CN101659926A - 非毒性、非产毒性、非致病性镰孢属表达***及所用启动子和终止子 - Google Patents

非毒性、非产毒性、非致病性镰孢属表达***及所用启动子和终止子 Download PDF

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CN101659926A
CN101659926A CN200710096070A CN200710096070A CN101659926A CN 101659926 A CN101659926 A CN 101659926A CN 200710096070 A CN200710096070 A CN 200710096070A CN 200710096070 A CN200710096070 A CN 200710096070A CN 101659926 A CN101659926 A CN 101659926A
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J·C·洛耶
D·L·墨耶
W·约德
J·R·舒斯特
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Abstract

本发明涉及一种非毒性、非产毒性、非致病性重组镰孢(Fusarium)，如禾本科镰孢(Fusarium)如禾本科镰孢(Fusarium graminearum)宿主细胞，它包括一个与一个启动子可操纵连接的编码一种异源蛋白质的核酸序列。本发明还涉及一种利用这种镰孢宿主细胞生产重组蛋白的方法。本发明还涉及可用于这种细胞的一个启动子和终止子序列。

Description

非毒性、非产毒性、非致病性镰孢属表达***及所用启动子和终止子
本案是分案申请，母案申请号为95193875.4(国际申请号PCT/US95/07743)，申请日为1995年6月15日，发明名称为“非毒性、非产毒性、非致病性镰孢属表达***及所用启动子和终止子”。
本发明涉及用于重组蛋白生产的宿主细胞。具体地讲，本发明涉及非毒性、非产毒性、非致病性镰孢属(Fusarium)真菌宿主细胞，可将其用于重组蛋白，尤其是酶的高水平表达。本发明还涉及可用于该***的启动子序列和终止子序列。
近年来，由于将重组宿主细胞用于表达异源蛋白，已极大地简化了大量有商业价值的蛋白质的生产，这种蛋白质原来只能从其天然来源中纯化。目前，已有多种可选择的表达***，可从中选择生产任何特定蛋白质所需的表达***，包括原核宿主和真核宿主。适当表达***的选择通常不仅取决于宿主细胞产生适量产率所述蛋白质的能力，而且在很大程度上取决于预想的该蛋白质的最终用途。
尽管哺乳动物细胞和酵母细胞一直是最常用的真核宿主，但现在已开始认识到丝状真菌是可用于重组蛋白质生产的很有用的宿主细胞。现在被利用或被推荐用于上述方法的丝状真菌的例子有：粗糙脉孢霉(Neurospora crassa)、Acremoniumchrysogenum、Tolypocladium geodes、卷枝毛霉(Mucorcircinelloides)和Trichoderma reesei，构巢曲霉(Aspergillusnidulans)、黑曲霉(A.niger)和米曲霉(A.oryzae)。
镰孢属的某些种已被用作植物致病性研究和基因调控研究的模式***，如尖镰孢(Fusarium oxysporum)(Dioletz等，1993，Gene 131：61-67；Langin等，1990，CurrGenet.17：313-319；Malardier等，1989，Gene 78：147-156和Kistler及Benny，1988，Curr，Genet.13：145-149)、腐皮镰孢(F.solani)(Crowhurst等，1992，Curr，Gene.21：463-469)和大刀镰孢(Fusarium culmorum)(Curragh等，1992，Mycol.Res.97：313-317)。这些镰孢菌不适用于异源蛋白的商品化生产，因为它们具有一些不理想的特征，例如，是植物病原体，或是会产生超过安全水平的真菌毒素。Dickman和Leslie(1992，Mol.Gen.Genet.235：458-462)披露了用一种带有粗糙脉孢霉的nit-2的质粒转化玉蜀黍赤霉(Gibberella.zeae)的方法。由Dickman和Leslie披露的玉蜀黍赤霉菌株是一种植物病原体，并能产生玉米赤霉烯酮-一种***类真菌毒素。Sanchez-Fernandez等(191，Mol.Gen.Genet.225：231-233)披露了用一种带有黑曲霉niaD基团的质粒转化具有niaD突变的藤仓赤霉(Gibberellafujikoroi)的方法。
理想的表达***应当是这样的：它基本上不产生蛋白酶和真菌毒素，而且基本上没有大量其它内源产生的分泌性蛋白，它能以比现有宿主细胞更高的水平表达。本发明现在就提供满足上述要求的新型镰孢属表达***。
本发明提供了一种重组的非毒性、非产毒性和非致病性镰孢属宿主细胞，它包括一个与一个启动子可操纵连接的编码一种异源蛋白的核酸序列。这里所说的“非毒性”是指该宿主细胞不是植物或动物的毒物。例如，如果以大约20ml(1∶1稀释)培养3天的镰孢属培养基/kg体重/小鼠的剂量给大约5只小鼠注射镰孢属培养基约14天之后，这些小鼠中无一只因镰孢处理而死亡的话，就认为这种镰孢宿主细胞是非毒性的。这里所说的“非产毒性”是指经标准分析方法，如HPLC分析测定基本没有真菌毒素的宿主细胞。例如，可用有机溶剂提取一定量的生长在盛有固体营养基的2×9cm培养皿上的镰孢菌，并将该提取物的0.5％加注到HPLC中进行分析。通过没有在真菌毒素标准的已知位置上出现可检测到的HPLC峰这一现象，可以推断没有已知的真菌毒素。这里所说的“非致病性”是指该宿主细胞不会导致健康植株或健康动物明显发病。例如，一种能使植物致病的镰孢菌，会出现对植物木质部组织的真菌性侵入，并导致以典型的枯萎症状为特征的病态。这里所说的“异源蛋白”是指一种对该宿主细胞来说不是天然的蛋白质，或是一种已通过修饰使其天然序列发生改变的天然蛋白质，或是一种对其表达做过定量改变的天然蛋白质，这种表达的定量改变可以是对表达该天然蛋白质所需的天然调节序列，如启动子、核糖体结合位点等进行操作的结果，或是用重组DNA技术对宿主细胞进行其它操作的结果。所述核酸序列可操纵连接在一个合适的启动子序列上，该启动子序列可指导所述核酸序列在所述选择的宿主细胞中转录。
本发明还涉及一种生产重组蛋白质的方法，该方法包括在有利于蛋白质表达的条件下培养上述菌种之一的宿主细胞，该宿主细胞带有一个编码一种异源蛋白质的核酸序列；以及从培养物中回收所述蛋白质。在一个优选实施方案中，所述蛋白质是一种真菌蛋白，最好是一种真菌酶。采用本发明的方法，至少能产生0.5g异源蛋白质/1宿主细胞。
出人意料的是，本发明的宿主细胞仅分泌少量蛋白酶，在下文的实施例1部分所公开的酪蛋白澄清试验证实了这一点；具体地讲，仅能检测到小的水解区或检测不到。本发明的宿主细胞和方法在某些真菌酶的生产方面出人意料地比其它已知真菌菌种，如黑曲霉、米曲霉或尖镰孢更为有效。
本发明还涉及一个由编码一种尖镰孢类胰蛋白酶蛋白酶的基因或其片断衍生的启动子序列，所述片断具有基本上与该序列相同的启动子活性。该启动子的序列如SEQ ID NO：5所示。
此外，本发明还涉及一个由编码一种尖镰孢类胰蛋白酶蛋白酶的基因或其片断衍生的终止子序列，所述片断具有基本上与该序列相同的终止子活性。该终止子的序列如SEQ ID NO：6所示。
图1表示由禾本科镰孢(Fusarium graminearum)(泳道1)、黑曲霉(泳道2)和米曲霉(泳道3)分泌的蛋白质的SDS凝胶电泳结果。泳道4表示分子量标记。
图2表示对下列样品进行的蛋白酶试验结果：米曲霉(孔1)；黑曲霉(孔2)；禾本科镰孢(孔3)；空白孔对照(孔4-6)。
图3表示质粒pJRoy6的构建。
图4表示对禾本科镰孢20334的转化体中所分泌的类胰蛋白酶蛋白酶进行SDS-PAGE分析的结果。泳道1：分子量标记：泳道2：空白；泳道3：纯化的类胰蛋白酶蛋白酶标准；泳道4：空白；泳道5：未转化的禾本科镰孢20334；泳道6：空白：泳道7：用质粒pJRoy6转化的禾本科镰孢菌株20334；泳道8：空白；泳道9：分子量标记。
图5表示pJRoy20的限制图。
图6表示pDM151的限制图。
图7表示pDM155的限制图。
图8A和8B表示当DSM151-4在禾本科镰孢中发酵20-160小时后
Figure A20071009607000081
在禾本科镰孢中的表达水平。图8A表示
Figure A20071009607000082
的分析结果。图8B表示对在所述禾本科镰孢中
Figure A20071009607000083
产生的SDS-PAGE分析结果。泳道1：分子量标记；泳道2：20小时；泳道3：50小时；泳道4：70小时；泳道5：90小时；泳道6：120小时；泳道7：140小时；泳道8：160小时。
图9A和9B表示当DSM 155-10在禾本科镰孢中发酵20-160小时后
Figure A20071009607000084
的表达水平。图9A表示
Figure A20071009607000085
分析结果。图9B表示对在所述禾本科镰孢中
Figure A20071009607000086
产生的SDS-PAGE分析结果。泳道1：分子量标记；泳道2：20小时；泳道3：50小时；泳道4：60小时；泳道5：90小时；泳道6：120小时；泳道7：140小时；泳道8：160小时。
图10表示pCaHj418的限制图。
图11表示pDM148的限制图。
图12表示pDM149的限制图。
图13表示pMHan37的限制图。
图14表示pDM154的限制图。
镰孢属的特征是：菌丝体伸展在培养中呈絮状，在固体培养基上的菌丝体常带有一些粉紫或黄色。分生孢子梗细长、长度不同而且是单的，或粗、短、不规则地分枝，或具有一个瓶梗螺环，单的或组合成分生孢子座。分生孢子主要有两类，通常藏在湿的小头中：几个细胞的大分生孢子，其尖尾略有卷曲或弯曲，一般为舟状；一个细胞的小分生孢子，卵圆形或长椭圆形，单生或呈链状。某些分生孢子为中等大小，2或3个细胞，长椭圆形或略有卷曲。
在一个具体实施方案中，本发明的宿主细胞是禾本科镰孢的种，它具有以下特征。分生孢子：无小分生孢子。大分生孢子明显有隔膜，厚壁，直至中等镰状，不均等弯曲，其腹面近于垂直而背面呈光滑的弧形。基细胞为特异足状。顶端细胞呈锥形或收缩成进口锥状。分生孢子梗：不分技或分枝的单瓶梗。厚壁孢子：在培养物中通常极缓慢地生成：当确实发生时，它们最常见的是生成于大分生孢子上，但也可生成于菌丝体上。菌落形态学：在PDA培养基上生长迅速，具有稠密的气生菌丝体，这些菌丝体几乎充满试管，通常为黄色至黄褐色，其边缘呈白色至洋红色。假如有的话，红棕至橙色分生孢子座也是稀疏的，只有当培养物在30天以上龄时才会出现。其下表面通常为洋红色。这种真菌能产生任何类型褪色部分的直筒状(背面与腹面平行)的大分生孢子。
在一个最具体的实施方案中，所述禾本科镰孢是Fusariumgraminearum Schwabe IMI145425，由美国典型培养物保藏所保藏，编号为ATCC 20334(US专利号4,041,189)，以及它的衍生物和突变体，它们同样是非毒性、非产毒性和非致病性的，例如在U.S.专利号4,041,189中所披露的。
应当理解，在本说明书及权利要求中，所用的“禾本科镰孢(Fusarium graminearum)”不但指包括在该种内的有机体，而且包括那些原先是该种的有机体的种或目前在另一种分类体系中被称为其它的种，但是仍具有与上文中所述的形态特征和培养特征相同的特征的种，而且可能与禾本科镰孢同义。这些种包括(但不限于)Fusarium roseum，F.roseum var.graminearum，玉蜀黍赤霉或Gibberella roseum，Gibberella roseum f.sp.cerealis。
本领域技术人员还知道，对这里所述宿主菌种的成功转化并不局限于采用本文具体列举的载体、启动子和选择标记。一般而言，用于转化尖镰孢、腐皮镰孢和大刀镰孢的技术也可用于转化本发明的宿主细胞。例如，尽管amdS选择标记是优选的，其它可用的选择标记包括arg B(构巢曲霉或黑曲霉)、trp C(黑曲霉或构巢曲霉)、pyrG(黑曲霉、米曲霉或构巢曲霉)、niaD(构巢曲霉、黑曲霉或尖镰孢)、及hyg B(大肠杆菌)标记。所述启动子可以是在这些种中具有强转录活性的任何DNA序列，并可以由编码胞外和胞内蛋白质，如淀粉酶、葡糖淀粉酶、蛋白酶、脂肪酶、纤维素酶和糖酵解酶的基因衍生而来。所述启动子的例子包括(但不限于)构巢曲霉amdS启动子或来自用于糖酵解酶和基因的启动子，如TPI、ADH、GAPDH和PGK。所述启动子还可以是同源启动子，即某一基因的启动子是所用宿主菌株先天固有的。启动子序列还可以带有接头，以便引入特异限制性位点，有利于该启动子序列与所选择的基因连接，或与所选择的信号肽或前导区(preregion)连接。
所述启动子序列可以由一个编码尖镰孢类胰蛋白酶蛋白酶的基因或其片断衍生而来，该片断具有与该序列基本相同的启动子活性。所述启动子的序列SEQ ID No：5所示。本发明还包括能在下列条件下与所述启动子序列杂交的核酸序列：在5×SSC中预浸泡，并在40℃下在由20％甲酰胺、5×Denhardt’s溶液、50mM磷酸钠、pH6.8和50μg变性的超声处理的小牛胸腺DNA组成的溶液中预杂交1小时，随后在大约40℃下在补充了100μM ATP的相同溶液中杂交18小时，接着在温度大约为45℃的0.4×SSC中洗涤；或者该核酸序列与SEQ ID No：5的同源性至少约为90％，最好约为95％，但它具有基本上与所述序列相同的启动子活性。在另一个实施方案中，所述启动子可以是包括大量AreA结合位点的一段序列，AreA是在氯限制期间表达的基因的正调节物；在粗糙脉孢霉中这些位点被称为nit-2(Fu和Marzlus，1990，Pro.Natl.Acad.Sci.U.S.A.87：5331-5335)。可通过增加或取代该AreA位点对上述启动子序列进行修饰。
终止子和聚腺苷酸化序列也可来自与所述启动子相同的序列。在一个特定实施方案中，终止子序列可以由一个编码尖镰孢类胰蛋白酶蛋白酶的基因或其片断衍生而来，所述片断具有基本上与该序列相同的终止子活性。该终止子的序列如SEQ ID No：6所示。本发明还包括能在下列条件下与所述终止子序列杂交的核酸序列：在5×SSC中预浸泡，并在大约40，在由20％甲酰胺、5×Denhandt’s溶液、50mM磷酸钠(pH6.8)和50μg变性的超声处理的小牛胸腺DNA组成的溶液中预杂交1小时，随后在40左右、在补充了100μM ATP的相同溶液中杂交18小时，接着在45℃左右的0.4×SSC中洗涤；或者该核酸序列与SEQ ID No：5的同源性至少约为90％，最好约为95％，但它具有基本上与所述序列相同的终止子活性。
还可以把增强子序列***该构建体中。
为了避免破坏细胞以获取表达产物，并减少表达产物在细胞内可能发生的降解，该产物最好能被分泌到细胞外面。为此，在一个优选实施方案中，将目的基因同一个前导区，如信号肽或前导肽连接，该前导区可引导表达产物进入细胞的分泌途径。所述前导区可由任何生物的用于任何分泌性蛋白的基因衍生而来，或者是天然前导区。该前导区的可选来源有：曲霉菌的葡糖淀粉酶或淀粉酶基因、芽孢杆菌的淀粉酶基因、Rhizomucor miehei的脂肪酶或蛋白酶基因、来自酿酒酵母(Saccharomyces cerevisiae)的α-因子编码基因或小牛前凝乳酶(prochymosin)基因。所述前导区可来自用于米曲酶TAKA淀粉酶、黑曲霉中性α-淀粉酶、黑曲霉酸稳定性α-淀粉酶、地衣型芽孢杆菌(B.licheniformis)α-淀粉酶、来自芽孢杆菌NCIB11837的生麦淀粉酶、嗜热脂肪芽孢杆菌(B.stearothermophilus)α-淀粉酶或地衣型芽孢杆菌枯草蛋白酶的基因。有效的信号序列是米曲霉TAKA淀粉酶信号、Rhizomucor miehei天冬氨酸蛋白酶信号和Rhizomucor miehei脂肪酶信号。另外，还可以采用被表达基因的天然前导区，例如SEQID No：4中氨基酸-24至-5之间的片段。
同启动子和终止子序列功能性地连接着的用于目的产物的基因可以***带有选择标记的一种载体中，或是携带在能够整合到宿主菌株的基因组里的一个单独的载体或质粒上。另外，所用的载体可以作为线状或环状染色体外因子在宿主细胞中复制。举例来说，这类载体包括质粒和微型染色体。所述载体***可以是单个的载体或质粒，或者是共同携带有待整合到基因组中去的全部DNA的两个或两个以上的载体或质粒。载体或质粒可以是线状或闭环分子。
可采用本领域已知的方法，如转化法和电穿孔法(例如，参见Fincham，1989，Microbial Rev.53：148-170)，用编码所述异源蛋白的核酸转化宿主细胞。
可采用本领域已知的方法培养本发明的重组宿主细胞。简单地说，是在标准的生长培养基上培养宿主细胞，例如，那些含有无机盐、维生素、适当的有机碳源(如葡萄糖或淀粉)、各种复合营养源中的任一种(酵母提取物、水解酪蛋白、大豆粉等)的生长培养基。例子之一是FP-1培养基(5％大豆粉、5％葡萄糖、2％K2HPO4、0.2％CaCl2、0.2％MgSO4·7H2O和0.1％Pluronicacid(BASF))。在pH约为4.5-8.0和温度大约为20-37℃的条件下发酵大约2-7天。
本发明的宿主细胞可用于表达任何感兴趣的原核或真核异源蛋白，但最好用于表达真核蛋白质。这种宿主细胞的特别有用的用途是用于表达异源蛋白，特别是真菌酶。这种新型表达***可用于表达各种酶，例如：过氧化氢酶、漆酶、酚氧化酶、氧化酶、氧化还原酶、纤维素酶、木聚糖酶、过氧化物酶、脂肪酶、水解酶、酯酶、角质酶、蛋白酶及其它蛋白酶、氨肽酶、羧肽酶、肌醇六磷酸酶、裂合酶、果胶酶及其它溶果胶酶、淀粉酶、葡糖淀粉酶、α-半乳糖苷酶、β-半乳糖苷酶、α-葡糖苷酶、β-葡糖苷酶、甘露糖苷酶、异构酶、转化酶、转移酶、核糖核酸酶、壳多糖酶、变位酶(mutanose)及脱氧核糖核酸酶。
在一个特定实施方案中，所述酶是一种碱性蛋白酶，例如尖镰孢前原胰蛋白酶(pre-pro-trypsin)基因。在一个最为具体的实施方案中，基因组序列如SEQ ID No：3所示，而蛋白质序列如SEQID No：4所示。
在另一个具体实施方案中，所述酶是一种碱性内切葡聚糖酶，它可与一种抗体发生免疫反应，这种抗体针对来自Humicolainsolens，DSM 1800的高度纯化的-43kD内切葡聚糖酶；或是具有纤维素酶活性的-43kD内切葡聚糖酶的衍生物(参见WO91/17243)。下文中称之为“
Figure A20071009607000131
”的内切葡聚糖酶可以由SEQ ID NO：7所示的基因编码，并可以具有SEQ ID NO：8所示的蛋白质序列。这种酶还可以是的变体。
在又一个具体实施方案中，所述酶是一种1，3-特异性脂肪酶，下文中称之为
Figure A20071009607000133
的变体。这种酶可以由SEQ ID NO：9所述的DNA序列编码，并可以具有SEQ ID NO：10所示的氨基酸序列。这种酶还可以是
Figure A20071009607000134
的变体，例如D96L，E210K、E210L(参见WO92/05249)。
本领域技术人员可以理解，“真菌酶”一词不仅包括天然真菌酶，而且还包括通过氨基酸取代、缺失、添加修饰过的真菌酶，或是对其进行其它修饰以增强其活性、热稳定性、pH耐受性等的真菌酶。本发明的宿主细胞还可用于表达有药用价值的异源蛋白，如激素、生长因子、受体等。
下面将通过非限定性实施例对本发明做进一步说明。
实施例
1.禾本科镰孢20334仅分泌少量的蛋白质
将禾本科镰孢菌株20334、米曲霉和黑曲霉的分生孢子悬浮液接种到盛在体积为125ml的摇瓶里的25ml YPD培养基(1％酵提取物(Difco)、2％细菌培养用胨(Difco)、2％葡萄糖)中，并在30℃、以300rpm的速度摇动的条件下培养5天。通过离心收集该培养物中的上清液。将每种样品共10μl与10μl 0.1M二硫苏糖醇(Sigma)和10μl加样缓冲液(40m M Tris碱、6％十二烷基硫酸钠、2.5mM EDTA、15％甘油、2mg/ml溴甲酚紫)混合。将上述样品煮沸5分钟，并在4-12％聚丙烯酰胺凝胶(Navex)上电泳5分钟。通过考马斯兰染色显现上述蛋白质。结果(图1)表明，禾本科镰孢菌株20334仅能产生极少的分泌性蛋白质。
2.禾本科镰孢20334仅分泌少量的蛋白酶
将来自禾本科镰孢菌株20334、米曲霉和黑曲霉的培养基(参见1.)共40μl各用移液管分别加注到在酪蛋白琼脂平皿(2％无脂奶粉(Lucerne)、50mM Tris-HCl pH＝7.5、1％惰性琼脂(Difco))上切出的孔中。将上述平皿在37℃培养5小时，并观察蛋白质水解范围。结果(图2)表明，禾本科菌株20334培养物只有极小的蛋白酶解活性。
3.尖镰孢基因组前原胰蛋白酶基因的克隆
由尖镰孢基因组DNA制备在λ噬菌体中的基因组文库，所采用的方法如Sambrook等在Molecular Cloning(A laboratoryManual，Cold Spring harber)N.Y.1989)一书中所述。将总计50μg基因组DNA放在含有10mM Tris(pH＝7.5)、50mM NaCl、7mMMgCl2、7mM2-巯基乙醇和4个单位的Sau3A限制酶的体积为200μl的溶液中，在37℃温度下消化1分钟。通过琼脂糖凝胶电泳分离分子大小为10-20kb的部分消化的DNA，接着电洗脱到透析膜上，并用一个Elutip-D柱(Schleicher和Schuell)进行浓缩。将1μg用限制酶BamHI切过、并用磷酸酶(Clonetech)处理过的EMBL4噬菌体的λ臂与300-400μg用Sau 3A切过的基因组DNA在25μl的体积中、在标准条件下连接(参见Sambrook等，1989，Molecular Cloning，A Laboratory Manual，cold Springharbor，NY)。采用一种商品化试剂盒(Gigapack Gold II，Stratagene)，按照生产商的说明由上述连接混合物制备λ噬菌体。
采用标准方法，铺平板培养大约15,000个重组λ噬菌体，并生产滤膜吸取物(吸至Hybond N+滤膜上，Amersham)(Sambrook，等，1989，Molecular Cloning，A LaboratoryManual，Cold Spring Harbar，NY)。按照标准方法(Sambrook等，1989，Molecular Cloning，A Laboratory Manual，ColdSpring Harbor，NY)处理该滤膜，与Genius试剂盒杂交，进行非放射性核酸检测。用作探针的DNA是用地高辛配基(DIG)标记的、存在于质粒pSX233上的尖镰孢类胰蛋白酶蛋白酶(以下称SP387)基因的完整编码区的0.75kb的PCR片段，该探针已由NRRL保藏，入藏号为NRRLB-21241。该PCR反应的引物是5′-tgcggatcc ATGGTCAAGTTCGCTTCCGTC(正向引物，SEQ ID NO：1)和5′-gacctcgag TTAAGCATAGGT GTCAATGAA(反向引物；SEQ ID NO：2)。在以上两个引物中，小写字母表示接头序列，而大写字母表示上述SP387基因的编码区。为了进行PCR，将25ng带有来自质粒pSX233的SP387基因的长度为907bp的BamH1/Xbal DNA片段，分别与68pmol的正向引物和反向引物混合。
将上述DNA片段与引物的混合物用1×Taq缓冲液/1×DIG标记Mix/5单位Taq(Boehringer Mannheim)调至80μl体积。反应条件为：95℃3分钟，然后进行这样的35个循环(95℃30秒，50℃1分钟，72℃1分钟)。通过PCR由尖镰孢类胰蛋白酶蛋白酶基因得到的DNA序列如SEQ ID NO：3所示。采用一种改进的(Engler和Blum，1993，Anal.Biochem.210：235-244)Genius试剂盒(Boehringer M-annheim)，用由DIG标记的探针筛选噬菌斑。分离阳性克隆并通过又一轮铺平皿培养和杂交进行纯化。制备带有尖镰孢类胰蛋白酶蛋白酶基因的重组λ噬菌体，并用Quiagenλmidi制备试剂盒(Quiagen)从该噬菌体中分离DNA。
4.表达质粒pJRoy6的构建
将限制作图、Southern印迹和杂交技术(Sambrook等，1989，Molecular Cloning，A Laboratory Manual，Cold SpringHarbor，NY)用于鉴定来自所述重组噬菌体之一的一个5.5kb的Pst1限制酶片段，它带有尖镰孢类胰蛋白酶蛋白酶编码基因及侧翼DNA序列。将这个5.5kb的Pst1片段亚克隆到Pst1消化的pUC118上，并将质粒命名为pJRoy4(见图3)。用限制酶EcoR1消化质粒pJRoy4，分离带有SP387基因和pUC118多接头的43bpEcoR1/Pst1区的3.5kb的EcoRI片段，并将其亚克隆到载体pToC90上，制成质粒pJRoy6(图3)。
5.SP387表达盒的构建
构建一种带有SP387启动子和终止子的表达盒(pJRoy20)，该表达盒是通过把所述启动子和终止子连接在pUC118上的BamH1位点而构成的。已将带有pJRoy20的大肠杆菌菌株保藏在NRRL。上述启动子片段是通过用EcoR1(切点为-1200)和Nco1(切点为翻译起始位点，见图5)消化SP387载体pJRoy6而产生的。上述终止子序列(图5中的碱基从2056-3107)是通过PCR扩增而产生的，扩增反应使用以下寡核苷酸：
正向
5′gcacaccatggtcgctggatccATACCTTGTTGGAAGCGTCG3′(SEQ ID N：11)
反向
5′atcggagcatgcggtaccgtttaaacgaatta AGGTAAACAAGATATAATTTTCTG3′(SEQ ID NO：12)
大写字母与SP387终止子DNA互补，而小写字母是带有工程限制位点的尾部。
在用Nco1和Sph1消化之后，分离所产生的带有终止子的扩增产物，在该终止子靠近5′末端的一侧有Nco1和BamH1位点，而在靠近其3′末端的一侧有EcoR1、Pme1、Kpn1和Sph1位点。连接上述启动子片段、终止子片段和用Kpn1/Sph1切过的pUC118三者，以制成pJRoy20(见图5)。
Figure A20071009607000161
构建体
将位于SP387启动子的-15位置上的EcoRV位点和位于
Figure A20071009607000162
编码区的+243位置上的Nco1位点用于在SP387启动子与基因之间产生正确融合。采用以下引物由载体pCaHj418(见图10)产生PCR片段，该片段带有SP387上-18至-1的片段，紧接着是基因上-1至+294的片段。
正向
EcoRV
5′ctcttggatatctatctcttcaccATGCGTTCCTCCCCCCTCCT3′(SEQ ID NO：13)
反向
5′CAATAGAGGTGGCAGCAAAA3′(SEQ ID NO：14)
正向引物里的小写字母是SP387启动子上从-24至-1处的bp，而大写字母是
Figure A20071009607000171
上从1至20处的bp。
所采用的PC R条件为：95℃5分钟，接着进行30个循环(95℃30秒，50℃1分钟，72℃1分钟)。用Invitrogen’s TA克隆试剂盒把所得到的0.32kb片段克隆到载体pCRII上，制成pDM148(见图11)。从pDM148中分离0.26kb的EcoRV/Nco1片段，并用来自pCaHj418的0.69kb Ncol/Bg lII片段连接，再将其克隆到用EcoRV/BamH1消化过的pJRoy20上，制成pDM149(见图12)。将3.2kb EcoR1
Figure A20071009607000172
表达盒(SP387启动子
Figure A20071009607000173
/SP387终止子)从pDM149上分离下来，并将其克隆到pToC90的EcoR1位点上，以制成pDM151(见图6)。表达构建体pDM151带有上述表达盒和amd S选择标记。已把带有pDM151的大肠杆菌菌株保藏在NRRL。
Figure A20071009607000174
构建体
将位于SP387启动子的-15位置上的EcoRV位点和位于
Figure A20071009607000175
编码区的+6位置上的Sacl位点用于在SP387启动子和
Figure A20071009607000176
基因之间产生正确融合。构建一个带有SP387启动子的最后15个bp、后接
Figure A20071009607000177
编码区的前6个bp的衔接头，该接头如下所示。
EcoRV               SacI
atctatctcttcaccATGAGGAGCT    (SEQ ID NO：15)
tagatagagaagtggTACTCC        (SEQ ID NO：16)
Figure A20071009607000178
cDNA基因的一个0.9kb SacI/BamHI片段是从米曲霉表达构建体pMHan37(见图13)上分离的。连接所述EcoRV/SacI衔接头和SacI/BamHI片段，并将其克隆到用EcoRV/BamHI消化的pJRoy20上，制成质粒pDM154(见图14)。从pDM154上分离3.2kb的KpnI
Figure A20071009607000182
表达盒(SP387启动子
Figure A20071009607000183
/SP387终止子)，并将其克隆到pToC90的KpnI位点上，制成质粒pDM155(见图7)。表达构建体pDM155带有
Figure A20071009607000184
表达盒和amdS选择标记。已将携带pDM151的大肠杆菌菌株保藏在NRRL。
8.禾本科镰孢的转化
在25℃下，将禾本科镰孢菌株ATCC20334培养物在加有1.5％葡萄糖和1.5％琼脂的Vogels培养基(Vogel，1964，Am.Nature98：435-446)的培养皿(100×15mm)上生长3周。用移菌环把分生孢子(每皿约108个)移至10ml无菌水中，并通过4层干酪包布、最后通过一层米拉布(miracloth)过滤而进行纯化。离心浓缩分生孢子。用108个分生孢子接种50mlYPG(1％酵母提取物(Difco)、2％细胞培养用胨(Difco)、2％葡萄糖)，并在20℃和150rpm的条件下培养14小时。把所得菌丝截留在无菌的0.4μm滤膜上，并用消毒蒸馏水和1.0M MgSO4连续洗涤。将上述菌丝重新悬浮在10ml234(Novo Nordisk)溶液(2-10mg/ml，溶于1.0MMgSO4中)里，并在34℃下以80rpm速度振荡的条件下消化15-30分钟。通过用4层干酪包布和米拉布进行连续过滤，将未消化的菌丝材料从所得原生质体中除去。让20ml1M山梨醇通过上述干酪包布和米拉布，并与原生质体溶液合并。混合以后，通过离心使原生质体(约5×108个)沉淀，并通过在20ml 1M山梨醇中和在20ml STC(0.8m山梨醇，50mM Tris-HCl pH-8.0，50mM CaCl2)中进行再悬浮和离心连续洗涤原生质体。将洗涤过的原生质体再悬浮于4份STC和1份SPTC(0.8M山梨醇，4％聚乙二醇4000(BDH)，50mM Tris-HCl pH＝8.0，50mMCaCl2)中，浓度为1-2×108/ml。将100μl原生质体悬浮液加进盛于聚丙烯试管(17×100mm)中的5μg pJRoy6和5μl肝素(以5mg/ml的浓度溶于STC中)里，并在冰上培养30分钟。将1ml SPTC轻轻混入上述原生质体悬浮液中，并在室温下继续培养20分钟。将原生质体铺在一种选择培养基上，该培养基由加有10mM乙酰胺的Cove盐(Cove，D.J.1966，Biochem.Biophys.Acta113：51-56)、15mMCsCl2/2.5％惰性琼脂(Difco)和1.0M蔗糖组成，并在该培养基上叠放一层含有0.6M蔗糖和1.0％低熔点琼脂糖(Sigma)的相同培养基。将培养皿在25℃温度下培养，转化子在第6-21天出现。
9.类胰蛋白酶蛋白酶在禾本科镰孢中的表达
将转化体转移到COVE2培养基(与上述COVE培养基相同，但没有氯化铯，并把1.0M的蔗糖的浓度改为30g/l)，并在25℃下生长3天或3天以上。用由COVE2平皿培养物切成的约为1cm的琼脂块接种盛在150ml烧瓶里的25ml FP1培养基(5％大豆粉、5％葡萄糖、2％K2HPO4、0.2％CaCl2、0.2％MgSO4·7H2O和0.1％Pluronic acid(BASF))等分试样，并在30℃、振荡(150rpm)下培养6天。离心后回收上清液样品，并进行以下SDS-PAGE分析。将每种上清液20μl与10μlSDS-PAGE样品缓冲液(1ml 0.5M Tris pH＝6.8、0.8ml甘油、1.6ml10％SDS、0.4ml 0.8M二硫苏糖醇、0.2ml 1％溴酚蓝)、2μl溶于异丙醇中的2％PMSF(Sigma)和2μl甘油混合。上述样品在沸水浴中放4分钟，并取每种样品40μl在10-27％聚丙烯酰胺凝胶(Novex)上电泳。按照标准方法，用考马斯染料对上述凝胶进行染色和脱色。经测定，类胰蛋白酶蛋白酶的表达水平≥0.5g/l。
10.酶分析
10.1
Figure A20071009607000191
缓冲剂：磷酸钠(50mM，pH 7.0)
底物：AZCL-HE纤维素(Megazyme)，2mg/ml缓冲液。
酶标准：将100mg
Figure A20071009607000192
标准(10,070ECu/g)溶于1ml缓冲液中，并在-20℃下保存。在用于酶分析之前，用缓冲液稀释上述储液(1∶100)。试验的范围为0.5-5.0ECU/ml。采用650,000ECU/g转换因子。
将底物溶液(990μl)加至24孔微量滴定板的样品孔中。将10μl样品(在缓冲液中稀释，以产生0.5-10ECU/ml的活性)加进上述底物中。在45℃下培养30分钟以进行反应，把上清液转移到96孔微量滴定板上，并测量在650nm处的光吸收。
Figure A20071009607000203
分析
缓冲液：0.1M MOPS，pH7.5，含有4mM CaCl2。
底物：将10ml丁酸对硝基苯酯(pNB)溶于1ml DMSO中；
将4ml缓冲液加进溶解在DMSO中的底物里
＊储液浓度＝11.5mM，溶于20％DMSO中
酶标准：以1000LU/ml的浓度把
Figure A20071009607000204
(23,100LU/g)溶于50％甘油中，并在-20℃下保存。在试验之前以1∶100的比例将该储液稀释于缓冲液中。试验范围为0.125-3.0LU/ml。
将100μl pNB储液加进100μl经适当稀释的酶样品中。在25℃下用5分钟时间测405nm处的活性(mOD/分钟)。
10.3SP387分析
在使用之前，通过把溶于二甲亚砜中的0.2M L-BAPNA(Sigma B3133)储液(冰冻保存)在缓冲液(0.01M谷氨酸二甲酯(Sigma)、0.2M硼酸和0.002M CaCl2，用NaOH把pH调至6.5)稀释为0.004M而制备L-BAPNA底物。对1μl培养物进行离心(14500xg，10分钟)。将100μl稀释过的培养物上清液等分试样加至96孔微量滴定板中的100μl底物中。采用一台ELISA计数器，在25℃温度下用5分钟时间以30秒的间隔测定405nm处的光吸收变化。结果是参比纯化的SP387标准计算出来的。
11.
Figure A20071009607000205
的表达
纯化23个pDM151转化体，在摇瓶里的大豆/葡萄糖培养基上培养，并于9天之后检验活性(见下面的表1)。4个转化体以大约50-100mg/L的水平表达将转化体pDM151-4在小型发酵罐中培养，采用生产SP387时所使用的条件(见9)。7天后，出现大约6.0g/L的
Figure A20071009607000212
以7天计，
Figure A20071009607000213
中含有高于90％的分泌性蛋白(图8A)。根据SDS凝胶电泳测得
Figure A20071009607000214
含有高于90％的分泌性蛋白(图8B)。
表I
  转化体   ECU/ml   mg/L
  pDM 151.3-4   58.2   90
  pDM 151.3-5   0   0
  pDM 151.3-6   0   0
  pDM 151.3-10   0   0
  pDM 151.3-11   2.46   4
  pDM 151.3-12   0   0
  pDM 151.3-13   12.2   19
  pDM 151.3-14   47.3   73
  pDM 151.3-15   22.7   35
  pDM 151.3-16   0   0
  pDM 151.3-17   0   0
  pDM 151.3-18   0   0
  pDM 151.3-19   0   0
  pDM 151.3-21   0   0
  pDM 151.3-22   43.7   67
  pDM 151.3-23   1.25   2
  pDM 151.3-24   17.8   27
  pDM 151.3-25   38   58
  pDM 151.3-26   0   0
  pDM 151.3-27   10.5   16
  pDM 151.3-28   49.3   76
  pDM 151.3-29   19.8   30
  pDM 151.3-30   22.7   35
Figure A20071009607000221
的表达
纯化15个pDM155转化体，在盛于摇瓶里的大豆/葡萄糖培养基中培养，并于9天之后检验
Figure A20071009607000222
活性(表2)。
表II
  转化体   LU/ml   mg/ml
  pDM 155-1   669   167
  pDM 155-2   45.2   11
  pDM 155-3   180   45
  pDM 155-4   0   0
  pDM 155-5   55.4   14
  pDM 155-6   116   29
  pDM 155-7   704   176
  pDM 155-8   214   54
  pDM 155-9   17.1   4
  pDM 155-10   712   178
  pDM 155-11   511   128
  pDM 155-12   0   0
  pDM 155-13   0   0
  pDM 155-14   0   0
  pDM 155-15   153   38
  pDM 155-16   0   0
  pDM 155-17   0   0
  pDM 155-18   0   0
  pDM 155-19   129   32
  pDM 155-20   378   95
  pDM 155-21   216   54
4个转化体以大约100-200mg/l的水平(基于pNB分析)表达在小型发酵罐中培养pDM155-10，采用生产SP387时所使用的条件(见9)。7天后，出现大约2.0g/l的
Figure A20071009607000232
(图8A)。根据SDS凝胶电泳测得
Figure A20071009607000233
含有90％以上的分泌性蛋白(图8B)。
7.微生物的保藏
已将下列生物材料保存在NRRL(Agricultural ResearchService Patent Culture Collection)，Northern Regional ResearchCenter，1815，University Street，Peoria，Illinois，61604，USA。
  菌株   入藏号   保藏日期
  带有pJRoy6的大肠杆菌   NRRL B-21285   6/20/94
  带有pJRoy20的大肠杆菌   NRRL B-21418   3/10/95
  带有pDM151的大肠杆菌   NRRL B-21419   3/10/95
  带有pDM155的大肠杆菌   NRRL B-21420   3/10/95
上述菌株在这样的条件下保藏：确保在该专利申请期间，由专利商标局长指定的人可以获得其培养物，是按照37C.R.R §1.14和35U.S.C.§122和布达佩斯条约的条件确定的。该保藏物代表每种保藏菌株的生物学纯培养物。可按照向其提交了该申请的副本或后续申请的外国国家的专利法要求，向其提供保藏物。不过，应当明白，可以得到保藏物并不意味着可以实施本发明而使由政府授予的专利权丧失效力。
在这里所披露和要求保护的本发明，并不局限于具体实施方案所限定的范围，因为这些实施方案只是用于说明本发明的几个方面的。任何类似实施方案都包括在本发明范围之内。实际上，通过以上说明，除本文所披露的内容外，对本发明所做的各种改进对本领域技术人员来说都是显而易见的。这样的改进同样落在所附权利要求的范围内。
这里所引用的各对比文件所披露的内容的整体，被作为本申请的参考。
序列表
(1)一般资料：
(I)申请人：
(A)名称：Novo Nordisk Biotech，Inc.
(B)街道：1445 Drew Avenue，Ste.105
(C)城市：Davis
(D)州：California
(E)国家：US
(F)邮政编码：95616-4880
(G)电话：(916)757-8100
(H)传真：(916)758-0317
(ii)发明名称：非毒性、非产毒性、非致病性镰孢属表达***及所用启动子和终止子
(iii)序列数：16
(iv)有关地址：
(A)地址：Novo Nordisk of North America，Inc.
(B)衔道：405 Lexington Avenue，64th Floor
(C)城市：New York
(D)州：New York
(E)国家：USA
(F)邮编：10174-6401
(v)计算机可读形式：
(A)介质类型：软盘
(B)计算机：IBM PC兼容机
(C)操作***：PC-DOS/MS-DOS
(D)软件：Patent IN Release #1.0，Version #1.30
(vi)本申请数据：
(A)申请号：待定
(B)申请日：1995年6月15日
(C)分类号：
(vii)优先权申请资料
(A)申请号：US08/269,449
(B)申请日：1994年6月30日
(vii)优先权申请资料
(A)申请号：US 08/404,678
(B)申请日：1995年3月15日
(viii)律师/代理人信息
(A)姓名：Agris Dr.，Cheryl H.
(B)登记号：34,086
(C)参考/案卷号：4216.204-WO
(ix)通讯信息：
(A)电话：212-867-0123
(B)传真：212-878-9655
(2)SEQ ID NO：1的资料
(i)序列特征：
(A)长度：30bp
(B)类型：核酸
(C)链型：单链
(D)拓扑学：线性
(xi)序列表述：SEQ ID NO：1
TGCGGATCCA TGGTCAAGTT CGCTTCCGTC                                            30
(2)SEQ ID NO：2的资料
(i)序列特征：
(A)长度：30bp
(B)类型：核酸
(C)链型：单链
(D)拓扑学：线性
(xi)序列表述：SEQ ID NO：2
GACCTCGAGT TAAGCATAGG TGTCAATGAA                                            30
(2)SEQ ID NO：3的资料
(i)序列特征：
(A)长度：998bp
(B)类型：核酸
(C)链型：单链
(D)拓扑学：线性
(xi)序列表述：SEQ ID NO：3：
ATCATCAACC ACTCTTCACT CTTCAACTCT CCTCTCTTGG ATATCTATCT CTTCACCATG     60
GTCAAGTTCG CTTCCGTCGT TGCACTTGTT GCTCCCCTGG CTGCTGCCGC TCCTCAGGAG    120
ATCCCCAACA TTGTTGGTGG CACTTCTGCC AGCGCTGGCG ACTTTCCCTT CATCGTGAGC    180
ATTAGCCGCA ACGGTGGCCC CTGGTGTGGA GGTTCTCTCC TCAACGCCAA CACCGTCTTG    240
ACTGCTGCCC ACTGCGTTTC CGGATACGCT CAGAGCGGTT TCCAGATTCG TGCTGGCAGT    300
CTGTCTCGCA CTTCTGGTGG TATTACCTCC TCGCTTTCCT CCGTCAGAGT TCACCCTAGC    360
TACAGCGGAA ACAACAACGA TCTTGCTATT CTGAAGCTCT CTACTTCCAT CCCCTCCGGC    420
GGAAACATCG GCTATGCTCG CCTGGCTGCT TCCGGCTCTG ACCCTGTCGC TGGATCTTCT    480
GCCACTGTTG CTGGCTGGGG CGCTACCTCT GAGGGCGGCA GCTCTACTCC CGTCAACCTT    540
CTGAAGGTTA CTGTCCCTAT CGTCTCTCGT GCTACCTGCC GAGCTCAGTA CGGCACCTCC    600
GCCATCACCA ACCAGATGTT CTGTGCTGGT GTTTCTTCCG GTGGCAAGGA CTCTTGCCAG    660
GGTGACAGCG GCGGCCCCAT CGTCGACAGC TCCAACACTC TTATCGGTGC TGTCTCTTGG    720
GGTAACGGAT GTGCCCGACC CAACTACTCT GGTGTCTATG CCAGCGTTGG TGCTCTCCGC    780
TCTTTCATTG ACACCTATGC TTAAATACCT TGTTGGAAGC GTCGAGATGT TCCTTGAATA    840
TTCTCTAGCT TGAGTCTTGG ATACGAAACC TGTTTGAGAA ATAGGTTTCA ACGAGTTAAG    900
AAGATATGAG TTGATTTCAG TTGGATCTTA GTCCTGGTTG CTCGTAATAG AGCAATCTAG    960
ATAGCCCAAA TTGAATATGA AATTTGATGA AAATATTC                            998
(2)SEQ ID NO：4的资料：
(i)序列特征
(A)长度：248个氨基酸
(B)类型：氨基酸
(C)链型：单链
(D)拓扑学：线性
(ix)特征：
(A)名称/关键词：蛋白质
(B)位置：1..224
(ix)特征：
(A)名称/关键词：肽
(B)位置：-24..0
(D)其它资料：/product＝“OTHER”/note＝“Label＝pre-propeptide”
(xi)序列表述：SEQ ID NO：4：
Met Val Lys Phe Ala Ser Val Val Ala Leu Val Ala Pro Leu Ala Ala
                -20                 -15                 -10
Ala Ala Pro Gln Glu Ile Pro Asn Ile Val Gly Gly Thr Ser Ala Ser
            -5                  15
Ala Gly Asp Phe Pro Phe Ile Val Ser Ile Ser Arg Asn Gly Gly Pro
    10                  15                  20
Trp Cys Gly Gly Ser Leu Leu Asn Ala Asn Thr Val Leu Thr Ala Ala
25                  30                  35                  40
His Cys Val Ser Gly Tyr Ala Gln Ser Gly Phe Gln Ile Arg Ala Gly
                45                  50                  55
Ser Leu Ser Arg Thr Ser Gly Gly Ile Thr Ser Ser Leu Ser Ser Val
            60                  65                  70
Arg Val His Pro Ser Tyr Ser Gly Asn Asn Asn Asp Leu Ala Ile Leu
        75                  80                  85
Lys Leu Ser Thr Ser Ile Pro Ser Gly Gly Asn Ile Gly Tyr Ala Arg
    90                  95                  100
Leu Ala Ala Ser Gly Ser Asp Pro Val Ala Gly Ser Ser Ala Thr Val
105                 110                 115                 120
Ala Gly Trp Gly Ala Thr Ser Glu Gly Gly Ser Ser Thr Pro Val Asn
                125                 130                 135
Leu Leu Lys Val Thr Val Pro Ile Val Ser Arg Ala Thr Cys Arg Ala
            140                 145                 150
Gln Tyr Gly Thr Ser Ala Ile Thr Asn Gln Met Phe Cys Ala Gly Val
        155                 160                 165
Ser Ser Gly Gly Lys Asp Ser Cys Gln Gly Asp Ser Gly Gly Pro Ile
    170                 175                 180
Val Asp Ser Ser Asn Thr Leu Ile Gly Ala Val Ser Trp Gly Asn Gly
185                 190                 195                 200
Cys Ala Arg Pro Asn Tyr Ser Gly Val Tyr Ala Ser Val Gly Ala Leu
                205                 210                 215
Arg Ser Phe Ile Asp Thr Tyr Ala
            220
(2)SEQ ID NO：5的资料
(i)序列特征：
(A)长度：1206bp
(B)类型：核酸
(C)链型：单链
(D)拓扑学：线性
(xi)序列表述：SEQ ID NO：5
GAATTCTTAC AAACCTTCAA CAGTGGAGAC TTCCGACACG ACATATCGAT CCTTTGAAGA     60
TACGGTGAGC GTCAGATCAT GAATTTCATA CATCCTCACG TCCTTCCTCT TTCAAACTAT    120
GCAAAGTCCT TCTAGTACCT CCCAAAACTT GATTTACGCG CTCTCCAATC AAAAGTACCT    180
TCCAAAAGTG ATCTACCTCA GCTCTAGATC AGGGCACCTA TTCGCAAAGA TCTACAAGCT    240
GAACTAGTAA GCATAGCGGG AGAATATCCC ACATCATTCG AGAAGGCCTT CGTATTAGAC    300
CTAGTGGGAT CGACAGAAAA GATAAGACGG AGATAGATGC TATGTTTGGA AGGTAGGGGA    360
TGGAATAGGA TGCAACAGGT ATTGGCATAA GCGATGCAAT AGGTGCATCT AGAAACTAGG    420
TGACAGACTG GCCACAGAGG TGTATCCTAT GCAGGTCGAT GCGTGCGTTA TCGCAGGGCT    480
GCTATTGCGT GGTGGTGGCT ACAAAAGTTC TATGTGGTTT CCAGTTTCAG AATATTGGGC    540
CATTGTGATT GATGGCGCAT GACCGAATTA TAGCAGTGAA CCCCGCCCAG AGTAGTAGTG    600
CAGATGCGCT TTGATGCTTG GCGATTCCTC GGGCTAAATA ACTCCGGTTG GTCTGTAGAA    660
TGCTGACGCG ATGATCCTTC GGCATTAATC GTAGATCTTG GGGGGGGATA AGCCGATCAA    720
AGACACACTG TAGATCAGCT CTTCGATGAC TCTTACCAGC TTTATAATAA CATTCATCTT    780
GAACGTCTTT TTCGTCCAGT GTTTACCTTT CGTCCTATTT ATCCGTCATA TCCACAGTGT    840
TATTGGCGAT AGAGTTATCG ACTTTCCTCA TCGGGATACT GGCCCCTGCT GCCAAGGGCC    900
TTATATGCCG ATCACTTTCA CGGGAGCATG ATAAGGTTAA TGCTTCTTCT GAATGCCGAA    960
CTAGACTACG GAACAACGGA GCTTAGTACC AGAAAGGCAG GTACGCCTAT TCGCAAACTC   1020
CGAAGATACA ACCAAGCAAG CTTATCGCGG GATAGTAACC AGAGAGGCAG GTAAGAAGAC   1080
ACAACAACAT CCATAGCTAT GTAGATTCTC GAATATAAAA GGACCAAGAT GGACTATTCG   1140
AAGTAGTCTA TCATCAACCA CTCTTCACTC TTCAACTCTC CTCTCTTGGA TATCTATCTC   1200
TTCACC                                                              1206
(2)SEQ ID NO：6的资料：
(i)序列特征
(A)长度：1188bp
(B)类型：核酸
(C)链型：单链
(D)拓扑学：线性
(xi)序列表述：SEQ ID NO：6：
TAAATACCTT GTTGGAAGCG TCGAGATGTT CCTTGAATAT TCTCTAGCTT GAGTCTTGGA     60
TACGAAACCT GTTTGAGAAA TAGGTTTCAA CGAGTTAAGA AGATATGAGT TGATTTCAGT    120
TGGATCTTAG TCCTGGTTGC TCGTAATAGA GCAATCTAGA TAGCCCAAAT TGAATATGAA    180
ATTTGATGGA AATATTCATT TCGATAGAAG CAACGTGAAA TGTCTAGCAG GACGAAAAGT    240
AGATCAAGGC TGTTATGTTC CCCGACCAAC CTACCTTGAT GTCAGTCTGC GAGTCGTGTG    300
CAGTGACCCA GAATGATGGA TTGACTTGGA CATTTTCTGT CTATGAAGTA TTATGAACAT    360
GAATATCGTT TCCTCATTAT CTATGTTGGC AGCCTAAAGT TTTACCATAT AGCTAGCAAT    420
CAGTCAAGTA TCTGCGTATG AAGGGTTGTT AAGCCAGGAC GGTATCAGCG TTGAATATTT    480
AAAGAATGAT ATGAGATAAT CAACATTGAC ATGATAAAAG AAAAGGGGAA ACAAATTGTG    540
CATATAGTAA AGACTTCAGG TCGACCCCTC AATAGACATA TGCGAACCGA AAACCAACAG    600
GATACAATTT ATAGATAAGT ATAACTACAG TTATCTGTCT GCCGAACAAA TACTCTTTTG    660
TGAAACAAAT GAAGAGTACA TAAGCTACAG TTCCTCAGTA GGAACATCCT TTACAATAAC    720
TCCCTTGACT TCCTTCAGCT TCTCAATAGC CTCCAAAGTC ATCGGTCTGC CATCAAGGCA    780
CGTCAGCTCT GGTGTAGCAT ACAGCAGTGC CATACTTACG GAGGATAGGA AGTGGGAGGA    840
ATCGTTCGTG TCTGCCTCCA AAAATCGACA CCAGTGTCCT TTTTGACGAT ACTGATATGG    900
TGGTAAGCTT GGGAGTCTAT TGTTGACGTT GCATCACTTA CTTAAGCACG GTTTCATTCC    960
TCTGCTGATA GTCCTCCAAC TTCTCGAAGT CGTAAACGAT GGCCTATAGT ATCTTATTGA   1020
GAAATATGTC TTCTCAGAAA ATTATATCTT GTTTACCTTT CGGTCCGCCA TGGCTGCTAA   1080
AACTGCTGGG AAATTCAAAA GCGCAGCACA AGCAGCAAGA GTGATGGGCA CAACGTGATA   1140
TGTTGATAAA AGCATCAGTA TCGATAAGTT CCACTCAGAA ACCTGCAG                1188
(2)SEQ ID NO：7的资料
(i)序列特征：
(A)长度：1060bp
(B)类型：核酸
(C)链型：单链
(D)拓扑学：线性
(ix)特征：
(A)名称/关键词：CDS
(B)位置：10..924
(ix)特征：
(A)名称/关键词：：mat-peptide
(B)位置：73..924
(ix)特征：
(A)名称/关键词：sig-peptide
(B)位置：10..72
(xi)序列表述：SEQ ID NO：7：
GGATCCAAG ATG CGT TCC TCC CCC CTC CTC CCG TCC GCC GTT GTG GCC       48
          Met Arg Ser Ser Pro Leu Leu Pro Ser Ala Val Val Ala
          -21 -20                 -15                 -10
GCC CTG CCG GTG TTG GCC CTT GCC GCT GAT GGC AGG TCC ACC CGC TAC     96
Ala Leu Pro Val Leu Ala Leu Ala Ala Asp Gly Arg Ser Thr Arg Tyr
             -5                   1               5
TGG GAC TGC TGC AAG CCT TCG TGC GGC TGG GCC AAG AAG GCT CCC GTG    144
Trp Asp Cys Cys Lys Pro Ser Cys Gly Trp Ala Lys Lys Ala Pro Val
     10                  15                  20
AAC CAG CCT GTC TTT TCC TGC AAC GCC AAC TTC CAG CGT ATC ACG GAC    192
Asn Gln Pro Val Phe Ser Cys Asn Ala Asn Phe Gln Arg Ile Thr Asp
 25                  30                  35                  40
TTC GAC GCC AAG TCC GGC TGC GAG CCG GGC GGT GTC GCC TAC TCG TGC    240
Phe Asp Ala Lys Ser Gly Cys Glu Pro Gly Gly Val Ala Tyr Ser Cys
                 45                  50                  55
GCC GAC CAG ACC CCA TGG GCT GTG AAC GAC GAC TTC GCG CTC GGT TTT    288
Ala Asp Gln Thr Pro Trp Ala Val Asn Asp Asp Phe Ala Leu Gly Phe
             60                  65                  70
GCT GCC ACC TCT ATT GCC GGC AGC AAT GAG GCG GGC TGG TGC TGC GCC    336
Ala Ala Thr Ser Ile Ala Gly Ser Asn Glu Ala Gly Trp Cys Cys Ala
         75                  80                  85
TGC TAC GAG CTC ACC TTC ACA TCC GGT CCT GTT GCT GGC AAG AAG ATG    384
Cys Tyr Glu Leu Thr Phe Thr Ser Gly Pro Val Ala Gly Lys Lys Met
     90                  95                 100
GTC GTC CAG TCC ACC AGC ACT GGC GGT GAT CTT GGC AGC AAC CAC TTC    432
Val Val Gln Ser Thr Ser Thr Gly Gly Asp Leu Gly Ser Asn His Phe
105                 110                 115                 120
GAT CTC AAC ATC CCC GGC GGC GGC GTC GGC ATC TTC GAC GGA TGC ACT    480
Asp Leu Asn Ile Pro Gly Gly Gly Val Gly Ile Phe Asp Gly Cys Thr
                125                 130                 135
CCC CAG TTC GGC GGT CTG CCC GGC CAG CGC TAC GGC GGC ATC TCG TCC    528
Pro Gln Phe Gly Gly Leu Pro Gly Gln Arg Tyr Gly Gly Ile Ser Ser
            140                 145                 150
CGC AAC GAG TGC GAT CGG TTC CCC GAC GCC CTC AAG CCC GGC TGC TAC    576
Arg Asn Glu Cys Asp Arg Phe Pro Asp Ala Leu Lys Pro Gly Cys Tyr
        155                 160                 165
TGG CGC TTC GAC TGG TTC AAG AAC GCC GAC AAT CCG AGC TTC AGC TTC    624
Trp Arg Phe Asp Trp Phe Lys Asn Ala Asp Asn Pro Ser Phe Ser Phe
    170                 175                 180
CGT CAG GTC CAG TGC CCA GCC GAG CTC GTC GCT CGC ACC GGA TGC CGC    672
Arg Gln Val Gln Cys Pro Ala Glu Leu Val Ala Arg Thr Gly Cys Arg
185                 190                 195                 200
CGC AAC GAC GAC GGC AAC TTC CCT GCC GTC CAG ATC CCC TCC AGC AGC    720
Arg Asn Asp Asp Gly Asn Phe Pro Ala Val Gln Ile Pro Ser Ser Ser
                205                 210                 215
ACC AGC TCT CCG GTC AAC CAG CCT ACC AGC ACC AGC ACC ACG TCC ACC    768
Thr Ser Ser Pro Val Asn Gln Pro Thr Ser Thr Ser Thr Thr Ser Thr
            220                 225                 230
TCC ACC ACC TCG AGC CCG CCA GTC CAG CCT ACG ACT CCC AGC GGC TGC    816
Ser Thr Thr Ser Ser Pro Pro Val Gln Pro Thr Thr Pro Ser Gly Cys
        235                 240                 245
ACT GCT GAG AGG TGG GCT CAG TGC GGC GGC AAT GGC TGG AGC GGC TGC    864
Thr Ala Glu Arg Trp Ala Gln Cys Gly Gly Asn Gly Trp Ser Gly Cys
    250                 255                 260
ACC ACC TGC GTC GCT GGC AGC ACT TGC ACG AAG ATT AAT GAC TGG TAC    912
Thr Thr Cys Val Ala Gly Ser Thr Cys Thr Lys Ile Ash Asp Trp Tyr
265                 270                 275                 280
CAT CAG TGC CTG TAGACGCAGG GCAGCTTGAG GGCCTTACTG GTGGCCGCAA        964
His Gln Cys Leu
CGAAATGACA CTCCCAATCA CTGTATTAGT TCTTGTACAT AATTTCGTCA TCCCTCCAGG 1024
GATTGTCACA TAAATGCAAT GAGGAACAAT GAGTAC                           1060
(2)SEQ ID NO：8的资料
(i)序列特征：
(A)长度：305个氨基酸
(B)类型：氨基酸
(D)拓扑学：线性
(ii)分子类型：蛋白质
(xi)序列表述：SEQ ID NO：8：
Met Arg Ser Ser Pro Leu Leu Pro Ser Ala Val Val Ala Ala Leu Pro
-21 -20                 -15                 -10
Val Leu Ala Leu Ala Ala Asp Gly Arg Ser Thr Arg Tyr Trp Asp Cys
 -5                   1               5                  10
Cys Lys Pro Ser Cys Gly Trp Ala Lys Lys Ala Pro Val Asn Gln Pro
             15                  20                  25
Val Phe Ser Cys Asn Ala Asn Phe Gln Arg Ile Thr Asp Phe Asp Ala
         30                  35                  40
Lys Ser Gly Cys Glu Pro Gly Gly Val Ala Tyr Ser Cys Ala Asp Gln
     45                  50                  55
Thr Pro Trp Ala Val Asn Asp Asp Phe Ala Leu Gly Phe Ala Ala Thr
 60                  65                  70                  75
Ser Ile Ala Gly Ser Asn Glu Ala Gly Trp Cys Cys Ala Cys Tyr Glu
                 80                  85                  90
Leu Thr Phe Thr Ser Gly Pro Val Ala Gly Lys Lys Met Val Val Gln
             95                 100                 105
Ser Thr Ser Thr Gly Gly Asp Leu Gly Ser Asn His Phe Asp Leu Asn
        110                 115                 120
Ile Pro Gly Gly Gly Val Gly Ile Phe Asp Gly Cys Thr Pro Gln Phe
    125                 130                 135
Gly Gly Leu Pro Gly Gln Arg Tyr Gly Gly Ile Ser Ser Arg Asn Glu
140                 145                 150                 155
Cys Asp Arg Phe Pro Asp Ala Leu Lys Pro Gly Cys Tyr Trp Arg Phe
                160                 165                 170
Asp Trp Phe Lys Asn Ala Asp Asn Pro Ser Phe Ser Phe Arg Gln Val
            175                 180                 185
Gln Cys Pro Ala Glu Leu Val Ala Arg Thr Gly Cys Arg Arg Asn Asp
        190                 195                 200
Asp Gly Asn Phe Pro Ala Val Gln Ile Pro Ser Ser Ser Thr Ser Ser
    205                 210                 215
Pro Val Asn Gln Pro Thr Ser Thr Ser Thr Thr Ser Thr Ser Thr Thr
220                 225                 230                 235
Ser Ser Pro Pro Val Gln Pro Thr Thr Pro Ser Gly Cys Thr Ala Glu
                240                 245                 250
Arg Trp Ala Gln Cys Gly Gly Asn Gly Trp Ser Gly Cys Thr Thr Cys
            255                 260                 265
Val Ala Gly Ser Thr Cys Thr Lys Ile Asn Asp Trp Tyr His Gln Cys
        270                 275                 280
Leu
(2)SEQ ID NO：9的资料：
(i)序列特征：
(A)长度：876bp
(B)类型：核酸
(C)链型：单链
(D)拓扑学：线性
(xi)序列表述：SEQ ID NO：9：
ATGAGGAGCT CCCTTGTGCT GTTCTTTGTC TCTGCGTGGA CGGCCTTGGC CAGTCCTATT     60
CGTCGAGAGG TCTCGCAGGA TCTGTTTAAC CAGTTCAATC TCTTTGCACA GTATTCTGCA    120
GCCGCATACT GCGGAAAAAA CAATGATGCC CCAGCTGGTA CAAACATTAC GTGCACGGGA    180
AATGCCTGCC CCGAGGTAGA GAAGGCGGAT GCAACGTTTC TCTACTCGTT TGAAGACTCT    240
GGAGTGGGCG ATGTCACCGG CTTCCTTGCT CTCGACAACA CGAACAAATT GATCGTCCTC    300
TCTTTCCGTG GCTCTCGTTC CATAGAGAAC TGGATCGGGA ATCTTAACTT CGACTTGAAA    360
GAAATAAATG ACATTTGCTC CGGCTGCAGG GGACATGACG GCTTCACTTC GTCCTGGAGG    420
TCTGTAGCCG ATACGTTAAG GCAGAAGGTG GAGGATGCTG TGAGGGAGCA TCCCGACTAT    480
CGCGTGGTGT TTACCGGACA TAGCTTGGGT GGTGCATTGG CAACTGTTGC CGGAGCAGAC    540
CTGCGTGGAA ATGGGTATGA TATCGACGTG TTTTCATATG GCGCCCCCCG AGTCGGAAAC    600
AGGGCTTTTG CAGAATTCCT GACCGTACAG ACCGGCGGAA CACTCTACCG CATTACCCAC    660
ACCAATGATA TTGTCCCTAG ACTCCCGCCG CGCGAATTCG GTTACAGCCA TTCTAGCCCA    720
GAGTACTGGA TCAAATCTGG AACCCTTGTC CCCGTCACCC GAAACGATAT CGTGAAGATA    780
GAAGGCATCG ATGCCACCGG CGGCAATAAC CAGCCTAACA TTCCGGATAT CCCTGCGCAC    840
CTATGGTACT TCGGGTTAAT TGGGACATGT CTTTAG                              876
(2)SEQ ID NO：10的资料
(i)序列特征：
(A)长度：291个氨基酸
(B)类型：氨基酸
(C)链型：单链
(D)拓扑学：线性
(xi)序列表述：SEQ ID NO：10：
Met Arg Ser Ser Leu Val Leu Phe Phe Val Ser Ala Trp Thr Ala Leu
1               5                   10                  15
Ala Ser Pro Ile Arg Arg Glu Val Ser Gln Asp Leu Phe Asn Gln Phe
            20                  25                  30
Asn Leu Phe Ala Gln Tyr Ser Ala Ala Ala Tyr Cys Gly Lys Asn Asn
        35                  40                  45
Asp Ala Pro Ala Gly Thr Asn Ile Thr Cys Thr Gly Asn Ala Cys Pro
    50                  55                  60
Glu Val Glu Lys Ala Asp Ala Thr Phe Leu Tyr Ser Phe Glu Asp Ser
65                  70                  75                  80
Gly Val Gly Asp Val Thr Gly Phe Leu Ala Leu Asp Asn Thr Asn Lys
                85                  90                  95
Leu Ile Val Leu Ser Phe Arg Gly Ser Arg Ser Ile Glu Asn Trp Ile
            100                 105                 110
Gly Asn Leu Asn Phe Asp Leu Lys Glu Ile Asn Asp Ile Cys Ser Gly
        115                 120                 125
Cys Arg Gly His Asp Gly Phe Thr Ser Ser Trp Arg Ser Val Ala Asp
    130                 135                 140
Thr Leu Arg Gln Lys Val Glu Asp Ala Val Arg Glu His Pro Asp Tyr
145                 150                 155                 160
Arg Val Val Phe Thr Gly His Ser Leu Gly Gly Ala Leu Ala Thr Val
                165                 170                 175
Ala Gly Ala Asp Leu Arg Gly Asn Gly Tyr Asp Ile Asp Val Phe Ser
            180                 185                 190
Tyr Gly Ala Pro Arg Val Gly Asn Arg Ala Phe Ala Glu Phe Leu Thr
        195                 200                 205
Val Gln Thr Gly Gly Thr Leu Tyr Arg Ile Thr His Thr Asn Asp Ile
    210                 215                 220
Val Pro Arg Leu Pro Pro Arg Glu Phe Gly Tyr Ser His Ser Ser Pro
225                 230                 235                 240
Glu Tyr Trp Ile Lys Ser Gly Thr Leu Val Pro Val Thr Arg Asn Asp
                245                 250                 255
Ile Val Lys Ile Glu Gly Ile Asp Ala Thr Gly Gly Asn Asn Gln Pro
            260                 265                 270
Asn Ile Pro Asp Ile Pro Ala His Leu Trp Tyr Phe Gly Leu Ile Gly
        275                 280                 285
Thr Cys Leu
    290
(2)SEQ ID NO：11的资料
(i)序列特征：
(A)长度：42bp
(B)类型：核酸
(C)链型：单链
(D)拓扑学：线性
(xi)序列表述：SEQ ID NO：11：
GCACACCATG GTCGCTGGAT CCATACCTTG TTGGAAGCGT CG                                   42
(2)SEQ ID NO：12的资料
(i)序列特征
(A)长度：56bp
(B)类型：核酸
(C)链型：单链
(D)拓扑学：线性
(xi)序列表述：SEQ ID NO：12：
ATCGGAGCAT GCGGTACCGT TTAAACGAAT TCAGGTAAAC AAGATATAAT TTTCTG                    56
(2)SEQ ID NO：13的资料
(i)序列特征
(A)长度：44bp
(B)类型：核酸
(C)链型：单链
(D)拓扑学：线性
(xi)序列表述：SEQ ID NO：13：
CTCTTGGATA TCTATCTCTT CACCATGCGT TCCTCCCCCC TCCT                                 44
(2)SEQ ID NO：14的资料
(i)序列特征
(A)长度：20bp
(B)类型：核酸
(C)链型：单链
(D)拓扑学：线性
(xi)序列表述：SEQ ID NO：14：
CAATAGAGGT GGCAGCAAAA                                                       20
(2)SEQ ID NO：15的资料
(i)序列特征：
(A)长度：25bp
(B)类型：核酸
(C)链型：单链
(D)拓扑学：线性
(xi)序列表述：SEQ ID NO：15：
ATCTATCTCT TCACCATGAG GAGCT                                                25
(2)SEQ ID NO：16的资料
(i)序列特征
(A)长度：21bp
(B)类型：核酸
(C)链型：单链
(D)拓扑学：线性
(xi)序列表述：SEQ ID NO：16：
TAGATAGAGA AGTGGTACTC C                                                    21

Claims (28)

1.一种非毒性、非产毒性、非致病性重组镰孢属(Fusarium)宿主细胞，它包括一个与一个启动子可操纵连接的编码一种分泌型异源蛋白的核酸序列。
2.如权利要求1的宿主细胞，其中，所述镰孢菌具有镰孢菌ATCC 20334的鉴定特征。
3.如权利要求1的宿主细胞，其中，所述分泌型异源蛋白是一种真菌蛋白。
4.如权利要求1的宿主细胞，其中，所述异源蛋白是一种真菌酶。
5.如权利要求4的宿主细胞，其中，所述真菌酶选自过氧化氢酶、漆酶、酚氧化酶、氧化酶、氧化还原酶、纤维素酶、木聚糖酶、过氧化物酶、脂肪酶、水解酶、酯酶、角质酶、蛋白酶、氨肽酶、羧肽酶、肌醇六磷酸酶、裂合酶、溶果胶酶、淀粉酶、葡糖淀粉酶、α-半乳糖苷酶、β-半乳糖苷酶、α-葡糖苷酶、β-葡糖苷酶、甘露糖苷酶、异构酶、转化酶、转移酶、核糖核酸酶、壳多糖酶和脱氧核糖核酸酶。
6.如权利要求4的宿主细胞，其中，所述真菌酶是一种蛋白酶。
7.如权利要求4的宿主细胞，其中，所述真菌酶是一种碱性蛋白酶。
8.如权利要求7的宿主细胞，其中，所述碱性蛋白酶是尖镰孢(Fusarium oxysporum)类胰蛋白酶蛋白酶。
9.如权利要求8的宿主细胞，其中，所述尖镰孢类胰蛋白酶蛋白酶的氨基酸序列如SEQ ID NO：4所示。
10.如权利要求4的宿主细胞，其中，所述真菌酶是一种内切葡聚糖酶。
11.如权利要求10的宿主细胞，其中，所述内切葡聚糖酶的氨基酸序列如SEQ ID NO：8所示。
12.如权利要求4的宿主细胞，其中，所述真菌酶是1，3-特异性脂肪酶。
13.如权利要求12的宿主细胞，其中，所述1，3-特异性脂肪酶的氨基酸序列如SEQ ID NO：10所示。
14.如权利要求1的宿主细胞，其中，所述异源蛋白选自激素，生长因子和受体。
15.如权利要求1的宿主细胞，其中，所述启动子是一种真菌启动子。
16.如权利要求15的宿主细胞，其中，所述启动子选自来自构巢曲霉(A.nidulans)amdS的启动子和来自编码TPI、ADH、GAPDH和PGK的基因的启动子。
17.如权利要求15的宿主细胞，其中，所述真菌启动子来自一个编码尖镰孢类胰蛋白酶蛋白酶的基因，其中编码所述蛋白酶的一般序列如SEQ ID NO：3所示。
18.如权利要求17的宿主细胞，其中，所述启动子序列如SEQID NO：5所示。
19.如权利要求1的宿主细胞，它还包括一个选择标记。
20.如权利要求19的宿主细胞，其中，所述选择标记选自arg B、trpC、pyrG、amdS、niaD和hygB。
21.如权利要求1的宿主细胞，它还包括一个终止子。
22.如权利要求21的宿主细胞，其中，所述终止子来自一个编码尖镰孢类胰蛋白酶蛋白酶的基因，其中所述蛋白酶的一般序列如SEQ ID NO：3所示。
23.如权利要求22的宿主细胞，其中，所述终止子序列如SEQID NO：6所示。
24.一种生产目的蛋白质的方法，它包括培养一种非毒性、非产毒性、非致病性重组镰孢属宿主细胞，该宿主细胞包括一个与一个启动子可操纵连接的编码所述蛋白质的核酸，以及分离所述蛋白质。
25.一种来自一个编码尖镰孢类胰蛋白酶蛋白酶的基因的真菌启动子，其中，编码所述蛋白酶的一般序列如SEQ ID NO：3所示。
26.权利要求25的真菌启动子，其中所述启动子具有SEQ IDNO：5所示的DNA序列。
27.一种来自一个编码尖镰孢类胰蛋白酶蛋白酶的基因的终止子序列，其中，编码所述蛋白酶的一般序列如SEQ ID NO：3所示。
28.权利要求27的真菌终止子，其中所述终止子具有SEQ IDNO：6所示的DNA序列。
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