在本发明的上下文中，“突变(型)”多肽被定义为通过引入一个或多个突变而获得的多肽。所述突变可选自替换、添加和缺失。术语“替换”在本文中是指多肽序列中的氨基酸残基被另一个氨基酸残基替代。“突变(型)”多肽、“突变的”多肽和“基因工程”多肽具有相同的含义并可互换使用。
在本文中，“对应的位置”是指SEQ ID NO:39和与SEQ ID NO:39同源的序列之间的氨基酸序列比对中的垂直栏，对应于SEQ ID NO:39中的特定位置，并显示出在其他比对的同源物中的该位置出现的氨基酸。
在本发明的上下文中，“对应的突变”是指在SEQ ID NO:39中“对应的位置”出现的氨基酸残基的突变。例如，“对应的替换”是指在SEQ ID NO:39中“对应的位置”出现的氨基酸残基被另一个氨基酸残基替换。
在一些另外的实施方式中，具有苹果酸脱氢酶活性的突变多肽可以进一步包含对应于SEQ ID NO:39中的第35、36、37、38、39和/或40位氨基酸中的任何的一个或多个的另外的突变。所述突变通常选自替换、添加和缺失。更优选地，一个或多个另外的突变是替换。
对应于第35位氨基酸(参照SEQ ID NO:39限定)的氨基酸的替换通常是替换为小氨基酸。合适的小氨基酸包括苏氨酸(T)、丝氨酸(S)、甘氨酸(G)、丙氨酸(A)和脯氨酸(P)。或者，对应于第35位氨基酸(参照SEQ ID NO:39限定)的氨基酸的替换是替换为疏水性氨基酸，例如异亮氨酸(I)。对应于第35位氨基酸(参照SEQ ID NO:39限定)的氨基酸的优选替换是替换为丝氨酸(S)或异亮氨酸(I)。
对应于第36位氨基酸(参照SEQ ID NO:39限定)的氨基酸的替换通常是替换为极性氨基酸。合适的极性氨基酸包括精氨酸(R)、谷氨酰胺(Q)、谷氨酸(E)和丝氨酸(S)。或者，对应于第36位氨基酸(参照SEQ ID NO:39限定)的氨基酸的替换是替换为小氨基酸，例如丙氨酸(A)或脯氨酸(P)。对应于第36位氨基酸(参照SEQ ID NO:39限定)的氨基酸的优选替换是替换为精氨酸(R)、谷氨酰胺(Q)、谷氨酸(E)、丝氨酸(S)、丙氨酸(A)或脯氨酸(P)。
对应于第37位氨基酸(参照SEQ ID NO:39限定)的氨基酸的替换通常是替换为小氨基酸。合适的小氨基酸包括甘氨酸(G)、天冬酰胺(N)和丙氨酸(A)。或者，对应于第37位氨基酸(参照SEQ ID NO:39限定)的氨基酸的替换是替换为极性氨基酸，例如精氨酸(R)或谷氨酰胺(Q)。对应于第37位氨基酸(参照SEQ ID NO:39限定)的氨基酸的优选替换是替换为(G)、天冬酰胺(N)、丙氨酸(A)、丙氨酸精氨酸(R)或谷氨酰胺(Q)。
对应于第38位氨基酸(参照SEQ ID NO:39限定)的氨基酸的替换通常是替换为小氨基酸。合适的小氨基酸包括缬氨酸(V)、苏氨酸(T)、丝氨酸(S)、甘氨酸(G)、丙氨酸(A)和脯氨酸(P)。对应于第38位氨基酸(参照SEQ ID NO:39限定)的氨基酸的优选替换是替换为丙氨酸(A)、缬氨酸(V)、苏氨酸(T)或丝氨酸(S)。
对应于第39位氨基酸(参照SEQ ID NO:39限定)的氨基酸的替换通常是替换为小氨基酸。合适的小氨基酸包括脯氨酸(P)。或者，对应于第39位氨基酸(参照SEQ ID NO:39限定)的氨基酸的替换是替换为疏水性氨基酸，例如赖氨酸(K)、苯丙氨酸(F)或亮氨酸(L)。或者，对应于第39位氨基酸(参照SEQ ID NO:39限定)的氨基酸的替换是替换为极性氨基酸，例如谷氨酸(E)。对应于第39位氨基酸(参照SEQ ID NO:39限定)的氨基酸的优选替换是替换为谷氨酸(E)、赖氨酸(K)、苯丙氨酸(F)、亮氨酸(L)或脯氨酸(P)。
对应于第40位氨基酸(参照SEQ ID NO:39限定)的氨基酸的替换通常是替换为小氨基酸。合适的小氨基酸包括甘氨酸(G)。或者，对应于第40位氨基酸(参照SEQ ID NO:39限定)的氨基酸的替换是替换为极性氨基酸，例如谷氨酰胺(Q)。对应于第40位氨基酸(参照SEQ ID NO:39限定)的氨基酸的优选替换是替换为甘氨酸(G)或谷氨酰胺(Q)。
以上各种类型的氨基酸参照例如Betts和Russell，Bioinformatics forGeneticists,Barnes和Gray编,Wiley 2003进行分类。
更详细地，在本发明的上下文中，具有苹果酸脱氢酶活性的突变多肽包含参照SEQID NO:39限定的第34位的G或S；
以及，任选的
参照SEQ ID NO:39限定的第35位的I或S；和/或
参照SEQ ID NO:39限定的第36位的R、Q、A、E、P或S；和/或
参照SEQ ID NO:39限定的第37位的A、N、G、R或Q；和/或
参照SEQ ID NO:39限定的38位的A、V、T或S；和/或
参照SEQ ID NO:39限定的第39位的E、K、P、F或L；和/或
参照SEQ ID NO:39限定的第40位的G或Q。
在一个具体实施方式中，具有苹果酸脱氢酶活性的突变多肽包含在第34位的小氨基酸(参照SEQ ID NO:39限定)和在第36位的小氨基酸或极性氨基酸(参照SEQ ID NO:39限定)。在所述实施方式中，第34位的优选的小氨基酸可选自G或S。在所述实施方式中，第36位的优选的小氨基酸或极性氨基酸可选自R、Q、A、E、P或S。任选地，在所述实施方式中，突变多肽包含
参照SEQ ID NO:39限定的第35位的I或S；和/或
参照SEQ ID NO:39限定的第37位的A、N、G、R或Q；和/或
参照SEQ ID NO:39限定的第38位的A、V、T或S；和/或
参照SEQ ID NO:39限定的第39位的E、K、P、F或L；和/或
参照SEQ ID NO:39限定的第40位的G或Q。
具有MDH活性的突变多肽还可以包含除上文限定的七个位置之外的另外的突变，例如，一个或多个另外的替换、添加或缺失。
具有MDH活性的突变多肽可包含不同类型的此类修饰的组合。具有MDH活性的突变多肽可包含一个、两个、三个、四个、至少5个、至少10个、至少15个、至少20个、至少25个、至少30个或更多个这样的修饰(它们可以全部是相同类型的修饰或可以是不同类型的修饰)。通常，另外的修饰可以是替换。
在另外的实施方式中，具有苹果酸脱氢酶活性的突变多肽参照表1(实施例4)限定，并且其中对应于SEQ ID NO:39中的第34位氨基酸的氨基酸残基选自甘氨酸(G)或丝氨酸(S)。也就是说，与合适的参照序列(例如SEQ ID NO:39中所示的参照序列)相比，突变多肽可以包含表1中所示的替换的任何组合，并且其中对应于SEQ ID NO:39中的第34位氨基酸的氨基酸残基选自甘氨酸(G)或丝氨酸(S)。
通常，突变多肽可以包含SEQ ID NO:39的序列，其在第34位具有一个替换，并且任选地在第35、36、37、38、39和/或40位具有一个或多个替换。也就是说，突变多肽在第34位具有不同于天冬氨酸的氨基酸，并且任选地在第35位具有不同于异亮氨酸的氨基酸，且/或在第36位具有不同于精氨酸的氨基酸，且/或在第37位具有不同于丙氨酸的氨基酸，且/或第38位具有不同于丙氨酸的氨基酸，且/或在第39位具有不同于谷氨酸的氨基酸，且/或在第40位具有不同于甘氨酸的氨基酸。
此外，通常，突变多肽可以包含SEQ ID NO:39的序列，其在第34位具有一个替换，在第36位具有一个替换，并且任选地在第35、37、38、39和/或40位具有一个或多个替换。也就是说，突变多肽在第34位具有不同于天冬氨酸的氨基酸，在第36位具有不同于精氨酸的氨基酸，并且任选地在第35位具有不同于异亮氨酸的氨基酸，且/或在第37位具有不同于丙氨酸的氨基酸，且/或第38位具有不同于丙氨酸的氨基酸，且/或在第39位具有不同于谷氨酸的氨基酸，且/或在第40位具有不同于甘氨酸的氨基酸。
在本发明的一个单独的实施方式中，与参照MDH多肽相比，具有苹果酸脱氢酶活性的突变多肽的NADP(H)-依赖型活性与NAD(H)-依赖型活性之比提高。在所述实施方式中，所述突变多肽可以是包含这样的氨基酸序列的突变多肽，当与包含SEQ ID NO:39所示序列的苹果酸脱氢酶比对时，所述氨基酸序列包含对应于SEQ ID NO:39中的第34位氨基酸的氨基酸残基的一个突变(例如一个替换)。关于所述突变多肽的氨基酸序列的其他实施方式如上所述。
在本发明的上下文中，具有苹果酸脱氢酶活性的参照多肽(也称为“参照MDH多肽”)可以是NAD-MDH(EC 1.1.1.37)。具有苹果酸脱氢酶活性的参照多肽可以是来自微生物来源例如酵母(例如Saccharomyces cerevisiae)的苹果酸脱氢酶。具有SEQ ID NO:39所示氨基酸序列的苹果酸脱氢酶可以是具有MDH活性的合适参照多肽。
表述“NADP(H)-依赖型活性与NAD(H)-依赖型活性之比提高”在本文中通常是指：与参照MDH多肽相比，例如与SEQ ID NO:39相比，突变多肽显示出NADP(H)-依赖型活性与NAD(H)-依赖型活性之比提高的性质。也就是说，与参照多肽相比，突变多肽可以显示出NADP(H)-依赖型活性与NAD(H)-依赖型活性的比值提高。在实施例5中，该比值也被称为“NADPH特异性:NADH特异性的比值”。
在本发明的上下文中，术语“NADP(H)-依赖型活性”和“NADP(H)-特异性活性”在本文中具有相同含义并可互换使用。这同样适用于术语“NAD(H)-依赖型活性”和“NAD(H)-特异性活性”。
术语“NADP(H)-依赖型活性”在本文中是指酶使用NADP(H)作为氧化还原辅因子的性质。酶的NADP(H)-依赖型活性可以通过酶活性试验(例如实施例5中所述)测定。
术语“NAD(H)-依赖型活性”在本文中是指酶使用NAD(H)作为氧化还原辅因子的性质。酶的NAD(H)-依赖型活性可以通过酶活性试验(例如实施例5中所述)测定。
平均NADPH特异性:NADH特异性比值的提高值可以指示，例如，降低的NAD(H)-依赖型活性、提高的NADP(H)-依赖型活性或两者的组合。在一些情况下，与参照MDH相比，使用具有相似或提高的NAD(H)-依赖型活性的MDH突变体可以获得所述比值的提高值。在后一情况下，MDH突变体可显示出提高的NAD(H)-依赖型活性和提高的NADP(H)-依赖型活性。
就改变的辅因子依赖型而言，突变MDH多肽通常具有改变的MDH活性。该NAD(H)-依赖型活性或NADP(H)-依赖型活性可以彼此独立地改变，例如降低至少10％、至少20％、至少30％、至少40％、至少50％、至少60％、至少70％、至少80％、至少90％、至少95％或至少99％。或者，该性质可以提高至少10％、至少25％、至少50％、至少70％、至少100％、至少200％、至少500％、至少700％、至少1000％、至少3000％、至少5000％或至少6000％。
在一个具体实施方式中，突变MDH多肽的NAD(H)-依赖型活性和NADP(H)-依赖型活性均提高。所述突变MDH多肽的NAD(H)-依赖型活性提高至少10％、至少25％、至少50％、至少70％、至少100％、至少200％或至少300％。所述突变MDH多肽的NADP(H)-依赖型活性提高至少10％、至少25％、至少50％、至少70％、至少100％、至少200％、至少500％、至少700％、至少1000％、至少3000％、至少5000％或至少6000％。
在另一个实施方式中，突变MDH多肽的NAD(H)-依赖型活性大约相同或降低至多5％、至多10％、至多20％、至多30％、至多40％、至多50％、至多60％、至多70％、至多80％，并且所述突变MDH多肽的NADP(H)-依赖型活性提高至少10％、至少25％、至少50％、至少70％、至少100％、至少200％、至少500％、至少700％、至少1000％、至少3000％、至少5000％或至少6000％。
在另一个实施方式中，突变型MDH多肽的NADPH特异性:NADH特异性的比值增加至少10％、至少50％、至少100％、至少200％、至少500％、至少1000％、至少3000％、至少5000％、至少7000％或至少8000％。
在该上下文中百分比的降低或提高表示与参照MDH多肽(例如SEQ ID NO:39的参照MDH多肽)相比的百分比降低或提高。本领域技术人员公知如何测量这样的百分比变化–其是如实施例中所述测量的参照MDH和突变体MDH的活性(例如NAD(H)-依赖型活性或NADP(H)-依赖型活性)的比较。
在本发明的上下文中，本所述的MDH突变多肽可以是突变型NAD(P)-苹果酸脱氢酶，例如突变型线粒体NAD-MDH、突变型胞质NAD-MDH、突变型乙醛酸循环体NAD-MDH、突变型过氧化物酶体NAD-MDH或突变型叶绿体NADP-MDH。也就是说，具有苹果酸脱氢酶活性的突变多肽可以通过在下述NAD(P)-苹果酸脱氢酶中引入一个或多个突变来获得，所述NAD(P)-苹果酸脱氢酶例如线粒体NAD-MDH、胞质NAD-MDH、乙醛酸循环体NAD-MDH、过氧化物酶体NAD-MDH或叶绿体NADP-MDH，(后者被称为用于引入所述一个或多个突变型的模板MDH多肽。优选地，具有苹果酸脱氢酶活性的突变多肽是突变型NAD(H)-苹果酸脱氢酶(EC 1.1.1.37)。更优选地，具有苹果酸脱氢酶活性的突变多肽是突变型过氧化物酶体NAD(H)-苹果酸脱氢酶。甚至更优选地，具有苹果酸脱氢酶活性的突变多肽是来自酵母或真菌的突变型NAD(H)-苹果酸脱氢酶。甚至更优选地，具有苹果酸脱氢酶活性的突变多肽是来自酵母或真菌的突变型NAD(H)-苹果酸脱氢酶，所示酵母或真菌例如S.cerevisiae、Torulaspora delbrueckii、Zygosaccharomyces bailii、Naumovozyma castellii、Naumovozyma dairenensis、Lachancea lanzarotensis、Zygosaccharomyces rouxii、Kazachstania Africana、Candida tropicalis、Kluyveromyces marxianus、Scheffersomyces stipites、Talaromyces marneffei、Rasamsonia emersonii、Aspergillus niger或Trametesversicolor。以下Uniprot数据库代码是指合适的酵母和真菌模板MDH多肽(http:// www.uniprot.org)：E7NGH7、G8ZXS3、G0V668、W0VUI8、G0WB63、A0A0W0D4X6、A0A0C7MME9、C5DQ42、C5DI45、H2AWW6、A0A090C493、J7R0C8、Q6CJP3、Q759M4、I2H037、C5M546、A7TL95、A0A0L0P3G3、Q6BM17、S8AW17、V5FMV2、A3LW84、A0A109UZS1、G8BVW8、G8BJ12、M3HPK4、A0A093UW53、B8MTP0、A0A0F4YPR0、C8V0H6、W6QNU3、A5DZ33、U1GAT6、G3B7S5、C4JPI7、A0A0F8UZY9、Q4WDM0、A0A093UPX3、B8ND04、A0A0M9VRI4、G7XZ98、Q5A5S6、M7S9E4、E4UYX5、A5DE02、A0A0J7BIJ5、A0A017SKI1、G8Y7A1、A0A0G2EFQ2、R7S165、I1RFM4、R1EVC8、U4L6K9、A0A0L0P507、W7HM94、A5DGY9、F2QY33、A0A0G2JA24、UPI000462180C、C7Z9W6、E5AAQ2、B2VVR8、A0A0H2RCV1、A8Q524、A0A0E9N879、N1JA02、A0A0D1ZEE3、J5T1X5、W6MY07、C4Y826、G3AJA2、G9N6G5、A0A0K8L6L9、A3GH28、A8P7W6、K5W0T4、G1XT67、A0A0B7K175、B8MTP5、A0A0D1ZAS7、A0A0C3BQC4、K5Y2Q9、A0A0C3DSW4、A0A068S518、W2RPL2、A0A0H5C453、A0A074WKG5、G8JRX4、A0A0U1M134、H6C0V9、A0A0H5BZ30、M2UXR7、A0A0C9YJV6、A0A0C3S6T1、W1QK02、A0A0C2YFC4、A0A061AJ54、A0A086T183、W2RVT1、UPI0004623914、A0A0C9X7U1、UPI0001F26169、G7E054、A0A0C2S9T3、A0A067NIX1、L8FPM0、G3ALW4、A0A0D0AYX2、G0VJG3、A0A0D2NGY0、C1GLB8、W9X415、A0A0D0CDE8、S7Q8G0、A0A0C9TB51、R7YP89、A0A0C2W4H8、UPI000455FA04、A0A067NL73、A0A067STP4、W9WWP5、A0A0D7A9T7、A0A0D6R2E1、M2YLX9、G0W7D4、N1QJ61、G4TRY5、F0XJ10、A0A063BQQ6、A0A061HBX5、A0A0A1PCT2、M1WIC4、A8QAQ2、A0A0C3N631、F2QTL7、A0A060S7U3、A0A0L0HTQ9、Q0CKY1、A0A0C2ZJ90、K5W527、I4Y5C3、R7YXZ2、F9XI12、A0A061J968、F9XL74、A0A0D0BEW9、A0A0C9WB72、F7WA21、A8NJ67、M2P8M6、W4KHW3、A0A0L1I2T4、UPI0004449A9D、P83778、C4Y9Q7、A0A0D7BHV7、A0A068RWX9、M2YWQ3、A0A137QV51、A0A0D0B6C2、I1BQQ7、S3DA07、Q4DXL5、G8Y022、A0A0C9W9B3、A0A0L6WJ49、A0A0L9SL52、D5GA85、A0A0N1J4Z6、J7S1G2、A0A0F4X5C6、Q6CIK3、A0A067QNN0、Q0UGT7、F8ND69、U5HIM0、J3NKC7、A0A061ATZ7、A5DSY0、Q9Y750、UPI0004F4119A、A0A086TL69、A0A0J0XSS4、UPI0003F496E1、UPI000455F0EA、S7RXX1、A0A067NAG9、A0A0B7N3M5、E6ZKH0、C8V1V3、A7UFI6、T5AEM1、A0A072PGA9、A0A094EKH6、S8ADX4、G8C073、Q6BXI8、G2R9I6和U9TUL6。甚至更优选地，具有苹果酸脱氢酶活性的突变多肽是突变型S.cerevisiae过氧化物酶体NAD-MDH(MDH3)。
另外，在本发明的重组宿主细胞中，具有苹果酸脱氢酶活性的突变多肽可以是同源或异源NAD(P)-苹果酸脱氢酶的突变体。在一个优选的实施方式中，MDH突变体是同源NAD(P)-苹果酸脱氢酶的突变体。更优选地，MDH突变体是同源NAD(H)-苹果酸脱氢酶(EC1.1.1.37)的突变体。更优选地，具有苹果酸脱氢酶活性的突变多肽是同源过氧化物酶体NAD(H)-苹果酸脱氢酶的突变体。
在该上下文中，当用于指出给定的(重组)核酸或多肽分子与给定的宿主生物或宿主细胞之间的相互关系时，术语“同源的”或“内源的”应被理解为表示该核酸或多肽分子天然地由相同物种、优选地相同品种或株系的宿主细胞或生物生产。
在本文中使用时，术语“异源的”是指非天然存在于宿主细胞中的核酸或氨基酸序列。换句话说，该核酸或氨基酸序列与宿主细胞中天然存在的序列不同。
优选地，在本发明的重组宿主细胞中，编码具有苹果酸脱氢酶活性的所述突变多肽的核酸序列在胞质中表达，并且具有苹果酸脱氢酶活性的突变多肽在胞质中有活性。在一些情况下，可通过缺失过氧化物酶体或线粒体靶向信号来获得胞质表达。过氧化物酶体或线粒体靶向信号的存在可以例如通过Schlüter et al.,Nucleid Acid Research 2007,35,D815-D822所公开的方法确定。当MDH突变体是突变型S.cerevisiae过氧化物酶体NAD-MDH(例如突变体MDH3)时，优选缺失其C-末端SKL，使其在胞质中有活性。
通常，具有苹果酸脱氢酶活性的突变多肽可与参照MDH多肽(例如SEQ ID NO:53或SEQ ID NO:39的MDH)具有至少约40％、50％、60％、70％、80％的序列同一性，例如与参照MDH多肽具有至少85％的序列同一性，例如与参照MDH多肽具有至少约90％的序列同一性，与参照MDH多肽具有至少95％的序列同一性，与参照MDH多肽具有至少98％的序列同一性或与参照MDH多肽具有至少99％的序列同一性。
出乎意料地发现：与(过)表达具有MDH活性的参照多肽的等同重组宿主细胞的二羧酸生产水平相比，当在重组宿主细胞中(过)表达上文所述的突变型MDH多肽时，所述突变型MDH多肽导致所述重组宿主细胞中所述二羧酸的生产提高；“参照MDH多肽”通常是NAD-MDH(EC1.1.1.37)，例如具有SEQ ID NO:39所示氨基酸序列的苹果酸脱氢酶。
因此，提供了一种重组宿主细胞，其生产或能够生产二羧酸，并且包含编码如上所述的具有苹果酸脱氢酶活性的突变多肽的核酸序列。
本发明的重组宿主细胞生产或能够生产二羧酸，例如苹果酸、延胡索酸和/或琥珀酸。
术语“二羧酸”和“二羧酸盐/酯”(例如“琥珀酸”和“琥珀酸盐/酯”)在本文中具有相同的含义并可互换使用，前者是后者的氢化形式。
通常，与表达参照MDH多肽(例如SEQ ID NO:39的参照MDH多肽)的重组宿主细胞相比，本发明的重组宿主细胞生产提高量的二羧酸。二羧酸生产可提高至少5％、10％、至少20％、至少30％、至少40％、至少50％、至少60％、至少70％、至少80％、至少90％、至少95％或至少100％或更多。生产水平可以用g/L表示，因此二羧酸生产水平的增加通过以g/L为单位的更高的生产水平来表示。
本发明的重组宿主细胞或所述宿主细胞的亲本可以是任何类型的宿主细胞。因此，包括原核细胞和真核细胞。宿主细胞还可包括但不限于哺乳动物细胞系，例如CHO、VERO、BHK、HeLa、COS、MDCK、293、3T3、WI38和脉络丛细胞系。
合适的本发明宿主细胞可以是原核细胞。优选地，所述原核细胞是细菌细胞。术语“细菌细胞”包括革兰氏阴性微生物和革兰氏阳性微生物。
合适的细菌可以选自例如Escherichia、Actinobacillus、Anabaena、Caulobactert、Gluconobacter、Mannheimia、Basfia、Rhodobacter、Pseudomonas、Paracoccus、Bacillus、Brevibacterium、Corynebacterium、Rhizobium(Sinorhizobium)、Flavobacterium、Klebsiella、Enterobacter、Lactobacillus、Lactococcus、Methylobacterium、Staphylococcus或Actinomycetes例如Streptomyces和Actinoplanes种。优选地，细菌细胞选自Bacillus subtilis、B.amyloliquefaciens、B.licheniformis、B.puntis、B.megaterium、B.halodurans、B.pumilus、Actinobacillus succinogenes、Gluconobacter oxydans、Caulobacter crescentus CB 15、Methylobacteriumextorquens、Rhodobacter sphaeroides、Pseudomonas zeaxanthinifaciens、Pseudomonasputida、Pseudomonas fluorescens、Paracoccus denitrificans、Escherichia coli、Corynebacterium glutamicum、Mannheimia succinoproducens、Basfiasuccinoproducens、Staphylococcus carnosus、Streptomyces lividans、Streptomycesclavuligerus、Sinorhizobium melioti和Rhizobium radiobacter。
根据本发明的宿主细胞可以是真核宿主细胞。优选地，所述真核细胞是哺乳动物、昆虫、植物、真菌或藻类细胞。更优选地，真核细胞是真菌细胞。合适的真菌细胞可以例如属于Saccharomyces、Schizosaccharomyces、Aspergillus、Penicillium、Pichia、Kluyveromyces、Yarrowia、Candida、Hansenula、Humicola、Pichia、Issatchenkia、Kloeckera、Schwanniomyces、Torulaspora、Trichosporon、Brettanomyces、Rhizopus、Zygosaccharomyces、Pachysolen或Yamadazyma属。真菌细胞可以例如属于Saccharomycescerevisiae、S.uvarum、S.bayanus S.pastorianus、S.carlsbergensis、Aspergillusniger、Penicilliumchrysogenum、Pichia stipidis、P.pastoris、Kluyveromycesmarxianus、K.lactis、K.thermotolerans、Yarrowia lipolytica、Candida sonorensis、C.revkaufi、C.pulcherrima、C.tropicalis、C.utilis、C.kruisei、C.glabrata、Hansenulapolymorpha、Issatchenkia orientalis、Torulaspora delbrueckii、Brettanomycesbruxellensis、Rhizopus oryzae或Zygosaccharomyces bailii种。在一种实施方式中，本发明的真菌细胞是酵母，例如属于Saccharomyces sp.，例如Saccharomyces cerevisiae。
具体宿主酵母细胞的实例包括C.sonorensis、K.marxianus、K.thermotolerans、C.methanesorbosa、Saccharomyces bulderi(S.bulderi)、P.kudriavzevii、I.orientalis、C.lambica、C.sorboxylosa、C.zemplinina、C.geochares、P.membranifaciens、Z.kombuchaensis、C.sorbosivorans、C.vanderwaltii、C.sorbophila、Z.bisporus、Z.lentus、Saccharomyces bayanus(S.bayanus)、D.castellii、C、boidinii、C.etchellsii、K.lactis、P.jadinii、P.anomala、Saccharomyces cerevisiae(S.cerevisiae)、Pichia galeiformis、Pichia sp.YB-4149(NRRL命名)、Candida ethanolica、P.deserticola、P.membranifaciens、P.fermentans和Saccharomycopsis crataegensis(S.crataegensis)。K.marxianus和C.sonorensis的合适菌株包括WO 00/71738Al、WO 02/42471A2、WO 03/049525A2、WO 03/102152A2和WO 03/102201A2中描述的那些。I.orientalis的合适菌株是ATCC菌株32196和ATCC菌株PTA-6648。在本发明中，宿主细胞可以是克拉布特里(Crabtree)阴性的野生型菌株。克拉布特里效应被定义为：当以高比生长速率(长期效应)或在存在高浓度葡萄糖的情况下(短期效应)培养时，由于微生物氧消耗的抑制在有氧条件下发生的发酵性代谢。克拉布特里阴性表型不展示出这种效应，因此即使在存在高浓度葡萄糖的情况下或在高生长速率下也能够消耗氧。
真核细胞可以是丝状真菌细胞。丝状真菌包括真菌(Eumycota)和卵菌(Oomycota)亚门的所有丝状形式(如Hawksworth等人在Ainsworth and Bisby's Dictionary of TheFungi,第8版,1995,CAB International,University Press,Cambridge,UK中定义)。丝状真菌的特征为由几丁质、纤维素、葡聚糖、壳聚糖、甘露聚糖和其它复杂多糖构成的菌丝壁。营养生长通过菌丝伸长进行且碳分解代谢专性好氧。丝状真菌菌株包括但不限于Acremonium、Aspergillus、Agaricus、Aureobasidium、Cryptococcus、Corynascus、Chrysosporium、Filibasidium、Fusarium、Humicola、Magnaporthe、Monascus、Mucor、Myceliophthora、Mortierella、Neocallimastix、Neurospora、Paecilomyces、Penicillium、Piromyces、Phanerochaete Podospora、Pycnoporus、Rhizopus、Schizophyllum、Sordaria、Talaromyces、Rasamsonia、Thermoascus、Thielavia、Tolypocladium、Trametes和Trichoderma的菌株。可作为宿主细胞的优选丝状真菌菌株属于Aspergillus niger、Aspergillus oryzae、Aspergillus fumigatus、Penicilliumchrysogenum、Penicillium citrinum、Acremoniumchrysogenum、Trichoderma reesei、Rasamsonia emersonii(以前被称为Talaromyces emersonii)、Aspergillus sojae、Chrysosporium lucknowense、Myceliophtora thermophyla种。用于比较经转化细胞和未转化细胞的发酵特征的参照宿主细胞包括例如Aspergillus niger CBS120.49、CBS513.88、Aspergillus oryzae ATCC16868、ATCC 20423、IFO 4177、ATCC 1011、ATCC 9576、ATCC14488-14491、ATCC 11601、ATCC12892、Aspergillus fumigatus AF293(CBS101355)、P.chrysogenum CBS 455.95、Penicilliumcitrinum ATCC 38065、Penicilliumchrysogenum P2、Thielavia terrestris NRRL8126、Talaromyces emersonii CBS124.902、Rasamsonia emersonii CBS393.64、Acremonium chrysogenum ATCC 36225、ATCC48272、Trichoderma reesei ATCC 26921、ATCC 56765、ATCC 26921、Aspergillus sojaeATCC11906、Chrysosporium lucknowense ATCC44006以及所有这些菌株的衍生株。
一种更优选的宿主细胞属于Aspergillus属，更优选地，宿主细胞属于Aspergillus niger种。当根据本发明的宿主细胞是Aspergillus niger宿主细胞时，宿主细胞优选是CBS 513.88、CBS124.903或其衍生株。
在一种优选的实施方式中，根据本发明的宿主细胞是选自Candida、Hansenula、Issatchenkia、Kluyveromyces、Pichia、Saccharomyces、Schizosaccharomyces或Yarrowia菌株的酵母细胞，或者选自属于Acremonium、Aspergillus、Chrysosporium、Myceliophthora、Penicillium、Talaromyces、Rasamsonia、Thielavia、Fusarium或Trichoderma种的丝状真菌细胞的丝状真菌细胞。
本发明的宿主细胞可以是生产二羧酸的任何野生型菌株。此外，合适的宿主细胞可以是通过对感兴趣的亲本或野生型细胞进行经典诱变处理或重组核酸转化而获得和/或改善的细胞。因此，合适的宿主细胞可以已经能够产生二羧酸。然而，细胞也可以具有同源或异源表达构建体，所述构建体编码参与二羧酸生产的一种或多种多肽。
因此，在一些实施方式中，本发明的重组宿主细胞可包含MDH突变型多肽和从磷酸烯醇式丙酮酸或丙酮酸至琥珀酸的活性还原性三羧酸(TCA)通路。
因此，除了编码MDH突变多肽的核酸之外，本发明的宿主细胞可以包含这样的核苷酸序列，所述核苷酸序列包含编码以下之中的一种或多种的序列：丙酮酸羧化酶、磷酸烯醇式丙酮酸羧激酶、磷酸烯醇式丙酮酸羧化酶、苹果酸脱氢酶、延胡索酸酶、异柠檬酸裂解酶、苹果酸合酶、延胡索酸还原酶和/或二羧酸转运体。优选地，一种或多种这样的酶(过)表达并且在胞质中有活性。
因此，本发明的重组宿主细胞可过表达合适的同源或异源核苷酸序列，所述核苷酸序列编码催化细胞内反应的内源和/或异源酶，从而导致朝向二羧酸(例如苹果酸、延胡索酸和/或琥珀酸)的通量增加。
本发明的重组宿主细胞可过表达如下文所述的内源或异源核酸序列。
本发明的重组宿主细胞可包含丙酮酸羧化酶(PYC)的遗传修饰，丙酮酸羧化酶催化从丙酮酸盐/酯到草酰乙酸盐/酯的反应(EC 6.4.1.1)。丙酮酸羧化酶例如可在基因表达后在胞质中有活性。例如，宿主细胞过表达丙酮酸羧化酶，例如内源性或同源丙酮酸羧化酶被过表达。可以用与SEQ ID NO:46的核酸序列所编码的氨基酸序列具有至少70％，优选至少75％、80％、85％、90％、91％、92％、93％、94％、95％、96％、97％、98％、99％或100％序列同一性的丙酮酸羧化酶遗传修饰根据本发明的重组真菌宿主细胞。
优选地，重组宿主细胞表达编码胞质中的磷酸烯醇式丙酮酸(PEP)羧激酶的核苷酸序列。优选地，编码PEP羧激酶的核苷酸序列被过表达。PEP羧激酶(EC 4.1.1.49)优选地是异源酶，优选地来自细菌，更优选地，具有PEP羧激酶活性的酶来自Escherichia coli、Mannheimia sp.、Actinobacillus sp.或Anaerobiospirillum sp.，更优选地Mannheimiasucciniciproducens。编码PEP羧激酶的基因可被过表达，并且在真菌细胞的胞质中有活性。优选地，用与SEQ ID NO:47的氨基酸序列具有至少70％，优选至少75％、80％、85％、90％、91％、92％、93％、94％、95％、96％、97％、98％、99％或100％序列同一性的PEP羧激酶遗传修饰根据本发明的重组宿主细胞。
优选地，重组宿主细胞表达编码胞质中的磷酸烯醇式丙酮酸(PEP)羧化酶的核苷酸序列。优选地，编码PEP羧化酶的核苷酸序列被过表达。PEP羧化酶(EC 4.1.1.31)优选地是异源酶，优选地来自细菌。
在一种实施方式中，用编码在基因表达之后在胞质中有活性的苹果酸脱氢酶(MDH)的基因进一步遗传修饰重组宿主细胞。胞质表达可以通过缺失过氧化物酶体靶向信号来实现。苹果酸脱氢酶可被过表达。胞质MDH可以是催化从草酰乙酸盐/酯到苹果酸盐/酯的反应的任何合适的同源或异源苹果酸脱氢酶(EC 1.1.1.37)，例如来自S.cerevisiae。
优选地，MDH是S.cerevisiae MDH3，更优选具有C-末端SKL缺失从而使得其在胞质中有活性。优选地，根据本发明的重组真菌细胞包含编码与SEQ ID NO:39的氨基酸序列具有至少70％，优选至少75％、80％、85％、90％、91％、92％、93％、94％、95％、96％、97％、98％、99％或100％序列同一性的苹果酸脱氢酶的核苷酸序列。
在另一种实施方式中，用编码催化从苹果酸到延胡索酸的反应的延胡索酸酶(EC4.2.1.2)的基因进一步遗传修饰本公开的重组宿主细胞。编码延胡索酸酶的基因可来自于任何合适的来源，优选地来自微生物来源，例如酵母(例如Saccharomyces)或丝状真菌(例如Rhizopus oryzae)或细菌(例如Escherichia coli)。本公开的宿主细胞可过表达编码延胡索酸酶的核苷酸序列。在核苷酸序列表达后，延胡索酸酶可在胞质中有活性，例如通过缺失过氧化物酶体靶向信号来实现。已发现延胡索酸酶的胞质活性导致真菌细胞对二羧酸的高生产力。
优选地，本发明的重组宿主细胞过表达编码与SEQ ID NO:48、SEQ ID NO:49或SEQID NO:67的氨基酸序列具有至少70％，优选至少75％、80％、85％、90％、91％、92％、93％、94％、95％、96％、97％、98％、99％或100％序列同一性的延胡索酸酶的核苷酸序列。
在另一种实施方式中，用编码催化从延胡索酸盐/酯到琥珀酸盐/酯的反应的NAD(H)依赖型延胡索酸还原酶(EC 1.3.1.6)的任何合适的同源或异源基因遗传修饰重组宿主细胞。NAD(H)-依赖型延胡索酸还原酶可以是异源酶，其可来自任何合适的来源，例如细菌、真菌、原生动物或植物。本公开的真菌细胞包含异源NAD(H)依赖型延胡索酸还原酶，优选地来自Trypanosoma sp，例如Trypanosoma brucei。在一种实施方式中，NAD(H)依赖型延胡索酸还原酶被表达并在胞质中有活性，例如通过缺失过氧化物酶体靶向信号来实现。宿主细胞可过表达编码NAD(H)依赖型延胡索酸还原酶的基因。
优选地，用与SEQ ID NO:8的氨基酸序列具有至少70％，优选至少75％、80％、85％、90％、91％、92％、93％、94％、95％、96％、97％、98％、99％或100％序列同一性的NAD(H)依赖型延胡索酸还原酶遗传修饰根据本发明的重组宿主细胞。还优选地，用WO2015/086839中所公开的具有延胡索酸还原酶活性的变体多肽遗传修饰根据本发明的宿主细胞。
在另一种实施方式中，本发明的重组宿主细胞表达编码二羧酸转运蛋白的核苷酸序列。优选地，二羧酸转运蛋白被过表达。二羧酸转运蛋白可以是任何合适的同源或异源蛋白。优选地，二羧酸转运蛋白是异源蛋白。二羧酸转运蛋白可以来自任何合适的生物体，优选来自酵母或真菌，例如Schizosaccharomyces pombe或Aspergillus niger。优选地，二羧酸转运蛋白是这样的二羧酸转运蛋白/苹果酸转运蛋白，例如来自Aspergillus niger，其与SEQ ID NO:50的氨基酸序列具有至少70％，优选至少75％、80％、85％、90％、91％、92％、93％、94％、95％、96％、97％、98％、99％或100％序列同一性。
重组宿主细胞可以进一步包含编码异柠檬酸裂合酶(EC 4.1.3.1)的基因的遗传修饰，所述异柠檬酸裂合酶可以是任何合适的同源或异源酶。异柠檬酸裂合酶可例如获自Kluyveromyces lactis或Escherichia coli。
用与SEQ ID NO:51的核酸序列所编码的氨基酸序列具有至少70％，优选至少75％、80％、85％、90％、91％、92％、93％、94％、95％、96％、97％、98％、99％或100％序列同一性的异柠檬酸裂合酶遗传修饰根据本发明的重组宿主。
重组宿主细胞可以进一步包含苹果酸合酶(EC 2.3.3.9)的遗传修饰。苹果酸合酶可被过表达和/或在胞质中有活性，例如通过缺失过氧化物酶体靶向信号来实现。在苹果酸合酶是S.cerevisiae苹果酸合酶的情况下，例如通过缺失SKL羧基端序列来改变天然苹果酸合酶。
用与SEQ ID NO:52的核酸序列所编码的氨基酸序列具有至少70％，优选至少75％、80％、85％、90％、91％、92％、93％、94％、95％、96％、97％、98％、99％或100％序列同一性的苹果酸合酶遗传修饰本发明的重组宿主细胞。
在另一种实施方式中，本文所公开的本发明的重组宿主细胞包含编码丙酮酸脱羧酶(EC 4.1.1.1)的基因的破坏，所述丙酮酸脱羧酶催化从丙酮酸盐/酯到乙醛的反应。
在另一种实施方式中，本发明的重组宿主细胞可以包含编码乙醇发酵通路的酶的基因的破坏。编码乙醇发酵通路的酶的基因可以是丙酮酸脱羧酶(EC 4.1.1.1)，其催化从丙酮酸盐/酯到乙醛的反应；或醇脱氢酶(EC1.1.1.1)，其催化从乙醛到乙醇的反应。优选地，本发明的宿主细胞包含一种、两种或更多种编码醇脱氢酶的基因的破坏。如果真菌细胞是酵母，例如S.cerevisiae，则酵母优选包含一种或多种醇脱氢酶基因(adh1、adh2、adh3、adh4、adh5、adh6)的破坏。
替代性地或另外，本发明的重组宿主细胞可包含至少一种无功能的编码甘油-3-磷酸脱氢酶的基因。无功能的甘油-3-磷酸脱氢酶基因在本文中用于描述这样的真核细胞，其包含降低的甘油-3-磷酸脱氢酶活性，例如通过突变、破坏或缺失编码甘油-3-磷酸脱氢酶的基因，从而导致与野生型酵母相比甘油形成降低。如果真菌细胞是酵母，例如S.cerevisiae，则酵母优选包含一种或多种甘油-3-磷酸脱氢酶基因(gpd1、gpd2、gut2)的破坏。
替代性地或另外，本发明的重组宿主细胞可包含至少一种无功能的编码线粒体外NADH脱氢酶的基因。无功能的线粒体外NADH脱氢酶基因在本文中用于描述这样的真核细胞，其包含降低的NADH脱氢酶活性，例如通过突变、破坏或缺失编码线粒体外NADH脱氢酶的基因。如果真菌细胞是酵母，例如S.cerevisiae，则酵母优选包含一种或多种线粒体外NADH脱氢酶基因(nde1、nde2)的破坏。
替代性地或另外，本发明的重组宿主细胞可包含至少一种无功能的编码醛脱氢酶的基因。无功能的醛脱氢酶基因在本文中用于描述这样的真核细胞，其包含降低的醛脱氢酶活性，例如通过突变、破坏或缺失编码醛脱氢酶的基因。如果真菌细胞是酵母，例如S.cerevisiae，则酵母优选包含一种或多种醛脱氢酶基因(ald2、ald3、ald4、ald5、ald6)的破坏。
优选地，本发明的重组宿主细胞是重组真菌细胞。更优选地，本发明的宿主细胞是重组酵母细胞。重组真菌细胞或重组酵母细胞的优选实施方式如上文关于重组宿主细胞所述。
在本发明的一些实施方式中，重组宿主细胞是重组酵母细胞，其能够生产如上所述的二羧酸，并且包含编码具有如上文详述的苹果酸脱氢酶活性的突变多肽的核酸序列。所述MDH突变体可以是同源或异源野生型MDH多肽的突变体。在一个优选的实施方式中，所述MDH突变体是同源MDH多肽的突变体。在一个甚至更优选的实施方式中，重组酵母细胞是重组Saccharomyces，例如S.cerevisiae，并且MDH突变体是同源MDH(例如MDH2或MDH3)的突变体。在一个更具体的实施方式中，所述重组酵母细胞包含编码如表1中所限定的具有苹果酸脱氢酶活性的突变多肽的核酸序列，并且其中对应于SEQ ID NO:39中的第34位氨基酸的氨基酸残基选自甘氨酸(G)或丝氨酸(S)。
标准的遗传学技术例如宿主细胞中酶的过表达、宿主细胞的遗传修饰或杂交技术是本领域已知的方法，例如Sambrook等人(Molecular Cloning:A Laboratory Manual,第三版,Cold Spring Harbor Laboratory Press,Cold Spring Harbor Laboratory,NY,2001)或Ausubel等人(Current protocols in molecular biology,Green Publishingand Wiley Interscience,NY,1987)中所述。用于真菌宿主细胞的遗传修饰、转化的方法从例如EP-A-0 635 574、WO 98/46772、WO 99/60102和WO 00/37671、WO90/14423、EP-A-0481008、EP-A-0635 574和US 6,265,186中已知。
在本文中使用时，术语“核酸”、“多核苷酸”或“核酸分子”旨在包括DNA分子(例如cDNA或基因组DNA)和RNA分子(例如mRNA)以及使用核苷酸类似物产生的DNA或RNA的类似物。核酸分子可以是单链或双链的，但优选是双链DNA。可以使用寡核苷酸类似物或衍生物(例如肌苷或硫代磷酸酯核苷酸)合成核酸。这种寡核苷酸可用于例如制备具有改变的碱基配对能力或增加的对核酸酶的抗性的核酸。
术语“核酸构建体”在本文中被称为单链或双链的核酸分子，其是从天然存在的基因分离的，或者更典型地，其已被修饰为含有以不会在自然中存在的方式组合和并置的核酸的区段。当核酸构建体含有在宿主细胞中表达编码序列所需的全部控制序列时，术语核酸构建体与术语“表达盒”的含义相同，其中所述控制序列与所述编码序列可操作地连接。
在本文中使用时，术语“可操作地连接”是指处于功能关系的多核苷酸元件(或编码序列或核酸序列)的连接。当核酸序列被置于与另一核酸序列的功能关系中时，其是“可操作地连接”的。例如，如果启动子或增强子影响编码序列的转录，则其与编码序列可操作地连接。
在本文中使用时，术语“启动子”是指这样的核酸片段，其功能为控制位于该基因的转录起始位点的转录方向上游的一个或多个基因的转录，并且在结构上通过DNA依赖性RNA聚合酶的结合位点、转录起始位点和本领域技术人员已知的任何其他DNA序列的存在来鉴定。“组成型”启动子是在大多数环境和发育条件下有活性的启动子。“诱导型”启动子是在环境调控或发育调控下有活性的启动子。
能够用于实现编码酶(例如延胡索酸还原酶或本发明的宿主细胞中引入的任何其它酶)的核苷酸序列表达的启动子对编码待表达的酶的核苷酸序列而言可以不是天然的，即对与之可操作地连接的核苷酸序列(编码序列)而言异源的启动子。优选地，启动子对宿主细胞而言是同源的，即内源的。
该上下文中合适的启动子既包括组成型又包括诱导型的天然启动子以及经改造的启动子，所述启动子是本领域技术人员公知的。真核宿主细胞中合适的启动子可以是GAL7、GAL10或GAL 1、CYC1、HIS3、ADH1、PGL、PH05、GAPDH、ADC1、TRP1、URA3、LEU2、ENO、TPI和AOX1。其它合适的启动子包括PDC、GPD1、PGK1、TEF1和TDH。
通常，编码酶的核苷酸序列包含“终止子”。本发明中可以使用在真核细胞中有功能的任何终止子。优选的终止子得自宿主细胞的天然基因。合适的终止子序列是本领域公知的。优选地，这类终止子与下述突变组合，所述突变防止本发明宿主细胞中无义介导的mRNA衰退(decay)(见例如Shirley等,2002,Genetics 161:1465-1482)。
核酸构建体可被并入“载体”(例如表达载体)且/或并入宿主细胞中，以实现待表达多肽的表达。
表达载体可以是可方便地进行重组DNA程序并且可引起编码具有苹果酸脱氢酶活性的多肽的多核苷酸表达的任何载体(例如质粒或病毒)。载体的选择通常取决于载体与待引入所述载体的宿主细胞的相容性。载体可以是线性质粒或闭环质粒。载体可以是自主复制型载体，即作为染色体外实体而存在、其复制独立于染色体复制的载体，例如，质粒、染色体外元件、微型染色体或人工染色体。如果打算用于真菌来源的宿主细胞中，则合适的附加型核酸构建体可以例如基于酵母2μ或pKD1质粒(Gleer等,1991,Biotechnology 9:968-975)或AMA质粒(Fierro等,1995,Curr Genet.29:482-489)。
或者，表达载体可以是当被引入宿主细胞时整合至基因组中并与其已整合至其中的染色体一起复制的载体。整合型克隆载体可随机地或在预定靶基因座处整合在宿主细胞的染色体中。在本发明的一个优选实施方式中，整合型克隆载体包含与宿主细胞基因组中预定靶基因座中的DNA序列同源的DNA片段，以将克隆载体靶向整合到该预定基因座。为了促进靶向整合，克隆载体优选在转化细胞之前线性化。优选进行线性化，使得克隆载体的至少一端但优选两端的侧翼是与靶基因座同源的序列。靶基因座侧翼的同源序列的长度优选为至少20bp、至少30bp、至少50bp、至少0.1kb、至少0.2kb、至少0.5kb、至少1kb、至少2kb或更长。宿主细胞增强的同源重组能力提高靶向整合到宿主细胞基因组中(即整合到预定的靶基因座中)的效率。
与靶基因座同源的克隆载体中的同源侧翼DNA序列源自高表达的基因座，这意味着它们源自在宿主细胞中能够具有高表达水平的基因。能够具有高表达水平的基因(即高表达基因)在本文中被定义为：在例如经诱导条件下，其mRNA可以构成总细胞mRNA的至少0.5％(w/w)的基因，或者，其基因产物可以构成总细胞蛋白质的至少1％(w/w)的基因，或者在分泌的基因产物的情况下，可以分泌至至少0.1g/l的水平。
核酸构建体或表达载体可以在本发明的宿主细胞体内装配，并且任选地，在单个步骤中整合到细胞的基因组中(参见例如WO2013/076280)。
可将多于一个拷贝的本发明的核酸构建体或表达载体插入宿主细胞中以增加由包含在核酸构建体内的核酸序列编码的具有苹果酸脱氢酶活性的多肽的产生(过表达)。这可以优选地通过将两个或更多个拷贝的核酸整合到其基因组中来完成，更优选地通过使核酸靶向整合到上文所限定的高表达基因座来完成。
本领域技术人员会理解：表达载体的设计可取决于诸如待转化的宿主细胞的选择、所期望蛋白质的表达水平等因素。
可以通过常规转化或转染技术将本发明的核酸构建体和/或表达载体引入原核或真核细胞中。在本文中使用时，术语“转化”和“转染”旨在指用于将外来核酸(例如DNA)引入本领域技术人员公知的宿主细胞中的多种本领域公认的技术。用于转化或转染宿主细胞的合适方法可见于Sambrook等(Molecular Cloning:A Laboratory Manual,第3版,ColdSpring Harbor Laboratory Press,Cold Spring Harbor Laboratory,NY,2001)，Davis等(Basic Methods in Molecular Biology,第1版,Elsevier,1986)和其他实验室手册。
上述酶的胞质表达可通过缺失过氧化物酶体或线粒体靶向信号来实现。过氧化物酶体或线粒体靶向信号的存在可例如通过Schlüter等人(Schlüter等人,2007,NucleicAcid Research 35:D815-D822)所公开的方法来确定。
两个序列之间的序列比较和序列同一性百分比的确定可使用数学算法来完成。本领域的技术人员将意识到以下事实：若干不同的计算机程序可用于比对两个序列并确定两个序列之间的同一性(Kruskal,J.B.(1983)An overview of sequence comparison，D.Sankoff和J.B.Kruskal,(编),Time warps,string edits and macromolecules:thetheory and practice of sequence comparison,第1-44页Addison Wesley)。两个氨基酸序列之间或者两个核苷酸序列之间的序列同一性百分比可使用用于比对两个序列的Needleman和Wunsch算法来确定(Needleman,S.B.和Wunsch,C.D.(1970)J.Mol.Biol.48,443-453)。氨基酸序列和核苷酸序列两者均可通过算法进行比对。Needleman-Wunsch算法已在计算机程序NEEDLE中实现。出于本发明的目的，使用了来自EMBOSS包的NEEDLE程序(2.8.0版或更高版本，EMBOSS:The European Molecular Biology Open Software Suite(2000)Rice,P.Longden,I.和Bleasby,A.Trends in Genetics 16,(6)第276—277页,http://emboss.bioinformatics.nl/)。对于蛋白质序列而言，EBLOSUM62用于替换矩阵。对于核苷酸序列而言，使用EDNAFULL。所使用的任选参数是空位开放罚分为10，以及空位延伸罚分为0.5。技术人员将理解的是，所有这些不同的参数将产生稍微不同的结果，但是当使用不同的算法时，两个序列的总体同一性百分比没有显著改变。
在通过如上所述的程序NEEDLE进行比对后，查询序列与本发明的序列之间的序列同一性的百分比计算如下：在两个序列中显示相同氨基酸或相同核苷酸的比对中的相应位置的数目除以在减去比对中的总空位数后比对的总长度。如本文定义的同一性可通过使用NOBRIEF选项从NEEDLE获得，并且在程序的输出中标记为“最长同一性”。
本发明的核酸和蛋白质序列可进一步用作“查询序列”以针对公共数据库进行搜索，以例如鉴定其它家族成员或相关序列。此类搜索可使用Altschul等人(1990)J.Mol.Biol.215:403—10的blastn和blastx程序(2.2.31版或更高版本)进行。BLAST核苷酸搜索可用blastn程序(得分＝100、字长＝11)来进行，以获得与本发明的核酸分子同源的核苷酸序列。BLAST蛋白质搜索可用blastx程序(得分＝50、字长＝3)来进行，以获得与本发明的蛋白分子同源的氨基酸序列。为了获得用于比较目的的空位比对，可如在Altschul等人,(1997)Nucleic Acids Res.25(17):3389-3402中描述利用空位BLAST。当利用BLAST和空位BLAST程序时，可使用相应程序(例如blastx和blastn)的默认参数。参见http://www.ncbi.nlm.nih.gov/的国家生物技术信息中心的主页。
本发明还提供了生产二羧酸(例如琥珀酸)的方法，所述方法包括：在适于生产二羧酸的条件下发酵如上文所述的本发明的重组宿主细胞，和任选地从发酵培养基中回收二羧酸。
在所述方法中，在包含合适发酵培养基的容器中发酵本发明的重组宿主细胞。本文所使用的术语发酵指的是化合物的微生物生产，在此处是由碳水化合物生产二羧酸。
优选地，发酵产物是二羧酸，优选地是苹果酸、延胡索酸和/或琥珀酸，优选地是琥珀酸。
分批发酵在本文中被定义为这样的发酵，其中所有营养物在发酵开始时加入。
补料分批发酵是这样的分批发酵，其中营养物在发酵期间加入。分批和补料分批发酵中的产物可在合适的时机收获，例如在一种或更多种营养物耗尽时收获。
连续发酵是这样的发酵，其中营养物被连续加入到发酵中且其中产物被连续地移出发酵。
在一种实施方式中，在本发明的方法中发酵宿主细胞是在碳水化合物限制条件下进行的。当在本文中使用时，碳水化合物限制条件被定义为维持碳水化合物浓度低于10g/l，例如约5g/l。
根据本发明的生产二羧酸的方法可以以任何合适的体积和规模进行，优选地以工业规模进行。工业规模在本文中被定义为至少10升或100升，优选地至少1m3，优选地至少10m3或100m3，优选地至少1000m3，通常低于10,000m3的体积。
在本发明的方法中发酵重组宿主细胞可在任何合适的发酵培养基中进行，所述发酵培养基包含合适的氮源、碳水化合物和本发明所述方法中生长和生产二羧酸所需的其它营养物。根据本发明的发酵方法中合适的碳水化合物可以是葡萄糖、半乳糖、木糖、阿拉伯糖、蔗糖或麦芽糖。
在一种实施方式中，所述发酵方法在CO2分压为5％-60％、优选地约50％下进行。
在所述生产二羧酸的方法中，pH通常在二羧酸生产期间降低。优选地，在所述生产二羧酸的方法中，pH在1和5之间、优选地1.5和4.5之间、更优选地2和4之间的范围中。
在另一优选实施方式中，根据本发明的方法包括以下步骤：在碳水化合物的存在下在需氧条件下预培养宿主细胞。优选地，在预培养期间宿主细胞的发酵在4-6之间的pH下进行。优选地，预培养期间的碳水化合物是非抑制性碳水化合物、优选半乳糖。已发现在非抑制性碳水化合物上预培养宿主细胞是有利的，因为这防止了葡萄糖抑制的发生，而葡萄糖抑制可负面影响所产生生物质的量。此外，还发现：在需氧条件下预培养宿主细胞的步骤导致更高的生物质产率和更快的生长。优选地，所述预培养以分批模式进行。
通常进行用于生产增加的生物质的增殖步骤，优选地在碳水化合物限制条件下进行。
生产二羧酸的方法可在任何合适的温度下进行。合适的温度可例如在约10℃和约40℃之间，例如在约15℃和约30℃之间。
在一种实施方式中，以至少部分宿主细胞被重复利用(即，再循环)的方式进行本发明的方法。细胞可被再循环回原始容器或者再循环入第二容器。优选地，用维生素和/或微量元素补充引入再循环宿主细胞的培养基。
在一种优选的实施方式中，发酵培养基包含1-150g/l的琥珀酸，优选5-100g/l、更优选10-80g/l或15-60g/l的琥珀酸。在任何情况下，与用参照MDH多肽(例如SEQ ID NO:39的参照MDH多肽)修饰的宿主细胞相比，本发明的重组宿主细胞通常能够在发酵培养基中积累更多琥珀酸。
生产二羧酸的方法可以进一步包括回收二羧酸。可通过本领域已知的任何合适方法来回收二羧酸，例如通过结晶、铵沉淀、离子交换技术、离心或过滤或这些方法的任意合适组合。
在一种优选的实施方式中，回收二羧酸包括：使二羧酸结晶和形成二羧酸晶体。优选地，使二羧酸结晶包括除去部分发酵培养基，优选地通过蒸发，以获得浓缩的培养基。
根据本发明，可以通过从pH为1-5且包含二羧酸(例如琥珀酸)的水性溶液结晶所述二羧酸(例如琥珀酸)来回收二羧酸(例如琥珀酸)，包括蒸发部分水性溶液以获得浓缩溶液，将浓缩的溶液的温度降低至5-35℃之间的值，其中琥珀酸晶体形成。优选地，结晶包括：使浓缩的培养基的温度达到10-30℃，优选地15-25℃的温度。优选地，发酵培养基的pH为1.5-4.5，优选地2-4。
已发现，相较于在低于10度的低温下结晶的二羧酸(例如琥珀酸)的晶体，使二羧酸(例如琥珀酸)在较高温度(例如，10-30℃之间)下结晶产生杂质(例如，有机酸、蛋白质、颜色和/或气味)量更低的二羧酸(例如琥珀酸)的晶体。
相较于在低于10℃或5℃的温度下进行的二羧酸结晶，在更高温度下结晶琥珀酸的另一个优点是：冷却水性溶液需要更少量的能量，从而导致更加经济和可持续的方法。
优选地，结晶二羧酸(例如琥珀酸)包括洗涤二羧酸晶体的步骤。可由pH为1-5的发酵培养基直接将二羧酸(例如琥珀酸)结晶至纯度为至少90重量/重量％，优选地至少95重量/重量％、96重量/重量％、97重量/重量％、或至少98重量/重量％、或99-100重量/重量％。
在一种优选的实施方式中，生产二羧酸的方法还包括：在工业方法中使用二羧酸。
优选地，在根据本发明的方法中制备的二羧酸(例如琥珀酸)被进一步转变为期望的产品。期望的产品可以是例如聚合物，例如聚丁二酸丁二醇酯(PBS)，除冰剂(deicingagent)，食品添加剂，化妆品添加剂或表面活性剂。也就是说，本发明提供了生产产品(例如聚合物，例如聚丁二酸丁二醇酯(PBS)，除冰剂，食品添加剂，化妆品添加剂或表面活性剂)的方法，所述方法包括：如本文所述生产二羧酸；以及在所述产品的生产中使用所述二羧酸。
本文中作为现有技术所给出的专利文件或其它材料的引用并不应被认为是承认所述文件或材料在任何一项权利要求的优先权日时是已知的或者其所包含的信息是公知常识的一部分。
本文中陈述的每个引用的公开内容通过引用以整体并入本文中。
通过以下实施例进一步阐释本发明：
实施例
通用的材料和方法
DNA程序
除非另有说明，否则如别处(Sambrook等人,1989,Molecular cloning:alaboratory manual,第2版,Cold Spring Harbor Laboratory Press,Cold SpringHarbor,New York)所述进行标准DNA程序。使用校正酶Phusion聚合酶(New EnglandBiolabs,USA)根据制造商的说明来扩增DNA。限制酶来自Invitrogen或New EnglandBiolabs。
二羧酸生产菌株的微量滴定板(MTP)发酵
为了确定二羧酸生产，使菌株一式三份地在湿度摇床(Infors)中的微量滴定板中在30度、550rpm和80％湿度下生长3天。培养基基于Verduyn培养基(Verduyn C,Postma E,Scheffers WA,Van Dijken JP.Yeast,1992Jul；8(7):501-517)，但如下文所述对碳源和氮源进行更改。
表.MTP预培养基组成
| 原材料 | 浓度(g/L) | |
| 半乳糖 | C6H12O6.H2O | 40.0 |
| 尿素 | (NH2)2CO | 2.3 |
| 磷酸二氢钾 | KH2PO4 | 3.0 |
| 硫酸镁 | MgSO4.7H2O | 0.5 |
| 微量元素溶液a | 1 | |
| 维生素溶液b | 1 |
a微量元素溶液
| 组分 | 式 | 浓度(g/kg) |
| EDTA | C10H14N2Na2O8.2H2O | 15.00 |
| 硫酸锌.7H2O | ZnSO4.7H2O | 4.50 |
| 氯化锰.2H2O | MnCl2.2H2O | 0.84 |
| 氯化钴(II).6H2O | CoCl2.6H2O | 0.30 |
| 硫酸铜(II).5H2O | CuSO4.5H2O | 0.30 |
| 钼酸钠.2H2O | Na2MoO4.2H2O | 0.40 |
| 氯化钙.2H2O | CaCl2.2H2O | 4.50 |
| 硫酸铁.7H2O | FeSO4.7H2O | 3.00 |
| 硼酸 | H3BO3 | 1.00 |
| 碘化钾 | KI | 0.10 |
b维生素溶液
| 组分 | 式 | 浓度(g/kg) |
| 生物素(D-) | C10H16N2O3S | 0.05 |
| Ca D(+)泛酸 | C18H32CaN2O10 | 1.00 |
| 烟酸 | C6H5NO2 | 1.00 |
| 肌醇 | C6H12O6 | 25.00 |
| 盐酸氯化硫胺 | C12H18Cl2N4OS.xH2O | 1.00 |
| 盐酸吡哆醇 | C8H12ClNO3 | 1.00 |
| 对氨基苯甲酸 | C7H7NO2 | 0.20 |
使用80μl预培养物接种24孔板中含有1.5％半乳糖作为碳源的2.5ml培养基。使培养物在30℃、550rpm、80％湿度的湿度摇床(Infors)中生长72小时。生成生物质后，通过将细胞重悬于2.5ml含有葡萄糖作为碳源的矿质培养基中来开始生产实验。在30℃、550rpm、80％湿度的湿度摇床(Infors)中孵育主要培养物，并在培养48小时后取样。
通过NMR的MTP样品的代谢物分析
对于MTP样品的代谢物分析，将90μl发酵样品上清液与10μl NMR标准品(20g/l马来酸)和100μl 10％D2O溶液混合。将样品冻干，随后溶在1mL D2O中。
使用不具有水抑制的脉冲程序(ZG)，在300K的温度下，使用90度激发脉冲、2.0s的采集时间和40s的放松延迟，在配备有He冷却低温探头、以500MHz的质子频率运行的BESTBruker Avance III光谱仪上记录1D 1H NMR谱。扫描次数设置为8。
基于以下信号(δ相对于4,4-二甲基-4-硅戊烷-1-磺酸)计算苹果酸浓度[以g/l计]：
苹果酸：根据pH值以及α-CH2和CH(OH)信号与其他化合物的重叠，选择以下三种苹果酸信号之一进行定量：α-CH(A)(2.92ppm,n＝1H,双峰(double doublet)或dd),α-CH(X)(2.85ppm,n＝1H,dd)或者CH(OH)(4.6ppm,n＝1H,dd)。
基于以下信号(δ相对于4,4-二甲基-4-硅戊烷-1-磺酸)计算琥珀酸浓度[以g/L计]：
琥珀酸：2.67ppm处的琥珀酸信号(s，4H)。
用于标准品的信号：约6.3ppm处的马来酸峰(S，2H)。
Bharti等人,2012,TrAC Trends in Analytical Chemistry 35:5-26描述了通过NMR定量。
实施例1：构建酵母菌株SUC-1029
使用菌株CEN.PK 113-7D(MATa HIS3 LEU2 TRP1 MAL2-8 SUC2)作为构建菌株SUC-1029的起点。如下所述将Rhyzopus oryzae的延胡索酸酶基因(FUMR)转化到菌株CEN.PK113-7D中。
生成PCR片段
使用扩增SEQ ID NO:34的完整核苷酸序列的引物，通过PCR扩增SEQ ID NO:34获得PCR片段9。SEQ ID NO:34描述了如专利申请WO2009/065779中所公开的含有来自Rhyzopus oryzae的延胡索酸酶(FUMR)核苷酸序列的合成多核苷酸。如专利申请WO2008/000632中所公开，针对在S.cerevisiae中表达对基因序列进行密码子对优化。FUMR基因的表达由TDH1启动子(紧接在TDH1基因的起始密码子之前的600bp)和TDH1终止子(紧接在TDH1基因的终止密码子之后的300bp)控制。控制FUMR表达的TDH1启动子和TDH1终止子序列是来自Saccharomyces cerevisiae S288C的天然序列。在TDH1启动子上游的SEQ ID NO:34的5’端的599bp区域是与YPRCtau3基因座同源的区域。
使用扩增SEQ ID NO:35的完整核苷酸序列的引物，通过PCR扩增SEQ ID NO:35获得PCR片段10。SEQ ID NO:35描述了含有部分pSUC227质粒序列的合成多核苷酸，其描述于PCT/EP2013/055047中。SEQ ID NO:35的5’端与SEQ ID NO:34的3’端具有重叠。SEQ ID NO:35的3’端与SEQ ID NO:36的5’端具有重叠。
使用扩增SEQ ID NO:36的完整核苷酸序列的引物，通过PCR扩增SEQ ID NO:36获得PCR片段11。SEQ ID NO:36描述了含有部分pSUC225质粒序列的合成多核苷酸，其描述于PCT/EP2013/055047中。SEQ ID NO:36的3’端与SEQ ID NO:37的5’端具有重叠。
使用扩增SEQ ID NO:37的完整核苷酸序列的引物，通过PCR扩增SEQ ID NO:37获得PCR片段12。SEQ ID NO:37描述了与YPRCtau3基因座同源的合成多核苷酸。
使用DNA Clean&ConcentratorTM-25试剂盒(Zymo Research,Irvine,CA,USA)根据制造商的说明纯化PCR片段9至12。
转化到CEN.PK113-7D中以构建菌株SUC-1029
通过本领域技术人员已知的方法进行酵母转化。用纯化的PCR片段9至12转化S.cerevisiae菌株CEN.PK113-7D。PCR片段10和11在其5’端和3’端具有重叠，PCR片段9和12在其3’端和5’端具有重叠，从而使得允许所有4个PCR片段的同源重组(图1)。PCR片段9的5’端和PCR片段12的3’端与YPRCtau3基因座同源，使得所有4个PCR片段能够整合在YPRCtau3基因座(图1)。这产生了一个由整合在YPRCtau3基因座的PCR片段9至12组成的线性片段，其位于染色体XVI上。
将转化混合物涂布在含有200μg G418(Sigma Aldrich,Zwijndrecht,荷兰)/ml的YEPhD-琼脂(每升：10g酵母提取物，20g PhytonePeptone，20g葡萄糖，20g琼脂)上。在30℃下生长三天后，将各个转化体重新划线在含有20g葡萄糖/升和200μg G418/ml的YEPh-琼脂板上。
随后，使用PCT/EP2013/055047中的所述方法，藉由CRE重组酶通过在KanMX标记和编码CRE重组酶的CRE基因侧翼的lox66和lox71位点之间的重组移除整合的PCR片段10和11上存在的标记盒和Cre-重组酶基因，从而导致KanMX标记和CRE基因被移除，并留下lox72位点作为lox66和lox71位点之间重组的结果。所产生的无标记菌株被命名为SUC-1029。
使用PCR来证实所引入的FUMR基因的存在，其中使用可以与SEQ ID NO:34编码的ORF的编码序列退火的引物序列。通过PCR证实KanMX标记的正确整合和移除，其中使用来自YPRCtau3基因座的5’和3’引物，所述引物不与PCR片段9和12上存在的YPRCtau3同源区域杂交。
实施例2：构建菌株SUC-1112
生成PCR片段
采用Saccharomyces cerevisiae菌株CEN.PK 113-7D(MATa HIS3 LEU2 TRP1MAL2至8SUC2)的基因组DNA作为模板，使用SEQ ID NO:9和SEQ ID NO:10中所述的引物序列生成由5’INT59整合位点组成的PCR片段1。
采用SEQ ID NO:1作为模板，使用SEQ ID NO:11和SEQ ID NO:12中所述的引物序列生成PCR片段2。SEQ ID NO:1编码来自Actinobacillus succinogenes的磷酸烯醇式丙酮酸羧激酶(PCKa)，如专利申请WO2009/065780中所公开。该合成序列包括启动子-基因-终止子序列，包括合适的限制性位点，由DNA 2.0(Menlo Park,California,USA)合成。如专利申请WO2008/000632中所公开，针对在S.cerevisiae中表达对基因序列进行密码子对优化。合成基因受来自S.cerevisiae的启动子控制(或与来自S.cerevisiae的启动子可操作地连接)，即TPI1启动子控制PCKα基因的表达。恰当的终止由来自S.cerevisiae的终止子序列(即GND2终止子)控制。
采用SEQ ID NO:2作为模板，使用SEQ ID NO:13和SEQ ID NO:14中所述的引物序列生成PCR片段3。SEQ ID NO:2编码来自Saccharomyces cerevisiae的丙酮酸羧化酶(PYC2)，如专利申请WO2009/065780中所公开。该合成序列包括启动子-基因-终止子序列，包括合适的限制性位点，由DNA 2.0(Menlo Park,California,USA)合成。如专利申请WO2008/000632中所公开，针对在S.cerevisiae中表达对基因序列进行密码子对优化。合成基因受来自S.cerevisiae的启动子控制(或与来自S.cerevisiae的启动子可操作地连接)，即PGK1启动子控制PYC2基因的表达。恰当的终止由来自S.cerevisiae的终止子序列(即ADH1终止子)控制。
采用SEQ ID NO:3作为模板，使用SEQ ID NO:15和SEQ ID NO:16中所述的引物序列生成PCR片段4。SEQ ID NO:3编码在Saccharomyces cerevisiae中有功能的KanMX选择标记，其是从质粒pUG7-EcoRV扩增的。pUG7-EcoRV是Gueldener等人(Nucleic AcidsRes.1996年7月；24(13):2519-24)所述的质粒pUG6的变体，其中pUG6中存在的loxP位点变为lox66和lox71位点(Lambert等人,Appl.Environ.Microbiol.2007年2月；73(4):1126-35.Epub 2006年12月1日)。
采用SEQ ID NO:4作为模板，使用SEQ ID NO:17和SEQ ID NO:18中所述的引物序列生成PCR片段5。SEQ ID NO:4编码来自Aspergillus niger的推定的二羧酸转运体，如EP2495304中所公开。该合成序列包括启动子-基因-终止子序列，包括合适的限制性位点，由DNA 2.0(Menlo Park,California,USA)合成。如专利申请WO2008/000632中所公开，针对在S.cerevisiae中表达对基因序列进行密码子对优化。合成基因受来自S.cerevisiae的启动子控制(或与来自S.cerevisiae的启动子可操作地连接)，即ENO1启动子控制DCT_02-基因的表达。恰当的终止由来自S.cerevisiae的终止子序列(即TEF2终止子)控制。
采用SEQ ID NO:5作为模板，使用SEQ ID NO:19和SEQ ID NO:20中所述的引物序列生成PCR片段6。SEQ ID NO:5编码来自Saccharomyces cerevisiae的苹果酸脱氢酶(MDH3)，如专利申请WO2009/065778中所公开。该合成序列包括启动子-基因-终止子序列，包括合适的限制性位点，由DNA 2.0(Menlo Park,California,USA)合成。如专利申请WO2008/000632中所公开，针对在S.cerevisiae中表达对基因序列进行密码子对优化。合成基因受来自Kluyveromyces lactis的启动子控制(或与来自Kluyveromyces lactis的启动子可操作地连接)，即ORF KLLA0_F20031g(uniprot检索号为Q6CJA9)的启动子控制MDH3基因的表达。恰当的终止由来自S.cerevisiae的终止子序列(即GPM1终止子)控制。
采用SEQ ID NO:6作为模板，使用SEQ ID NO:21和SEQ ID NO:22中所述的引物序列生成PCR片段7。SEQ ID NO:6编码来自Escherichia coli的延胡索酸酶(fumB)(E.C.4.2.1.2，UniProt检索号为P14407)。如专利申请WO2008/000632中所公开，针对在S.cerevisiae中表达对基因序列进行密码子对优化。合成序列包括启动子-基因-终止子序列，包括合适的限制性位点，由DNA 2.0(Menlo Park,California,USA)合成。合成基因受来自S.cerevisiae的启动子控制(或与来自S.cerevisiae的启动子可操作地连接)，即TDH3启动子控制fumB基因的表达。恰当的终止由来自S.cerevisiae的终止子序列(即TDH1终止子)控制。
采用SEQ ID NO:7作为模板，使用SEQ ID NO:23和SEQ ID NO:24中所述的引物序列生成PCR片段8。如专利申请WO2009/065778中所公开，SEQ ID NO:7编码来自Trypanosomabrucei的延胡索酸还原酶(FRDg)。如专利申请WO2008/000632中所公开，针对在S.cerevisiae中表达对基因序列进行密码子对优化。合成序列包括启动子-基因-终止子序列，包括合适的限制性位点，由DNA 2.0(Menlo Park,California,USA)合成。合成基因受来自S.cerevisiae的启动子控制(或与来自S.cerevisiae的启动子可操作地连接)，即TEF1启动子控制fumB基因的表达。恰当的终止由来自S.cerevisiae的终止子序列(即TAL1终止子)控制。
采用菌株CEN.PK 113-7D的基因组DNA作为模板，使用SEQ ID NO:40和SEQ ID NO:41中所述的引物序列生成由3’INT59整合位点组成的PCR片段113。
使用DNA Clean&ConcentratorTM-25试剂盒(Zymo Research,Irvine,CA,USA)根据制造商的说明纯化PCR片段1至8和PCR片段113。
转化到SUC-1029中以构建菌株SUC-1112
通过本领域技术人员已知的方法进行酵母转化。用纯化的PCR片段1至8和PCR片段113转化S.cerevisiae菌株SUC-1029。PCR片段2至8在其5’端和3’端具有重叠，PCR片段1和113在其3’端和5’端具有重叠，从而使得允许所有8个PCR片段的同源重组。PCR片段1的5’端和PCR片段113的3’端与INT59基因座同源，使得所有9个PCR片段能够整合在INT59基因座(参见图2)。这产生了一个由整合在INT59基因座的PCR片段2至8组成的线性片段。专利申请WO2013076280中描述了这种整合方法。INT59基因座位于染色体XI上，在YKR092C下游923bp且YKR093W上游922bp。
将转化混合物涂布在含有20g半乳糖/升和200μg G418(Sigma Aldrich,Zwijndrecht,荷兰)/ml的YEPh-琼脂(每升：10g酵母提取物，20g PhytonePeptone，20g琼脂)上。在30℃下生长三天后，将各个转化体重新划线在含有20g半乳糖/升和200μg G418/ml的YEPh-琼脂板上。使用PCR来证实所有引入基因的存在，其中使用可以与SEQ ID NO:1至SEQ ID NO:7编码的ORF的编码序列退火的引物序列。所得菌株被命名为SUC-1112。如果需要，可以移除菌株SUC-1112中存在的KanMX标记。
实施例3：将苹果酸脱氢酶基因转化到菌株SUC-1112中并在所得转化体中生产苹
果酸
生成PCR片段
采用菌株CEN.PK 113-7D的基因组DNA作为模板，使用SEQ ID NO:25和SEQ ID NO:26中所述的引物序列生成由5’INT1整合位点组成的PCR片段13。
通过采用SEQ ID NO:42作为模板，使用SEQ ID NO:43和SEQ ID NO:44中所述的引物序列生成PCR片段114。SEQ ID NO:42含有编码葡萄糖-6-磷酸脱氢酶(G6PD)的ZWF1基因。该合成序列包括启动子-基因-终止子序列，包括合适的限制性位点，由DNA 2.0(MenloPark,California,USA)合成。如专利申请WO2008/000632中所公开，针对在S.cerevisiae中表达对基因序列进行密码子对优化。合成基因受来自Kluyveromyces lactis的启动子控制(或与来自Kluyveromyces lactis的启动子可操作地连接)，即ORF KLLA0C05566g(uniprot检索号为Q6CUE2)的启动子控制ZWF1基因的表达。恰当的终止由来自S.cerevisiae的终止子序列(即TEF1终止子)控制。
通过采用SEQ ID NO:38作为模板，使用SEQ ID NO:45和SEQ ID NO:28中所述的引物序列生成PCR片段115。SEQ ID NO:38编码在Saccharomyces cerevisiae中有功能的诺尔丝菌素选择标记，其是从质粒pUG7-Nat的修改形式扩增的。pUG7-Nat是Gueldener等人(Nucleic Acids Res.1996年7月；24(13):2519-24)所述的质粒pUG6的变体，其中pUG6中存在的loxP位点变为lox66和lox71位点(Lambert等人,Appl.Environ.Microbiol.2007年2月；73(4):1126-35.Epub 2006年12月1日)，并且其中KanMX标记被诺尔丝菌素标记替代(Goldstein和McCusker,Yeast.1999年10月；15(14):1541-53)。
通过采用SEQ ID NO:31作为模板，使用SEQ ID NO:29和SEQ ID NO:30中所述的引物序列生成PCR片段15。SEQ ID NO:31编码来自S.cerevisiae的苹果酸脱氢酶(MDH3)。如专利申请WO2013/004670 A1中所公开，通过缺失SKL羧基末端序列来改变MDH3。该合成序列包括启动子-基因-终止子序列，包括合适的限制性位点，由DNA 2.0(Menlo Park,California,USA)合成。如专利申请WO2008/000632中所公开，针对在S.cerevisiae中表达对基因序列进行密码子对优化。合成基因受来自S.cerevisiae的启动子控制(或与来自S.cerevisiae的启动子可操作地连接)，即TDH3启动子控制MDH3基因的表达。恰当的终止由来自S.cerevisiae的终止子序列(即GPM1终止子)控制。
采用菌株CEN.PK 113-7D的基因组DNA作为模板，使用SEQ ID NO:32和SEQ ID NO:33中所述的引物序列生成由3’INT1整合位点组成的PCR片段16。
使用DNA Clean&ConcentratorTM-25试剂盒(Zymo Research,Irvine,CA,USA)根据制造商的说明纯化PCR片段13-16。
转化到SUC-1112中
通过本领域技术人员已知的方法进行酵母转化。用纯化的PCR片段13、114、115、15和16转化S.cerevisiae菌株SUC-1112。PCR片段114和115和15在其5’端和3’端具有重叠，PCR片段13和16在其3’端和5’端具有重叠，从而使得允许所有5个PCR片段的同源重组。PCR片段13的5’端和PCR片段16的3’端与INT1基因座同源，使得所有4个PCR片段能够整合在INT1基因座(参见图3)。这产生了一个由整合在INT1基因座的PCR片段13和16组成的线性片段。专利申请WO2013/076280中描述了这种整合方法。INT1基因座位于染色体XV上，在YOR071c下游659bp且YOR070c上游998bp。该方法导致如SEQ ID NO:39所示的340个氨基酸的苹果酸脱氢酶蛋白的表达，SEQ ID NO:39与来自S.cerevisiae的天然序列相比缺少C-末端氨基酸SKL。
将转化混合物涂布在含有20g半乳糖/升和200μg诺尔丝菌素(Jena Bioscience，德国)/ml的YEPh-琼脂(每升：10g酵母提取物，20g PhytonePeptone，20g琼脂)上。在30℃下生长三天后，将各个转化体重新划线在含有20g半乳糖/升和200μg诺尔丝菌素/ml的YEPh-琼脂板上。使用PCR来证实所引入基因的存在，其中使用可以与PCR片段114、115和15上存在的编码ORF的编码序列退火的引物序列。为了证实PCR片段13、114、115、15和16在正确基因座的整合，使用与INT1基因座的5’和3’区域退火并且不与PCR片段13和16上的INT1区域结合的引物和与PCR片段114和15上的ORF退火的引物的组合，使得只有PCR片段114和15整合在INT1基因座时，才能形成PCR产物。得到了三种单菌落：SUC-1112+MDH3#1、SUC-1112+MDH3#2、SUC-1112+MDH3#3。如果需要，可以移除菌株SUC-1112+MDH3#1、SUC-1112+MDH3#2、SUC-1112+MDH3#中存在的KanMX和诺尔丝菌素标记。
二羧酸生产
为了确定二羧酸生产，如“通用的材料和方法”中所述进行MTP发酵和NMR测量。
在含有存在于PCR片段15上的额外MDH3基因拷贝的SUC-1112+MDH 3菌株(SUC-1112+MDH3#1、SUC-1112+MDH3#2、SUC-1112+MDH3#3)的上清液中，测得苹果酸的平均滴度为8.7g/L。琥珀酸水平低于预期；这些菌株看起来已经失去了FRDg基因，从而导致苹果酸盐/酯朝向琥珀酸盐/酯的转变有限。
实施例4：将编码苹果酸脱氢酶突变体的基因转化到菌株SUC-1112中并在所得转
化体中生产苹果酸
生成PCR片段
如实施例3中所述生成PCR片段13和16。
采用SEQ ID NO:38作为模板，使用SEQ ID NO:27和SEQ ID NO:28中所述引物序列生成PCR片段14。SEQ ID NO:38编码在Saccharomyces cerevisiae中有功能的诺尔丝菌素选择标记，其是从质粒pUG7-Nat的修改形式扩增的。pUG7-Nat是Gueldener等人(NucleicAcids Res.1996年7月；24(13):2519-24)所述的质粒pUG6的变体，其中pUG6中存在的loxP位点变为lox66和lox71位点(Lambert等人,Appl.Environ.Microbiol.2007年2月；73(4):1126-35.Epub 2006年12月1日)，并且其中KanMX标记被诺尔丝菌素标记替代(Goldstein和McCusker,Yeast.1999年10月；15(14):1541-53)。
编码SEQ ID NO:39中所述的参照苹果酸脱氢酶序列的不同蛋白质突变体的合成核苷酸序列由DNA 2.0(Menlo Park,California,USA)合成。合成的核苷酸序列编码相对于参照MDH3序列(SEQ ID NO:39)在第34至40位突变的氨基酸序列，如表1中所示。除了编码表1中所示的突变氨基酸之外，合成的核苷酸序列突变体与SEQ ID NO:31相同。合成基因受来自S.cerevisiae的启动子控制(或与来自S.cerevisiae的启动子可操作地连接)，即TDH3启动子控制突变MDH基因的表达。恰当的终止由来自S.cerevisiae的终止子序列(即GPM1终止子)控制。
合成的基因序列包含TDH3启动子-突变的MDH-GPM1终止子及其它，其使用SEQ IDNO:29和SEQ ID NO:30中所述引物序列通过PCR扩增，以生成PCR片段17至108(参见表1)。
使用DNA Clean&ConcentratorTM-25试剂盒(Zymo Research,Irvine,CA,USA)根据制造商的说明纯化PCR片段13、14、16和17至108。
转化到SUC-1112中
用纯化的PCR片段13、14和16与单独的PCR片段17至108的组合转化菌株SUC-1112。PCR片段14和PCR片段17至108在其5’端和3’端具有重叠，PCR片段13和16在其3’端和5’端具有重叠，从而使得允许所有4个PCR片段的同源重组。PCR片段13的5’端和PCR片段16的3’端与INT1基因座同源，使得所有4个PCR片段能够整合在INT1基因座(图4)。转化体的转化和选择描述于实施例2中。
二羧酸生产
为了确定二羧酸生产，如“通用的材料和方法”中所述使四个独立的表达突变的苹果酸脱氢酶序列的SUC-1112转化体在微量滴定板中生长。由于FRDg的丢失，产生了少量的琥珀酸(参见实施例3)，但仍可通过测量苹果酸盐/酯水平来确定MDH3活性。平均苹果酸滴度如表1所示。表达突变的苹果酸脱氢酶序列的几种SUC-1112转化体的苹果酸的平均产量超过10g/L苹果酸。有趣的是，在第34位用甘氨酸或丝氨酸残基替换天冬氨酸的突变体显示出提高的苹果酸生产。这显著高于实施例3中所述的用参照MDH3序列转化的SUC-1112的平均苹果酸滴度。显著高于是指：具有参照苹果酸脱氢酶序列和经改进的突变型苹果酸脱氢酶序列的菌株的苹果酸滴度的95％置信区间不重叠。用参照MDH3序列转化的SUC-1112的苹果酸滴度的95％置信区间的上限低于10g/L。
表1：培养菌株SUC-1112转化体4天后在生产培养基的上清液中测量的平均苹果酸
滴度，所述SUC-1112转化体表达磷酸烯醇式丙酮酸羧激酶(PCKa)、丙酮酸羧化酶(PYC2)、苹
果酸脱氢酶(MDH3)、延胡索酸酶(FUMR和fumB)、二羧酸转运体(DCT_02)，并用编码参照苹果
酸脱氢酶(SEQ ID NO:39)或苹果酸脱氢酶突变体(MUT_001-MUT_94)的核苷酸序列转化，与
参照序列相比，所述苹果酸脱氢酶突变体在如下所示的氨基酸位置含有突变。
实施例5：测量苹果酸脱氢酶突变体的NADH特异性活性和NADPH特异性活性
从表1中选择共19种突变体，并如“通用的材料和方法”中所述进行再培养。通过离心(4000rpm，10分钟，4℃)收获生物质，并用PBS(磷酸盐缓冲盐水，Sigma Aldrich)洗涤两次，之后将细胞沉淀物在-20℃冷冻。使用0.5mm酸洗玻璃珠与来自Qiagen的TissueLyserII(3000rpm，2x10秒，循环之间在冰上冷却1分钟)的组合，在方孔96深孔微量滴定板(MTP)中实现细胞破碎。将体积为600μl的玻璃珠加入细胞沉淀物中，然后加入1ml体内样试验培养基(描述于van Eunen等人,FEBS Journal 277:749-760中，调整为含有0.5mM DTT(二硫苏糖醇，Sigma-Aldrich)和0.1mM PMSF(苯基甲磺酰氟，Amresco)。通过下述方法加入玻璃珠：在其中每个孔充满了玻璃珠(＝体积为300μl)的标准MTP上翻转含有冷冻沉淀物的深孔MTP，然后翻转两个板，使玻璃珠落到细胞沉淀物上。重复该过程，以在细胞沉淀物中得到600μl玻璃珠。然后加入1ml上述体内样试验培养基。细胞破碎后，通过离心(4000rpm，30分钟，4℃)使细胞碎片沉淀。收集上清液(可溶性细胞提取物)并储存在冰上。使用牛血清白蛋白(BSA)作为标准品，通过Bradford测定提取物的蛋白质浓度。
采用分光光度法通过340nm处吸光度的降低测定苹果酸脱氢酶(MDH)活性，340nm处吸光度的降低是由NADH或NADPH分别被氧化成NAD+或NADP+引起的。在体内样试验培养基中，试验含有400μM NADH或400μM NADPH、2mM草酰乙酸(Sigma Aldrich)和约0.0625mg蛋白质ml-1可溶性细胞提取物。以200μl的终体积进行试验。在NADH依赖型试验和NADPH依赖型试验中加入等体积的可溶性细胞提取物。通过加入100μl草酰乙酸储备溶液(4mM)开始反应，然后在30摄氏度下9分钟，斜率被用作NADH或NADPH依赖型MDH活性的度量。使用MicrosoftExcel中的“斜率”函数确定斜率(以ΔA340/min计)，其中吸光度值为“y”值，以分钟为单位的时间为“x”值。Microsoft Excel中的“RSQ”函数被用于检查斜率拟合的质量(标准>0.9)。对含有体内样试验培养基而非底物的空白反应的斜率校正斜率。使用Tecan InfiniteM1000读板仪测量吸光度。将每种突变体的NADH依赖型活性与NADPH活性进行比较。通过以下方法确定NADPH依赖型活性:NADH依赖型活性的比值或NADPH特异性:NADH特异性的比值：
1)确定NADPH依赖型MDH活性的斜率(以ΔA340/min计)
2)确定NADH依赖型MDH活性的斜率(以ΔA340/min计)
3)将NADPH依赖型MDH活性的斜率除以NADH依赖型MDH活性的斜率。
为了确定比值，使用斜率，而不针对总蛋白的量进行均一化，因为在NADH依赖型MDH活性试验和NADPH依赖型MDH活性试验中使用等体积的每种突变体。
计算19种突变体的比值(图5D)。与参照相比，该比值的值增加指示辅因子特异性已经改变。
如“通用的材料和方法”中所述，分析19种培养的突变体和参照菌株的上清液的苹果酸滴度。如上所述测量NADPH特异性活性和NADH特异性活性，并通过除以试验中的总蛋白浓度针对总蛋白进行均一化。结果显示在图5B-5C中。显然，19种所选择的突变体具有增强的NADPH特异性苹果酸脱氢酶活性。与参照相比，19种突变体中大多数的NADH特异性苹果酸脱氢酶活性降低(图5B)。有趣的是，在6种突变体中，NADH特异性活性提高(图5B)，从而指示：在这些突变体中，NADH特异性活性和NADPH特异性活性均提高。在所有突变体中，NADPH特异性:NADH特异性比值均增加(图5D)。
出乎意料地，在第34位用甘氨酸或丝氨酸残基替换天冬氨酸对用这些苹果酸脱氢酶突变体转化的SUC-1112菌株的苹果酸滴度具有正面影响(图5A)。
实施例6：构建菌株REV-0001
生成PCR片段
采用菌株CEN.PK 113-7D的基因组DNA作为模板，使用SEQ ID NO:54和SEQ ID NO:55中所述的引物序列生产由5’INT1整合位点组成的PCR片段116。
通过采用SEQ ID NO:38作为模板，使用SEQ ID NO:56和SEQ ID NO:57中所述的引物序列生成PCR片段117。SEQ ID NO:38编码在Saccharomyces cerevisiae中有功能的诺尔丝菌素选择标记，其是从质粒pUG7-Nat的修改形式扩增的。pUG7-Nat是Gueldener等人(Nucleic Acids Res.1996年7月；24(13):2519-24)所述的质粒pUG6的变体，其中pUG6中存在的loxP位点变为lox66和lox71位点(Lambert等人,Appl.Environ.Microbiol.2007年2月；73(4):1126-35.Epub 2006年12月1日)，并且其中KanMX标记被诺尔丝菌素标记替代(Goldstein和McCusker,Yeast.1999年10月；15(14):1541-53)。
通过采用SEQ ID NO:62作为模板，使用SEQ ID NO:60和SEQ ID NO:61中所述的引物序列生成PCR片段118。SEQ ID NO:62编码来自Trypanosoma brucei的延胡索酸还原酶(FRDg)，如专利申请WO2009/065778中所公开。该合成序列包括启动子-基因-终止子序列，包括适当的限制性位点，由DNA 2.0(Menlo Park,California,USA)合成。如专利申请WO2008/000632中所公开，针对在S.cerevisiae中表达对基因序列进行密码子对优化。合成基因受来自S.cerevisiae的启动子控制(或与来自S.cerevisiae的启动子可操作地连接)，即TDH3启动子控制FRDg基因的表达。恰当的终止由来自S.cerevisiae的终止子序列(即TAL1终止子)控制。采用菌株CEN.PK 113-7D的基因组DNA作为模板，使用SEQ ID NO:58和SEQ ID NO:59中所述的引物序列生成由3’INT1整合位点组成的PCR片段119。
使用DNA Clean&ConcentratorTM-25试剂盒(Zymo Research,Irvine,CA,USA)根据制造商的说明纯化PCR片段116至119。
转化SUC-1029
通过本领域技术人员已知的方法进行酵母转化。用纯化的PCR片段116至119转化S.cerevisiae菌株SUC-1029(实施例1)。PCR片段117和118在其5’端和3’端具有重叠，PCR片段116和119在其3’端和5’端具有重叠，从而使得允许所有4个PCR片段的同源重组(图1)。PCR片段116的5’端和PCR片段119的3’端与INT1基因座同源，使得所有4个PCR片段能够整合在INT1基因座(参见图6)。这产生了一个由整合在INT1基因座的PCR片段116至119组成的线性片段。专利申请WO2013076280中描述了这种整合方法。INT1基因座位于染色体XV上，在YOR071c下游659bp且YOR070c上游998bp。该方法导致如SEQ ID NO:8所示的1139个氨基酸的延胡索酸还原酶蛋白的表达，与来自T.brucei的天然序列相比，所述延胡索酸还原酶蛋白缺少C-末端氨基酸SKI。
将转化混合物涂布在含有100μg诺尔丝菌素(Jena Bioscience，德国)/ml的YEPh-琼脂(每升：10g酵母提取物，20g PhytonePeptone，20g半乳糖，20g琼脂)上。在30℃下生长三天后，将各个转化体重新划线在含有20g半乳糖/升和100μg诺尔丝菌素/ml的YEPh-琼脂板上。使用PCR来证实所引入基因的存在，其中使用可以与SEQ ID NO:38和SEQ ID NO:62编码的ORF的编码序列退火的引物序列。
实施例7：将编码苹果酸脱氢酶突变体的基因转化到菌株REV-0001中并在所得转
化体中生产琥珀酸
为了确定表达MDH突变体的菌株中琥珀酸盐/酯水平是否提高，将MDH突变体引入其中表达FRDg的菌株REV-0001中(实施例6)。基于苹果酸生产结果(实施例4)和体外活性试验结果(实施例5)，选择3种MDH突变体：MUT_014、MUT_015和MUT_034。
生成PCR片段
PCR片段1、2、3、4和5的扩增描述于实施例2中。为了引入野生型和多样化的MDH突变体，使用PCR片段120(野生型MDH3，SEQ ID NO:31)、片段121(MUT_014，SEQ ID NO:64)、片段122(MUT_015，SEQ ID NO:65)或片段123(MUT_034，SEQ ID NO:66)。野生型和突变型MDH3基因由S.cerevisiae TDH3终止子驱动，终止由S.cerevisiae GPM1终止子控制。使用SEQID NO:19和SEQ ID NO:20中所述的引物序列通过PCR扩增盒，以生成PCR片段120至123。
采用菌株CEN.PK 113-7D的基因组DNA作为模板，使用SEQ ID NO:63和SEQ ID NO:41中所述的引物序列生成由3’INT59整合位点组成的PCR片段124。
使用DNA Clean&ConcentratorTM-25试剂盒(Zymo Research,Irvine,CA,USA)根据制造商的说明纯化PCR片段1-5/120-123和PCR片段124。
转化到REV-0001中
通过本领域技术人员已知的方法进行酵母转化。用纯化的PCR片段1至5和124与PCR片段120、121、122或123的组合转化S.cerevisiae菌株REV-0001。PCR片2至5/120-123在其5’端和3’端具有重叠，PCR片段1和124在其3’端和5’端具有重叠，从而使得允许所有7个PCR片段的同源重组。PCR片段1的5’端和PCR片段124的3’端与INT59基因座同源，使得所有7个PCR片段能够整合在INT59基因座(参见图7)。这产生了一个由整合在INT59基因座的PCR片段2至5/120-123组成的线性片段。专利申请WO2013076280中描述了这种整合方法。INT59基因座位于染色体XI上，在YKR092C下游923bp且YKR093W上游922bp。
将转化混合物涂布在含有20g半乳糖/升和200μg G418(Sigma Aldrich,Zwijndrecht,荷兰)/ml的YEPh-琼脂(每升：10g酵母提取物，20g PhytonePeptone，20g琼脂)上。在30℃下生长三天后，将各个转化体重新划线在含有20g半乳糖/升和200μg G418/ml的YEPh-琼脂板上。通过PCR来证实所有引入基因的存在。
二羧酸生产
为了确定二羧酸生产，如“通用的材料和方法”中所述，使具有野生型MDH3或单个MDH突变体的表达琥珀酸产生通路的源自REV-0001的转化体在微量滴定板中生长，并测定二羧酸浓度。
表达MUT_014、MUT_015和MUT_34的菌株的琥珀酸滴度分别比表达MDH3的菌株的琥珀酸滴度高1.3倍、1.2倍和1.4倍，从而指示：在表达MDH突变体的菌株中，琥珀酸生产也得到改善。
序列表
<110> 帝斯曼知识产权资产管理有限公司
<120> 苹果酸脱氢酶
<160> 67
<170> BiSSAP 1.2
<210> 1
<211> 10504
<212> DNA
<213> Artificial Sequence
<220>
<221> source
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<223> /organism="Artificial Sequence"
/note="Artificial"
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ttgcccatcg aacgtacaag tactcctctg ttctctcctt cctttgcttt gtgcgacacc 60
taactacata gtgtttaaag attacggata tttaacttac ttagaataat gccatttttt 120
tgagttataa taatcctacg ttagtgtgag cgggatttaa actgtgagga ccttaataca 180
ttcagacact tctgcggtat caccctactt attcccttcg agattatatc taggaaccca 240
tcaggttggt ggaagattac ccgttctaag acttttcagc ttcctctatt gatgttacac 300
ctggacaccc cttttctggc atccagtttt taatcttcag tggcatgtga gattctccga 360
aattaattaa agcaatcaca caattctctc ggataccacc tcggttgaaa ctgacaggtg 420
gtttgttacg catgctaatg caaaggagcc tatatacctt tggctcggct gctgtaacag 480
ggaatataaa gggcagcata atttaggagt ttagtgaact tgcaacattt actattttcc 540
cttcttacgt aaatattttt ctttttaatt ctaaatcaat ctttttcaat tttttgtttg 600
tattcttttc ttgcttaaat ctataactac aaaaaacaca tacataaact aaaaatgact 660
gatttgaaca aattggtcaa ggaattgaat gatttgggtt tgactgacgt caaggaaatt 720
gtctacaacc catcttacga acaattattc gaagaagaaa ccaagccagg tttggaaggt 780
ttcgacaagg gtactttgac cactttaggt gctgttgctg ttgacaccgg tattttcacc 840
ggtcgttctc caaaggacaa atacattgtt tgtgatgaaa ccaccaagga caccgtctgg 900
tggaactctg aagctgccaa gaacgataac aagccaatga ctcaagaaac ctggaaatct 960
ttgagagaat tggttgccaa gcaattgtct ggtaagagat tattcgttgt tgacgctttc 1020
tgtggtgctt ctgaaaagca cagaattggt gtcagaatgg tcactgaagt tgcttggcaa 1080
gctcatttcg tcaagaacat gttcatcaga ccaactgacg aagaattgaa gaacttcaag 1140
gctgacttca ccgttttgaa tggtgccaag tgtaccaacc caaactggaa ggaacaaggt 1200
ttgaactctg aaaactttgt tgctttcaac atcactgaag gtatccaatt gattggtggt 1260
acttggtacg gtggtgaaat gaagaagggt atgttctcca tgatgaacta tttcttgcca 1320
ttgaaaggtg ttgcttccat gcactgttct gccaatgtcg gtaaggatgg tgacgttgcc 1380
atcttcttcg gtctatccgg tactggtaag accactctat ccactgaccc aaagagacaa 1440
ttgattggtg atgacgaaca cggttgggac gaatctggtg tctttaactt tgaaggtggt 1500
tgttacgcca agaccatcaa cttatctcaa gaaaacgaac cagatatcta cggtgccatc 1560
cgtcgtgatg ctttgttgga aaacgttgtt gtcagagctg acggttctgt tgacttcgac 1620
gacggttcca agactgaaaa caccagagtt tcttacccaa tctaccacat tgacaacatt 1680
gtcagacctg tttccaaggc tggtcacgct accaaggtta tcttcttgac tgctgatgct 1740
ttcggtgtct tgccacctgt ttccaaattg actccagaac aaaccgaata ctacttcttg 1800
tccggtttca ctgccaaatt ggctggtact gaaagaggtg tcactgaacc aactccaact 1860
ttctctgctt gtttcggtgc tgctttctta tctttgcacc caatccaata cgctgatgtc 1920
ttggttgaaa gaatgaaggc ttctggtgct gaagcttact tggtcaacac cggttggaac 1980
ggtactggta agagaatctc catcaaggat accagaggta tcattgatgc tatcttggac 2040
ggttccattg aaaaggctga aatgggtgaa ttgccaatct tcaacttggc cattccaaag 2100
gctttgccag gtgttgaccc agccatctta gatccaagag acacctacgc tgacaaggct 2160
caatggcaag tcaaggctga agatttggct aacagattcg tcaagaactt tgtcaaatac 2220
actgctaacc cagaagctgc caaattggtt ggtgctggtc caaaggctta aaggagttaa 2280
aggcaaagtt ttcttttcta gagccgttcc cacaaataat tatacgtata tgcttctttt 2340
cgtttactat atatctatat ttacaagcct ttattcactg atgcaatttg tttccaaata 2400
cttttttgga gatctcataa ctagatatca tgatggcgca acttggcgct atcttaatta 2460
ctctggctgc caggcccgtg tagagggccg caagaccttc tgtacgccat atagtctcta 2520
agaacttgaa caagtttcta gacctattgc cgcctttcgg atcgctattg ttcctccgga 2580
tcgatgtaca caaccgactg cacccaaacg aacacaaatc ttagcattgc ccatcgaacg 2640
tacaagtact cctctgttct ctccttcctt tgctttgtgc gacacctaac tacatagtgt 2700
ttaaagatta cggatattta acttacttag aataatgcca tttttttgag ttataataat 2760
cctacgttag tgtgagcggg atttaaactg tgaggacctt aatacattca gacacttctg 2820
cggtatcacc ctacttattc ccttcgagat tatatctagg aacccatcag gttggtggaa 2880
gattacccgt tctaagactt ttcagcttcc tctattgatg ttacacctgg acaccccttt 2940
tctggcatcc agtttttaat cttcagtggc atgtgagatt ctccgaaatt aattaaagca 3000
atcacacaat tctctcggat accacctcgg ttgaaactga caggtggttt gttacgcatg 3060
ctaatgcaaa ggagcctata tacctttggc tcggctgctg taacagggaa tataaagggc 3120
agcataattt aggagtttag tgaacttgca acatttacta ttttcccttc ttacgtaaat 3180
atttttcttt ttaattctaa atcaatcttt ttcaattttt tgtttgtatt cttttcttgc 3240
ttaaatctat aactacaaaa aacacataca taaactaaaa atgactgatt tgaacaaatt 3300
ggtcaaggaa ttgaatgatt tgggtttgac tgacgtcaag gaaattgtct acaacccatc 3360
ttacgaacaa ttattcgaag aagaaaccaa gccaggtttg gaaggtttcg acaagggtac 3420
tttgaccact ttaggtgctg ttgctgttga caccggtatt ttcaccggtc gttctccaaa 3480
ggacaaatac attgtttgtg atgaaaccac caaggacacc gtctggtgga actctgaagc 3540
tgccaagaac gataacaagc caatgactca agaaacctgg aaatctttga gagaattggt 3600
tgccaagcaa ttgtctggta agagattatt cgttgttgac gctttctgtg gtgcttctga 3660
aaagcacaga attggtgtca gaatggtcac tgaagttgct tggcaagctc atttcgtcaa 3720
gaacatgttc atcagaccaa ctgacgaaga attgaagaac ttcaaggctg acttcaccgt 3780
tttgaatggt gccaagtgta ccaacccaaa ctggaaggaa caaggtttga actctgaaaa 3840
ctttgttgct ttcaacatca ctgaaggtat ccaattgatt ggtggtactt ggtacggtgg 3900
tgaaatgaag aagggtatgt tctccatgat gaactatttc ttgccattga aaggtgttgc 3960
ttccatgcac tgttctgcca atgtcggtaa ggatggtgac gttgccatct tcttcggtct 4020
atccggtact ggtaagacca ctctatccac tgacccaaag agacaattga ttggtgatga 4080
cgaacacggt tgggacgaat ctggtgtctt taactttgaa ggtggttgtt acgccaagac 4140
catcaactta tctcaagaaa acgaaccaga tatctacggt gccatccgtc gtgatgcttt 4200
gttggaaaac gttgttgtca gagctgacgg ttctgttgac ttcgacgacg gttccaagac 4260
tgaaaacacc agagtttctt acccaatcta ccacattgac aacattgtca gacctgtttc 4320
caaggctggt cacgctacca aggttatctt cttgactgct gatgctttcg gtgtcttgcc 4380
acctgtttcc aaattgactc cagaacaaac cgaatactac ttcttgtccg gtttcactgc 4440
caaattggct ggtactgaaa gaggtgtcac tgaaccaact ccaactttct ctgcttgttt 4500
cggtgctgct ttcttatctt tgcacccaat ccaatacgct gatgtcttgg ttgaaagaat 4560
gaaggcttct ggtgctgaag cttacttggt caacaccggt tggaacggta ctggtaagag 4620
aatctccatc aaggatacca gaggtatcat tgatgctatc ttggacggtt ccattgaaaa 4680
ggctgaaatg ggtgaattgc caatcttcaa cttggccatt ccaaaggctt tgccaggtgt 4740
tgacccagcc atcttagatc caagagacac ctacgctgac aaggctcaat ggcaagtcaa 4800
ggctgaagat ttggctaaca gattcgtcaa gaactttgtc aaatacactg ctaacccaga 4860
agctgccaaa ttggttggtg ctggtccaaa ggcttaaagg agttaaaggc aaagttttct 4920
tttctagagc cgttcccaca aataattata cgtatatgct tcttttcgtt tactatatat 4980
ctatatttac aagcctttat tcactgatgc aatttgtttc caaatacttt tttggagatc 5040
tcataactag atatcatgat ggcgcaactt ggcgctatct taattactct ggctgccagg 5100
cccgtgtaga gggccgcaag accttctgta cgccatatag tctctaagaa cttgaacaag 5160
tttctagacc tattgccgcc tttcggatcg ctattgttcc tccggatcga tgtacacaac 5220
cgactgcacc caaacgaaca caaatcttag cattgcccat cgaacgtaca agtactcctc 5280
tgttctctcc ttcctttgct ttgtgcgaca cctaactaca tagtgtttaa agattacgga 5340
tatttaactt acttagaata atgccatttt tttgagttat aataatccta cgttagtgtg 5400
agcgggattt aaactgtgag gaccttaata cattcagaca cttctgcggt atcaccctac 5460
ttattccctt cgagattata tctaggaacc catcaggttg gtggaagatt acccgttcta 5520
agacttttca gcttcctcta ttgatgttac acctggacac cccttttctg gcatccagtt 5580
tttaatcttc agtggcatgt gagattctcc gaaattaatt aaagcaatca cacaattctc 5640
tcggatacca cctcggttga aactgacagg tggtttgtta cgcatgctaa tgcaaaggag 5700
cctatatacc tttggctcgg ctgctgtaac agggaatata aagggcagca taatttagga 5760
gtttagtgaa cttgcaacat ttactatttt cccttcttac gtaaatattt ttctttttaa 5820
ttctaaatca atctttttca attttttgtt tgtattcttt tcttgcttaa atctataact 5880
acaaaaaaca catacataaa ctaaaaatga ctgatttgaa caaattggtc aaggaattga 5940
atgatttggg tttgactgac gtcaaggaaa ttgtctacaa cccatcttac gaacaattat 6000
tcgaagaaga aaccaagcca ggtttggaag gtttcgacaa gggtactttg accactttag 6060
gtgctgttgc tgttgacacc ggtattttca ccggtcgttc tccaaaggac aaatacattg 6120
tttgtgatga aaccaccaag gacaccgtct ggtggaactc tgaagctgcc aagaacgata 6180
acaagccaat gactcaagaa acctggaaat ctttgagaga attggttgcc aagcaattgt 6240
ctggtaagag attattcgtt gttgacgctt tctgtggtgc ttctgaaaag cacagaattg 6300
gtgtcagaat ggtcactgaa gttgcttggc aagctcattt cgtcaagaac atgttcatca 6360
gaccaactga cgaagaattg aagaacttca aggctgactt caccgttttg aatggtgcca 6420
agtgtaccaa cccaaactgg aaggaacaag gtttgaactc tgaaaacttt gttgctttca 6480
acatcactga aggtatccaa ttgattggtg gtacttggta cggtggtgaa atgaagaagg 6540
gtatgttctc catgatgaac tatttcttgc cattgaaagg tgttgcttcc atgcactgtt 6600
ctgccaatgt cggtaaggat ggtgacgttg ccatcttctt cggtctatcc ggtactggta 6660
agaccactct atccactgac ccaaagagac aattgattgg tgatgacgaa cacggttggg 6720
acgaatctgg tgtctttaac tttgaaggtg gttgttacgc caagaccatc aacttatctc 6780
aagaaaacga accagatatc tacggtgcca tccgtcgtga tgctttgttg gaaaacgttg 6840
ttgtcagagc tgacggttct gttgacttcg acgacggttc caagactgaa aacaccagag 6900
tttcttaccc aatctaccac attgacaaca ttgtcagacc tgtttccaag gctggtcacg 6960
ctaccaaggt tatcttcttg actgctgatg ctttcggtgt cttgccacct gtttccaaat 7020
tgactccaga acaaaccgaa tactacttct tgtccggttt cactgccaaa ttggctggta 7080
ctgaaagagg tgtcactgaa ccaactccaa ctttctctgc ttgtttcggt gctgctttct 7140
tatctttgca cccaatccaa tacgctgatg tcttggttga aagaatgaag gcttctggtg 7200
ctgaagctta cttggtcaac accggttgga acggtactgg taagagaatc tccatcaagg 7260
ataccagagg tatcattgat gctatcttgg acggttccat tgaaaaggct gaaatgggtg 7320
aattgccaat cttcaacttg gccattccaa aggctttgcc aggtgttgac ccagccatct 7380
tagatccaag agacacctac gctgacaagg ctcaatggca agtcaaggct gaagatttgg 7440
ctaacagatt cgtcaagaac tttgtcaaat acactgctaa cccagaagct gccaaattgg 7500
ttggtgctgg tccaaaggct taaaggagtt aaaggcaaag ttttcttttc tagagccgtt 7560
cccacaaata attatacgta tatgcttctt ttcgtttact atatatctat atttacaagc 7620
ctttattcac tgatgcaatt tgtttccaaa tacttttttg gagatctcat aactagatat 7680
catgatggcg caacttggcg ctatcttaat tactctggct gccaggcccg tgtagagggc 7740
cgcaagacct tctgtacgcc atatagtctc taagaacttg aacaagtttc tagacctatt 7800
gccgcctttc ggatcgctat tgttcctccg gatcgatgta cacaaccgac tgcacccaaa 7860
cgaacacaaa tcttagcatt gcccatcgaa cgtacaagta ctcctctgtt ctctccttcc 7920
tttgctttgt gcgacaccta actacatagt gtttaaagat tacggatatt taacttactt 7980
agaataatgc catttttttg agttataata atcctacgtt agtgtgagcg ggatttaaac 8040
tgtgaggacc ttaatacatt cagacacttc tgcggtatca ccctacttat tcccttcgag 8100
attatatcta ggaacccatc aggttggtgg aagattaccc gttctaagac ttttcagctt 8160
cctctattga tgttacacct ggacacccct tttctggcat ccagttttta atcttcagtg 8220
gcatgtgaga ttctccgaaa ttaattaaag caatcacaca attctctcgg ataccacctc 8280
ggttgaaact gacaggtggt ttgttacgca tgctaatgca aaggagccta tatacctttg 8340
gctcggctgc tgtaacaggg aatataaagg gcagcataat ttaggagttt agtgaacttg 8400
caacatttac tattttccct tcttacgtaa atatttttct ttttaattct aaatcaatct 8460
ttttcaattt tttgtttgta ttcttttctt gcttaaatct ataactacaa aaaacacata 8520
cataaactaa aaatgactga tttgaacaaa ttggtcaagg aattgaatga tttgggtttg 8580
actgacgtca aggaaattgt ctacaaccca tcttacgaac aattattcga agaagaaacc 8640
aagccaggtt tggaaggttt cgacaagggt actttgacca ctttaggtgc tgttgctgtt 8700
gacaccggta ttttcaccgg tcgttctcca aaggacaaat acattgtttg tgatgaaacc 8760
accaaggaca ccgtctggtg gaactctgaa gctgccaaga acgataacaa gccaatgact 8820
caagaaacct ggaaatcttt gagagaattg gttgccaagc aattgtctgg taagagatta 8880
ttcgttgttg acgctttctg tggtgcttct gaaaagcaca gaattggtgt cagaatggtc 8940
actgaagttg cttggcaagc tcatttcgtc aagaacatgt tcatcagacc aactgacgaa 9000
gaattgaaga acttcaaggc tgacttcacc gttttgaatg gtgccaagtg taccaaccca 9060
aactggaagg aacaaggttt gaactctgaa aactttgttg ctttcaacat cactgaaggt 9120
atccaattga ttggtggtac ttggtacggt ggtgaaatga agaagggtat gttctccatg 9180
atgaactatt tcttgccatt gaaaggtgtt gcttccatgc actgttctgc caatgtcggt 9240
aaggatggtg acgttgccat cttcttcggt ctatccggta ctggtaagac cactctatcc 9300
actgacccaa agagacaatt gattggtgat gacgaacacg gttgggacga atctggtgtc 9360
tttaactttg aaggtggttg ttacgccaag accatcaact tatctcaaga aaacgaacca 9420
gatatctacg gtgccatccg tcgtgatgct ttgttggaaa acgttgttgt cagagctgac 9480
ggttctgttg acttcgacga cggttccaag actgaaaaca ccagagtttc ttacccaatc 9540
taccacattg acaacattgt cagacctgtt tccaaggctg gtcacgctac caaggttatc 9600
ttcttgactg ctgatgcttt cggtgtcttg ccacctgttt ccaaattgac tccagaacaa 9660
accgaatact acttcttgtc cggtttcact gccaaattgg ctggtactga aagaggtgtc 9720
actgaaccaa ctccaacttt ctctgcttgt ttcggtgctg ctttcttatc tttgcaccca 9780
atccaatacg ctgatgtctt ggttgaaaga atgaaggctt ctggtgctga agcttacttg 9840
gtcaacaccg gttggaacgg tactggtaag agaatctcca tcaaggatac cagaggtatc 9900
attgatgcta tcttggacgg ttccattgaa aaggctgaaa tgggtgaatt gccaatcttc 9960
aacttggcca ttccaaaggc tttgccaggt gttgacccag ccatcttaga tccaagagac 10020
acctacgctg acaaggctca atggcaagtc aaggctgaag atttggctaa cagattcgtc 10080
aagaactttg tcaaatacac tgctaaccca gaagctgcca aattggttgg tgctggtcca 10140
aaggcttaaa ggagttaaag gcaaagtttt cttttctaga gccgttccca caaataatta 10200
tacgtatatg cttcttttcg tttactatat atctatattt acaagccttt attcactgat 10260
gcaatttgtt tccaaatact tttttggaga tctcataact agatatcatg atggcgcaac 10320
ttggcgctat cttaattact ctggctgcca ggcccgtgta gagggccgca agaccttctg 10380
tacgccatat agtctctaag aacttgaaca agtttctaga cctattgccg cctttcggat 10440
cgctattgtt cctccggatc gatgtacaca accgactgca cccaaacgaa cacaaatctt 10500
agca 10504
<210> 2
<211> 4552
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..4552
<223> /organism="Artificial Sequence"
/note="Artificial"
/mol_type="unassigned DNA"
<400> 2
cggatcgatg tacacaaccg actgcaccca aacgaacaca aatcttagca gtgcgggcca 60
gaaaaaggaa gtgtttccct ccttcttgaa ttgatgttac cctcataaag cacgtggcct 120
cttatcgaga aagaaattac cgtcgctcgt gatttgtttg caaaaagaac aaaactgaaa 180
aaacccagac acgctcgact tcctgtcttc ctattgattg cagcttccaa tttcgtcaca 240
caacaaggtc ctagcgacgg ctcacaggtt ttgtaacaag caatcgaagg ttctggaatg 300
gcgggaaagg gtttagtacc acatgctatg atgcccactg tgatctccag agcaaagttc 360
gttcgatcgt actgttactc tctctctttc aaacagaatt gtccgaatcg tgtgacaaca 420
acagcctgtt ctcacacact cttttcttct aaccaagggg gtggtttagt ttagtagaac 480
ctcgtgaaac ttacatttac atatatataa acttgcataa attggtcaat gcaagaaata 540
catatttggt cttttctaat tcgtagtttt tcaagttctt agatgctttc tttttctctt 600
ttttacagat catcaaggaa gtaattatct actttttaca acaaatataa aacaatgtcc 660
tcttccaaga tcttggctgg tttgagagac aacttttctt tgttgggtga aaagaacaag 720
attttggtcg ccaacagagg tgaaatccca atcagaattt tcagatctgc tcacgaattg 780
tctatgagaa ctatcgccat ctactctcac gaagatagat tatccatgca cagattgaag 840
gctgatgaag cctacgttat cggtgaagaa ggtcaataca ccccagtcgg tgcttacttg 900
gccatggacg aaatcatcga aattgccaag aagcacaagg tcgatttcat ccacccaggt 960
tacggtttct tgtctgaaaa ctctgaattt gctgacaagg ttgttaaggc tggtattacc 1020
tggattggtc caccagctga agtcattgaa tctgttggtg acaaggtttc tgccagacat 1080
ttggctgctc gtgccaacgt tccaactgtc ccaggtactc caggtcctat cgaaaccgtt 1140
caagaagctc tagatttcgt caatgaatac ggttacccag ttatcatcaa ggctgctttc 1200
ggtggtggtg gtcgtggtat gagagttgtc agagaaggtg acgatgtcgc tgatgctttc 1260
caaagagcca cttctgaagc tagaactgct ttcggtaacg gtacttgttt cgtcgaaaga 1320
ttcttggaca agccaaagca cattgaagtt caattattag ctgacaacca cggtaacgtt 1380
gtccacttgt tcgaaagaga ctgttccgtc caaagacgtc accaaaaggt tgtcgaagtt 1440
gctccagcta agactttacc aagagaagtt agagatgcta tcttgaccga tgccgttaag 1500
ttggctaagg tttgtggtta cagaaacgct ggtactgctg aattcttggt tgacaaccaa 1560
aacagacatt acttcattga aatcaaccca agaattcaag tcgaacacac catcactgaa 1620
gaaatcactg gtattgacat tgtctccgct caaatccaaa tcgccgctgg tgctactttg 1680
actcaattag gtctattaca agacaaaatc accaccagag gtttctctat ccaatgtcgt 1740
atcaccactg aagatccatc caagaacttc caaccagaca ctggtcgttt ggaagtctac 1800
agatccgctg gtggtaacgg tgtcagattg gacggtggta acgcctacgc tggtgctacc 1860
atctctccac actacgactc catgttggtt aagtgttcct gttctggttc tacctacgaa 1920
attgtcagaa gaaagatgat cagagctttg attgaattca gaatcagagg tgtcaagacc 1980
aacatcccat tcttgttgac tttgttgacc aacccagttt tcattgaagg tacctactgg 2040
accactttca tcgatgacac tccacaattg ttccaaatgg tttcctctca aaacagagct 2100
caaaaattgt tgcactactt ggctgacttg gccgtcaacg gttcctctat caagggtcaa 2160
atcggtttac caaagttgaa gtccaaccct tccgttccac atttgcacga tgctcaaggt 2220
aatgtcatca acgttaccaa atctgcccca ccatccggtt ggagacaagt cttgttggaa 2280
aagggtccat ccgaatttgc caagcaagtc agacaattca acggtacttt gttgatggac 2340
accacctgga gagatgctca ccaatctttg ctagctacca gagtcagaac tcacgatttg 2400
gccaccattg ctccaaccac tgctcacgct ttggctggtg cctttgcttt ggaatgttgg 2460
ggtggtgcta ctttcgatgt cgccatgaga ttcttgcatg aggacccatg ggaaagattg 2520
agaaaattga gatctttggt cccaaacatt ccattccaaa tgttgttgag aggtgctaac 2580
ggtgttgctt actcctcttt gccagacaac gccattgacc atttcgttaa gcaagccaag 2640
gacaatggtg ttgacatttt cagagtcttt gacgctttga acgacttgga acaattgaag 2700
gttggtgtta atgctgtcaa gaaggctggt ggtgttgtcg aagctaccgt ttgttactct 2760
ggtgacatgt tgcaaccagg taagaaatac aacttggact actacttaga agttgtcgaa 2820
aagatcgttc aaatgggtac tcacatcttg ggtatcaagg acatggctgg taccatgaag 2880
ccagctgctg ccaaattgtt gattggttct ttacgtacca gatacccaga cttgccaatc 2940
cacgttcact ctcatgactc cgctggtact gctgttgctt ccatgactgc ttgtgctttg 3000
gccggtgctg atgttgttga cgttgccatt aactccatgt ccggtttgac ctctcaacca 3060
tctattaacg ctttgttggc ctccttggaa ggtaacattg acactggtat caacgtcgaa 3120
cacgttagag aattggacgc ttactgggct gaaatgagat tattatactc ttgtttcgaa 3180
gctgacttga agggtccaga ccctgaagtt taccaacacg aaattccagg tggtcaattg 3240
accaacttgt tgttccaagc tcaacaatta ggtctaggtg aacaatgggc tgaaaccaag 3300
agagcttaca gagaagctaa ctacttgttg ggtgacattg ttaaggtcac cccaacttct 3360
aaggtcgttg gtgatttggc tcaattcatg gtttctaaca aattgacttc tgatgacatc 3420
agaagattag ctaactcttt ggacttccca gactccgtta tggacttctt cgaaggtttg 3480
atcggtcaac catacggtgg tttcccagaa ccattgagat ccgatgtttt gagaaacaag 3540
cgtcgtaaat tgacttgtag accaggttta gaattggaac cattcgattt ggaaaagatc 3600
agagaagatt tgcaaaacag attcggtgat atcgatgaat gtgatgttgc ctcctacaac 3660
atgtatcctc gtgtctacga agatttccaa aagattagag aaacttacgg tgacttgtct 3720
gtcttaccaa ccaagaactt cttggctcca gctgaaccag acgaagaaat cgaagtcacc 3780
attgaacaag gtaagacttt gattatcaaa ttacaagctg ttggtgattt gaacaagaaa 3840
accggtcaaa gagaagtcta cttcgaattg aacggtgaat tgagaaagat cagagttgct 3900
gacaaatctc aaaacattca atctgttgcc aagccaaagg ctgatgtcca cgacacccac 3960
caaatcggtg ctccaatggc tggtgtcatc attgaagtca aggttcacaa gggttctttg 4020
gtcaagaagg gtgaatctat cgccgttttg tctgctatga agatggaaat ggttgtttcc 4080
tctccagctg atggtcaagt caaagatgtc tttatccgtg acggtgaatc cgtcgatgct 4140
tctgacttgt tggttgtttt ggaagaagaa actctaccac cttctcaaaa gaaataaagc 4200
gaatttctta tgatttatga tttttattat taaataagtt ataaaaaaaa taagtgtata 4260
caaattttaa agtgactctt aggttttaaa acgaaaattc ttattcttga gtaactcttt 4320
cctgtaggtc aggttgcttt ctcaggtata gcatgaggtc gctcttattg accacacctc 4380
taccggcatg ccgagcaaat gcctgcaaat cgctccccat ttcacccaat tgtagatatg 4440
ctaactccag caatgagttg atgaatctcg gtgtgtattt tatgtcctca gaggacaacc 4500
tcacgctttc cggcatcttc cagaccacag tatatccatc cgcctcctgt tg 4552
<210> 3
<211> 1572
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..1572
<223> /organism="Artificial Sequence"
/note="Artificial"
/mol_type="unassigned DNA"
<400> 3
tcgtacgctg caggtcgacg aattctaccg ttcgtataat gtatgctata cgaagttata 60
gatctgttta gcttgcctcg tccccgccgg gtcacccggc cagcgacatg gaggcccaga 120
ataccctcct tgacagtctt gacgtgcgca gctcaggggc atgatgtgac tgtcgcccgt 180
acatttagcc catacatccc catgtataat catttgcatc catacatttt gatggccgca 240
cggcgcgaag caaaaattac ggctcctcgc tgcagacctg cgagcaggga aacgctcccc 300
tcacagacgc gttgaattgt ccccacgccg cgcccctgta gagaaatata aaaggttagg 360
atttgccact gaggttcttc tttcatatac ttccttttaa aatcttgcta ggatacagtt 420
ctcacatcac atccgaacat aaacaaccat gggtaaggaa aagactcacg tttcgaggcc 480
gcgattaaat tccaacatgg atgctgattt atatgggtat aaatgggctc gcgataatgt 540
cgggcaatca ggtgcgacaa tctatcgatt gtatgggaag cccgatgcgc cagagttgtt 600
tctgaaacat ggcaaaggta gcgttgccaa tgatgttaca gatgagatgg tcagactaaa 660
ctggctgacg gaatttatgc ctcttccgac catcaagcat tttatccgta ctcctgatga 720
tgcatggtta ctcaccactg cgatccccgg caaaacagca ttccaggtat tagaagaata 780
tcctgattca ggtgaaaata ttgttgatgc gctggcagtg ttcctgcgcc ggttgcattc 840
gattcctgtt tgtaattgtc cttttaacag cgatcgcgta tttcgtctcg ctcaggcgca 900
atcacgaatg aataacggtt tggttgatgc gagtgatttt gatgacgagc gtaatggctg 960
gcctgttgaa caagtctgga aagaaatgca taagcttttg ccattctcac cggattcagt 1020
cgtcactcat ggtgatttct cacttgataa ccttattttt gacgagggga aattaatagg 1080
ttgtattgat gttggacgag tcggaatcgc agaccgatac caggatcttg ccatcctatg 1140
gaactgcctc ggtgagtttt ctccttcatt acagaaacgg ctttttcaaa aatatggtat 1200
tgataatcct gatatgaata aattgcagtt tcatttgatg ctcgatgagt ttttctaatc 1260
agtactgaca ataaaaagat tcttgttttc aagaacttgt catttgtata gtttttttat 1320
attgtagttg ttctatttta atcaaatgtt agcgtgattt atattttttt tcgcctcgac 1380
atcatctgcc cagatgcgaa gttaagtgcg cagaaagtaa tatcatgcgt caatcgtatg 1440
tgaatgctgg tcgctatact gctgtcgatt cgatactaac gccgccatcc agtgtcgaaa 1500
acgagctcat aacttcgtat aatgtatgct atacgaacgg tagaattcga tatcagatcc 1560
actagtggcc ta 1572
<210> 4
<211> 2260
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..2260
<223> /organism="Artificial Sequence"
/note="Artificial"
/mol_type="unassigned DNA"
<400> 4
aacgttgtcc aggtttgtat ccacgtgtgt ccgttccgcc aatattccgc gtgcccgcgg 60
aaccgccaga tattcattac ttgacgcaaa agcgtttgaa ataatgacga aaaagaagga 120
agaaaaaaaa agaaaaatac cgcttctagg cgggttatct actgatccga gcttccacta 180
ggatagcacc caaacacctg catatttgga cgacctttac ttacaccacc aaaaaccact 240
ttcgcctctc ccgcccctga taacgtccac taattgagcg attacctgag cggtcctctt 300
ttgtttgcag catgagactt gcatactgca aatcgtaagt agcaacgtgt caaggtcaaa 360
actgtatgga aaccttgtca cctcacttaa ttctagctag cctaccctgc aagtcaagag 420
gtgtccgtga ttcctagcca cctcaaggta tgcctctccc cggaaactgt ggccttttct 480
ggcacacatg atctccacga tttcaacata taaatagctt ttgataatgg caatattaat 540
caaatttatt ttacttcttt cttgtaacat ctctcttgta atcccttatt ccttctagct 600
atttttcata aaaaaccaag caactgctta tcaacacaca aacactaaat caaaatgaac 660
gttgaaactt ctttgccagg ttcttctggt tctgacttgg aaactttcca ccacgaaacc 720
aagaagcatg ccaaccacga ctctggtatt tccgtcaacc atgaagctga aattggtgtt 780
aaccacactt tcgaaaagcc aggtccagtt ggtatcagag aaagattacg tcacttcacc 840
tgggcttggt acactttgac catgtcctgt ggtggtttgg ctttgttgat tgtcaaccaa 900
ccacacgact tcaagggttt gaaagatatt gccagagttg tctactgttt gaacttggct 960
ttctttgtta tcgttacctc tttgatggcc atcagattca tcttgcacaa gaacatgtgg 1020
gaatccttgg gtcacgacag agaaggtttg tttttcccaa ctttctggtt atccattgct 1080
accatgatca ctggtttgta caagtgtttc ggtgatgatg ctaacgaaaa gttcaccaag 1140
tgtttgcaag ttttgttctg gatctactgt ggttgtacca tgatcactgc tgtcggtcaa 1200
tactctttcg tctttgctac ccacaaatac gaattgcaca ccatgatgcc atcctggatc 1260
ttgccagctt tcccagttat gttgtctggt actatcgcct ccgtcatcgg ttctggtcaa 1320
ccagcttccg atggtattcc aattattatt gctggtatca ctttccaagg tttaggtttc 1380
tccatctcct tcatgatgta cgctcactac attggtagat tgatggaagt tggtttacca 1440
tctccagaac acagaccagg tatgttcatc tgtgttggtc ctccagcttt caccgctttg 1500
gctttggtcg gtatggccaa ggctttacca gacgacttcc aaattgtcgg tgaccctcac 1560
gctgtcattg acggtcgtgt tatgttgttc ttggctgtct ctgctgccat cttcttatgg 1620
gctttgtctt tctggttctt ctgtatcgct gttgttgctg ttgtcagatc tccaccaaag 1680
ggtttccatt tgaactggtt tgccatggtt ttcccaaaca ctggtttcac cttggctacc 1740
atcactttgg ctaacatgtt cgaatctcca ggtgtcaagg gtgttgccac tgctatgtcc 1800
ctatgtgtca tcatcatgtt tattttcgtc ttggtttctg ccatcagagc tgtcatcaga 1860
aaggacatca tgtggccagg tcaagatgaa gatgtttctg aataaagagt aataattatt 1920
gcttccatat aatattttta tatacctctt atttttatgt attagttaat taagtatttt 1980
tatctatctg cttatcattt tcttttcata tagggggggt tggtgttttc ttgcccatca 2040
gattgatgtc ctccaactcg gcactatttt acaaagggtt tttttgtaag agaaggagaa 2100
gacagatact aaaccatacg ttactcgaaa caaaaaaaaa aaaaatggaa aaagctgcta 2160
tcaacaaaag acggcctcat caaacctaaa gaaaccatgt cagcgtcctc aaataaccac 2220
aaacatcctt cccatatgct cggtcgtgct tgttgtacct 2260
<210> 5
<211> 2432
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..2432
<223> /organism="Artificial Sequence"
/note="Artificial"
/mol_type="unassigned DNA"
<400> 5
aaataaccac aaacatcctt cccatatgct cggtcgtgct tgttgtacct gtgcttttct 60
ttttttgcgg tcacccccat gtggcgggga ggcagaggag taggtagagc aacgaatcct 120
actatttatc caaattagtc taggaactct ttttctagat tttttagatt tgagggcaag 180
cgctgttaac gactcagaaa tgtaagcact acggagtaga acgagaaatc cgccataggt 240
ggaaatccta gcaaaatctt gcttacccta gctagcctca ggtaagctag ccttagcctg 300
tcaaattttt ttcaaaattt ggtaagtttc tactagcaaa gcaaacacgg ttcaacaaac 360
cgaaaactcc actcattata cgtggaaacc gaaacaaaaa aacaaaaacc aaaatactcg 420
ccaatgagaa agttgctgcg tttctacttt cgaggaagag gaactgagag gattgactac 480
gaaaggggca aaaacgagtc gtattctccc attattgtct gctaccacgc ggtctagtag 540
aataagcaac cagtcaacgc taagacaggt aatcaaaata ccagtctgct ggctacgggc 600
tagtttttac ctcttttaga acccactgta aaagtccgtt gtaaagcccg ttctcactgt 660
tggcgttttt ttttttttgg tttagtttct tatttttcat ttttttcttt catgaccaaa 720
aacaaacaaa tctcgcgatt tgtactgcgg ccactggggc gtggccaaaa aaatgacaaa 780
tttagaaacc ttagtttctg atttttcctg ttatgaggag atatgataaa aaatattact 840
gctttattgt ttttttttta tctactgaaa tagagaaact tacccaagga ggaggcaaaa 900
aaaagagtat atatacagca ggtaccattc agattttaat atattctttt ctcttcttct 960
acactattat tataataatt ttactatatt catttttagc ttaaaacctc atagaatatt 1020
attcttcagt cactcgctta aatacttatc aaaaatggtt aaggttgcca tcttaggtgc 1080
ttctggtggt gtcggtcaac cattatctct attattgaaa ttgtctccat acgtttctga 1140
attggctttg tacgatatca gagctgctga aggtattggt aaggatttgt cccacatcaa 1200
caccaactcc tcttgtgttg gttacgacaa ggattccatc gaaaacactt tgtccaatgc 1260
tcaagttgtc ttgattccag ctggtgttcc aagaaagcca ggtttgacca gagatgattt 1320
gttcaagatg aacgctggta tcgttaagtc tttggttact gctgtcggta aatttgcccc 1380
aaacgctcgt atcttagtca tctccaaccc tgttaactct ttggttccaa ttgccgttga 1440
aactttgaag aagatgggta agttcaagcc aggtaacgtt atgggtgtca ccaacttgga 1500
tttggtcaga gctgaaactt tcttggttga ctacttgatg ttgaagaacc caaagatcgg 1560
tcaagaacaa gacaagacca ccatgcacag aaaggtcacc gtcatcggtg gtcactctgg 1620
tgaaaccatc attccaatca tcactgacaa atccttggtt ttccaattgg acaagcaata 1680
cgaacatttc atccacagag tccaattcgg tggtgacgaa attgtcaagg ccaagcaagg 1740
tgccggttct gctaccttgt ccatggcttt cgctggtgcc aaatttgctg aagaagtctt 1800
acgttctttc cacaacgaaa agccagaaac tgaatctttg tctgctttcg tctacttgcc 1860
aggtttgaag aacggtaaga aggctcaaca attagtcggt gacaactcca ttgaatactt 1920
ctctttgcca attgttttga gaaacggttc cgttgtttcc attgacactt ctgttttgga 1980
aaaattgtct ccaagagaag aacaattggt caacactgct gtcaaggaat tgagaaagaa 2040
cattgaaaag ggtaagtctt tcatcttgga cagttaaagt ctgaagaatg aatgatttga 2100
tgatttcttt ttccctccat ttttcttact gaatatatca atgatataga cttgtatagt 2160
ttattatttc aaattaagta gctatatata gtcaagataa cgtttgtttg acacgattac 2220
attattcgtc gacatctttt ttcagcctgt cgtggtagca atttgaggag tattattaat 2280
tgaataggtt cattttgcgc tcgcataaac agttttcgtc agggacagta tgttggaatg 2340
agtggtaatt aatggtgaca tgacatgtta tagcaatacc tcgaaacctt cgaatccagc 2400
cagcatgtcg acacccacaa gatgtagtgc ac 2432
<210> 6
<211> 2656
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..2656
<223> /organism="Artificial Sequence"
/note="Artificial"
/mol_type="unassigned DNA"
<400> 6
gaaaccttcg aatccagcca gcatgtcgac acccacaaga tgtagtgcac gtgcttagtc 60
aaaaaattag ccttttaatt ctgctgtaac ccgtacatgc ccaaaatagg gggcgggtta 120
cacagaatat ataacatcgt aggtgtctgg gtgaacagtt tattcctggc atccactaaa 180
tataatggag cccgcttttt aagctggcat ccagaaaaaa aaagaatccc agcaccaaaa 240
tattgttttc ttcaccaacc atcagttcat aggtccattc tcttagcgca actacagaga 300
acaggggcac aaacaggcaa aaaacgggca caacctcaat ggagtgatgc aacctgcctg 360
gagtaaatga tgacacaagg caattgaccc acgcatgtat ctatctcatt ttcttacacc 420
ttctattacc ttctgctctc tctgatttgg aaaaagctga aaaaaaaggt tgaaaccagt 480
tccctgaaat tattccccta cttgactaat aagtatataa agacggtagg tattgattgt 540
aattctgtaa atctatttct taaacttctt aaattctact tttatagtta gtcttttttt 600
tagttttaaa acaccaagaa cttagtttcg aataaacaca cataaacaaa caaaatgtcc 660
aacaagcctt tcatctacca agctccattc ccaatgggta aggacaacac tgaatactac 720
ttgttgactt ctgactacgt ttccgttgct gatttcgatg gtgaaaccat cttgaaggtt 780
gaaccagaag ccttgacttt gttggctcaa caagccttcc acgatgcttc tttcatgttg 840
cgtccagctc accaaaagca agttgctgcc attttgcacg acccagaagc ctccgaaaac 900
gacaaatacg ttgctttgca attcttgaga aactctgaaa ttgctgccaa gggtgtctta 960
ccaacttgtc aagacactgg tactgccatc attgtcggta agaagggtca aagagtctgg 1020
accggtggtg gtgacgaaga aactctatcc aagggtgttt acaacactta cattgaagat 1080
aatttacgtt actctcaaaa tgctgctttg gacatgtaca aggaagtcaa cactggtact 1140
aacttgccag ctcaaatcga cttatacgct gttgacggtg acgaatacaa gttcttgtgt 1200
gttgccaagg gtggtggttc tgctaacaag acctacttgt accaagaaac caaggctttg 1260
ttgactccag gtaaattgaa gaacttcttg gtcgaaaaga tgagaacttt gggtactgct 1320
gcttgtccac cataccacat tgctttcgtt atcggtggta cttccgctga aaccaacttg 1380
aaaaccgtca aattggcttc cgctcactac tacgatgaat tgccaactga aggtaacgaa 1440
cacggtcaag ccttcagaga tgtccaattg gaacaagaat tgttggaaga agctcaaaaa 1500
ttaggtttgg gtgctcaatt tggtggtaaa tactttgctc acgatatcag agttatcaga 1560
ttaccaagac atggtgcttc ttgtccagtt ggtatgggtg tttcctgttc tgctgacaga 1620
aacatcaagg ccaagatcaa cagagaaggt atctggattg aaaaattgga acacaaccca 1680
ggtcaataca tcccacaaga attgagacaa gctggtgaag gtgaagctgt caaggttgac 1740
ttgaacagac caatgaagga aatcttggct caattatctc aatacccagt ttccaccaga 1800
ttatctttga ccggtactat cattgtcggt cgtgacattg ctcatgccaa gttgaaggaa 1860
ttgattgatg ctggtaagga attgcctcaa tacatcaagg accatccaat ctactacgct 1920
ggtccagcca agaccccagc tggttaccca tctggttctt tgggtccaac caccgctggt 1980
agaatggact cttacgttga cttgctacaa tctcacggtg gttccatgat catgttggct 2040
aagggtaaca gatctcaaca agtcaccgat gcttgtcaca agcacggtgg tttctatttg 2100
ggttccattg gtggtccagc tgctgtcttg gctcaacaat ctatcaagca cttggaatgt 2160
gttgcttacc cagaattggg tatggaagcc atctggaaga ttgaagtcga agatttccca 2220
gctttcatct tagtcgatga caagggtaac gacttcttcc aacaaattgt caacaagcaa 2280
tgtgccaact gtaccaagta aaataaagca atcttgatga ggataatgat ttttttttga 2340
atatacataa atactaccgt ttttctgcta gattttgtga agacgtaaat aagtacatat 2400
tactttttaa gccaagacaa gattaagcat taactttacc cttttctctt ctaagtttca 2460
atactagtta tcactgttta aaagttatgg cgagaacgtc ggcggttaaa atatattacc 2520
ctgaacgtgg tgaattgaag ttctaggatg gtttaaagat ttttcctttt tgggaaataa 2580
gtaaacaata tattgctgcc ttcctcaaag ccaaagttcg cgttccgacc ttgcctccca 2640
aatccgagtt gcgatt 2656
<210> 7
<211> 4429
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..4429
<223> /organism="Artificial Sequence"
/note="Artificial"
/mol_type="unassigned DNA"
<400> 7
aaagccaaag ttcgcgttcc gaccttgcct cccaaatccg agttgcgatt gtgcttggct 60
gataatagcg tataaacaat gcatactttg tacgttcaaa atacaatgca gtagatatat 120
ttatgcatat tacatataat acatatcaca taggaagcaa caggcgcgtt ggacttttaa 180
ttttcgagga ccgcgaatcc ttacatcaca cccaatcccc cacaagtgat cccccacaca 240
ccatagcttc aaaatgtttc tactcctttt ttactcttcc agattttctc ggactccgcg 300
catcgccgta ccacttcaaa acacccaagc acagcatact aaatttcccc tctttcttcc 360
tctagggtgt cgttaattac ccgtactaaa ggtttggaaa agaaaaaaga caccgcctcg 420
tttctttttc ttcgtcgaaa aaggcaataa aaatttttat cacgtttctt tttcttgaaa 480
attttttttt ttgatttttt tctctttcga tgacctccca ttgatattta agttaataaa 540
cggtcttcaa tttctcaagt ttcagtttca tttttcttgt tctattacaa ctttttttac 600
ttcttgctca ttagaaagaa agcatagcaa tctaatctaa gttttaatta caaaatggtt 660
gatggtagat cttctgcttc cattgttgcc gttgacccag aaagagctgc cagagaaaga 720
gatgctgctg ccagagcttt gttgcaagac tctccattgc acaccaccat gcaatacgct 780
acctctggtt tggaattgac tgttccatac gctttgaagg ttgttgcttc tgctgacact 840
ttcgacagag ccaaggaagt tgctgatgaa gtcttgagat gtgcctggca attggctgac 900
accgttttga actctttcaa cccaaactct gaagtctctt tagtcggtag attaccagtc 960
ggtcaaaagc atcaaatgtc tgctccattg aaacgtgtca tggcttgttg tcaaagagtc 1020
tacaactcct ctgctggttg tttcgaccca tccactgctc cagttgccaa ggctttgaga 1080
gaaattgctt tgggtaagga aagaaacaat gcttgtttgg aagctttgac tcaagcttgt 1140
accttgccaa actctttcgt cattgatttc gaagctggta ctatctccag aaagcacgaa 1200
cacgcttctt tggatttggg tggtgtttcc aagggttaca tcgtcgatta cgtcattgac 1260
aacatcaatg ctgctggttt ccaaaacgtt ttctttgact ggggtggtga ctgtcgtgcc 1320
tccggtatga acgccagaaa cactccatgg gttgtcggta tcactagacc tccttccttg 1380
gacatgttgc caaaccctcc aaaggaagct tcttacatct ccgtcatctc tttggacaat 1440
gaagctttgg ctacctctgg tgattacgaa aacttgatct acactgctga cgataaacca 1500
ttgacctgta cctacgattg gaaaggtaag gaattgatga agccatctca atccaatatc 1560
gctcaagttt ccgtcaagtg ttactctgcc atgtacgctg acgctttggc taccgcttgt 1620
ttcatcaagc gtgacccagc caaggtcaga caattgttgg atggttggag atacgttaga 1680
gacaccgtca gagattaccg tgtctacgtc agagaaaacg aaagagttgc caagatgttc 1740
gaaattgcca ctgaagatgc tgaaatgaga aagagaagaa tttccaacac tttaccagct 1800
cgtgtcattg ttgttggtgg tggtttggct ggtttgtccg ctgccattga agctgctggt 1860
tgtggtgctc aagttgtttt gatggaaaag gaagccaagt tgggtggtaa ctctgccaag 1920
gctacctctg gtatcaacgg ttggggtact agagcccaag ctaaggcttc cattgtcgat 1980
ggtggtaagt acttcgaaag agatacctac aagtctggta tcggtggtaa caccgatcca 2040
gctttggtta agactttgtc catgaaatct gctgacgcta tcggttggtt gacttctcta 2100
ggtgttccat tgactgtttt gtcccaatta ggtggtcact ccagaaagag aactcacaga 2160
gccccagaca agaaggatgg tactccattg ccaattggtt tcaccatcat gaaaacttta 2220
gaagatcatg ttagaggtaa cttgtccggt agaatcacca tcatggaaaa ctgttccgtt 2280
acctctttgt tgtctgaaac caaggaaaga ccagacggta ctaagcaaat cagagttacc 2340
ggtgtcgaat tcactcaagc tggttctggt aagaccacca ttttggctga tgctgttatc 2400
ttggccaccg gtggtttctc caacgacaag actgctgatt ctttgttgag agaacatgcc 2460
ccacacttgg ttaacttccc aaccaccaac ggtccatggg ctactggtga tggtgtcaag 2520
ttggctcaaa gattaggtgc tcaattggtc gatatggaca aggttcaatt gcacccaact 2580
ggtttgatca acccaaagga cccagccaac ccaaccaaat tcttgggtcc agaagctcta 2640
agaggttctg gtggtgtttt gttgaacaaa caaggtaaga gatttgtcaa cgaattggat 2700
ttgagatctg ttgtttccaa ggccatcatg gaacaaggtg ctgaataccc aggttctggt 2760
ggttccatgt ttgcttactg tgtcttgaac gctgctgctc aaaaattgtt tggtgtttcc 2820
tctcacgaat tctactggaa gaagatgggt ttgttcgtca aggctgacac catgagagac 2880
ttggctgctt tgattggttg tccagttgaa tccgttcaac aaactttaga agaatacgaa 2940
agattatcca tctctcaaag atcttgtcca attaccagaa aatctgttta cccatgtgtt 3000
ttgggtacta aaggtccata ctatgtcgcc tttgtcactc catctatcca ctacaccatg 3060
ggtggttgtt tgatttctcc atctgctgaa atccaaatga agaacacttc ttccagagcc 3120
ccattgtccc actccaaccc aatcttgggt ttattcggtg ctggtgaagt caccggtggt 3180
gtccacggtg gtaacagatt aggtggtaac tctttgttgg aatgtgttgt tttcggtaga 3240
attgccggtg acagagcttc taccattttg caaagaaagt cctctgcttt gtctttcaag 3300
gtctggacca ctgttgtttt gagagaagtc agagaaggtg gtgtctacgg tgctggttcc 3360
cgtgtcttga gattcaactt accaggtgct ctacaaagat ctggtctatc cttgggtcaa 3420
ttcattgcca tcagaggtga ctgggacggt caacaattga ttggttacta ctctccaatc 3480
actttgccag acgatttggg tatgattgac attttggcca gatctgacaa gggtacttta 3540
cgtgaatgga tctctgcttt ggaaccaggt gacgctgtcg aaatgaaggc ttgtggtggt 3600
ttggtcatcg aaagaagatt atctgacaag cacttcgttt tcatgggtca cattatcaac 3660
aagctatgtt tgattgctgg tggtactggt gttgctccaa tgttgcaaat catcaaggcc 3720
gctttcatga agccattcat cgacactttg gaatccgtcc acttgatcta cgctgctgaa 3780
gatgtcactg aattgactta cagagaagtt ttggaagaac gtcgtcgtga atccagaggt 3840
aaattcaaga aaactttcgt tttgaacaga cctcctccat tatggactga cggtgtcggt 3900
ttcatcgacc gtggtatctt gaccaaccac gttcaaccac catctgacaa cttattggtt 3960
gccatctgtg gtccaccagt tatgcaaaga attgtcaagg ccactttaaa gactttaggt 4020
tacaacatga acttggtcag aaccgttgac gaaactgaac catctggaag ttaaaggaag 4080
tatctcggaa atattaattt aggccatgtc cttatgcacg tttcttttga tacttacggg 4140
tacatgtaca caagtatatc tatatatata aattaatgaa aatcccctat ttatatatat 4200
gactttaacg agacagaaca gttttttatt ttttatccta tttgatgaat gatacagttt 4260
cttattcacg tgttataccc acaccaaatc caatagcaat accggccatc acaatcactg 4320
tttcggcagc ccctaagatc agacaaaaca tccggaacca ccttaaatca acgtccctca 4380
gaaagcctgt atgcgaagcc acaatccttt ccaacagacc atactaagt 4429
<210> 8
<211> 1138
<212> PRT
<213> Trypanosoma brucei
<220>
<223> Fumarate reductase (FRDg)
<400> 8
Met Val Asp Gly Arg Ser Ser Ala Ser Ile Val Ala Val Asp Pro Glu
1 5 10 15
Arg Ala Ala Arg Glu Arg Asp Ala Ala Ala Arg Ala Leu Leu Gln Asp
20 25 30
Ser Pro Leu His Thr Thr Met Gln Tyr Ala Thr Ser Gly Leu Glu Leu
35 40 45
Thr Val Pro Tyr Ala Leu Lys Val Val Ala Ser Ala Asp Thr Phe Asp
50 55 60
Arg Ala Lys Glu Val Ala Asp Glu Val Leu Arg Cys Ala Trp Gln Leu
65 70 75 80
Ala Asp Thr Val Leu Asn Ser Phe Asn Pro Asn Ser Glu Val Ser Leu
85 90 95
Val Gly Arg Leu Pro Val Gly Gln Lys His Gln Met Ser Ala Pro Leu
100 105 110
Lys Arg Val Met Ala Cys Cys Gln Arg Val Tyr Asn Ser Ser Ala Gly
115 120 125
Cys Phe Asp Pro Ser Thr Ala Pro Val Ala Lys Ala Leu Arg Glu Ile
130 135 140
Ala Leu Gly Lys Glu Arg Asn Asn Ala Cys Leu Glu Ala Leu Thr Gln
145 150 155 160
Ala Cys Thr Leu Pro Asn Ser Phe Val Ile Asp Phe Glu Ala Gly Thr
165 170 175
Ile Ser Arg Lys His Glu His Ala Ser Leu Asp Leu Gly Gly Val Ser
180 185 190
Lys Gly Tyr Ile Val Asp Tyr Val Ile Asp Asn Ile Asn Ala Ala Gly
195 200 205
Phe Gln Asn Val Phe Phe Asp Trp Gly Gly Asp Cys Arg Ala Ser Gly
210 215 220
Met Asn Ala Arg Asn Thr Pro Trp Val Val Gly Ile Thr Arg Pro Pro
225 230 235 240
Ser Leu Asp Met Leu Pro Asn Pro Pro Lys Glu Ala Ser Tyr Ile Ser
245 250 255
Val Ile Ser Leu Asp Asn Glu Ala Leu Ala Thr Ser Gly Asp Tyr Glu
260 265 270
Asn Leu Ile Tyr Thr Ala Asp Asp Lys Pro Leu Thr Cys Thr Tyr Asp
275 280 285
Trp Lys Gly Lys Glu Leu Met Lys Pro Ser Gln Ser Asn Ile Ala Gln
290 295 300
Val Ser Val Lys Cys Tyr Ser Ala Met Tyr Ala Asp Ala Leu Ala Thr
305 310 315 320
Ala Cys Phe Ile Lys Arg Asp Pro Ala Lys Val Arg Gln Leu Leu Asp
325 330 335
Gly Trp Arg Tyr Val Arg Asp Thr Val Arg Asp Tyr Arg Val Tyr Val
340 345 350
Arg Glu Asn Glu Arg Val Ala Lys Met Phe Glu Ile Ala Thr Glu Asp
355 360 365
Ala Glu Met Arg Lys Arg Arg Ile Ser Asn Thr Leu Pro Ala Arg Val
370 375 380
Ile Val Val Gly Gly Gly Leu Ala Gly Leu Ser Ala Ala Ile Glu Ala
385 390 395 400
Ala Gly Cys Gly Ala Gln Val Val Leu Met Glu Lys Glu Ala Lys Leu
405 410 415
Gly Gly Asn Ser Ala Lys Ala Thr Ser Gly Ile Asn Gly Trp Gly Thr
420 425 430
Arg Ala Gln Ala Lys Ala Ser Ile Val Asp Gly Gly Lys Tyr Phe Glu
435 440 445
Arg Asp Thr Tyr Lys Ser Gly Ile Gly Gly Asn Thr Asp Pro Ala Leu
450 455 460
Val Lys Thr Leu Ser Met Lys Ser Ala Asp Ala Ile Gly Trp Leu Thr
465 470 475 480
Ser Leu Gly Val Pro Leu Thr Val Leu Ser Gln Leu Gly Gly His Ser
485 490 495
Arg Lys Arg Thr His Arg Ala Pro Asp Lys Lys Asp Gly Thr Pro Leu
500 505 510
Pro Ile Gly Phe Thr Ile Met Lys Thr Leu Glu Asp His Val Arg Gly
515 520 525
Asn Leu Ser Gly Arg Ile Thr Ile Met Glu Asn Cys Ser Val Thr Ser
530 535 540
Leu Leu Ser Glu Thr Lys Glu Arg Pro Asp Gly Thr Lys Gln Ile Arg
545 550 555 560
Val Thr Gly Val Glu Phe Thr Gln Ala Gly Ser Gly Lys Thr Thr Ile
565 570 575
Leu Ala Asp Ala Val Ile Leu Ala Thr Gly Gly Phe Ser Asn Asp Lys
580 585 590
Thr Ala Asp Ser Leu Leu Arg Glu His Ala Pro His Leu Val Asn Phe
595 600 605
Pro Thr Thr Asn Gly Pro Trp Ala Thr Gly Asp Gly Val Lys Leu Ala
610 615 620
Gln Arg Leu Gly Ala Gln Leu Val Asp Met Asp Lys Val Gln Leu His
625 630 635 640
Pro Thr Gly Leu Ile Asn Pro Lys Asp Pro Ala Asn Pro Thr Lys Phe
645 650 655
Leu Gly Pro Glu Ala Leu Arg Gly Ser Gly Gly Val Leu Leu Asn Lys
660 665 670
Gln Gly Lys Arg Phe Val Asn Glu Leu Asp Leu Arg Ser Val Val Ser
675 680 685
Lys Ala Ile Met Glu Gln Gly Ala Glu Tyr Pro Gly Ser Gly Gly Ser
690 695 700
Met Phe Ala Tyr Cys Val Leu Asn Ala Ala Ala Gln Lys Leu Phe Gly
705 710 715 720
Val Ser Ser His Glu Phe Tyr Trp Lys Lys Met Gly Leu Phe Val Lys
725 730 735
Ala Asp Thr Met Arg Asp Leu Ala Ala Leu Ile Gly Cys Pro Val Glu
740 745 750
Ser Val Gln Gln Thr Leu Glu Glu Tyr Glu Arg Leu Ser Ile Ser Gln
755 760 765
Arg Ser Cys Pro Ile Thr Arg Lys Ser Val Tyr Pro Cys Val Leu Gly
770 775 780
Thr Lys Gly Pro Tyr Tyr Val Ala Phe Val Thr Pro Ser Ile His Tyr
785 790 795 800
Thr Met Gly Gly Cys Leu Ile Ser Pro Ser Ala Glu Ile Gln Met Lys
805 810 815
Asn Thr Ser Ser Arg Ala Pro Leu Ser His Ser Asn Pro Ile Leu Gly
820 825 830
Leu Phe Gly Ala Gly Glu Val Thr Gly Gly Val His Gly Gly Asn Arg
835 840 845
Leu Gly Gly Asn Ser Leu Leu Glu Cys Val Val Phe Gly Arg Ile Ala
850 855 860
Gly Asp Arg Ala Ser Thr Ile Leu Gln Arg Lys Ser Ser Ala Leu Ser
865 870 875 880
Phe Lys Val Trp Thr Thr Val Val Leu Arg Glu Val Arg Glu Gly Gly
885 890 895
Val Tyr Gly Ala Gly Ser Arg Val Leu Arg Phe Asn Leu Pro Gly Ala
900 905 910
Leu Gln Arg Ser Gly Leu Ser Leu Gly Gln Phe Ile Ala Ile Arg Gly
915 920 925
Asp Trp Asp Gly Gln Gln Leu Ile Gly Tyr Tyr Ser Pro Ile Thr Leu
930 935 940
Pro Asp Asp Leu Gly Met Ile Asp Ile Leu Ala Arg Ser Asp Lys Gly
945 950 955 960
Thr Leu Arg Glu Trp Ile Ser Ala Leu Glu Pro Gly Asp Ala Val Glu
965 970 975
Met Lys Ala Cys Gly Gly Leu Val Ile Glu Arg Arg Leu Ser Asp Lys
980 985 990
His Phe Val Phe Met Gly His Ile Ile Asn Lys Leu Cys Leu Ile Ala
995 1000 1005
Gly Gly Thr Gly Val Ala Pro Met Leu Gln Ile Ile Lys Ala Ala Phe
1010 1015 1020
Met Lys Pro Phe Ile Asp Thr Leu Glu Ser Val His Leu Ile Tyr Ala
1025 1030 1035 1040
Ala Glu Asp Val Thr Glu Leu Thr Tyr Arg Glu Val Leu Glu Glu Arg
1045 1050 1055
Arg Arg Glu Ser Arg Gly Lys Phe Lys Lys Thr Phe Val Leu Asn Arg
1060 1065 1070
Pro Pro Pro Leu Trp Thr Asp Gly Val Gly Phe Ile Asp Arg Gly Ile
1075 1080 1085
Leu Thr Asn His Val Gln Pro Pro Ser Asp Asn Leu Leu Val Ala Ile
1090 1095 1100
Cys Gly Pro Pro Val Met Gln Arg Ile Val Lys Ala Thr Leu Lys Thr
1105 1110 1115 1120
Leu Gly Tyr Asn Met Asn Leu Val Arg Thr Val Asp Glu Thr Glu Pro
1125 1130 1135
Ser Gly
<210> 9
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..20
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 9
cattatatcg aggaaagccc 20
<210> 10
<211> 75
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..75
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 10
aaagcaaagg aaggagagaa cagaggagta cttgtacgtt cgatgggcaa agaaagagac 60
acaaaactac gtggg 75
<210> 11
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..20
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 11
ttgcccatcg aacgtacaag 20
<210> 12
<211> 23
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..23
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 12
tgctaagatt tgtgttcgtt tgg 23
<210> 13
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..20
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 13
cggatcgatg tacacaaccg 20
<210> 14
<211> 23
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..23
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 14
caacaggagg cggatggata tac 23
<210> 15
<211> 79
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..79
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 15
acgctttccg gcatcttcca gaccacagta tatccatccg cctcctgttg tcgtacgctg 60
caggtcgacg aattctacc 79
<210> 16
<211> 75
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..75
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 16
gcggaatatt ggcggaacgg acacacgtgg atacaaacct ggacaacgtt taggccacta 60
gtggatctga tatcg 75
<210> 17
<211> 22
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..22
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 17
aacgttgtcc aggtttgtat cc 22
<210> 18
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..20
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 18
aggtacaaca agcacgaccg 20
<210> 19
<211> 23
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..23
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 19
aaataaccac aaacatcctt ccc 23
<210> 20
<211> 23
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..23
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 20
gtgcactaca tcttgtgggt gtc 23
<210> 21
<211> 22
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..22
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 21
gaaaccttcg aatccagcca gc 22
<210> 22
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..20
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 22
aatcgcaact cggatttggg 20
<210> 23
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..20
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 23
aaagccaaag ttcgcgttcc 20
<210> 24
<211> 24
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..24
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 24
acttagtatg gtctgttgga aagg 24
<210> 25
<211> 25
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..25
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 25
cggcattatt gtgtatggct caata 25
<210> 26
<211> 75
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..75
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 26
gaacttcgac ctgttgcaat acttcgggtt cggcacaaac gtgtacggat agggtttcaa 60
agatccatac ttctc 75
<210> 27
<211> 79
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..79
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 27
atccgtacac gtttgtgccg aacccgaagt attgcaacag gtcgaagttc tcgtacgctg 60
caggtcgacg aattctacc 79
<210> 28
<211> 74
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..74
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 28
aggtacaaca agcacgaccg agcatatggg aaggatgttt gtggttattt aggccactag 60
tggatctgat atcg 74
<210> 29
<211> 23
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..23
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 29
aaataaccac aaacatcctt ccc 23
<210> 30
<211> 23
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..23
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 30
gtgcactaca tcttgtgggt gtc 23
<210> 31
<211> 2032
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..2032
<223> /organism="Artificial Sequence"
/note="Artificial"
/mol_type="unassigned DNA"
<400> 31
aaataaccac aaacatcctt cccatatgct cggtcgtgct tgttgtacct gtgcttagtc 60
aaaaaattag ccttttaatt ctgctgtaac ccgtacatgc ccaaaatagg gggcgggtta 120
cacagaatat ataacatcgt aggtgtctgg gtgaacagtt tattcctggc atccactaaa 180
tataatggag cccgcttttt aagctggcat ccagaaaaaa aaagaatccc agcaccaaaa 240
tattgttttc ttcaccaacc atcagttcat aggtccattc tcttagcgca actacagaga 300
acaggggcac aaacaggcaa aaaacgggca caacctcaat ggagtgatgc aacctgcctg 360
gagtaaatga tgacacaagg caattgaccc acgcatgtat ctatctcatt ttcttacacc 420
ttctattacc ttctgctctc tctgatttgg aaaaagctga aaaaaaaggt tgaaaccagt 480
tccctgaaat tattccccta cttgactaat aagtatataa agacggtagg tattgattgt 540
aattctgtaa atctatttct taaacttctt aaattctact tttatagtta gtcttttttt 600
tagttttaaa acaccaagaa cttagtttcg aataaacaca cataaacaaa caaaatggtt 660
aaggttgcca tcttaggtgc ttctggtggt gtcggtcaac cattatctct attattgaaa 720
ttgtctccat acgtttctga attggctttg tacgatatca gagctgctga aggtattggt 780
aaggatttgt cccacatcaa caccaactcc tcttgtgttg gttacgacaa ggattccatc 840
gaaaacactt tgtccaatgc tcaagttgtc ttgattccag ctggtgttcc aagaaagcca 900
ggtttgacca gagatgattt gttcaagatg aacgctggta tcgttaagtc tttggttact 960
gctgtcggta aatttgcccc aaacgctcgt atcttagtca tctccaaccc tgttaactct 1020
ttggttccaa ttgccgttga aactttgaag aagatgggta agttcaagcc aggtaacgtt 1080
atgggtgtca ccaacttgga tttggtcaga gctgaaactt tcttggttga ctacttgatg 1140
ttgaagaacc caaagatcgg tcaagaacaa gacaagacca ccatgcacag aaaggtcacc 1200
gtcatcggtg gtcactctgg tgaaaccatc attccaatca tcactgacaa atccttggtt 1260
ttccaattgg acaagcaata cgaacatttc atccacagag tccaattcgg tggtgacgaa 1320
attgtcaagg ccaagcaagg tgccggttct gctaccttgt ccatggcttt cgctggtgcc 1380
aaatttgctg aagaagtctt acgttctttc cacaacgaaa agccagaaac tgaatctttg 1440
tctgctttcg tctacttgcc aggtttgaag aacggtaaga aggctcaaca attagtcggt 1500
gacaactcca ttgaatactt ctctttgcca attgttttga gaaacggttc cgttgtttcc 1560
attgacactt ctgttttgga aaaattgtct ccaagagaag aacaattggt caacactgct 1620
gtcaaggaat tgagaaagaa cattgaaaag ggtaagtctt tcatcttgga cagttaaagt 1680
ctgaagaatg aatgatttga tgatttcttt ttccctccat ttttcttact gaatatatca 1740
atgatataga cttgtatagt ttattatttc aaattaagta gctatatata gtcaagataa 1800
cgtttgtttg acacgattac attattcgtc gacatctttt ttcagcctgt cgtggtagca 1860
atttgaggag tattattaat tgaataggtt cattttgcgc tcgcataaac agttttcgtc 1920
agggacagta tgttggaatg agtggtaatt aatggtgaca tgacatgtta tagcaatacc 1980
tcgaaacctt cgaatccagc cagcatgtcg acacccacaa gatgtagtgc ac 2032
<210> 32
<211> 78
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..78
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 32
gaaaccttcg aatccagcca gcatgtcgac acccacaaga tgtagtgcac acttttttta 60
gaatgacctg ttcccgac 78
<210> 33
<211> 24
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..24
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 33
cacaagctta ttcttccaaa aatc 24
<210> 34
<211> 3534
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..3534
<223> /organism="Artificial Sequence"
/note="Artificial"
/mol_type="unassigned DNA"
<400> 34
aaaggaggtg cacgcattat ggagaccact acgatacgat agctgcgttg ttgttgaagg 60
ggtttcttaa ggttgttttc gttgaaggta aatattggtc gtttttgtgc agcatattgt 120
cctctagatg caaactctgc aggtccattt gcagtaaagt gagttgcctc tcgaagaatc 180
attaatttcg tataaccgtc actattaaag tcagaaaata aattctgtcg tagacaatgt 240
taccataatg ttcttgtcca ttttgcatac actttaaata ttcatttgat ttctcagggt 300
tcatgatcat aataaattgc gcattcgcaa ggcggtagta ttataatggg gtccatcatt 360
ctgtagcaag aagttacagt acgctgttca agcgttaaac aagataagta atctcgaatg 420
aaacattcat atttcgcatg agccaacata cagttgctga gtaatcttca ttgcgcttat 480
ttatcggcat tgagattgta aaggaagtaa aacgcatttt tgcagatctg ttctcttatg 540
tatttttaat cgtccttgta tggaagtatc aaaggggacg ttcttcacct ccttggaagg 600
atcccttccc ttttacagtg cttcggaaaa gcacagcgtt gtccaaggga acaatttttc 660
ttcaagttaa tgcataagaa atatcttttt ttatgtttag ctaagtaaaa gcagcttgga 720
gtaaaaaaaa aaatgagtaa atttctcgat ggattagttt ctcacaggta acataacaaa 780
aaccaagaaa agcccgcttc tgaaaactac agttgacttg tatgctaaag ggccagacta 840
atgggaggag aaaaagaaac gaatgtatat gctcatttac actctatatc accatatgga 900
ggataagttg ggctgagctt ctgatccaat ttattctatc cattagttgc tgatatgtcc 960
caccagccaa cacttgatag tatctactcg ccattcactt ccagcagcgc cagtagggtt 1020
gttgagctta gtaaaaatgt gcgcaccaca agcctacatg actccacgtc acatgaaacc 1080
acaccgtggg gccttgttgc gctaggaata ggatatgcga cgaagacgct tctgcttagt 1140
aaccacacca cattttcagg gggtcgatct gcttgcttcc tttactgtca cgagcggccc 1200
ataatcgcgc ttttttttta aaaggcgcga gacagcaaac aggaagctcg ggtttcaacc 1260
ttcggagtgg tcgcagatct ggagactgga tctttacaat acagtaaggc aagccaccat 1320
ctgcttctta ggtgcatgcg acggtatcca cgtgcagaac aacatagtct gaagaagggg 1380
gggaggagca tgttcattct ctgtagcagt aagagcttgg tgataatgac caaaactgga 1440
gtctcgaaat catataaata gacaatatat tttcacacaa tgagatttgt agtacagttc 1500
tattctctct cttgcataaa taagaaattc atcaagaact tggtttgata tttcaccaac 1560
acacacaaaa aacagtactt cactaaattt acacacaaaa caaaatgtcc tctgcttctg 1620
ctgctttgca aaaattcaga gctgaaagag ataccttcgg tgacttgcaa gttccagctg 1680
accgttactg gggtgctcaa actcaaagat ctttgcaaaa ctttgacatt ggtggtccaa 1740
ctgaaagaat gccagaacca ttaatcagag ctttcggtgt tttgaagaag gctgctgcca 1800
ccgtcaacat gacctacggt ttggacccaa aggttggtga agccatccaa aaggctgctg 1860
acgaagttat cgatggttct ttgattgacc atttcccatt ggttgtctgg caaaccggtt 1920
ctggtactca aaccaagatg aacgtcaatg aagtcatctc caacagagcc attgaattgt 1980
tgggtggtga attaggttcc aaggctccag tccacccaaa cgatcatgtc aacatgtctc 2040
aatcttccaa cgacactttc ccaactgcca tgcacgttgc tgccgttgtt gaaattcacg 2100
gtagattgat tccagctttg accactttga gagatgcttt gcaagccaaa tctgctgaat 2160
tcgaacacat catcaagatt ggtagaaccc acttgcaaga tgctacccca ttgactttag 2220
gtcaagaatt ctccggttac actcaacaat tgacctacgg tattgctcgt gttcaaggta 2280
ctttggaaag attatacaac ttggctcaag gtggtactgc tgtcggtact ggtttgaaca 2340
ccagaaaggg tttcgatgcc aaggttgctg aagccattgc ttccatcact ggtttaccat 2400
tcaagaccgc tccaaacaaa ttcgaagctt tggctgctca cgacgctttg gttgaagctc 2460
acggtgcttt gaacaccgtt gcttgttctt tgatgaagat tgccaacgat atccgttact 2520
tgggttctgg tccaagatgt ggtttaggtg aattgtctct accagaaaac gaaccaggtt 2580
cttccatcat gccaggtaag gtcaacccaa ctcaatgtga agctatgacc atggtttgtg 2640
ctcaagtcat gggtaacaac actgccatct ctgttgctgg ttccaacggt caattcgaat 2700
tgaatgtctt taaaccagtc atgatcaaga acttgatcca atccatcaga ttaatctctg 2760
acgcttccat ctctttcacc aagaactgtg ttgtcggtat tgaagctaac gaaaagaaga 2820
tctcctccat catgaacgaa tctttgatgt tggtcactgc tttgaaccct cacattggtt 2880
acgacaaggc tgccaagtgt gccaagaagg ctcacaagga aggtaccact ttgaaagaag 2940
ctgctctatc tttgggttac ttgacctctg aagaattcga ccaatgggtt agacctgagg 3000
acatgatttc tgccaaggat taaggcccgg gcataaagca atcttgatga ggataatgat 3060
ttttttttga atatacataa atactaccgt ttttctgcta gattttgtga agacgtaaat 3120
aagtacatat tactttttaa gccaagacaa gattaagcat taactttacc cttttctctt 3180
ctaagtttca atactagtta tcactgttta aaagttatgg cgagaacgtc ggcggttaaa 3240
atatattacc ctgaacgtgg tgaattgaag ttctaggatg gtttaaagat ttttcctttt 3300
tgggaaataa gtaaacaata tattgctgcc tttgcaaaac gcacataccc acaatatgtg 3360
actattggca aagaacgcat tatcctttga agaggtggat actgatacta agagagtctc 3420
tattccggct ccacttttag tccagagatt acttgtcttc ttacgtatca gaacaagaaa 3480
gcatttccaa agtaattgca tttgcccttg agcagtatat atatactaag aagg 3534
<210> 35
<211> 2578
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..2578
<223> /organism="Artificial Sequence"
/note="Artificial"
/mol_type="unassigned DNA"
<400> 35
ttccaaagta attgcatttg cccttgagca gtatatatat actaagaagg tcgacctcga 60
gtaccgttcg tataatgtat gctatacgaa gttatattta aatcagtata gcgaccagca 120
ttcacatacg attgacgcat gatattactt tctgcgcact taacttcgca tctgggcaga 180
tgatgtcgag gcgaaaaaaa atataaatca cgctaacatt tgattaaaat agaacaacta 240
caatataaaa aaactataca aatgacaagt tcttgaaaac aagaatcttt ttattgtcag 300
tactgattag aaaaactcat cgagcatcaa atgaaactgc aatttattca tatcaggatt 360
atcaatacca tatttttgaa aaagccgttt ctgtaatgaa ggagaaaact caccgaggca 420
gttccatagg atggcaagat cctggtatcg gtctgcgatt ccgactcgtc caacatcaat 480
acaacctatt aatttcccct cgtcaaaaat aaggttatca agtgagaaat caccatgagt 540
gacgactgaa tccggtgaga atggcaaaag cttatgcatt tctttccaga cttgttcaac 600
aggccagcca ttacgctcgt catcaaaatc actcgcatca accaaaccgt tattcattcg 660
tgattgcgcc tgagcgagac gaaatacgcg atcgctgtta aaaggacaat tacaaacagg 720
aatcgaatgc aaccggcgca ggaacactgc cagcgcatca acaatatttt cacctgaatc 780
aggatattct tctaatacct ggaatgctgt tttgccgggg atcgcagtgg tgagtaacca 840
tgcatcatca ggagtacgga taaaatgctt gatggtcgga agaggcataa attccgtcag 900
ccagtttagt ctgaccatct catctgtaac atcattggca acgctacctt tgccatgttt 960
cagaaacaac tctggcgcat cgggcttccc atacaatcga tagattgtcg cacctgattg 1020
cccgacatta tcgcgagccc atttataccc atataaatca gcatccatgt tggaatttaa 1080
tcgcggcctc gaaacgtgag tcttttcctt acccatggtt gtttatgttc ggatgtgatg 1140
tgagaactgt atcctagcaa gattttaaaa ggaagtatat gaaagaagaa cctcagtggc 1200
aaatcctaac cttttatatt tctctacagg ggcgcggcgt ggggacaatt caacgcgtct 1260
gtgaggggag cgtttccctg ctcgcaggtc tgcagcgagg agccgtaatt tttgcttcgc 1320
gccgtgcggc catcaaaatg tatggatgca aatgattata catggggatg tatgggctaa 1380
atgtacgggc gacagtcaca tcatgcccct gagctgcgca cgtcaagact gtcaaggagg 1440
gtattctggg cctccatgtc atttaaatct agtacggatt agaagccgcc gagcgggtga 1500
cagccctccg aaggaagact ctcctccgtg cgtcctcgtc ttcaccggtc gcgttcctga 1560
aacgcagatg tgcctcgcgc cgcactgctc cgaacaataa agattctaca atactagctt 1620
ttatggttat gaagaggaaa aattggcagt aacctggccc cacaaacctt caaatgaacg 1680
aatcaaatta acaaccatag gatgataatg cgattagttt tttagcctta tttctggggt 1740
aattaatcag cgaagcgatg atttttgatc tattaacaga tatataaatg caaaaactgc 1800
ataaccactt taactaatac tttcaacatt ttcggtttgt attacttctt attcaaatgt 1860
aataaaagta tcaacaaaaa attgttaata tacctctata ctttaacgtc aaggagaaaa 1920
aaccccggat tctagaacta gtggatcccc cgggctgcag gaattcgata tcaagcttat 1980
cgataccgtc gaggggcaga gccgatcctg tacactttac ttaaaaccat tatctgagtg 2040
ttaaatgtcc aatttactga ccgtacacca aaatttgcct gcattaccgg tcgatgcaac 2100
gagtgatgag gttcgcaaga acctgatgga catgttcagg gatcgccagg cgttttctga 2160
gcatacctgg aaaatgcttc tgtccgtttg ccggtcgtgg gcggcatggt gcaagttgaa 2220
taaccggaaa tggtttcccg cagaacctga agatgttcgc gattatcttc tatatcttca 2280
ggcgcgcggt ctggcagtaa aaactatcca gcaacatttg ggccagctaa acatgcttca 2340
tcgtcggtcc gggctgccac gaccaagtga cagcaatgct gtttcactgg ttatgcggcg 2400
gatccgaaaa gaaaacgttg atgccggtga acgtgcaaaa caggctctag cgttcgaacg 2460
cactgatttc gaccaggttc gttcactcat ggaaaatagc gatcgctgcc aggatatacg 2520
taatctggca tttctgggga ttgcttataa caccctgtta cgtatagccg aaattgcc 2578
<210> 36
<211> 1263
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..1263
<223> /organism="Artificial Sequence"
/note="Artificial"
/mol_type="unassigned DNA"
<400> 36
cgttcactca tggaaaatag cgatcgctgc caggatatac gtaatctggc atttctgggg 60
attgcttata acaccctgtt acgtatagcc gaaattgcca ggatcagggt taaagatatc 120
tcacgtactg acggtgggag aatgttaatc catattggca gaacgaaaac gctggttagc 180
accgcaggtg tagagaaggc acttagcctg ggggtaacta aactggtcga gcgatggatt 240
tccgtctctg gtgtagctga tgatccgaat aactacctgt tttgccgggt cagaaaaaat 300
ggtgttgccg cgccatctgc caccagccag ctatcaactc gcgccctgga agggattttt 360
gaagcaactc atcgattgat ttacggcgct aaggatgact ctggtcagag atacctggcc 420
tggtctggac acagtgcccg tgtcggagcc gcgcgagata tggcccgcgc tggagtttca 480
ataccggaga tcatgcaagc tggtggctgg accaatgtaa atattgtcat gaactatatc 540
cgtaccctgg atagtgaaac aggggcaatg gtgcgcctgc tggaagatgg cgattagcca 600
ttaacgcgta aatgattgct ataattattt gatatttatg gtgacatatg agaaaggatt 660
tcaacatcga cggaaaatat gtagtgctgt ctgtaagcac taatattcag tcgccagccg 720
tcattgtcac tgtaaagctg agcgatagaa tgcctgatat tgactcaata tccgttgcgt 780
ttcctgtcaa aagtatgcgt agtgctgaac atttcgtgat gaatgccacc gaggaagaag 840
cacggcgcgg ttttgcttaa agtgatgtct gagtttggcg aactcttggg taaggttgga 900
attgtcgacc tcgagtcatg taattagtta tgtcacgctt acattcacgc cctcccccca 960
catccgctct aaccgaaaag gaaggagtta gacaacctga agtctaggtc cctatttatt 1020
tttttatagt tatgttagta ttaagaacgt tatttatatt tcaaattttt cttttttttc 1080
tgtacagacg cgtgtacgca tgtaacatta tactgaaaac cttgcttgag aaggttttgg 1140
gacgctcgaa ggctttaatt tgcggccggt acataacttc gtataatgta tgctatacga 1200
acggtaggat ccggatggga cgtcagcact gtacttgttt ttgcgactag attgtaaatc 1260
att 1263
<210> 37
<211> 631
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..631
<223> /organism="Artificial Sequence"
/note="Artificial"
/mol_type="unassigned DNA"
<400> 37
gatgggacgt cagcactgta cttgtttttg cgactagatt gtaaatcatt ctttatttaa 60
tctctttctt taactactgc ttaaagtata atttggtccg tagtttaata actatactaa 120
gcgtaacaat gcatactgac attataagcc tgaacattac gagtttaagt tgtatgtagg 180
cgttctgtaa gaggttactg cgtaaattat caacgaatgc attggtgtat ttgcgaaagc 240
tacttctttt aacaagtatt tacataagaa taatggtgat ctgctcaact gatttggtga 300
taactctaac ttttttagca acaatttaaa agataattcg aacatatata acagtaggaa 360
gaatttgtgt acgtcaaatt aagataattt agcattacca aagttattaa cctaaacata 420
aaatatatat gagacacatg tggaaatcgt atgaaacaac tgttatgaaa ctgacaagaa 480
tgaatatata gagtaagctc cgcttgtaaa gaggaatcac ttaagtgtat aaatgtctcg 540
acgattactt tagatccaag attgatgatt gatattactc tgtaatactt aagctctttt 600
aatagctcac tgttgtatta cgggctcgag t 631
<210> 38
<211> 1334
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..1334
<223> /organism="Artificial Sequence"
/note="Artificial"
/mol_type="unassigned DNA"
<400> 38
tcgtacgctg caggtcgacg aattctaccg ttcgtataat gtatgctata cgaagttata 60
gatctgttta gcttgcctcg tccccgccgg gtcacccggc cagcgacatg gaggcccaga 120
ataccctcct tgacagtctt gacgtgcgca gctcaggggc atgatgtgac tgtcgcccgt 180
acatttagcc catacatccc catgtataat catttgcatc catacatttt gatggccgca 240
cggcgcgaag caaaaattac ggctcctcgc tgcagacctg cgagcaggga aacgctcccc 300
tcacagacgc gttgaattgt ccccacgccg cgcccctgta gagaaatata aaaggttagg 360
atttgccact gaggttcttc tttcatatac ttccttttaa aatcttgcta ggatacagtt 420
ctcacatcac atccgaacat aaacaaccat gggtaccact cttgacgaca cggcttaccg 480
gtaccgcacc agtgtcccgg gggacgccga ggccatcgag gcactggatg ggtccttcac 540
caccgacacc gtcttccgcg tcaccgccac cggggacggc ttcaccctgc gggaggtgcc 600
ggtggacccg cccctgacca aggtgttccc cgacgacgaa tcggacgacg aatcggacga 660
cggggaggac ggcgacccgg actcccggac gttcgtcgcg tacggggacg acggcgacct 720
ggcgggcttc gtggtcgtct cgtactccgg ctggaaccgc cggctgaccg tcgaggacat 780
cgaggtcgcc ccggagcacc gggggcacgg ggtcgggcgc gcgttgatgg ggctcgcgac 840
ggagttcgcc cgcgagcggg gcgccgggca cctctggctg gaggtcacca acgtcaacgc 900
accggcgatc cacgcgtacc ggcggatggg gttcaccctc tgcggcctgg acaccgccct 960
gtacgacggc accgcctcgg acggcgagca ggcgctctac atgagcatgc cctgccccta 1020
atcagtactg acaataaaaa gattcttgtt ttcaagaact tgtcatttgt atagtttttt 1080
tatattgtag ttgttctatt ttaatcaaat gttagcgtga tttatatttt ttttcgcctc 1140
gacatcatct gcccagatgc gaagttaagt gcgcagaaag taatatcatg cgtcaatcgt 1200
atgtgaatgc tggtcgctat actgctgtcg attcgatact aacgccgcca tccagtgtcg 1260
aaaacgagct cataacttcg tataatgtat gctatacgaa cggtagaatt cgatatcaga 1320
tccactagtg gcct 1334
<210> 39
<211> 340
<212> PRT
<213> Artificial Sequence
<220>
<223> MDH3 lacking 3C-terminal peroxisomal targeting sequence
<400> 39
Met Val Lys Val Ala Ile Leu Gly Ala Ser Gly Gly Val Gly Gln Pro
1 5 10 15
Leu Ser Leu Leu Leu Lys Leu Ser Pro Tyr Val Ser Glu Leu Ala Leu
20 25 30
Tyr Asp Ile Arg Ala Ala Glu Gly Ile Gly Lys Asp Leu Ser His Ile
35 40 45
Asn Thr Asn Ser Ser Cys Val Gly Tyr Asp Lys Asp Ser Ile Glu Asn
50 55 60
Thr Leu Ser Asn Ala Gln Val Val Leu Ile Pro Ala Gly Val Pro Arg
65 70 75 80
Lys Pro Gly Leu Thr Arg Asp Asp Leu Phe Lys Met Asn Ala Gly Ile
85 90 95
Val Lys Ser Leu Val Thr Ala Val Gly Lys Phe Ala Pro Asn Ala Arg
100 105 110
Ile Leu Val Ile Ser Asn Pro Val Asn Ser Leu Val Pro Ile Ala Val
115 120 125
Glu Thr Leu Lys Lys Met Gly Lys Phe Lys Pro Gly Asn Val Met Gly
130 135 140
Val Thr Asn Leu Asp Leu Val Arg Ala Glu Thr Phe Leu Val Asp Tyr
145 150 155 160
Leu Met Leu Lys Asn Pro Lys Ile Gly Gln Glu Gln Asp Lys Thr Thr
165 170 175
Met His Arg Lys Val Thr Val Ile Gly Gly His Ser Gly Glu Thr Ile
180 185 190
Ile Pro Ile Ile Thr Asp Lys Ser Leu Val Phe Gln Leu Asp Lys Gln
195 200 205
Tyr Glu His Phe Ile His Arg Val Gln Phe Gly Gly Asp Glu Ile Val
210 215 220
Lys Ala Lys Gln Gly Ala Gly Ser Ala Thr Leu Ser Met Ala Phe Ala
225 230 235 240
Gly Ala Lys Phe Ala Glu Glu Val Leu Arg Ser Phe His Asn Glu Lys
245 250 255
Pro Glu Thr Glu Ser Leu Ser Ala Phe Val Tyr Leu Pro Gly Leu Lys
260 265 270
Asn Gly Lys Lys Ala Gln Gln Leu Val Gly Asp Asn Ser Ile Glu Tyr
275 280 285
Phe Ser Leu Pro Ile Val Leu Arg Asn Gly Ser Val Val Ser Ile Asp
290 295 300
Thr Ser Val Leu Glu Lys Leu Ser Pro Arg Glu Glu Gln Leu Val Asn
305 310 315 320
Thr Ala Val Lys Glu Leu Arg Lys Asn Ile Glu Lys Gly Lys Ser Phe
325 330 335
Ile Leu Asp Ser
340
<210> 40
<211> 75
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..75
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 40
agaaagcctg tatgcgaagc cacaatcctt tccaacagac catactaagt acaggtgatt 60
gtatgtgggc ttatg 75
<210> 41
<211> 29
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..29
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 41
acattattgt aaaaacggag tagaaaggg 29
<210> 42
<211> 2927
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..2927
<223> /organism="Artificial Sequence"
/note="Artificial"
/mol_type="unassigned DNA"
<400> 42
atccgtacac gtttgtgccg aacccgaagt attgcaacag gtcgaagttc gtgccaaagg 60
gggggcaggg acagggatac gacaagggct ggggaaaaaa aaaaagatag atacgattgg 120
ccgggtaagc ctggggaaat gtagcaagtg cgggtaagtt aaaaggtaac cacgtgactc 180
cggaagagtc acgtggttac ggactttttt ctctagatct cagcttttta tcggtcttac 240
cctgccctcc tgccccctgc cccttccctt tgccccaaaa agaaaggaaa tctgttggat 300
ttcgctcagg ccatcccttt cgttaatatc ggttatcgct ttacacactg cacatccttc 360
tgtccaaaag gaatccagaa gtttagcttt tccttccttt cccacagaca ttagcctagg 420
ccctctctca tcatttgcat gcctcagcca atgtaccaag aataacgcaa cgaggttggg 480
aaattttaac ccaacaatcg atgcagatgt gacaagagat tagacacgtt ccagatacca 540
gattacacag cttgtgctag cagagtgaca tatggtggtg ttgtgtctcg tttagtacct 600
gtaatcgaga gtgttcaaat cagtcgattt gaacaccctt actgccactg aatattgatt 660
gaataccgtt tattgaaggt tttatgagtg atcttctttc ggtccaggac aatttgttga 720
gctttttcta tgtagagttc cgtccctttt tttttttttt ttgctttctc gcacttacta 780
gcactatttt tttttcacac actaaaacac tttattttaa tctatatata tatatatata 840
tatatgtagg aatggaatca cagacatttg atactcatcc tcatccttat taattcttgt 900
tttaatttgt ttgacttagc caaaccacca atctcaaccc atcgtatttc aggtattgtg 960
tgtctagtgt gtctctggta tacggaaata agtgccagaa gtaaggaaga aacaaagaac 1020
aagtgtctga atactactag cctctctttt cataatgagt gaaggccccg tcaaattcga 1080
aaaaaatacc gtcatatctg tctttggtgc gtcaggtgat ctggcaaaga agaagacttt 1140
tcccgcctta tttgggcttt tcagagaagg ttaccttgat ccatctacca agatcttcgg 1200
ttatgcccgg tccaaattgt ccatggagga ggacctgaag tcccgtgtcc taccccactt 1260
gaaaaaacct cacggtgaag ccgatgactc taaggtcgaa cagttcttca agatggtcag 1320
ctacatttcg ggaaattacg acacagatga aggcttcgac gaattaagaa cgcagatcga 1380
gaaattcgag aaaagtgcca acgtcgatgt cccacaccgt ctcttctatc tggccttgcc 1440
gccaagcgtt tttttgacgg tggccaagca gatcaagagt cgtgtgtacg cagagaatgg 1500
catcacccgt gtaatcgtag agaaaccttt cggccacgac ctggcctctg ccagggagct 1560
gcaaaaaaac ctggggcccc tctttaaaga agaagagttg tacagaattg accattactt 1620
gggtaaagag ttggtcaaga atcttttagt cttgaggttc ggtaaccagt ttttgaatgc 1680
ctcgtggaat agagacaaca ttcaaagcgt tcagatttcg tttaaagaga ggttcggcac 1740
cgaaggccgt ggcggctatt tcgactctat aggcataatc agagacgtga tgcagaacca 1800
tctgttacaa atcatgactc tcttgactat ggaaagaccg gtgtcttttg acccggaatc 1860
tattcgtgac gaaaaggtta aggttctaaa ggccgtggcc cccatcgaca cggacgacgt 1920
cctcttgggc cagtacggta aatctgagga cgggtctaag cccgcctacg tggatgatga 1980
cactgtagac aaggactcta aatgtgtcac ttttgcagca atgactttca acatcgaaaa 2040
cgagcgttgg gagggcgtcc ccatcatgat gcgtgccggt aaggctttga atgagtccaa 2100
ggtggagatc agactgcagt acaaagcggt cgcatcgggt gtcttcaaag acattccaaa 2160
taacgaactg gtcatcagag tgcagcccga tgccgctgtg tacctaaagt ttaatgctaa 2220
gacccctggt ctgtcaaatg ctacccaagt cacagatctg aatctaactt acgcaagcag 2280
gtaccaagac ttttggattc cagaggctta cgaggtgttg ataagagacg ccctactggg 2340
tgaccattcc aactttgtca gagatgacga attggatatc agttggggca tattcacccc 2400
attactgaag cacatagagc gtccggacgg tccaacaccg gaaatttacc cctacggatc 2460
aagaggtcca aagggattga aggaatatat gcaaaaacac aagtatgtta tgcccgaaaa 2520
gcacccttac gcttggcccg tgactaagcc agaagatacg aaggataatt aaaggagatt 2580
gataagactt ttctagttgc atatctttta tatttaaatc ttatctatta gttaattttt 2640
tgtaatttat ccttatatat agtctggtta ttctaaaata tcatttcagt atctaaaaat 2700
tcccctcttt tttcagttat atcttaacag gcgacagtcc aaatgttgat ttatcccagt 2760
ccgattcatc agggttgtga agcattttgt caatggtcga aatcacatca gtaatagtgc 2820
ctcttacttg cctcatagaa tttctttctc ttaacgtcac cgtttggtct tttcctcacc 2880
cgatcccgaa gaaagtccaa ctccagttcg attccctgct ttcgagt 2927
<210> 43
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..20
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 43
atccgtacac gtttgtgccg 20
<210> 44
<211> 22
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..22
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 44
actcgaaagc agggaatcga ac 22
<210> 45
<211> 79
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..79
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 45
acccgatccc gaagaaagtc caactccagt tcgattccct gctttcgagt tcgtacgctg 60
caggtcgacg aattctacc 79
<210> 46
<211> 1180
<212> PRT
<213> Saccharomyces cerevisiae
<400> 46
Met Ser Ser Ser Lys Ile Leu Ala Gly Leu Arg Asp Asn Phe Ser Leu
1 5 10 15
Leu Gly Glu Lys Asn Lys Ile Leu Val Ala Asn Arg Gly Glu Ile Pro
20 25 30
Ile Arg Ile Phe Arg Ser Ala His Glu Leu Ser Met Arg Thr Ile Ala
35 40 45
Ile Tyr Ser His Glu Asp Arg Leu Ser Met His Arg Leu Lys Ala Asp
50 55 60
Glu Ala Tyr Val Ile Gly Glu Glu Gly Gln Tyr Thr Pro Val Gly Ala
65 70 75 80
Tyr Leu Ala Met Asp Glu Ile Ile Glu Ile Ala Lys Lys His Lys Val
85 90 95
Asp Phe Ile His Pro Gly Tyr Gly Phe Leu Ser Glu Asn Ser Glu Phe
100 105 110
Ala Asp Lys Val Val Lys Ala Gly Ile Thr Trp Ile Gly Pro Pro Ala
115 120 125
Glu Val Ile Glu Ser Val Gly Asp Lys Val Ser Ala Arg His Leu Ala
130 135 140
Ala Arg Ala Asn Val Pro Thr Val Pro Gly Thr Pro Gly Pro Ile Glu
145 150 155 160
Thr Val Gln Glu Ala Leu Asp Phe Val Asn Glu Tyr Gly Tyr Pro Val
165 170 175
Ile Ile Lys Ala Ala Phe Gly Gly Gly Gly Arg Gly Met Arg Val Val
180 185 190
Arg Glu Gly Asp Asp Val Ala Asp Ala Phe Gln Arg Ala Thr Ser Glu
195 200 205
Ala Arg Thr Ala Phe Gly Asn Gly Thr Cys Phe Val Glu Arg Phe Leu
210 215 220
Asp Lys Pro Lys His Ile Glu Val Gln Leu Leu Ala Asp Asn His Gly
225 230 235 240
Asn Val Val His Leu Phe Glu Arg Asp Cys Ser Val Gln Arg Arg His
245 250 255
Gln Lys Val Val Glu Val Ala Pro Ala Lys Thr Leu Pro Arg Glu Val
260 265 270
Arg Asp Ala Ile Leu Thr Asp Ala Val Lys Leu Ala Lys Val Cys Gly
275 280 285
Tyr Arg Asn Ala Gly Thr Ala Glu Phe Leu Val Asp Asn Gln Asn Arg
290 295 300
His Tyr Phe Ile Glu Ile Asn Pro Arg Ile Gln Val Glu His Thr Ile
305 310 315 320
Thr Glu Glu Ile Thr Gly Ile Asp Ile Val Ser Ala Gln Ile Gln Ile
325 330 335
Ala Ala Gly Ala Thr Leu Thr Gln Leu Gly Leu Leu Gln Asp Lys Ile
340 345 350
Thr Thr Arg Gly Phe Ser Ile Gln Cys Arg Ile Thr Thr Glu Asp Pro
355 360 365
Ser Lys Asn Phe Gln Pro Asp Thr Gly Arg Leu Glu Val Tyr Arg Ser
370 375 380
Ala Gly Gly Asn Gly Val Arg Leu Asp Gly Gly Asn Ala Tyr Ala Gly
385 390 395 400
Ala Thr Ile Ser Pro His Tyr Asp Ser Met Leu Val Lys Cys Ser Cys
405 410 415
Ser Gly Ser Thr Tyr Glu Ile Val Arg Arg Lys Met Ile Arg Ala Leu
420 425 430
Ile Glu Phe Arg Ile Arg Gly Val Lys Thr Asn Ile Pro Phe Leu Leu
435 440 445
Thr Leu Leu Thr Asn Pro Val Phe Ile Glu Gly Thr Tyr Trp Thr Thr
450 455 460
Phe Ile Asp Asp Thr Pro Gln Leu Phe Gln Met Val Ser Ser Gln Asn
465 470 475 480
Arg Ala Gln Lys Leu Leu His Tyr Leu Ala Asp Leu Ala Val Asn Gly
485 490 495
Ser Ser Ile Lys Gly Gln Ile Gly Leu Pro Lys Leu Lys Ser Asn Pro
500 505 510
Ser Val Pro His Leu His Asp Ala Gln Gly Asn Val Ile Asn Val Thr
515 520 525
Lys Ser Ala Pro Pro Ser Gly Trp Arg Gln Val Leu Leu Glu Lys Gly
530 535 540
Pro Ser Glu Phe Ala Lys Gln Val Arg Gln Phe Asn Gly Thr Leu Leu
545 550 555 560
Met Asp Thr Thr Trp Arg Asp Ala His Gln Ser Leu Leu Ala Thr Arg
565 570 575
Val Arg Thr His Asp Leu Ala Thr Ile Ala Pro Thr Thr Ala His Ala
580 585 590
Leu Ala Gly Ala Phe Ala Leu Glu Cys Trp Gly Gly Ala Thr Phe Asp
595 600 605
Val Ala Met Arg Phe Leu His Glu Asp Pro Trp Glu Arg Leu Arg Lys
610 615 620
Leu Arg Ser Leu Val Pro Asn Ile Pro Phe Gln Met Leu Leu Arg Gly
625 630 635 640
Ala Asn Gly Val Ala Tyr Ser Ser Leu Pro Asp Asn Ala Ile Asp His
645 650 655
Phe Val Lys Gln Ala Lys Asp Asn Gly Val Asp Ile Phe Arg Val Phe
660 665 670
Asp Ala Leu Asn Asp Leu Glu Gln Leu Lys Val Gly Val Asn Ala Val
675 680 685
Lys Lys Ala Gly Gly Val Val Glu Ala Thr Val Cys Tyr Ser Gly Asp
690 695 700
Met Leu Gln Pro Gly Lys Lys Tyr Asn Leu Asp Tyr Tyr Leu Glu Val
705 710 715 720
Val Glu Lys Ile Val Gln Met Gly Thr His Ile Leu Gly Ile Lys Asp
725 730 735
Met Ala Gly Thr Met Lys Pro Ala Ala Ala Lys Leu Leu Ile Gly Ser
740 745 750
Leu Arg Thr Arg Tyr Pro Asp Leu Pro Ile His Val His Ser His Asp
755 760 765
Ser Ala Gly Thr Ala Val Ala Ser Met Thr Ala Cys Ala Leu Ala Gly
770 775 780
Ala Asp Val Val Asp Val Ala Ile Asn Ser Met Ser Gly Leu Thr Ser
785 790 795 800
Gln Pro Ser Ile Asn Ala Leu Leu Ala Ser Leu Glu Gly Asn Ile Asp
805 810 815
Thr Gly Ile Asn Val Glu His Val Arg Glu Leu Asp Ala Tyr Trp Ala
820 825 830
Glu Met Arg Leu Leu Tyr Ser Cys Phe Glu Ala Asp Leu Lys Gly Pro
835 840 845
Asp Pro Glu Val Tyr Gln His Glu Ile Pro Gly Gly Gln Leu Thr Asn
850 855 860
Leu Leu Phe Gln Ala Gln Gln Leu Gly Leu Gly Glu Gln Trp Ala Glu
865 870 875 880
Thr Lys Arg Ala Tyr Arg Glu Ala Asn Tyr Leu Leu Gly Asp Ile Val
885 890 895
Lys Val Thr Pro Thr Ser Lys Val Val Gly Asp Leu Ala Gln Phe Met
900 905 910
Val Ser Asn Lys Leu Thr Ser Asp Asp Ile Arg Arg Leu Ala Asn Ser
915 920 925
Leu Asp Phe Pro Asp Ser Val Met Asp Phe Phe Glu Gly Leu Ile Gly
930 935 940
Gln Pro Tyr Gly Gly Phe Pro Glu Pro Leu Arg Ser Asp Val Leu Arg
945 950 955 960
Asn Lys Arg Arg Lys Leu Thr Cys Arg Pro Gly Leu Glu Leu Glu Pro
965 970 975
Phe Asp Leu Glu Lys Ile Arg Glu Asp Leu Gln Asn Arg Phe Gly Asp
980 985 990
Ile Asp Glu Cys Asp Val Ala Ser Tyr Asn Met Tyr Pro Arg Val Tyr
995 1000 1005
Glu Asp Phe Gln Lys Ile Arg Glu Thr Tyr Gly Asp Leu Ser Val Leu
1010 1015 1020
Pro Thr Lys Asn Phe Leu Ala Pro Ala Glu Pro Asp Glu Glu Ile Glu
1025 1030 1035 1040
Val Thr Ile Glu Gln Gly Lys Thr Leu Ile Ile Lys Leu Gln Ala Val
1045 1050 1055
Gly Asp Leu Asn Lys Lys Thr Gly Gln Arg Glu Val Tyr Phe Glu Leu
1060 1065 1070
Asn Gly Glu Leu Arg Lys Ile Arg Val Ala Asp Lys Ser Gln Asn Ile
1075 1080 1085
Gln Ser Val Ala Lys Pro Lys Ala Asp Val His Asp Thr His Gln Ile
1090 1095 1100
Gly Ala Pro Met Ala Gly Val Ile Ile Glu Val Lys Val His Lys Gly
1105 1110 1115 1120
Ser Leu Val Lys Lys Gly Glu Ser Ile Ala Val Leu Ser Ala Met Lys
1125 1130 1135
Met Glu Met Val Val Ser Ser Pro Ala Asp Gly Gln Val Lys Asp Val
1140 1145 1150
Phe Ile Arg Asp Gly Glu Ser Val Asp Ala Ser Asp Leu Leu Val Val
1155 1160 1165
Leu Glu Glu Glu Thr Leu Pro Pro Ser Gln Lys Lys
1170 1175 1180
<210> 47
<211> 538
<212> PRT
<213> Artificial Sequence
<220>
<223> Artificial
<400> 47
Met Thr Asp Leu Asn Lys Leu Val Lys Glu Leu Asn Asp Leu Gly Leu
1 5 10 15
Thr Asp Val Lys Glu Ile Val Tyr Asn Pro Ser Tyr Glu Gln Leu Phe
20 25 30
Glu Glu Glu Thr Lys Pro Gly Leu Glu Gly Phe Asp Lys Gly Thr Leu
35 40 45
Thr Thr Leu Gly Ala Val Ala Val Asp Thr Gly Ile Phe Thr Gly Arg
50 55 60
Ser Pro Lys Asp Lys Tyr Ile Val Cys Asp Glu Thr Thr Lys Asp Thr
65 70 75 80
Val Trp Trp Asn Ser Glu Ala Ala Lys Asn Asp Asn Lys Pro Met Thr
85 90 95
Gln Glu Thr Trp Lys Ser Leu Arg Glu Leu Val Ala Lys Gln Leu Ser
100 105 110
Gly Lys Arg Leu Phe Val Val Asp Ala Phe Cys Gly Ala Ser Glu Lys
115 120 125
His Arg Ile Gly Val Arg Met Val Thr Glu Val Ala Trp Gln Ala His
130 135 140
Phe Val Lys Asn Met Phe Ile Arg Pro Thr Asp Glu Glu Leu Lys Asn
145 150 155 160
Phe Lys Ala Asp Phe Thr Val Leu Asn Gly Ala Lys Cys Thr Asn Pro
165 170 175
Asn Trp Lys Glu Gln Gly Leu Asn Ser Glu Asn Phe Val Ala Phe Asn
180 185 190
Ile Thr Glu Gly Ile Gln Leu Ile Gly Gly Thr Trp Tyr Gly Gly Glu
195 200 205
Met Lys Lys Gly Met Phe Ser Met Met Asn Tyr Phe Leu Pro Leu Lys
210 215 220
Gly Val Ala Ser Met His Cys Ser Ala Asn Val Gly Lys Asp Gly Asp
225 230 235 240
Val Ala Ile Phe Phe Gly Leu Ser Gly Thr Gly Lys Thr Thr Leu Ser
245 250 255
Thr Asp Pro Lys Arg Gln Leu Ile Gly Asp Asp Glu His Gly Trp Asp
260 265 270
Glu Ser Gly Val Phe Asn Phe Glu Gly Gly Cys Tyr Ala Lys Thr Ile
275 280 285
Asn Leu Ser Gln Glu Asn Glu Pro Asp Ile Tyr Gly Ala Ile Arg Arg
290 295 300
Asp Ala Leu Leu Glu Asn Val Val Val Arg Ala Asp Gly Ser Val Asp
305 310 315 320
Phe Asp Asp Gly Ser Lys Thr Glu Asn Thr Arg Val Ser Tyr Pro Ile
325 330 335
Tyr His Ile Asp Asn Ile Val Arg Pro Val Ser Lys Ala Gly His Ala
340 345 350
Thr Lys Val Ile Phe Leu Thr Ala Asp Ala Phe Gly Val Leu Pro Pro
355 360 365
Val Ser Lys Leu Thr Pro Glu Gln Thr Glu Tyr Tyr Phe Leu Ser Gly
370 375 380
Phe Thr Ala Lys Leu Ala Gly Thr Glu Arg Gly Val Thr Glu Pro Thr
385 390 395 400
Pro Thr Phe Ser Ala Cys Phe Gly Ala Ala Phe Leu Ser Leu His Pro
405 410 415
Ile Gln Tyr Ala Asp Val Leu Val Glu Arg Met Lys Ala Ser Gly Ala
420 425 430
Glu Ala Tyr Leu Val Asn Thr Gly Trp Asn Gly Thr Gly Lys Arg Ile
435 440 445
Ser Ile Lys Asp Thr Arg Gly Ile Ile Asp Ala Ile Leu Asp Gly Ser
450 455 460
Ile Glu Lys Ala Glu Met Gly Glu Leu Pro Ile Phe Asn Leu Ala Ile
465 470 475 480
Pro Lys Ala Leu Pro Gly Val Asp Pro Ala Ile Leu Asp Pro Arg Asp
485 490 495
Thr Tyr Ala Asp Lys Ala Gln Trp Gln Val Lys Ala Glu Asp Leu Ala
500 505 510
Asn Arg Phe Val Lys Asn Phe Val Lys Tyr Thr Ala Asn Pro Glu Ala
515 520 525
Ala Lys Leu Val Gly Ala Gly Pro Lys Ala
530 535
<210> 48
<211> 548
<212> PRT
<213> Escherichia coli
<400> 48
Met Ser Asn Lys Pro Phe Ile Tyr Gln Ala Pro Phe Pro Met Gly Lys
1 5 10 15
Asp Asn Thr Glu Tyr Tyr Leu Leu Thr Ser Asp Tyr Val Ser Val Ala
20 25 30
Asp Phe Asp Gly Glu Thr Ile Leu Lys Val Glu Pro Glu Ala Leu Thr
35 40 45
Leu Leu Ala Gln Gln Ala Phe His Asp Ala Ser Phe Met Leu Arg Pro
50 55 60
Ala His Gln Lys Gln Val Ala Ala Ile Leu His Asp Pro Glu Ala Ser
65 70 75 80
Glu Asn Asp Lys Tyr Val Ala Leu Gln Phe Leu Arg Asn Ser Glu Ile
85 90 95
Ala Ala Lys Gly Val Leu Pro Thr Cys Gln Asp Thr Gly Thr Ala Ile
100 105 110
Ile Val Gly Lys Lys Gly Gln Arg Val Trp Thr Gly Gly Gly Asp Glu
115 120 125
Glu Thr Leu Ser Lys Gly Val Tyr Asn Thr Tyr Ile Glu Asp Asn Leu
130 135 140
Arg Tyr Ser Gln Asn Ala Ala Leu Asp Met Tyr Lys Glu Val Asn Thr
145 150 155 160
Gly Thr Asn Leu Pro Ala Gln Ile Asp Leu Tyr Ala Val Asp Gly Asp
165 170 175
Glu Tyr Lys Phe Leu Cys Val Ala Lys Gly Gly Gly Ser Ala Asn Lys
180 185 190
Thr Tyr Leu Tyr Gln Glu Thr Lys Ala Leu Leu Thr Pro Gly Lys Leu
195 200 205
Lys Asn Phe Leu Val Glu Lys Met Arg Thr Leu Gly Thr Ala Ala Cys
210 215 220
Pro Pro Tyr His Ile Ala Phe Val Ile Gly Gly Thr Ser Ala Glu Thr
225 230 235 240
Asn Leu Lys Thr Val Lys Leu Ala Ser Ala His Tyr Tyr Asp Glu Leu
245 250 255
Pro Thr Glu Gly Asn Glu His Gly Gln Ala Phe Arg Asp Val Gln Leu
260 265 270
Glu Gln Glu Leu Leu Glu Glu Ala Gln Lys Leu Gly Leu Gly Ala Gln
275 280 285
Phe Gly Gly Lys Tyr Phe Ala His Asp Ile Arg Val Ile Arg Leu Pro
290 295 300
Arg His Gly Ala Ser Cys Pro Val Gly Met Gly Val Ser Cys Ser Ala
305 310 315 320
Asp Arg Asn Ile Lys Ala Lys Ile Asn Arg Glu Gly Ile Trp Ile Glu
325 330 335
Lys Leu Glu His Asn Pro Gly Gln Tyr Ile Pro Gln Glu Leu Arg Gln
340 345 350
Ala Gly Glu Gly Glu Ala Val Lys Val Asp Leu Asn Arg Pro Met Lys
355 360 365
Glu Ile Leu Ala Gln Leu Ser Gln Tyr Pro Val Ser Thr Arg Leu Ser
370 375 380
Leu Thr Gly Thr Ile Ile Val Gly Arg Asp Ile Ala His Ala Lys Leu
385 390 395 400
Lys Glu Leu Ile Asp Ala Gly Lys Glu Leu Pro Gln Tyr Ile Lys Asp
405 410 415
His Pro Ile Tyr Tyr Ala Gly Pro Ala Lys Thr Pro Ala Gly Tyr Pro
420 425 430
Ser Gly Ser Leu Gly Pro Thr Thr Ala Gly Arg Met Asp Ser Tyr Val
435 440 445
Asp Leu Leu Gln Ser His Gly Gly Ser Met Ile Met Leu Ala Lys Gly
450 455 460
Asn Arg Ser Gln Gln Val Thr Asp Ala Cys His Lys His Gly Gly Phe
465 470 475 480
Tyr Leu Gly Ser Ile Gly Gly Pro Ala Ala Val Leu Ala Gln Gln Ser
485 490 495
Ile Lys His Leu Glu Cys Val Ala Tyr Pro Glu Leu Gly Met Glu Ala
500 505 510
Ile Trp Lys Ile Glu Val Glu Asp Phe Pro Ala Phe Ile Leu Val Asp
515 520 525
Asp Lys Gly Asn Asp Phe Phe Gln Gln Ile Val Asn Lys Gln Cys Ala
530 535 540
Asn Cys Thr Lys
545
<210> 49
<211> 472
<212> PRT
<213> Artificial Sequence
<220>
<223> Artificial
<400> 49
Met Ser Ser Ala Ser Ala Ala Leu Gln Lys Phe Arg Ala Glu Arg Asp
1 5 10 15
Thr Phe Gly Asp Leu Gln Val Pro Ala Asp Arg Tyr Trp Gly Ala Gln
20 25 30
Thr Gln Arg Ser Leu Gln Asn Phe Asp Ile Gly Gly Pro Thr Glu Arg
35 40 45
Met Pro Glu Pro Leu Ile Arg Ala Phe Gly Val Leu Lys Lys Ala Ala
50 55 60
Ala Thr Val Asn Met Thr Tyr Gly Leu Asp Pro Lys Val Gly Glu Ala
65 70 75 80
Ile Gln Lys Ala Ala Asp Glu Val Ile Asp Gly Ser Leu Ile Asp His
85 90 95
Phe Pro Leu Val Val Trp Gln Thr Gly Ser Gly Thr Gln Thr Lys Met
100 105 110
Asn Val Asn Glu Val Ile Ser Asn Arg Ala Ile Glu Leu Leu Gly Gly
115 120 125
Glu Leu Gly Ser Lys Ala Pro Val His Pro Asn Asp His Val Asn Met
130 135 140
Ser Gln Ser Ser Asn Asp Thr Phe Pro Thr Ala Met His Val Ala Ala
145 150 155 160
Val Val Glu Ile His Gly Arg Leu Ile Pro Ala Leu Thr Thr Leu Arg
165 170 175
Asp Ala Leu Gln Ala Lys Ser Ala Glu Phe Glu His Ile Ile Lys Ile
180 185 190
Gly Arg Thr His Leu Gln Asp Ala Thr Pro Leu Thr Leu Gly Gln Glu
195 200 205
Phe Ser Gly Tyr Thr Gln Gln Leu Thr Tyr Gly Ile Ala Arg Val Gln
210 215 220
Gly Thr Leu Glu Arg Leu Tyr Asn Leu Ala Gln Gly Gly Thr Ala Val
225 230 235 240
Gly Thr Gly Leu Asn Thr Arg Lys Gly Phe Asp Ala Lys Val Ala Glu
245 250 255
Ala Ile Ala Ser Ile Thr Gly Leu Pro Phe Lys Thr Ala Pro Asn Lys
260 265 270
Phe Glu Ala Leu Ala Ala His Asp Ala Leu Val Glu Ala His Gly Ala
275 280 285
Leu Asn Thr Val Ala Cys Ser Leu Met Lys Ile Ala Asn Asp Ile Arg
290 295 300
Tyr Leu Gly Ser Gly Pro Arg Cys Gly Leu Gly Glu Leu Ser Leu Pro
305 310 315 320
Glu Asn Glu Pro Gly Ser Ser Ile Met Pro Gly Lys Val Asn Pro Thr
325 330 335
Gln Cys Glu Ala Met Thr Met Val Cys Ala Gln Val Met Gly Asn Asn
340 345 350
Thr Ala Ile Ser Val Ala Gly Ser Asn Gly Gln Phe Glu Leu Asn Val
355 360 365
Phe Lys Pro Val Met Ile Lys Asn Leu Ile Gln Ser Ile Arg Leu Ile
370 375 380
Ser Asp Ala Ser Ile Ser Phe Thr Lys Asn Cys Val Val Gly Ile Glu
385 390 395 400
Ala Asn Glu Lys Lys Ile Ser Ser Ile Met Asn Glu Ser Leu Met Leu
405 410 415
Val Thr Ala Leu Asn Pro His Ile Gly Tyr Asp Lys Ala Ala Lys Cys
420 425 430
Ala Lys Lys Ala His Lys Glu Gly Thr Thr Leu Lys Glu Ala Ala Leu
435 440 445
Ser Leu Gly Tyr Leu Thr Ser Glu Glu Phe Asp Gln Trp Val Arg Pro
450 455 460
Glu Asp Met Ile Ser Ala Lys Asp
465 470
<210> 50
<211> 416
<212> PRT
<213> Aspergillus niger
<400> 50
Met Asn Val Glu Thr Ser Leu Pro Gly Ser Ser Gly Ser Asp Leu Glu
1 5 10 15
Thr Phe His His Glu Thr Lys Lys His Ala Asn His Asp Ser Gly Ile
20 25 30
Ser Val Asn His Glu Ala Glu Ile Gly Val Asn His Thr Phe Glu Lys
35 40 45
Pro Gly Pro Val Gly Ile Arg Glu Arg Leu Arg His Phe Thr Trp Ala
50 55 60
Trp Tyr Thr Leu Thr Met Ser Cys Gly Gly Leu Ala Leu Leu Ile Val
65 70 75 80
Asn Gln Pro His Asp Phe Lys Gly Leu Lys Asp Ile Ala Arg Val Val
85 90 95
Tyr Cys Leu Asn Leu Ala Phe Phe Val Ile Val Thr Ser Leu Met Ala
100 105 110
Ile Arg Phe Ile Leu His Lys Asn Met Trp Glu Ser Leu Gly His Asp
115 120 125
Arg Glu Gly Leu Phe Phe Pro Thr Phe Trp Leu Ser Ile Ala Thr Met
130 135 140
Ile Thr Gly Leu Tyr Lys Cys Phe Gly Asp Asp Ala Asn Glu Lys Phe
145 150 155 160
Thr Lys Cys Leu Gln Val Leu Phe Trp Ile Tyr Cys Gly Cys Thr Met
165 170 175
Ile Thr Ala Val Gly Gln Tyr Ser Phe Val Phe Ala Thr His Lys Tyr
180 185 190
Glu Leu His Thr Met Met Pro Ser Trp Ile Leu Pro Ala Phe Pro Val
195 200 205
Met Leu Ser Gly Thr Ile Ala Ser Val Ile Gly Ser Gly Gln Pro Ala
210 215 220
Ser Asp Gly Ile Pro Ile Ile Ile Ala Gly Ile Thr Phe Gln Gly Leu
225 230 235 240
Gly Phe Ser Ile Ser Phe Met Met Tyr Ala His Tyr Ile Gly Arg Leu
245 250 255
Met Glu Val Gly Leu Pro Ser Pro Glu His Arg Pro Gly Met Phe Ile
260 265 270
Cys Val Gly Pro Pro Ala Phe Thr Ala Leu Ala Leu Val Gly Met Ala
275 280 285
Lys Ala Leu Pro Asp Asp Phe Gln Ile Val Gly Asp Pro His Ala Val
290 295 300
Ile Asp Gly Arg Val Met Leu Phe Leu Ala Val Ser Ala Ala Ile Phe
305 310 315 320
Leu Trp Ala Leu Ser Phe Trp Phe Phe Cys Ile Ala Val Val Ala Val
325 330 335
Val Arg Ser Pro Pro Lys Gly Phe His Leu Asn Trp Phe Ala Met Val
340 345 350
Phe Pro Asn Thr Gly Phe Thr Leu Ala Thr Ile Thr Leu Ala Asn Met
355 360 365
Phe Glu Ser Pro Gly Val Lys Gly Val Ala Thr Ala Met Ser Leu Cys
370 375 380
Val Ile Ile Met Phe Ile Phe Val Leu Val Ser Ala Ile Arg Ala Val
385 390 395 400
Ile Arg Lys Asp Ile Met Trp Pro Gly Gln Asp Glu Asp Val Ser Glu
405 410 415
<210> 51
<211> 542
<212> PRT
<213> Kluyveromyces lactis
<400> 51
Met Val Ser Val Lys Ala Ser Ala Ala Glu Lys Lys Glu Phe Leu Gln
1 5 10 15
Ser Gln Ile Asp Glu Ile Glu Lys Trp Trp Ser Glu Pro Arg Trp Lys
20 25 30
Asp Thr Lys Arg Ile Tyr Ser Ala Tyr Glu Ile Ala Lys Arg Arg Gly
35 40 45
Ser Val Lys Pro Asn Thr Phe Pro Ser Thr Val Met Ser Gln Lys Leu
50 55 60
Phe Lys Ile Leu Gly Glu His Ala Lys Asn Gly Thr Val Ser Lys Thr
65 70 75 80
Phe Gly Ala Leu Asp Pro Val Gln Val Thr Gln Met Ser Lys Tyr Leu
85 90 95
Asp Thr Ile Tyr Val Ser Gly Trp Gln Cys Ser Ser Thr Ala Ser Thr
100 105 110
Ser Asn Glu Pro Gly Pro Asp Leu Ala Asp Tyr Pro Met Asp Thr Val
115 120 125
Pro Asn Lys Val Glu His Leu Phe Lys Ala Gln Gln Phe His Asp Arg
130 135 140
Lys Gln Trp Glu Arg Ile Cys Asp Gly Thr Ile Glu Glu Ser Glu Ile
145 150 155 160
Ile Asp Tyr Leu Thr Pro Ile Val Ala Asp Gly Asp Ala Gly His Gly
165 170 175
Gly Leu Thr Ala Val Phe Lys Leu Thr Lys Met Phe Ile Glu Arg Gly
180 185 190
Ala Ala Gly Ile His Ile Glu Asp Gln Thr Ser Thr Asn Lys Lys Cys
195 200 205
Gly His Met Ala Gly Arg Cys Val Ile Pro Val Gln Glu His Ile Asn
210 215 220
Arg Leu Ile Thr Cys Arg Met Ala Ala Asp Val Leu Gly Ser Asp Leu
225 230 235 240
Ile Leu Val Ala Arg Thr Asp Ser Glu Ala Ala Thr Leu Leu Ser Ser
245 250 255
Thr Ala Asp Ser Arg Asp His Tyr Phe Ile Leu Gly Ala Ser Asn Pro
260 265 270
Ala Val Lys Gly Lys Pro Leu Asn Asp Leu Leu Asn Lys Ala Ile Leu
275 280 285
Asp Gly Ala Thr Ile Asp Asp Leu Gln Thr Ile Glu Lys Glu Trp Leu
290 295 300
Ala Lys Ala Asp Val Lys Leu Phe His Glu Val Phe Ala Asp Ala Ala
305 310 315 320
Lys Ala Ala Gly Lys Asp Gln Ser Val Ile Asp Gln Phe Asn Ser Lys
325 330 335
Val Asn Pro Leu Ser Glu Thr Ser Ile Tyr Glu Met Gln Ala Leu Ala
340 345 350
Lys Glu Leu Leu Gly Thr Glu Leu Phe Phe Asp Trp Asp Leu Pro Arg
355 360 365
Gly Arg Glu Gly Leu Tyr Arg Tyr Gln Gly Gly Thr Gln Cys Ser Val
370 375 380
Met Arg Ala Arg Ala Phe Ala Pro Tyr Ala Asp Leu Cys Trp Met Glu
385 390 395 400
Ser Asn Tyr Pro Asp Tyr Glu Gln Ala Lys Glu Phe Ala Glu Gly Val
405 410 415
Thr Ala Lys Phe Pro Gly Lys Trp Met Ala Tyr Asn Leu Ser Pro Ser
420 425 430
Phe Asn Trp Thr Lys Ala Met Ser Val Asp Glu Gln Glu Thr Phe Ile
435 440 445
Gln Arg Leu Gly Asp Leu Gly Tyr Ile Trp Gln Phe Ile Thr Leu Ala
450 455 460
Gly Leu His Thr Ser Gly Leu Ala Ile Glu Gln Phe Ser Lys Asn Phe
465 470 475 480
Ala Lys Leu Gly Met Lys Ala Tyr Ala Gln Asp Ile Gln Lys Lys Glu
485 490 495
Leu Asp Asn Gly Ile Asp Met Val Lys His Gln Lys Trp Ser Gly Ala
500 505 510
Glu Tyr Ile Asp Gly Leu Leu Arg Leu Ala Gln Gly Gly Leu Ala Ala
515 520 525
Thr Ala Ala Met Gly Gln Gly Val Thr Glu Asp Gln Phe Lys
530 535 540
<210> 52
<211> 551
<212> PRT
<213> Artificial Sequence
<220>
<223> Artificial
<400> 52
Met Val Lys Val Ser Leu Asp Asn Val Lys Leu Leu Val Asp Val Asp
1 5 10 15
Lys Glu Pro Phe Phe Lys Pro Ser Ser Thr Thr Val Gly Asp Ile Leu
20 25 30
Thr Lys Asp Ala Leu Glu Phe Ile Val Leu Leu His Arg Thr Phe Asn
35 40 45
Asn Lys Arg Lys Gln Leu Leu Glu Asn Arg Gln Val Val Gln Lys Lys
50 55 60
Leu Asp Ser Gly Ser Tyr His Leu Asp Phe Leu Pro Glu Thr Ala Asn
65 70 75 80
Ile Arg Asn Asp Pro Thr Trp Gln Gly Pro Ile Leu Ala Pro Gly Leu
85 90 95
Ile Asn Arg Ser Thr Glu Ile Thr Gly Pro Pro Leu Arg Asn Met Leu
100 105 110
Ile Asn Ala Leu Asn Ala Pro Val Asn Thr Tyr Met Thr Asp Phe Glu
115 120 125
Asp Ser Ala Ser Pro Thr Trp Asn Asn Met Val Tyr Gly Gln Val Asn
130 135 140
Leu Tyr Asp Ala Ile Arg Asn Gln Ile Asp Phe Asp Thr Pro Arg Lys
145 150 155 160
Ser Tyr Lys Leu Asn Gly Asn Val Ala Asn Leu Pro Thr Ile Ile Val
165 170 175
Arg Pro Arg Gly Trp His Met Val Glu Lys His Leu Tyr Val Asp Asp
180 185 190
Glu Pro Ile Ser Ala Ser Ile Phe Asp Phe Gly Leu Tyr Phe Tyr His
195 200 205
Asn Ala Lys Glu Leu Ile Lys Leu Gly Lys Gly Pro Tyr Phe Tyr Leu
210 215 220
Pro Lys Met Glu His His Leu Glu Ala Lys Leu Trp Asn Asp Val Phe
225 230 235 240
Cys Val Ala Gln Asp Tyr Ile Gly Ile Pro Arg Gly Thr Ile Arg Ala
245 250 255
Thr Val Leu Ile Glu Thr Leu Pro Ala Ala Phe Gln Met Glu Glu Ile
260 265 270
Ile Tyr Gln Leu Arg Gln His Ser Ser Gly Leu Asn Cys Gly Arg Trp
275 280 285
Asp Tyr Ile Phe Ser Thr Ile Lys Arg Leu Arg Asn Asp Pro Asn His
290 295 300
Ile Leu Pro Asn Arg Asn Gln Val Thr Met Thr Ser Pro Phe Met Asp
305 310 315 320
Ala Tyr Val Lys Arg Leu Ile Asn Thr Cys His Arg Arg Gly Val His
325 330 335
Ala Met Gly Gly Met Ala Ala Gln Ile Pro Ile Lys Asp Asp Pro Ala
340 345 350
Ala Asn Glu Lys Ala Met Thr Lys Val Arg Asn Asp Lys Ile Arg Glu
355 360 365
Leu Thr Asn Gly His Asp Gly Ser Trp Val Ala His Pro Ala Leu Ala
370 375 380
Pro Ile Cys Asn Glu Val Phe Ile Asn Met Gly Thr Pro Asn Gln Ile
385 390 395 400
Tyr Phe Ile Pro Glu Asn Val Val Thr Ala Ala Asn Leu Leu Glu Thr
405 410 415
Lys Ile Pro Asn Gly Glu Ile Thr Thr Glu Gly Ile Val Gln Asn Leu
420 425 430
Asp Ile Gly Leu Gln Tyr Met Glu Ala Trp Leu Arg Gly Ser Gly Cys
435 440 445
Val Pro Ile Asn Asn Leu Met Glu Asp Ala Ala Thr Ala Glu Val Ser
450 455 460
Arg Cys Gln Leu Tyr Gln Trp Val Lys His Gly Val Thr Leu Lys Asp
465 470 475 480
Thr Gly Glu Lys Val Thr Pro Glu Leu Thr Glu Lys Ile Leu Lys Glu
485 490 495
Gln Val Glu Arg Leu Ser Lys Ala Ser Pro Leu Gly Asp Lys Asn Lys
500 505 510
Phe Ala Leu Ala Ala Lys Tyr Phe Leu Pro Glu Ile Arg Gly Glu Lys
515 520 525
Phe Ser Glu Phe Leu Thr Thr Leu Leu Tyr Asp Glu Ile Val Ser Thr
530 535 540
Lys Ala Thr Pro Thr Asp Leu
545 550
<210> 53
<211> 343
<212> PRT
<213> Saccharomyces cerevisiae
<400> 53
Met Val Lys Val Ala Ile Leu Gly Ala Ser Gly Gly Val Gly Gln Pro
1 5 10 15
Leu Ser Leu Leu Leu Lys Leu Ser Pro Tyr Val Ser Glu Leu Ala Leu
20 25 30
Tyr Asp Ile Arg Ala Ala Glu Gly Ile Gly Lys Asp Leu Ser His Ile
35 40 45
Asn Thr Asn Ser Ser Cys Val Gly Tyr Asp Lys Asp Ser Ile Glu Asn
50 55 60
Thr Leu Ser Asn Ala Gln Val Val Leu Ile Pro Ala Gly Val Pro Arg
65 70 75 80
Lys Pro Gly Leu Thr Arg Asp Asp Leu Phe Lys Met Asn Ala Gly Ile
85 90 95
Val Lys Ser Leu Val Thr Ala Val Gly Lys Phe Ala Pro Asn Ala Arg
100 105 110
Ile Leu Val Ile Ser Asn Pro Val Asn Ser Leu Val Pro Ile Ala Val
115 120 125
Glu Thr Leu Lys Lys Met Gly Lys Phe Lys Pro Gly Asn Val Met Gly
130 135 140
Val Thr Asn Leu Asp Leu Val Arg Ala Glu Thr Phe Leu Val Asp Tyr
145 150 155 160
Leu Met Leu Lys Asn Pro Lys Ile Gly Gln Glu Gln Asp Lys Thr Thr
165 170 175
Met His Arg Lys Val Thr Val Ile Gly Gly His Ser Gly Glu Thr Ile
180 185 190
Ile Pro Ile Ile Thr Asp Lys Ser Leu Val Phe Gln Leu Asp Lys Gln
195 200 205
Tyr Glu His Phe Ile His Arg Val Gln Phe Gly Gly Asp Glu Ile Val
210 215 220
Lys Ala Lys Gln Gly Ala Gly Ser Ala Thr Leu Ser Met Ala Phe Ala
225 230 235 240
Gly Ala Lys Phe Ala Glu Glu Val Leu Arg Ser Phe His Asn Glu Lys
245 250 255
Pro Glu Thr Glu Ser Leu Ser Ala Phe Val Tyr Leu Pro Gly Leu Lys
260 265 270
Asn Gly Lys Lys Ala Gln Gln Leu Val Gly Asp Asn Ser Ile Glu Tyr
275 280 285
Phe Ser Leu Pro Ile Val Leu Arg Asn Gly Ser Val Val Ser Ile Asp
290 295 300
Thr Ser Val Leu Glu Lys Leu Ser Pro Arg Glu Glu Gln Leu Val Asn
305 310 315 320
Thr Ala Val Lys Glu Leu Arg Lys Asn Ile Glu Lys Gly Lys Ser Phe
325 330 335
Ile Leu Asp Ser Ser Lys Leu
340
<210> 54
<211> 25
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..25
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 54
cggcattatt gtgtatggct caata 25
<210> 55
<211> 75
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..75
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 55
gaacttcgac ctgttgcaat acttcgggtt cggcacaaac gtgtacggat agggtttcaa 60
agatccatac ttctc 75
<210> 56
<211> 79
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..79
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 56
atccgtacac gtttgtgccg aacccgaagt attgcaacag gtcgaagttc tcgtacgctg 60
caggtcgacg aattctacc 79
<210> 57
<211> 74
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..74
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 57
aatcgcaact cggatttggg aggcaaggtc ggaacgcgaa ctttggcttt aggccactag 60
tggatctgat atcg 74
<210> 58
<211> 90
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..90
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 58
agaaagcctg tatgcgaagc cacaatcctt tccaacagac catactaagt attttatttt 60
acttttttta gaatgacctg ttcccgacac 90
<210> 59
<211> 24
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..24
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 59
cacaagctta ttcttccaaa aatc 24
<210> 60
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..20
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 60
aaagccaaag ttcgcgttcc 20
<210> 61
<211> 24
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..24
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 61
acttagtatg gtctgttgga aagg 24
<210> 62
<211> 4429
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..4429
<223> /organism="Artificial Sequence"
/note="Artificial"
/mol_type="unassigned DNA"
<400> 62
aaagccaaag ttcgcgttcc gaccttgcct cccaaatccg agttgcgatt gtgcttagtc 60
aaaaaattag ccttttaatt ctgctgtaac ccgtacatgc ccaaaatagg gggcgggtta 120
cacagaatat ataacatcgt aggtgtctgg gtgaacagtt tattcctggc atccactaaa 180
tataatggag cccgcttttt aagctggcat ccagaaaaaa aaagaatccc agcaccaaaa 240
tattgttttc ttcaccaacc atcagttcat aggtccattc tcttagcgca actacagaga 300
acaggggcac aaacaggcaa aaaacgggca caacctcaat ggagtgatgc aacctgcctg 360
gagtaaatga tgacacaagg caattgaccc acgcatgtat ctatctcatt ttcttacacc 420
ttctattacc ttctgctctc tctgatttgg aaaaagctga aaaaaaaggt tgaaaccagt 480
tccctgaaat tattccccta cttgactaat aagtatataa agacggtagg tattgattgt 540
aattctgtaa atctatttct taaacttctt aaattctact tttatagtta gtcttttttt 600
tagttttaaa acaccaagaa cttagtttcg aataaacaca cataaacaaa caaaatggtt 660
gatggtagat cttctgcttc cattgttgcc gttgacccag aaagagctgc cagagaaaga 720
gatgctgctg ccagagcttt gttgcaagac tctccattgc acaccaccat gcaatacgct 780
acctctggtt tggaattgac tgttccatac gctttgaagg ttgttgcttc tgctgacact 840
ttcgacagag ccaaggaagt tgctgatgaa gtcttgagat gtgcctggca attggctgac 900
accgttttga actctttcaa cccaaactct gaagtctctt tagtcggtag attaccagtc 960
ggtcaaaagc atcaaatgtc tgctccattg aaacgtgtca tggcttgttg tcaaagagtc 1020
tacaactcct ctgctggttg tttcgaccca tccactgctc cagttgccaa ggctttgaga 1080
gaaattgctt tgggtaagga aagaaacaat gcttgtttgg aagctttgac tcaagcttgt 1140
accttgccaa actctttcgt cattgatttc gaagctggta ctatctccag aaagcacgaa 1200
cacgcttctt tggatttggg tggtgtttcc aagggttaca tcgtcgatta cgtcattgac 1260
aacatcaatg ctgctggttt ccaaaacgtt ttctttgact ggggtggtga ctgtcgtgcc 1320
tccggtatga acgccagaaa cactccatgg gttgtcggta tcactagacc tccttccttg 1380
gacatgttgc caaaccctcc aaaggaagct tcttacatct ccgtcatctc tttggacaat 1440
gaagctttgg ctacctctgg tgattacgaa aacttgatct acactgctga cgataaacca 1500
ttgacctgta cctacgattg gaaaggtaag gaattgatga agccatctca atccaatatc 1560
gctcaagttt ccgtcaagtg ttactctgcc atgtacgctg acgctttggc taccgcttgt 1620
ttcatcaagc gtgacccagc caaggtcaga caattgttgg atggttggag atacgttaga 1680
gacaccgtca gagattaccg tgtctacgtc agagaaaacg aaagagttgc caagatgttc 1740
gaaattgcca ctgaagatgc tgaaatgaga aagagaagaa tttccaacac tttaccagct 1800
cgtgtcattg ttgttggtgg tggtttggct ggtttgtccg ctgccattga agctgctggt 1860
tgtggtgctc aagttgtttt gatggaaaag gaagccaagt tgggtggtaa ctctgccaag 1920
gctacctctg gtatcaacgg ttggggtact agagcccaag ctaaggcttc cattgtcgat 1980
ggtggtaagt acttcgaaag agatacctac aagtctggta tcggtggtaa caccgatcca 2040
gctttggtta agactttgtc catgaaatct gctgacgcta tcggttggtt gacttctcta 2100
ggtgttccat tgactgtttt gtcccaatta ggtggtcact ccagaaagag aactcacaga 2160
gccccagaca agaaggatgg tactccattg ccaattggtt tcaccatcat gaaaacttta 2220
gaagatcatg ttagaggtaa cttgtccggt agaatcacca tcatggaaaa ctgttccgtt 2280
acctctttgt tgtctgaaac caaggaaaga ccagacggta ctaagcaaat cagagttacc 2340
ggtgtcgaat tcactcaagc tggttctggt aagaccacca ttttggctga tgctgttatc 2400
ttggccaccg gtggtttctc caacgacaag actgctgatt ctttgttgag agaacatgcc 2460
ccacacttgg ttaacttccc aaccaccaac ggtccatggg ctactggtga tggtgtcaag 2520
ttggctcaaa gattaggtgc tcaattggtc gatatggaca aggttcaatt gcacccaact 2580
ggtttgatca acccaaagga cccagccaac ccaaccaaat tcttgggtcc agaagctcta 2640
agaggttctg gtggtgtttt gttgaacaaa caaggtaaga gatttgtcaa cgaattggat 2700
ttgagatctg ttgtttccaa ggccatcatg gaacaaggtg ctgaataccc aggttctggt 2760
ggttccatgt ttgcttactg tgtcttgaac gctgctgctc aaaaattgtt tggtgtttcc 2820
tctcacgaat tctactggaa gaagatgggt ttgttcgtca aggctgacac catgagagac 2880
ttggctgctt tgattggttg tccagttgaa tccgttcaac aaactttaga agaatacgaa 2940
agattatcca tctctcaaag atcttgtcca attaccagaa aatctgttta cccatgtgtt 3000
ttgggtacta aaggtccata ctatgtcgcc tttgtcactc catctatcca ctacaccatg 3060
ggtggttgtt tgatttctcc atctgctgaa atccaaatga agaacacttc ttccagagcc 3120
ccattgtccc actccaaccc aatcttgggt ttattcggtg ctggtgaagt caccggtggt 3180
gtccacggtg gtaacagatt aggtggtaac tctttgttgg aatgtgttgt tttcggtaga 3240
attgccggtg acagagcttc taccattttg caaagaaagt cctctgcttt gtctttcaag 3300
gtctggacca ctgttgtttt gagagaagtc agagaaggtg gtgtctacgg tgctggttcc 3360
cgtgtcttga gattcaactt accaggtgct ctacaaagat ctggtctatc cttgggtcaa 3420
ttcattgcca tcagaggtga ctgggacggt caacaattga ttggttacta ctctccaatc 3480
actttgccag acgatttggg tatgattgac attttggcca gatctgacaa gggtacttta 3540
cgtgaatgga tctctgcttt ggaaccaggt gacgctgtcg aaatgaaggc ttgtggtggt 3600
ttggtcatcg aaagaagatt atctgacaag cacttcgttt tcatgggtca cattatcaac 3660
aagctatgtt tgattgctgg tggtactggt gttgctccaa tgttgcaaat catcaaggcc 3720
gctttcatga agccattcat cgacactttg gaatccgtcc acttgatcta cgctgctgaa 3780
gatgtcactg aattgactta cagagaagtt ttggaagaac gtcgtcgtga atccagaggt 3840
aaattcaaga aaactttcgt tttgaacaga cctcctccat tatggactga cggtgtcggt 3900
ttcatcgacc gtggtatctt gaccaaccac gttcaaccac catctgacaa cttattggtt 3960
gccatctgtg gtccaccagt tatgcaaaga attgtcaagg ccactttaaa gactttaggt 4020
tacaacatga acttggtcag aaccgttgac gaaactgaac catctggaag ttaaaggaag 4080
tatctcggaa atattaattt aggccatgtc cttatgcacg tttcttttga tacttacggg 4140
tacatgtaca caagtatatc tatatatata aattaatgaa aatcccctat ttatatatat 4200
gactttaacg agacagaaca gttttttatt ttttatccta tttgatgaat gatacagttt 4260
cttattcacg tgttataccc acaccaaatc caatagcaat accggccatc acaatcactg 4320
tttcggcagc ccctaagatc agacaaaaca tccggaacca ccttaaatca acgtccctca 4380
gaaagcctgt atgcgaagcc acaatccttt ccaacagacc atactaagt 4429
<210> 63
<211> 75
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..75
<223> /organism="Artificial Sequence"
/note="Primer"
/mol_type="unassigned DNA"
<400> 63
gaaaccttcg aatccagcca gcatgtcgac acccacaaga tgtagtgcac acaggtgatt 60
gtatgtgggc ttatg 75
<210> 64
<211> 2032
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..2032
<223> /organism="Artificial Sequence"
/note="Artificial"
/mol_type="unassigned DNA"
<400> 64
aaataaccac aaacatcctt cccatatgct cggtcgtgct tgttgtacct gtgcttagtc 60
aaaaaattag ccttttaatt ctgctgtaac ccgtacatgc ccaaaatagg gggcgggtta 120
cacagaatat ataacatcgt aggtgtctgg gtgaacagtt tattcctggc atccactaaa 180
tataatggag cccgcttttt aagctggcat ccagaaaaaa aaagaatccc agcaccaaaa 240
tattgttttc ttcaccaacc atcagttcat aggtccattc tcttagcgca actacagaga 300
acaggggcac aaacaggcaa aaaacgggca caacctcaat ggagtgatgc aacctgcctg 360
gagtaaatga tgacacaagg caattgaccc acgcatgtat ctatctcatt ttcttacacc 420
ttctattacc ttctgctctc tctgatttgg aaaaagctga aaaaaaaggt tgaaaccagt 480
tccctgaaat tattccccta cttgactaat aagtatataa agacggtagg tattgattgt 540
aattctgtaa atctatttct taaacttctt aaattctact tttatagtta gtcttttttt 600
tagttttaaa acaccaagaa cttagtttcg aataaacaca cataaacaaa caaaatggtt 660
aaggttgcca tcttaggtgc ttctggtggt gtcggtcaac cattatctct attattgaaa 720
ttgtctccat acgtttctga attggctttg tacggtatct ctgctgctga aggtattggt 780
aaggatttgt cccacatcaa caccaactcc tcttgtgttg gttacgacaa ggattccatc 840
gaaaacactt tgtccaatgc tcaagttgtc ttgattccag ctggtgttcc aagaaagcca 900
ggtttgacca gagatgattt gttcaagatg aacgctggta tcgttaagtc tttggttact 960
gctgtcggta aatttgcccc aaacgctcgt atcttagtca tctccaaccc tgttaactct 1020
ttggttccaa ttgccgttga aactttgaag aagatgggta agttcaagcc aggtaacgtt 1080
atgggtgtca ccaacttgga tttggtcaga gctgaaactt tcttggttga ctacttgatg 1140
ttgaagaacc caaagatcgg tcaagaacaa gacaagacca ccatgcacag aaaggtcacc 1200
gtcatcggtg gtcactctgg tgaaaccatc attccaatca tcactgacaa atccttggtt 1260
ttccaattgg acaagcaata cgaacatttc atccacagag tccaattcgg tggtgacgaa 1320
attgtcaagg ccaagcaagg tgccggttct gctaccttgt ccatggcttt cgctggtgcc 1380
aaatttgctg aagaagtctt acgttctttc cacaacgaaa agccagaaac tgaatctttg 1440
tctgctttcg tctacttgcc aggtttgaag aacggtaaga aggctcaaca attagtcggt 1500
gacaactcca ttgaatactt ctctttgcca attgttttga gaaacggttc cgttgtttcc 1560
attgacactt ctgttttgga aaaattgtct ccaagagaag aacaattggt caacactgct 1620
gtcaaggaat tgagaaagaa cattgaaaag ggtaagtctt tcatcttgga cagttaaagt 1680
ctgaagaatg aatgatttga tgatttcttt ttccctccat ttttcttact gaatatatca 1740
atgatataga cttgtatagt ttattatttc aaattaagta gctatatata gtcaagataa 1800
cgtttgtttg acacgattac attattcgtc gacatctttt ttcagcctgt cgtggtagca 1860
atttgaggag tattattaat tgaataggtt cattttgcgc tcgcataaac agttttcgtc 1920
agggacagta tgttggaatg agtggtaatt aatggtgaca tgacatgtta tagcaatacc 1980
tcgaaacctt cgaatccagc cagcatgtcg acacccacaa gatgtagtgc ac 2032
<210> 65
<211> 2032
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..2032
<223> /organism="Artificial Sequence"
/note="Artificial"
/mol_type="unassigned DNA"
<400> 65
aaataaccac aaacatcctt cccatatgct cggtcgtgct tgttgtacct gtgcttagtc 60
aaaaaattag ccttttaatt ctgctgtaac ccgtacatgc ccaaaatagg gggcgggtta 120
cacagaatat ataacatcgt aggtgtctgg gtgaacagtt tattcctggc atccactaaa 180
tataatggag cccgcttttt aagctggcat ccagaaaaaa aaagaatccc agcaccaaaa 240
tattgttttc ttcaccaacc atcagttcat aggtccattc tcttagcgca actacagaga 300
acaggggcac aaacaggcaa aaaacgggca caacctcaat ggagtgatgc aacctgcctg 360
gagtaaatga tgacacaagg caattgaccc acgcatgtat ctatctcatt ttcttacacc 420
ttctattacc ttctgctctc tctgatttgg aaaaagctga aaaaaaaggt tgaaaccagt 480
tccctgaaat tattccccta cttgactaat aagtatataa agacggtagg tattgattgt 540
aattctgtaa atctatttct taaacttctt aaattctact tttatagtta gtcttttttt 600
tagttttaaa acaccaagaa cttagtttcg aataaacaca cataaacaaa caaaatggtt 660
aaggttgcca tcttaggtgc ttctggtggt gtcggtcaac cattatctct attattgaaa 720
ttgtctccat acgtttctga attggctttg tacggttcca gagctgctga aggtattggt 780
aaggatttgt cccacatcaa caccaactcc tcttgtgttg gttacgacaa ggattccatc 840
gaaaacactt tgtccaatgc tcaagttgtc ttgattccag ctggtgttcc aagaaagcca 900
ggtttgacca gagatgattt gttcaagatg aacgctggta tcgttaagtc tttggttact 960
gctgtcggta aatttgcccc aaacgctcgt atcttagtca tctccaaccc tgttaactct 1020
ttggttccaa ttgccgttga aactttgaag aagatgggta agttcaagcc aggtaacgtt 1080
atgggtgtca ccaacttgga tttggtcaga gctgaaactt tcttggttga ctacttgatg 1140
ttgaagaacc caaagatcgg tcaagaacaa gacaagacca ccatgcacag aaaggtcacc 1200
gtcatcggtg gtcactctgg tgaaaccatc attccaatca tcactgacaa atccttggtt 1260
ttccaattgg acaagcaata cgaacatttc atccacagag tccaattcgg tggtgacgaa 1320
attgtcaagg ccaagcaagg tgccggttct gctaccttgt ccatggcttt cgctggtgcc 1380
aaatttgctg aagaagtctt acgttctttc cacaacgaaa agccagaaac tgaatctttg 1440
tctgctttcg tctacttgcc aggtttgaag aacggtaaga aggctcaaca attagtcggt 1500
gacaactcca ttgaatactt ctctttgcca attgttttga gaaacggttc cgttgtttcc 1560
attgacactt ctgttttgga aaaattgtct ccaagagaag aacaattggt caacactgct 1620
gtcaaggaat tgagaaagaa cattgaaaag ggtaagtctt tcatcttgga cagttaaagt 1680
ctgaagaatg aatgatttga tgatttcttt ttccctccat ttttcttact gaatatatca 1740
atgatataga cttgtatagt ttattatttc aaattaagta gctatatata gtcaagataa 1800
cgtttgtttg acacgattac attattcgtc gacatctttt ttcagcctgt cgtggtagca 1860
atttgaggag tattattaat tgaataggtt cattttgcgc tcgcataaac agttttcgtc 1920
agggacagta tgttggaatg agtggtaatt aatggtgaca tgacatgtta tagcaatacc 1980
tcgaaacctt cgaatccagc cagcatgtcg acacccacaa gatgtagtgc ac 2032
<210> 66
<211> 2032
<212> DNA
<213> Artificial Sequence
<220>
<221> source
<222> 1..2032
<223> /organism="Artificial Sequence"
/note="Artificial"
/mol_type="unassigned DNA"
<400> 66
aaataaccac aaacatcctt cccatatgct cggtcgtgct tgttgtacct gtgcttagtc 60
aaaaaattag ccttttaatt ctgctgtaac ccgtacatgc ccaaaatagg gggcgggtta 120
cacagaatat ataacatcgt aggtgtctgg gtgaacagtt tattcctggc atccactaaa 180
tataatggag cccgcttttt aagctggcat ccagaaaaaa aaagaatccc agcaccaaaa 240
tattgttttc ttcaccaacc atcagttcat aggtccattc tcttagcgca actacagaga 300
acaggggcac aaacaggcaa aaaacgggca caacctcaat ggagtgatgc aacctgcctg 360
gagtaaatga tgacacaagg caattgaccc acgcatgtat ctatctcatt ttcttacacc 420
ttctattacc ttctgctctc tctgatttgg aaaaagctga aaaaaaaggt tgaaaccagt 480
tccctgaaat tattccccta cttgactaat aagtatataa agacggtagg tattgattgt 540
aattctgtaa atctatttct taaacttctt aaattctact tttatagtta gtcttttttt 600
tagttttaaa acaccaagaa cttagtttcg aataaacaca cataaacaaa caaaatggtt 660
aaggttgcca tcttaggtgc ttctggtggt gtcggtcaac cattatctct attattgaaa 720
ttgtctccat acgtttctga attggctttg tactcttctt ccaacgtcaa gggtattggt 780
aaggatttgt cccacatcaa caccaactcc tcttgtgttg gttacgacaa ggattccatc 840
gaaaacactt tgtccaatgc tcaagttgtc ttgattccag ctggtgttcc aagaaagcca 900
ggtttgacca gagatgattt gttcaagatg aacgctggta tcgttaagtc tttggttact 960
gctgtcggta aatttgcccc aaacgctcgt atcttagtca tctccaaccc tgttaactct 1020
ttggttccaa ttgccgttga aactttgaag aagatgggta agttcaagcc aggtaacgtt 1080
atgggtgtca ccaacttgga tttggtcaga gctgaaactt tcttggttga ctacttgatg 1140
ttgaagaacc caaagatcgg tcaagaacaa gacaagacca ccatgcacag aaaggtcacc 1200
gtcatcggtg gtcactctgg tgaaaccatc attccaatca tcactgacaa atccttggtt 1260
ttccaattgg acaagcaata cgaacatttc atccacagag tccaattcgg tggtgacgaa 1320
attgtcaagg ccaagcaagg tgccggttct gctaccttgt ccatggcttt cgctggtgcc 1380
aaatttgctg aagaagtctt acgttctttc cacaacgaaa agccagaaac tgaatctttg 1440
tctgctttcg tctacttgcc aggtttgaag aacggtaaga aggctcaaca attagtcggt 1500
gacaactcca ttgaatactt ctctttgcca attgttttga gaaacggttc cgttgtttcc 1560
attgacactt ctgttttgga aaaattgtct ccaagagaag aacaattggt caacactgct 1620
gtcaaggaat tgagaaagaa cattgaaaag ggtaagtctt tcatcttgga cagttaaagt 1680
ctgaagaatg aatgatttga tgatttcttt ttccctccat ttttcttact gaatatatca 1740
atgatataga cttgtatagt ttattatttc aaattaagta gctatatata gtcaagataa 1800
cgtttgtttg acacgattac attattcgtc gacatctttt ttcagcctgt cgtggtagca 1860
atttgaggag tattattaat tgaataggtt cattttgcgc tcgcataaac agttttcgtc 1920
agggacagta tgttggaatg agtggtaatt aatggtgaca tgacatgtta tagcaatacc 1980
tcgaaacctt cgaatccagc cagcatgtcg acacccacaa gatgtagtgc ac 2032
<210> 67
<211> 499
<212> PRT
<213> Arabidopsis thaliana
<400> 67
Met Ala Ala Leu Thr Met Gln Phe Glu Gly Glu Lys Lys Asn Val Ser
1 5 10 15
Glu Val Ala Asp Val Thr Leu Lys Gln Glu Asp Glu Gln Gln Glu Arg
20 25 30
Arg Ser Tyr Ser Thr Pro Phe Arg Glu Glu Arg Asp Thr Phe Gly Pro
35 40 45
Ile Gln Val Pro Ser Asp Lys Leu Trp Gly Ala Gln Thr Gln Arg Ser
50 55 60
Leu Gln Asn Phe Glu Ile Gly Gly Asp Arg Glu Arg Met Pro Glu Pro
65 70 75 80
Ile Val Arg Ala Phe Gly Val Leu Lys Lys Cys Ala Ala Lys Val Asn
85 90 95
Met Glu Tyr Gly Leu Asp Pro Met Ile Gly Glu Ala Ile Met Glu Ala
100 105 110
Ala Gln Glu Val Ala Glu Gly Lys Leu Asn Asp His Phe Pro Leu Val
115 120 125
Val Trp Gln Thr Gly Ser Gly Thr Gln Ser Asn Met Asn Ala Asn Glu
130 135 140
Val Ile Ala Asn Arg Ala Ala Glu Ile Leu Gly His Lys Arg Gly Glu
145 150 155 160
Lys Ile Val His Pro Asn Asp His Val Asn Arg Ser Gln Ser Ser Asn
165 170 175
Asp Thr Phe Pro Thr Val Met His Ile Ala Ala Ala Thr Glu Ile Thr
180 185 190
Ser Arg Leu Ile Pro Ser Leu Lys Asn Leu His Ser Ser Leu Glu Ser
195 200 205
Lys Ser Phe Glu Phe Lys Asp Ile Val Lys Ile Gly Arg Thr His Thr
210 215 220
Gln Asp Ala Thr Pro Leu Thr Leu Gly Gln Glu Phe Gly Gly Tyr Ala
225 230 235 240
Thr Gln Val Glu Tyr Gly Leu Asn Arg Val Ala Cys Thr Leu Pro Arg
245 250 255
Ile Tyr Gln Leu Ala Gln Gly Gly Thr Ala Val Gly Thr Gly Leu Asn
260 265 270
Thr Lys Lys Gly Phe Asp Val Lys Ile Ala Ala Ala Val Ala Glu Glu
275 280 285
Thr Asn Leu Pro Phe Val Thr Ala Glu Asn Lys Phe Glu Ala Leu Ala
290 295 300
Ala His Asp Ala Cys Val Glu Thr Ser Gly Ser Leu Asn Thr Ile Ala
305 310 315 320
Thr Ser Leu Met Lys Ile Ala Asn Asp Ile Arg Phe Leu Gly Ser Gly
325 330 335
Pro Arg Cys Gly Leu Gly Glu Leu Ser Leu Pro Glu Asn Glu Pro Gly
340 345 350
Ser Ser Ile Met Pro Gly Lys Val Asn Pro Thr Gln Cys Glu Ala Leu
355 360 365
Thr Met Val Cys Ala Gln Val Met Gly Asn His Val Ala Val Thr Ile
370 375 380
Gly Gly Ser Asn Gly His Phe Glu Leu Asn Val Phe Lys Pro Val Ile
385 390 395 400
Ala Ser Ala Leu Leu His Ser Ile Arg Leu Ile Ala Asp Ala Ser Ala
405 410 415
Ser Phe Glu Lys Asn Cys Val Arg Gly Ile Glu Ala Asn Arg Glu Arg
420 425 430
Ile Ser Lys Leu Leu His Glu Ser Leu Met Leu Val Thr Ser Leu Asn
435 440 445
Pro Lys Ile Gly Tyr Asp Asn Ala Ala Ala Val Ala Lys Arg Ala His
450 455 460
Lys Glu Gly Cys Thr Leu Lys His Ala Ala Met Lys Leu Gly Val Leu
465 470 475 480
Thr Ser Glu Glu Phe Asp Thr Leu Val Val Pro Glu Lys Met Ile Gly
485 490 495
Pro Ser Asp
Claims (10)
1.一种重组酵母宿主细胞，其能够生产二羧酸并且包含编码具有苹果酸脱氢酶活性的突变多肽的核酸序列，其中所述突变多肽在所述宿主细胞的胞质中表达，并且其中与在相同条件下培养的表达SEQ ID NO: 39的重组宿主细胞相比，所述宿主细胞产生增加的二羧酸；
其中所述突变多肽的氨基酸序列为在SEQ ID NO: 39的基础上在第34至40位发生突变的序列，所述在第34至40位突变的氨基酸序列选自由以下序列组成的组：
2.一种根据权利要求1所述的重组宿主细胞，其中相较于在残基34处为D并且在残基36处为R的SEQ ID NO: 39，所述突变多肽的NADP(H)-依赖型活性与NAD(H)-依赖型活性之比提高。
3.根据权利要求2所述的重组宿主细胞，其中相较于在残基34处为D并且在残基36处为R的SEQ ID NO: 39，所述具有苹果酸脱氢酶活性的突变多肽的NAD(H)-依赖型活性和NADP(H)-依赖型活性均提高。
4.根据权利要求1所述的重组宿主细胞，其中所述重组酵母细胞选自Candida、 Hansenula、Kluyveromyces、Pichia、Issatchenkia、Saccharomyces、Schizosaccharomyces或Yarrowia菌株的酵母细胞。
5.根据权利要求4所述的重组宿主细胞，其中所述酵母细胞是Saccharomyces cerevisiae。
6.根据权利要求1所述的重组宿主细胞，其中编码所述具有苹果酸脱氢酶活性的突变多肽的核酸序列在胞质中表达，并且所述具有苹果酸脱氢酶活性的突变多肽在胞质中有活性。
7.根据权利要求1所述的重组宿主细胞，其中所述重组宿主细胞还包含一个或多个拷贝的编码以下之中的一种或多种的核酸：磷酸烯醇式丙酮酸羧激酶、磷酸烯醇式丙酮酸羧化酶、丙酮酸羧化酶、延胡索酸酶、延胡索酸还原酶和/或琥珀酸转运体。
8.一种生产二羧酸的方法，其中所述方法包括在适于生产所述二羧酸的条件下发酵根据前述权利要求中任一项所述的重组宿主细胞。
9.根据权利要求8所述的方法，所述方法还包括从发酵培养基中回收所述二羧酸。
10.根据权利要求8或9所述的方法，其中所述二羧酸是琥珀酸、苹果酸和/或延胡索酸。
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| CN115819529A (zh) * | 2018-10-19 | 2023-03-21 | 中国科学院天津工业生物技术研究所 | 提高苹果酸产量的新突变蛋白 |
| WO2023093794A1 (zh) * | 2021-11-24 | 2023-06-01 | 中国科学院天津工业生物技术研究所 | 一种高产l-苹果酸的耐酸酵母菌株及其构建方法和应用 |
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| EP3919615A1 (en) | 2021-12-08 |
| WO2018011161A1 (en) | 2018-01-18 |
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