CN110527676A - 多肽,分离的其多核苷酸,以及包含多肽的添加剂、其用途及方法 - Google Patents
多肽,分离的其多核苷酸,以及包含多肽的添加剂、其用途及方法 Download PDFInfo
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- CN110527676A CN110527676A CN201910861350.XA CN201910861350A CN110527676A CN 110527676 A CN110527676 A CN 110527676A CN 201910861350 A CN201910861350 A CN 201910861350A CN 110527676 A CN110527676 A CN 110527676A
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Abstract
以水解方式裂解玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物的多肽,其是具有选自SEQ ID No.1‑15的氨基酸序列或其功能性变体的水解酶,其中在所述功能性变体与至少一个所述氨基酸序列之间的序列同一性为至少40%;以及包含所述多肽的添加剂;以及编码所述多肽的分离的多核苷酸;以及用于用所述多肽来以水解方式裂解玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物的方法。
Description
本申请是申请日为2014年8月27日、申请号为201480055221.7、发明名称为“用于以水解方式裂解玉米赤霉烯酮和/或玉米赤霉烯酮衍生物的多肽,分离的其多核苷酸,以及包含多肽的添加剂、其用途及方法”的发明专利申请的分案申请。
本发明涉及用于以水解方式裂解玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物的多肽,编码此类多肽的分离的多核苷酸,以及包含此类多肽的添加剂,还涉及此类多肽的用途,以及涉及用于以水解方式裂解玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物的方法。
真菌毒素是丝状真菌产生的次生代谢物。其一个重要的代表是世界范围传播的玉米赤霉烯酮(ZEN)(以前称为F-2毒素),其由许多镰孢菌真菌产生。这些真菌尤其侵染栽培植物,例如各种各样的谷类,其中真菌侵染通常出现在收获之前,其中真菌生长或真菌毒素产生可以在收获之前发生,或者在不适当的贮藏的情况下也可以在收获之后发生。FAO估计,在世界范围内25%的农产品被真菌毒素污染,这导致巨大的经济损失。在一项最近在世界范围内进行的研究中,从2009年1月至2011年12月分析了总共23,781个样品,其中81%经测试为对于至少一种真菌毒素来说阳性和45%对于ZEN来说阳性。可以在世界的所有地区中,同样地在所有经测试的谷物类别和饲料类别,例如玉米、大豆粉、小麦、麦麸、DDGS(干酒糟)中,以及在预制饲料混合物中,以直至100%的频率发现ZEN。
ZEN是一种非甾类的、雌激素的、大环的、通过聚酮化合物物质代谢途径合成的内酯,其具有下述结构式:
和IUPAC命名名称“(2E,11S)-15,17-二羟基-11–甲基-12-氧杂双环[12.4.0]十八碳-1(18),2,14,16-四烯-7,13-二酮”。
然而,在自然界中还存在有许多ZEN衍生物,其通过ZEN的酶促或化学修饰而形成。关于此的例子为糖苷式的或含硫酸酯的ZEN缀合物,其由真菌、植物或哺乳动物代谢来形成;以及ZEN代谢物,其尤其在人或动物机体中形成。在下文中,ZEN衍生物是指在自然界中存在的或者通过化学或生物化学合成而制备的ZEN缀合物或ZEN代谢物,但特别地是指α-玉米赤霉烯醇(α-ZEL;(2E,7R,11S)-7,15,17-三羟基-11-甲基-12-氧杂双环[12.4.0]十八碳-1(18),2,14,16-四烯-13-酮)、β-玉米赤霉烯醇(β-ZEL;(2E,7S,11S)-7,15,17-三羟基-11-甲基-12-氧杂双环[12.4.0]十八碳-1(18),2,14,16-四烯-13-酮)、α-玉米赤霉醇(α-ZAL;(7R,11S)-7,15,17-三羟基-11-甲基-12-氧杂双环[12.4.0]十八碳-1(18),14,16-三烯-13-酮)、β-玉米赤霉醇(β-ZAL;(7S,11S)-7,15,17-三羟基-11-甲基-12-氧杂双环[12.4.0]十八碳-1(14),15,17-三烯-13-酮)、玉米赤霉烯酮-14-硫酸酯(Z14S;[(2E,11S)-15-羟基-11-甲基-7,13-二氧代-12-氧杂双环[12.4.0]十八碳-1(18),2,14,16-四烯-17-基]硫酸氢酯)、玉米赤霉烯酮-14-糖苷(Z14G;(2E,11S)-15-羟基-11-甲基-17-[(3R,4S,5S,6R)-3,4,5-三羟基-6-(羟甲基)四氢吡喃-2-基]氧基-12-氧杂双环[12.4.0]十八碳-1(18),2,14,16-四烯-7,13-二酮)以及玉米赤霉酮(ZAN;(11S)-15,17-二羟基-11-甲基-12-氧杂双环[12.4.0]十八碳-1(18),14,16-三烯-7,13-二酮)。
ZEN,同样地还有ZEN衍生物,尤其是α-ZEL、β-ZEL、Z14S、α-ZAL、β-ZAL、Z14G和ZAN,由于其高的化学和物理稳定性而还可以在经加工的食品或饲料例如面包或啤酒中检测到。
ZEN与雌激素受体结合并且可以引起激素紊乱,其中它在经口摄取后立即被吸收,并且被哺乳动物转变为两个立体异构的代谢物,分别为α-ZEL和β-ZEL。在此,例如α-ZEL,还有α-ZAL或ZAN具有比ZEN强得多的动情效应。经缀合的ZEN衍生物有时具有比ZEN低的雌激素活性,然而如果情况可能,ZEN可以从这些ZEN衍生物中重新被释放到消化道中。
虽然ZEN具有相对低的急性毒性并且具有直至20,000mg/kg体重的口服LD50值,但是在较长期摄入的情况下在动物或人中出现亚急性和/或亚慢性的毒性效应,例如致畸形的、致癌的、动情的和免疫抑制的效应。被ZEN污染的饲料导致哺乳动物中的发育障碍,其中猪,特别是幼猪对于ZEN是极其敏感的。超过0.5ppm的在饲料中的ZEN浓度导致发育障碍,其中例如超过1.5ppm的浓度可以导致猪中的超雌激素活性,和12ppm ZEN的浓度被认为对牛中的流产负责。由于玉米赤霉烯酮通过粘膜,特别是通过胃粘膜,但也通过口腔粘膜被迅速吸收,因此立即的和尤其是定量的钝化是必要的。在ZEN的口服给药后30分钟已经可以在血液中检测到它们。在这种情况下,使用经分离的针对微生物的酶具有优点,例如更高的比活性或更快的作用。由于ZEN的有害效应,因而在欧盟存在有在食品中的强制性的ZEN上限以及对于在饲料中的ZEN上限的推荐(EC NO:1881/2006)。
为了减少食品或饲料的ZEN污染的最初策略是限制真菌生长,例如通过遵循“良好的农业实践经验”。这尤其包括,种子没有害虫和真菌侵染,或者及时从田地中移除农业废料。此外,还可以通过使用杀真菌剂来减少田地上的真菌生长。在收获之后,应当将收获物储藏在低于15%的残留水分含量和低的温度下,以阻止真菌生长。同样地,应当在再加工之前去除由于真菌侵染而被污染的物料。尽管有着这一系列措施,但I.Rodriges和K.Naehrer(2012)仍然报道了,即使在具有最高农业标准的地区,例如USA和中欧,在2009年至2011年中,分别有29%和39%的经测试的玉米样品被ZEN污染。
用于从饲料或食品中去除ZEN的其他可能性为真菌毒素的吸附或转化。为此所必需的是,真菌毒素与吸附剂的结合在宽的pH范围内是强的且特异的,并且在胃肠区域内的整个消化过程中保持稳定。虽然对于黄曲霉毒素可以有效地使用几种非生物吸附剂,例如活性炭、硅酸盐或合成的聚合物,例如消胆胺,但是它们用于其他真菌毒素则受到限制。吸附剂的主要缺点是其他有时候对于营养供给来说必需的分子的非特异性结合。在文献中同样也描述了生物吸附剂,例如酵母或酵母提取物,然而生物吸附剂像非生物吸附剂一样也具有类似的限制。
通过物理和化学处理来对ZEN进行解毒也受到限制。通过热处理,不能有效地使ZEN钝化,但是可以通过挤出和用氧化剂进行处理,例如在80℃下用10%的过氧化氢溶液处理16小时,来使ZEN含量降低83.9%。在饲料和食品的制备中挤出方法和氧化剂例如臭氧或过氧化氢的使用,由于高的成本、品质损失、有时候低的效率和低的特异性而受限。
借助于微生物例如噬真菌毒素丝孢酵母(Trichosporon mycotoxinivorans)、粉红粘帚霉(Gliocladium roseum)或枯草芽孢杆菌(Bacillus subtilis)菌株或者从其中分离出的酶例如水解酶或过氧化物酶来进行ZEN的生物转化也例如在E.Vekiru等人,Appl.and Environ.Microb.,2010,76,7,2353-2359中进行了描述。
从EP 0 938 575 B1中知晓了红球菌属(Rhodococcus)和诺卡氏菌属(Nocardia)的细菌,特别是球状红球菌(R.globerulus)、红串红球菌(R.erythropolis)和球形诺卡氏菌(N.globerula)的ZEN降解特性。
从WO 02/076205中可获知从粉红粘帚霉中分离出的酶的ZEN降解作用,所述酶尤其为α/β-水解酶、玉米赤霉烯酮水解酶1(ZHD1),其借助于催化三元体来催化ZEN的降解。
从WO 2012/113827中获知重组Zonase,即ZEN降解酶,其在胃肠道中保持稳定,特别地在其中描述了微生物例如褐色双歧嗜热菌(Thermobifidia fusca)、脱叶链霉菌(Streptomyces exfoliatus)、德氏食酸菌(Acidovorans delafieldii)和链霉菌属物种(Streptomyces sp.)。
能够水解ZEN和/或至少一种ZEN衍生物的多肽或酶也可以称为Zonase。
下面所使用的术语取自专业用语并且总是以传统的意义进行使用,除非另外指出。因此,术语“多核苷酸”涉及具有所有长度和序列的每一种类型的遗传物质,例如单链和双链DNA和RNA分子,包括调控元件、结构元件、基因群、质粒、全基因组及其片段。名称“多肽”包括蛋白质,例如酶、抗体,以及具有直至500个氨基酸的多肽,例如肽抑制剂,蛋白质的结构域,还有具有小的序列长度例如少于10个氨基酸的短多肽,例如受体、配体、肽激素、标签等。名称“在多核苷酸或多肽中的‘位置’”涉及在所述多核苷酸或多肽的序列中单个特定的碱基或氨基酸。
现在,本发明旨在提供这样的多肽以供使用,用所述多肽可成功实现将ZEN和/或至少一种ZEN衍生物快速地且可靠地转化为经水解的ZEN和/或经水解的ZEN衍生物。为了解决该任务,本发明的特征主要在于,所述多肽是具有选自SEQ ID No.1-15的氨基酸序列或其功能性变体的水解酶,其中在所述功能性变体与至少一个所述氨基酸序列之间的序列同一性为至少40%。
根据本发明,名称“序列同一性”涉及序列同一性百分比。对于氨基酸序列和核苷酸序列,序列同一性可以目视测定,但是优选地用计算机程序来计算出来。还在序列区段内进行序列比较,其中参考序列的连续序列被理解为区段,并且优选地包含该序列的保守区域。
在当前的情况下,序列同一性借助于程序NCBI BLAST(Basic Local AlignmentSearch Tool)来进行测定,特别地对于多肽,用BLASTP,和对于多核苷酸,用BLASTN,其提供在“National Center for Biotechnology Information”的主页(NCBI;http://www.ncbi.nlm.nih.gov/)上以供使用。因此,可能的是,根据Altschul等人,1997(NucleicAcids Res.,25:3389-3402)的算法,将两个或更多个序列互相进行比较。为了本发明的目的,使用2013年5月15日的那个版本的程序。作为程序设置,考虑基本设置,但是特别地,对于氨基酸序列比较:“最大靶序列(max target sequence)”=100;“期望阈值(expectedthreshold)”=10;“字长(word size)”=3;“矩阵(matrix)”=BLOSOM62;“缺口成本(gapcosts)”=“存在(Existence):11;延伸(Extention):1”;“计算调整(computationaladjustment)”=“条件式组成得分矩阵调整(Conditional compositional score matrixadjustment)”;以及对于核苷酸序列比较:字长:11;期望值(Expect value):10;缺口成本:存在=5,延伸=2;过滤器(Filter)=低复杂度激活的(low complexity activated);匹配/错配得分(Match/Mismatch Scores):2,-3;过滤器字符串(Filter String):L;m。
术语“功能性多肽变体”或“功能性变体”一方面涉及多肽的“等位基因变体”和多肽的“功能性片段”,另一方面涉及多肽的“修饰”,其中酶促功能基本上未改变。名称“等位基因变体”涉及这样的多肽,所述多肽通过在自然界中随机发生的核苷酸序列突变而产生并且引起氨基酸序列的改变,其中其酶促功能未受影响。“修饰”可以例如是与多肽的C-末端或N-末端融合物或者经突变的多肽,其中突变可以通过至少一个氨基酸的置换、插入或缺失来获得,这特别地通过位点特异性诱变或随机诱变、重组和/或任何其他蛋白质工程方法来进行。术语“置换”、“插入”和“缺失”以在基因技术中通常的和专业人员熟悉的含义来进行使用。术语“功能性片段”涉及多肽的部分或部分序列或者其功能性变体的部分或部分序列,其中酶促功能基本上得到保留。当酶促反应机制保持不变,即在相同的位置处水解该真菌毒素,并且“功能性变体”相对于原始多肽而言的比残余活性为至少5%,优选地至少10%,特别地至少50%时,那么基本上保留了酶促功能。具有有着SEQ ID No.1-15的氨基酸序列的多肽是另一个或者同一个酶的功能性等位基因变体,其中所述序列各自来源于不同的微生物。这从下列中可清楚地看出来:借助于序列同一性百分比而测量出的接近的相互亲缘关系,以及这样的事实,即所有多肽通过相同的降解机制作用于ZEN和ZEN衍生物。
由于具有SEQ ID No.1-15的多肽的氨基酸序列相互间的相似性,因而可能的是,这些多肽之一的功能性变体与多于一个所要求保护的具有SEQ ID No.1-15的多肽具有至少40%的序列同一性。
通过选择此类氨基酸序列或其功能性变体,查明了令人惊讶地快速且完全的ZEN和/或至少一种ZEN衍生物的水解。
如相应于本发明的一个优选的进一步发展方案那样,所述多肽具有这样的氨基酸序列,其包含至少一个保守氨基酸序列区段或其功能性变体,其中所述氨基酸序列区段的功能性变体具有至少70%,优选地至少84%,更优选地至少92%,和最优选地至少98%的序列同一性,并且所述至少一个保守氨基酸序列区段选自具有SEQ ID No.1的序列的氨基酸序列+24至+50、+52至+77、+79至+87、+89至+145、+150至+171、+177至+193、+223至+228、+230至+237、+239至+247、+249至+255、+257至+261、+263至+270、+272至+279、+297至+301、+303至+313、+24至328、+1至+328。通过至少一个此类保守氨基酸序列区段的存在,可成功提供这样的多肽以供使用,所述多肽除了ZEN和/或至少一种ZEN衍生物的快速且完全的水解外,还具有相比于迄今已知的ZEN降解多肽而言特别高的活性值。
当如相应于本发明的一个进一步改造方案那样,所述功能性变体具有至少一个选自一个或多个氨基酸的置换、缺失和插入的氨基酸修饰时,依然可以取得良好的结果。
通过如此地进一步改造本发明,即使得所述多肽具有至少0.01U/mg,优选地至少0.1U/mg,特别地至少1U/mg的比活性,和/或具有最高50μΜ,优选地最高3.5μΜ,特别地最高0.5μΜ的以水解方式裂解ZEN的KM值,和/或具有至少0.05s-1,优选地至少0.6s-1,特别地至少5s-1的以水解方式裂解ZEN的kcat值,和/或具有至少0.00001μΜ-1s-1,优选地至少0.0001μΜ-1s-1,特别地至少0.001μΜ-1s-1的以水解方式裂解ZEN的vmax值,ZEN和/或ZEN衍生物可以被特别快速且完全地水解,特别是解毒。
如相应于本发明的一个优选的进一步发展方案那样,所述多肽包含选自SEQ IDNo.2、5-7、9、11、12和15的氨基酸序列或其功能性变体,其中所述功能性变体与至少一个所述氨基酸序列具有至少40%的序列同一性,并且所述多肽在pH 5.0下的pH稳定性为至少15%,优选地50%,和特别优选地至少90%。通过这样的进一步发展方案可以确保,所述多肽在酸性介质中,因此例如在哺乳动物的胃中的情况下,也将玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物裂解或解毒。在此,将多肽的pH稳定性定义为所述多肽在pH 5.0下的残余活性百分比,相对于在各自最佳pH下的活性而言。
如相应于本发明的一个优选的进一步发展方案那样,所述多肽包含选自SEQ IDNo.1、2、5-7、9、11和15的氨基酸序列或其功能性变体,其中所述功能性变体与至少一个所述氨基酸序列具有至少40%的序列同一性,并且所述多肽在30℃和75℃之间,优选地38℃和55℃之间,特别优选地38℃和52℃之间的温度范围内具有最高的酶促活性。通过这样的本发明的进一步发展方案确保了,玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物在中温温度下,特别是在人和有用动物的体温下,也被所述多肽水解或解毒。将所述多肽具有最高酶促活性时所处的温度定义为所述多肽的最佳温度。
如相应于本发明的一个优选的进一步发展方案那样,所述多肽包含选自SEQ IDNo.1、5、6、9、11、12和15的氨基酸序列或其功能性变体,其中所述功能性变体与至少一个所述氨基酸序列具有至少40%的序列同一性,并且所述多肽是温度稳定的,直至90℃,优选地75℃,和特别优选地60℃的温度。由此确保了,所述多肽和其酶促功能即使在提高的温度负荷下(例如这可以是在容器中进行运输期间或在饲料粒化期间的情况)也基本上保持完整。多肽的温度稳定性被定义为这样的温度,在该温度下所述多肽在15分钟的预温育后具有50%的残余活性,相比于在各自最佳温度下的活性而言。
因此,所述多肽可以如此地来进行选择,从而它是对于不依赖于氧和无辅因子地以水解方式裂解玉米赤霉烯酮和/或ZEN衍生物的酯基团来说合适的α/β-水解酶,其具有催化所述水解式裂解的氨基酸三元体,该三元体由丝氨酸,一种选自谷氨酸和天冬氨酸的酸性氨基酸,特别是天冬氨酸,以及组氨酸组成,并且该催化三元体为例如S128、D264和H303,其中描述了相对于SEQ ID No.1的定位。
用SEQ ID No.1-15的多肽中的每一个可成功按照下述反应机制在玉米赤霉烯酮或其衍生物的酯基团处实现ZEN和ZEN衍生物的水解:
ZEN至无毒的经水解的玉米赤霉烯酮(HZEN)或经水解的ZEN衍生物的水解通过根据本发明的多肽,特别是所述α/β-水解酶来进行。HZEN至经脱羧的经水解的ZEN(DHZEN)或经脱羧的经水解的ZEN衍生物的进一步的脱羧通常自发地进行。
特别地,借助于上面提及的催化三元体,可成功实现完全水解ZEN和ZEN衍生物,其中该降解反应具有良好的pH稳定性,特别是在酸性范围内的pH值下。
令人惊讶地已证实,用在由上面提及的催化三元体的丝氨酸之前的3个氨基酸和之后的3个氨基酸组成的序列区段中包含至少一个选自Y、Q、N、T、K、R、E、D的极性氨基酸和至少一个选自F、M、L、I、V、A、G、P的非极性氨基酸的多肽,可成功取得依然良好的结果并且此外还改善至少一个酶动力学参数。
在本发明的一个优选的进一步改造方案中,所述多肽在至少一个下列位置处具有关于SEQ ID No.1的氨基酸序列的至少一个突变:22、23、25、26、27、29、31、32、35、37、42、43、46、51、53、54、57、60、69、72、73、78、80、84、88、95、97、99、114、118、119、123、132、141、146、148、149、154、163、164、165、169、170、172、176、180、182、183、190、191、194、196、197、198、201、204、205、206、207、208、209、210、212、213、214、216、217、220、221、222、229、231、233、238、240、244、245、246、248、249、251、254、256、260、262、263、266、269、271、277、280、281、282、283、284、285、286、287、292、296、298、302、307、308、309、311、314、317、319、321、323、325和326。这些位置得自具有SEQ ID No.1的多肽和与该序列具有高的同一性程度和特别地有活性的具有SEQ ID No.2-6的多肽之间的序列差异。通过在这些位置中的至少一个处如此地改变具有SEQ ID No.1的多肽,即在该位置处采用SEQ ID No.2-6的氨基酸变体,可成功显示,这些位置对于所述多肽的酶动力学参数具有显著的影响,并且此外,SEQID No.1与具有高的序列同一性程度的SEQ ID No.2-6的组合导致更高的活性。
根据本发明的一个进一步改造方案,所述多肽在关于SEQ ID No.1的氨基酸序列中具有至少一个选自下列的突变:D22A、S23Q、S23L、N25D、I26V、F27Y、F27H、S29P、R31A、F32Y、R35K、R35Q、V37A、V42I、V43T、F46Y、S51E、S51D、D53G、N54M、N54R、L57V、L60I、S69G、P72E、V73A、A78S、N80H、F84Y、I88L、T95S、T97A、R99K、I114M、I118V、K119R、V123I、L132V、A141S、I146V、I146L、A148G、A149V、A154P、P163T、A164T、Y165C、Y165H、V169I、L170R、A172G、A176M、A176V、Y180F、D182T、F183Y、I190V、G191S、K194T、K194E、F196Y、V197C、V197R、E198R、E198S、K201D、K201G、P204S、P204A、A205S、K206P、A207M、M208A、Q209R、L210A、L210S、ΔP212、T213V、P214A、E216T、E216G、A217I、N220H、L221M、K222R、K222Q、G229A、A231V、F233W、F233Y、F233H、A238G、H240N、H240S、D244E、R245Q、M246L、S248T、S248N、S248G、Q249R、K251N、I254V、I256L、A260M、T262D、T262G、I263T、E266D、E269H、E269N、L271V、L277E、E280A、E280L、H281R、H281Q、A282V、Q283R、D284L、D284R、I285L、I286M、R287E、R287D、R292K、R292T、Q296A、Q296E、H298V、L302S、L307Q、F308S、D309A、A311P、A314V、L317F、S319Q、S319P、S319R、S321A、S321T、T323A、P325A、A326P。用这样的多肽,可成功实现在短的时间内完全水解ZEN,特别是使其解毒,其中所述多肽的比活性为至少6.00U/mg,优选地至少7.00U/mg,特别地至少8.00U/mg。单位“U”或者“Unit”是绝对催化活性的量度,并且通过在32℃下在50mM Tris-HCl缓冲液(pH 8.2)中水解1μΜοl ZEN/分钟来定义,其中“催化活性”是指在确定的反应条件下底物的酶促转化,和“比活性”是指催化活性与多肽质量浓度(质量/体积单位)之比。
通过如此地产生所述多肽,即包含至少一个具有SEQ ID No.32-50的下列氨基酸基序,可成功提供这样的多肽以供使用,该多肽具有至少7.00U/mg,优选地至少8.00U/mg的比活性。令人惊讶地已显示,当包含至少一个具有有着SEQ ID No.51-58的序列的下列氨基酸基序时,所述多肽的酶促活性更进一步增加,例如相对于包含7个氨基酸的基序而言。当包含至少一个具有有着SEQ ID No.59-69的序列的下列氨基酸基序时,达到更加高的比活性。
根据本发明的一个进一步改造方案,所述多肽在至少一个位置处包含至少一个保守氨基酸置换,其中所述保守氨基酸置换选自:G至A;或A至G、S;或V至I、L、A、T、S;或I至V、L、M;或L至I、M、V;或M至L、I、V;或P至A、S、N;或F至Y、W、H;或Y至F、W、H;或W至Y、F、H;或R至K、E、D;或K至R、E、D;或H至Q、N、S;或D或N、E、K、R、Q;或E至Q、D、K、R、N;或S至T、A;或T至S、V、A;或C至S、T、A;或N至D、Q、H、S;或Q至E、N、H、K、R的置换。其中名称“保守氨基酸置换”涉及氨基酸由被专业人员视为保守的(即具有类似的特殊特性的)其他氨基酸来进行的置换。此类特殊特性为例如氨基酸的大小、极性、疏水性、电荷或pK值。保守突变是指例如将一个酸性氨基酸置换成另一个酸性氨基酸,将一个碱性氨基酸置换成另一个碱性氨基酸,或者将一个极性氨基酸置换成另一个极性氨基酸。
用这样的保守氨基酸置换,可成功制备出功能性多肽变体,其比活性相比于亲本多肽而言大约一样强,然而优选地增加了至少0.1U/mg。
此外,本发明还旨在,提供分离的多核苷酸以供使用,用所述多核苷酸可成功制备出用于快速且可靠地以水解方式裂解ZEN和/或至少一种ZEN-衍生物的多肽。
为了解决该任务,本发明的特征在于,所述分离的多核苷酸具有编码多肽的核苷酸序列,其中所述多肽具有水解玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物的特性;并且所述核苷酸序列编码至少一种根据权利要求1至11之一的多肽;和/或所述核苷酸序列与至少一种选自SEQ ID No.16-31的核苷酸序列具有序列同一性程度,其中所选择的核苷酸序列为至少40%;和/或所述核苷酸序列在中等严紧条件下与至少一种选自SEQ IDNo.16-31的核苷酸序列杂交,和/或与具有至少200个核苷酸,特别是至少100个核苷酸的其部分序列杂交,和/或与上述核苷酸序列或其部分序列的互补链杂交。
待表达的核苷酸序列,特别是其三联体(密码子),通常随着宿主细胞而变化,从而密码子偏倚性随着宿主细胞而优化。这导致,具有远低于80%,还有低于70%或低于60%的序列同一性程度的多核苷酸也可以编码同一种多肽。用于确定序列同一性程度的序列比较还必须在序列区段内进行,其中区段是指参考序列的连续序列。对于核苷酸序列,序列区段的长度通常为15至600。
借助于现有的分离的核苷酸序列或序列区段,可成功产生具有通常至少15、30或40个核苷酸的长度的核酸探针。借助于此类探针(其通常另外还例如用3H、32P、35S、生物素或抗生物素蛋白进行标记)可以通过使用标准方法来鉴定编码具有降解ZEN和/或ZEN衍生物的作用的多肽的核苷酸序列。作为用于鉴定此类序列的起始材料,可以考虑例如单个微生物的DNA、RNA或cDNA,基因组DNA文库,或者cDNA文库。
对于具有至少100个核苷酸的长度的核苷酸序列或核苷酸探针,中等严紧条件被定义为在42℃下在包含0.3%十二烷基硫酸钠(SDS)、200μg/ml经剪切和变性的鲑精DNA和35%甲酰胺的配备有5×NaCl的Na-EDTA缓冲液(SSPE,0.9M NaCl,60mM NaH2PO4,6mM EDTA)中的预杂交和杂交,随后为标准Southern印迹条件,其中最后将载体材料用2×的氯化钠-柠檬酸盐缓冲液(SSC,300mM NaCl和30mM柠檬酸三钠,0.2%SDS)在55℃下洗涤15分钟,进行三次。
对于具有15个核苷酸至100个核苷酸的长度的核苷酸序列或核苷酸探针,中等严紧条件被定义为在由0.9M NaCl、0.09M Tris-HCl pH=7.6、6mM EDTA、0.5%NP-40、1×Denhardt溶液、1mM焦磷酸钠、1mM磷酸二氢钠、0.1mM ATP和0.2mg/ml酵母RNA组成的缓冲液中的预杂交和杂交,其中在比计算出的解链温度(Tm)低5℃至10℃的温度下进行预杂交和杂交,其中Tm通过按照Bolton和McCarthy(1962,Proceedings of the National Academyof Sciences USA,48:1390)进行的计算来确定。随后,该试验在标准Southern印迹条件下继续进行(J.Sambrook,E.F.Fritsch和T.Maniatis,1989,Molecular Cloning,ALaboratory Manual,第二版,Cold Spring Harbor,New York)。最后,将载体材料用包含0.1%SDS的6×SCC缓冲液洗涤15分钟,进行一次;和在各自处于比计算出的Tm低5℃至10℃的温度下用6×SSC缓冲液洗涤15分钟,进行两次。
此外,本发明还旨在,提供添加剂以供使用,用所述添加剂可成功实现在确定的或复杂的基质中,例如在饲料或食品中,快速且可靠地以水解方式裂解ZEN和/或至少一种ZEN衍生物。
为了解决该任务,提供了以水解方式裂解玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物的添加剂以供使用,其中所述添加剂包含至少一种具有选自SEQ ID No.1-15的氨基酸序列或其功能性变体的多肽,其中在所述功能性变体与至少一个所述氨基酸序列之间的序列同一性为至少40%;并且任选地包含助剂。
用这样的添加剂,可成功实现ZEN和/或至少一种ZEN衍生物向经水解的ZEN和/或经水解的ZEN衍生物的生物化学转化。例如对于在工业过程中ZEN和/或ZEN衍生物的立体选择性水解,也可以考虑该添加剂。
在本发明的一个优选的进一步改造方案中,如此地产生所述添加剂,即所述助剂选自至少一种惰性载体,以及任选地,其他成分,例如维生素和/或矿物质和/或酶和/或其他用于使真菌毒素解毒的组分。通过使用这样的添加剂,可以例如在饲料或食品中确保,所任选地包含的ZEN和/或ZEN衍生物的量安全可靠地被水解特别是解毒至如此程度,从而对于摄取这些饲料或食品的受试者的机体没有有害效应。
在这种情况下,根据本发明的多肽也可以存在于酶制剂中,所述酶制剂除了包含至少一种根据本发明的多肽外,还包含至少一种酶,所述酶例如参与蛋白质的降解,例如蛋白酶,或者所述酶参与淀粉或纤维或脂肪或糖原的代谢,例如淀粉酶、纤维素酶或葡聚糖酶,以及例如水解酶、脂解酶、甘露糖苷酶、氧化酶、氧化还原酶、植酸酶、木聚糖酶和/或其组合。
本发明的其他使用范围为酶制剂,其除了包含至少一种根据本发明的多肽外,还包含至少一种用于使真菌毒素解毒的组分,例如真菌毒素降解酶,例如黄曲霉毒素氧化酶、麦角胺水解酶、麦角胺酰胺酶、玉米赤霉烯酮酯酶、玉米赤霉烯酮内酯酶、赭曲毒素酰胺酶、串珠镰孢菌素羧酸酯酶、串珠镰孢菌素氨基转移酶、氨基多元醇氨基氧化酶、脱氧瓜萎镰菌醇环氧化物水解酶;和/或至少一种降解真菌毒素的微生物,例如枯草芽孢杆菌;和/或至少一种结合真菌毒素的组分,例如微生物细胞壁或无机材料例如膨润土。
根据本发明的一个特别优选的进一步改造方案,在所述添加剂中,所述多肽以最高10,000U/g,优选地最高1,000U/g,更优选地最高100U/g,和最优选地最高10U/g的浓度包含在其中,由此可成功实现将ZEN和/或ZEN衍生物快速地,和特别地在其由消耗了被污染的饲料或食品的受试者(特别是哺乳动物)的身体吸收之前,已经转变为无毒或低毒的代谢物,特别是HZEN和DHZEN。
根据本发明的一个进一步改造方案,所述多肽以经包囊或经包衣的形式存在,其中对于所述包囊或包衣过程,可以考虑标准方法,例如在WO 92/12645中所描述的那些。通过所述包囊或包衣过程,可成功实现将所述多肽运输至其使用位点,而没有改变,特别是没有降解或损害,从而在保护壳溶解(例如在动物的消化道中)之后,所述多肽才开始起作用,由此在酸性的、富含蛋白酶的和缺氧的介质中也可以达到ZEN和/或ZEN衍生物的更加有针对性的、快速的和完全的降解。此外,通过所述包囊或包衣过程,还可成功提高在所述添加剂中所述多肽的温度稳定性。
此外,本发明还旨在,所述添加剂用于以水解方式裂解在饲料(其特别地用于猪、家禽和水产养殖)中、在食物中或在干酒糟中的玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物的用途。通过根据本发明来使用所述添加剂,可成功实现将在食物或饲料中或者在干酒糟中所包含的ZEN和/或ZEN衍生物水解或解毒,其中在大约1U/g被污染的饲料或食物的多肽浓度的情况下已经成功实现了这样的解毒。
此外,本发明还旨在,提供这样的方法以供使用,用所述方法使得ZEN和/或至少一种ZEN衍生物的快速且可靠的水解式裂解成为可能。
为了解决该任务,如此地来施行所述方法,从而玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物被具有选自SEQ ID No.1-15的氨基酸序列或其功能性变体的多肽水解,其中在所述功能性变体与至少一个所述氨基酸序列之间的序列同一性为至少40%。
根据本发明的一个进一步改造方案,如此地来施行所述方法,从而其中在相应于本发明的添加剂中使用所述多肽。
根据一个优选的进一步发展方案,如此地来施行所述方法,从而其中将所述多肽或添加剂与被玉米赤霉烯酮和/或被至少一种玉米赤霉烯酮衍生物污染的饲料或食物相掺混,使所述被污染的饲料或食物与湿气相接触,并且所述多肽或添加剂水解在所述被污染的饲料或食物中所包含的玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物。在湿润的饲料或食物例如麦芽浆或糊浆的情况下,玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物的水解在经口摄取之前在该湿润的饲料或食物中发生。通过该方法可以确保,最大程度地消除玉米赤霉烯酮和玉米赤霉烯酮衍生物对于人和动物的有害效应。在此,湿气是指水或含水液体的存在,其中例如唾液或其他在消化道中存在的液体也属于该范畴。消化道被定义为口腔、咽(咽喉)、食管和胃肠道或者其等价物,其中在动物中可以存在不同的名称或者个别成分可以不存在于动物的消化道中。
本发明的方法还可以如此地来施行,从而将所述饲料或食物在经口摄取之前进行粒化。
根据本发明的一个进一步改造方案,如此地来施行所述方法,从而至少70%,优选地至少80%,特别优选地至少90%的所述玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物被水解。由此可以预防在动物或人中的亚急性和/或亚慢性的毒性效应,例如致畸形的、致癌的、动情的和免疫抑制的效应。
下面将依据实施例以及附图更详细地阐明本发明。其中:
图1显示了通过具有SEQ ID No.1的多肽来进行的ZEN的随时间的降解以及代谢物HZEN和DHZEN的增加,其中在图1A中,所述多肽未加标签,在图1B中,所述多肽具有C-末端6×His标签,和在图1C中,所述多肽具有N-末端6×His标签。
图2显示了具有SEQ ID No.1的多肽的米-曼(Michaelis-Menten)动力学。
图3显示了通过经纯化的具有SEQ ID No.1(图3A)、SEQ ID No.2(图3B)、SEQ IDNo.5(图3C)、SEQ ID No.6(图3D)、SEQ ID No.7(图3E)、SEQ ID No.9(图3F)、SEQ ID No.11(图3G)、SEQ ID No.12(图3H)和SEQ ID No.15(图3I)的多肽来进行的ZEN的随时间的降解以及代谢物HZEN和DHZEN的增加,其中所有序列均具有C-末端6×His标签。
实施例1:编码能够以水解方式裂解ZEN和/或至少一种ZEN衍生物的多肽的多核苷酸的修饰、克隆和表达
按照说明书,使用“Quick-change Site-directed Mutagenesis Kits”(Stratagene),借助于PCR,通过核苷酸序列的突变来进行氨基酸置换、插入或缺失。备选地,为此还购进了完整的核苷酸序列(GeneArt)。借助于PCR诱变而产生的或从GeneArt购进的核苷酸序列在氨基酸水平上可选地另外还包含C-末端或N-末端6×His标签,并且借助于标准方法将其整合到用于在大肠杆菌(E.coli)或巴斯德毕赤酵母(P.pastoris)中进行表达的表达载体之中,转化到大肠杆菌或巴斯德毕赤酵母中,以及在大肠杆菌或巴斯德毕赤酵母中进行表达(J.M.Cregg,Pichia Protocols,第二版,ISBN-10:1588294293、2007;J.Sambrook等人,2012,Molecular Cloning,A Laboratory Manual,第4版,Cold SpringHarbor),其中也可以为该任务考虑任何其他合适的宿主细胞。
名称“表达载体”涉及能够在体内或在体外表达基因的DNA构建体。特别地,该名称涵盖这样的DNA构建体,其适合于将编码所述多肽的核苷酸序列转移到宿主细胞中,以便在那里整合到基因组中或者游离地存在于染色体外空间中,并且在细胞内表达编码所述多肽的核苷酸序列和任选地从细胞中提取出所述多肽。
名称“宿主细胞”涉及所有这样的细胞,其要么包含待表达的核苷酸序列,要么包含表达载体,并且能够制备根据本发明的多肽。特别地,该名称涵盖原核细胞和/或真核细胞,优选地巴斯德毕赤酵母,大肠杆菌,枯草芽孢杆菌,链霉菌属(Streptomyces),汉逊酵母属(Hansenula),木霉属(Trichoderma),乳杆菌属(Lactobacillus),曲霉属(Aspergillus),植物细胞,和/或芽孢杆菌属(Bacillus)、木霉属或曲霉属的孢子。
为了测定多肽的催化特性,在大肠杆菌的情况下考虑可溶的细胞裂解物,或者在巴斯德毕赤酵母的情况下考虑培养物上清液。为了测定KM值、vmax、kcat和比活性,借助于标准方法以色谱法方式通过镍-Sepharose柱选择性地富集所述多肽。蛋白质浓度的测定借助于标准方法来进行,要么采用BCA方法(Pierce BCA Protein Assay KitProd#23225),然而优选地采用用关于各自蛋白质的比消光系数来实施的光度法,所述比消光系数用在http://web.expasy.org/protparam上在线可用的程序“ProtParam”来计算(Gasteiger E.等人,Protein Identification and Analysis Tools on the ExPASy Server,JohnM.Walker(编辑):The Proteomics Protocols Handbook,Humana Press,2005,第571-607页)。
实施例2:序列同一性以及保守氨基酸序列区段的确定
在具有有着SEQ ID No.1-15的氨基酸序列的多肽的总多肽长度上的相互之间的序列同一性百分比(表1)的确定可以借助于程序BLAST(Basic Local Alignment SearchTool),特别是用可以在“国家生物技术信息中心(National Center for BiotechnologyInformation)”的主页(NCBI;http://www.ncbi.nlm.nih.gov/)上使用的BLASTP来进行。因此,可能的是,根据Altschul等人,1997(Nucleic Acids Res.(1997)25:3389-3402)的算法,将两个或更多个序列互相进行比较。作为程序设置,考虑基本设置,但是特别地:“最大靶序列”=100;“期望阈值”=10;“字长”=3;“矩阵”=BLOSOM62;“缺口成本”=“存在:11;延伸:1”;“计算调整”=“条件式组成得分矩阵调整”。
为了确定保守氨基酸序列区段,借助于软件COBALT(J.S.Papadopoulos和R.Agarwala,2007,COBALT:constraint-based alignment tool for multiple proteinsequences,Bioinformatics23:1073-79)来调节对准相互之间具有至少70%的序列同一性的具有SEQ ID No.1-6的多肽,其中考虑标准参数,特别是下列参数(“缺口罚分”:-11,-1;“末端缺口罚分”:-5,-1;“使用RPS BLAST(Use RPS BLAST)”:on;“Blast E-值”:0.003;“发现保守列和重新计算(Find Conserved columns and Recompute)”:on;“使用查询聚簇(use query clusters)”:on;“字长”:4;“可能聚簇距离”(may cluster distance):0.8;“字母表”:常规;“同源性保守设置(Homology conversation setting):3bits)。该分析的结果描绘出了保守氨基酸。保守氨基酸序列区段被定义为具有至少5个连续的保守氨基酸的下列区域:即关于具有SEQ ID No.1的序列,位置+24至位置+50的区段A、位置+52至位置+77的区段B、位置+79至位置+87的区段C、位置+89至位置+145的区段D、位置+150至位置+171的区段E、位置+177至位置+193的区段F、位置+223至位置+228的区段G、位置+230至位置+237的区段H、位置+239至位置+247的区段I、位置+249至位置+255的区段J、位置+257至位置+261的区段K、位置+263至位置+270的区段L、位置+272至位置+279的区段M、位置+297至位置+301的区段N和位置+303至位置+313的区段O。
如上面所描述的那样,进行多肽相互之间的以及独个多肽的保守氨基酸序列区段相对于具有SEQ ID No.1的序列的保守氨基酸序列区段而言的序列同一性百分比的测定。结果呈现在表1和2中。
表1:多肽相互之间的序列同一性百分比。
表2:保守氨基酸序列区段A至O的序列同一性百分比。
实施例3:通过在细胞裂解物中的多肽来水解ZEN
为了测定其将ZEN降解为非毒性或低毒性的代谢物HZEN和DHZEN的能力,如在实施例1中所描述的那样,在大肠杆菌中制备由具有SEQ ID No.17的核苷酸序列所编码的具有SEQ ID No.1的多肽,所述多肽就这样地和以具有C-末端或N-末端6×His标签的方式来进行制备。将有着由具有SEQ ID No.18-31的核苷酸序列所编码的具有SEQ ID No.2-15的氨基酸序列的多肽仅在C-末端标记上6×His。各将100ml的具有2.0-2.5的光密度(OD600nm)的大肠杆菌培养物通过在4℃下离心来进行收获,并且重悬浮在20ml Brunner矿质培养基(DSMZ微生物培养基编号462,2012)中。通过用弗氏压碎器在20,000psi下处理3次来将细胞悬浮液裂解。将如此获得的细胞裂解物以1:10、1:100或1:1,000的稀释物进行使用,所述稀释物在包含0.1mg/ml BSA(牛血清白蛋白)的Brunner矿质培养基中制备。对于ZEN降解试验,使用9.9ml的包含0.1mg/ml BSA的Brunner矿质培养基、0.1ml的经稀释的细胞裂解物和31μl的ZEN底物储液。总之,因此将细胞裂解物以1:1,000、1:10,000或1:100,000进行了稀释。2.08mMZEN溶液(40体积%的ACN+60体积%的H2O)用作ZEN底物储液。为了制备该溶液,相应地称量以晶体形式的ZEN(Romer Labs的生物纯标准品,商品编号001109,纯度为至少98%)并装瓶,并且将其溶解。每个降解批次在25ml玻璃小瓶中进行,并且在25℃下在以100rpm进行摇动的情况下温育总共120小时。在时间点0、0.5、1、2、5、24、47、72和120小时,各取1ml的样品,将多肽在99℃下加热失活10分钟并贮存于-20℃。在样品解冻后,通过离心分离开不溶性成分。借助于LC/MS/MS来分析ZEN、HZEN和DHZEN。为此,借助于尺寸为250mm×3mm和颗粒大小为5μm的Phenomenex Luna C18(2)柱通过色谱法来将代谢物分开。具有1ml/l的甲酸浓度的乙腈-水混合物用作流动相。在270nm下记录UV-信号。电喷雾离子化(ESI)用作离子化源。借助于QTrap/LC/MS/MS(三重四极,Applied Biosystems)以“增强模式”来对ZEN、HZEN和DHZEN进行定量。在最迟24小时后,在批料中不再检测到重大量的ZEN。大部分(超过80%)的ZEN被转变成HZEN或DHZEN。
在图1中看到,示例性地关于经1:10,000稀释的细胞裂解物溶液,对于未加标签的(图1A)以及对于加有C-末端6×His标签的(图1B)和加有N-末端6×His标签的(图1C)具有SEQ ID No.1的多肽,ZEN的随时间的降解和HZEN及DHZEN的随时间的增加。从中清楚地显而易见的是:1.ZEN的转化立即且完全地进行,这是因为在于试验开始后立即抽取的第一个样品(0小时)中,就已经几乎不再能够检测到ZEN;和2.通过附加C-末端或N-末端的标签,未出现值得注意的活性损失。
实施例4:通过在细胞裂解物中的多肽来水解ZEN衍生物
为了测定多肽将除了ZEN外还有ZEN衍生物转化为非毒性或低毒性的代谢物的能力,如在实施例3中所描述的那样,以具有C-末端His标签的方式制备具有SEQ ID No.1-15的多肽,并且作为在“降解15”中的细胞裂解物,考虑各自的具有有着SEQ ID No.17-31的序列的合成核苷酸序列。
降解试验如在实施例3中所描述的那样来进行,其中每种多肽用选自α-ZEL、β-ZEL、α-ZAL、β-ZAL、Z14G、Z14S和ZAN的每种ZEN衍生物来进行测试。以1:10,000的总稀释度来使用细胞裂解物。作为底物储液,使用等摩尔的(即2.08mM的)ZEN衍生物溶液,以代替2.08mM ZEN溶液(40体积%的ACN+60体积%的H2O)。α-ZEL、β-ZEL、α-ZAL、β-ZAL和ZAN从Sigma购进并且作为关于该分析的标准品来进行使用。按照诸如在P.Krenn等人,2007(Mykotoxin Research,23,4,180-184)和M.Sulyok等人,2007(Anal.Bioanal.Chem.289,1505-1523)中所描述的方法,以至少90%的纯度来制备Z14G和Z14S,并且将其作为关于该分析的标准品来进行使用。与实施例3的另一个差别是,仅取一个样品,即在24小时后。在降解试验期间ZEN衍生物浓度的降低借助于LC/MS/MS来进行定量。α-ZEL、β-ZEL、Z14G和Z14S按照M.Sulyok等人(2010,Food Chemistry,119,408-416)的方法来进行测量;α-ZAL、β-ZAL和ZAN按照P.Songsermaskul等人(2011,J.of Animal Physiol.and Animal Nutr.,97,155-161)的方法来进行测量。令人惊讶地表明,在所有降解试验中,在24小时的温育后,仅仅存在ZEN衍生物的起始量的0%至最大13%。
实施例5:多肽以及其变体的比活性以及酶动力学参数
多肽以及其变体的比活性的测定通过光度法来进行,其中所有所使用的多肽具有C-末端6×His标签。多肽或其变体的制备、富集和纯化如在实施例1中所描述的那样来进行。ZEN至HZEN的降解通过在315nm的波长下吸光度的下降来测量。ZEN和HZEN的摩尔消光系数[ε]通过实验来测定,并且为0.0078895L μmοl-1cm-1和0.0030857L μmοl-1cm-1。消光系数是强烈地pH依赖性的,并因此必须总是在精确相同的pH值下,优选地甚至在相同的基质中进行活性测量。在32℃下,在UV-VIS光度计(Hitachi U-2001)中,在200至2500nm的波长范围内,在石英比色皿中,在50mM Tris-HCl(pH=8.2)缓冲溶液中进行测量。
2.08mM ZEN溶液(40体积%的ACN+60体积%的H2O)用作ZEN底物储液。为了制备该溶液,相应地称量以晶体形式的ZEN(Romer Labs的生物纯标准品,商品编号001109,纯度为至少98%)并装瓶,并且将其溶解。用50mM Tris-HCl(pH=8.2)来制备ZEN底物稀释物(0.79μΜ、1.57μΜ、2.36μΜ、3.14μΜ、4.71μΜ、6.28μΜ、7.85μΜ、9.42μΜ、10.99μΜ、12.56μΜ、14.13μΜ、15.71μΜ、17.28μΜ、18.85μΜ)。将多肽溶液用50mM Tris-HCl缓冲液(pH=8.2)稀释至大约70ng/ml的终浓度。将ZEN底物稀释物在水浴中预热至32℃。
给100μl的各个ZEN底物稀释物掺入0.2μl的多肽溶液,并且测量吸光度5分钟,其中测量每个“多肽溶液-ZEN底物稀释物”组合至少两次。
通过考虑ZEN和HZEN的消光系数,经由随时间进程的吸光度的斜率,计算出关于每个底物浓度的反应速率。
名称“KM值”或“米-曼常数”涉及用于描述酶亲和力的参数,其具有单位[μΜ]或[mM],并且按照H.Bisswang(2002,EnzymeKinetics,ISBN 3-527-30343-X,第19页)借助于线性Hanes标绘图来计算,其中为此优选地使用程序SigmaPlot 12.0的函数“酶动力学,单一底物”。名称“酶反应的催化常数”或“kcat值”涉及用于描述多肽或酶的转化速度的参数,其以[s]-1给出,并且优选地借助于程序SigmaPlot 12.0的函数“酶动力学,单一底物”来计算。“最大酶速率”或“vmax值”以单位[μΜ/s]或[mM/s]给出,并且类似于KM值,借助于线性Hanes标绘图来测算,其中为此优选地使用程序SigmaPlot12.0的函数“酶动力学,单一底物”。
借助于vmax和所使用的酶浓度,根据下述公式来计算比活性,
其中一个单位被定义为在32℃下在50mM Tris-HCl缓冲溶液(pH=8.2)中每分钟水解1μmοl ZEN。
下面,示例性地对于具有SEQ ID NO:1的多肽,列举出关于酶参数KM、vmax、kcat以及比活性的测定的原始数据。表3显示了在各个ZEN底物浓度下的反应速率,图2显示了各自所属的米-曼图,和在表4中给出了相应的酶动力学参数。所使用的酶溶液具有68ng/l的浓度。
表3:在不同的ZEN浓度下,
具有SEQ ID NO:1的多肽的反应速率。
表4:具有SEQ ID No.1的多肽的酶动力学参数。
所研究的多肽的比活性为:8.25U/mg,对于SEQ ID No.1;10.56U/mg,对于SEQ IDNo.2;8.36U/mg,对于SEQ ID No.3;8.33U/mg,对于SEQ ID No.4;8.56U/mg,对于SEQ IDNo.5;9.95U/mg,对于SEQ ID No.6;3.83U/mg,对于SEQ ID No.7;2.57U/mg,对于SEQ IDNo.8;4.87U/mg,对于SEQ ID No.9;5.12U/mg,对于SEQ ID No.10;3.88U/mg,对于SEQ IDNo.11;2.78U/mg,对于SEQ ID No.12;6.43U/mg,对于SEQ ID No.13;3.33U/mg,对于SEQ IDNo.14;和7.76U/mg,对于SEQ ID No.15。
所研究的多肽变体的比活性在表5和表6中列出。
表5:具有SEQ ID No.1的多肽的功能性变体的比活性;突变所位于的保守氨基酸序列区段;和功能性变体相对于具有SEQ ID No.1的亲本序列而言的序列同一性。相对于具有SEQ ID No.1的氨基酸序列而言给出了突变的位置。如在实施例2中所描述的那样,
借助于BLAST来测算序列同一性。
表6:具有SEQ ID No.2的多肽的功能性变体的比活性。突变的位置是相对于具有SEQ ID No.2的氨基酸序列而言的。如在实施例2中所描述的那样,借助于BLAST来测算序列同一性。
实施例6:在被污染的玉米中的ZEN和ZEN衍生物的降解
为了测定多肽在复杂基质中和在低pH值下降解天然存在的ZEN和ZEN衍生物的能力,每次给被污染的玉米掺入不同浓度的具有SEQ ID No.1-6的多肽中的一种,并且追踪ZEN和ZEN衍生物的降解。
碾磨被污染的玉米并且将其在降解试验中使用,其中批料由1g经碾磨的被污染的玉米、8.9ml 100mM乙酸盐缓冲液(pH=4.0)和0.1ml多肽溶液组成。如在实施例5中所描述的那样,制备经富集和经纯化的多肽溶液,其中将其稀释至10mU/ml、100mU/ml或1,000mU/ml的浓度。因此,在批料中以绝对方式使用1mU(=1mU/克玉米)、10mU(=10mU/克玉米)或100mU(=100mU/克玉米)。每个降解批料以25ml来实施制备,并且在37℃下在以100rpm进行摇动的情况下进行温育。在酶添加之前和在1小时的温育后,分别取出1ml的样品,将多肽在99℃下加热失活10分钟,并且将样品贮存于-20℃。在样品解冻后,通过离心分离开不溶性成分。如在M.Sulyok等人(2007,Anal.Bioanal.Chem.,289,1505-1523)中所描述的那样,借助于LC/MS/MS来测量ZEN以及ZEN衍生物的浓度。在该玉米中ZEN和ZEN衍生物的含量为:238ppb,对于ZEN;15ppb,对于α-ZEL;23ppb,对于β-ZEL;32ppb,对于Z14G;和81ppb,对于Z14S。在表7中呈现了在所述降解试验中ZEN和ZEN衍生物含量下降的百分比。
表7:不同多肽和多肽量的降解试验的相对于起始含量而言的以百分比表示的ZEN和ZEN衍生物的减少。
实施例7:用于以水解方式裂解ZEN和/或ZEN衍生物的包含多肽的添加剂
为了制备用于以水解方式裂解ZEN的添加剂,借助于微量过滤和超滤(排阻极限:10kDa)在标准条件下纯化经由巴斯德毕赤酵母表达的具有SEQ ID No.1、2、6和13的多肽的发酵上清液,并且将其浓缩至大约9重量%的干物质浓度。随后,用喷雾干燥器(Büchi的Mini B290)同样在标准条件下将该含多肽的溶液进一步加工成干粉末。将这四种粉末相继地命名为Z1、Z2、Z6和Z13。此外,将Z1、Z2、Z6或Z13与平均粒度为大约1μm的膨润土,以1重量%的添加剂Z1、Z2、Z6或Z13和99重量%的膨润土的比例,在向上式摇动器中进行混合。将如此获得的添加剂命名为添加剂Z1.B、Z2.B、Z6.B和Z13.B。此外,将Z1、Z2、Z6和Z13与膨润土和维生素-微量元素浓缩物,以0.1重量%的Z1、Z2、Z6或Z13,0.9重量%的维生素-微量元素浓缩物,和99重量%的膨润土的比例,在向上式摇动器中进行混合。将如此获得的添加剂命名为添加剂Z1.BVS、Z2.BVS、Z6.BVS和Z13.BVS。100g的添加剂Z1.BVS、Z2.BVS、Z6.BVS和Z13.BVS包含200mg硫酸铁、50mg硫酸铜、130mg氧化锌、130mg氧化锰、2.55mg碳酸钙、160mg维生素E、6.5mg维生素K3、6.5mg维生素B1、14mg维生素B2、15mg维生素B6、0.15mg维生素B12、150mg烟酸、30mg泛酸和5.3mg叶酸。
将所述添加剂在50mM Tris-HCl缓冲液(pH=8.2)中抽提30分钟,并且在同一缓冲液中如此地进一步稀释,从而使得多肽的终浓度位于大约70ng/ml。
随后,如在实施例5中所描述的那样,测定这些溶液的降解玉米赤霉烯酮的效应。相应的活性为:8.230U/g,对于Z1;9.310U/g,对于Z2;9.214U/g,对于Z6;83U/g,对于Z1.B;92U/g,对于Z2.B;90U/g,对于Z2.C;57U/g,对于Z13.B;8U/g,对于Z1.BVS;9U/g,对于Z2.BVS;9U/g,对于Z6.BVS;和6U/g,对于Z13.BVS。
如在实施例4中所描述的那样来施行添加剂Z1、Z2、Z6、Z13、Z1.B、Z2.B、Z6.B、Z13.B、Z1.BVS、Z2.BVS、Z6.BVS和Z13.BVS降解ZEN衍生物α-ZEL、β-ZEL、α-ZAL、β-ZAL、Z14G、Z14S和ZAN的能力,但是使用100μl的具有大约70ng/ml的多肽浓度的多肽溶液来代替100μl的细胞裂解物。在6小时的温育后,起始量的仅仅最大15%作为未水解的ZEN衍生物存在。
实施例8:最佳温度
为了测定具有SEQ ID No.1、2、5、6、7、9、11、12和15的多肽的最佳温度,如在实施例1中所描述的那样,将它们以具有C-末端6×His标签的方式进行克隆,在大肠杆菌中表达,并纯化。在初步试验中,对于每种多肽,测算出那个浓度,在所述浓度下在试验条件(Teorell Stenhagen缓冲液(Teorell和Stenhagen,Ein Universalpuffer für den pH-Bereich 2.0bis 12.0.Biochem Ztschrft,1938,299:第416-419页),pH 7.5,具有0.1mg/ml BSA,在30℃下)下在3小时的试验持续时间后可以确保ZEN的完全转化。以所测算出的浓度在用于测定最佳温度的降解批料中使用所述制备物。该试验在PCR循环仪(Eppendorf)中,通过使用温度梯度函数,在20℃+/-10℃下,在40℃+/-10℃下,和如果需要,在60℃+/-10℃下(在各自范围内10个温度;预先确定的PCR循环仪的温度)来进行。对于批料,在各自最佳pH下给Teorell-Stenhagen缓冲液掺入相应的酶浓度以及0.1mg/ml BSA和5ppm ZEN。具有0.1mg/ml BSA和5ppm ZEN而没有酶添加的批料用作阴性对照。在0小时、0.5小时、1小时、2小时和3小时的温育时间后,对于每个温育温度各自抽取一个样品,在99℃下加热失活10分钟,并且贮存于-20℃。在解冻后,将样品转移到HPLC小瓶中。借助于HPLC-DAD来分析ZEN、HZEN和DHZEN。为此,借助于尺寸为4.6mm×150mm和颗粒大小为5μm的Zorbax SB-AqC18柱通过色谱法来将代谢物分开。具有5mM乙酸铵的甲醇-水混合物用作流动相。在274nm下记录UV-信号。代谢物的定量通过包括所携带的标准品系列来进行。最佳温度的测算通过测算出的降解曲线的斜率来进行,其中将斜率为最大时所处的温度定义为最佳温度。最佳温度显示在表8中。
表8:多肽的最佳温度。
实施例9:温度稳定性
为了测定具有SEQ ID No.1、2、5、6、7、9、11、12和15的多肽的温度稳定性,如在实施例1中所描述的那样,将它们以具有C-末端6×His标签的方式进行克隆,在大肠杆菌中表达,并纯化。将它们在具有梯度函数的PCR循环仪中在各自的最佳温度+/-10℃下进行温育。在0分钟、15分钟、30分钟和60分钟后,对于每个批料和每个温度抽取一个样品。随后,在降解试验中,在具有0.1mg/ml BSA和5ppm ZEN的处于各自最佳pH下的Teorell-Stenhagen缓冲液中,在批料中使用这些经预温育的样品。在初步试验中,对于每种多肽,测算出那个浓度,在所述浓度下在试验条件(Teorell Stenhagen缓冲液,pH 7.5,具有0.1mg/ml BSA,在30℃下)下在3小时的试验持续时间后可以确保ZEN的完全转化。在批料中使用各自所测算出的酶浓度。在30℃下温育所述降解批料。在0小时、0.5小时、1小时、2小时和3小时的温育时间后进行取样。随后,将多肽在99℃下加热失活10分钟,并且将样品贮存于-20℃。在解冻后,将样品转移到HPLC小瓶中,并且如在实施例8中所描述的那样借助于HPLC-DAD来进行分析。
将温度稳定性定义为这样的温度,在所述温度下多肽在15分钟的预温育后具有50%的残余活性,相比于最佳温度而言。作为对于活性的量度,考虑降解曲线的斜率。温度稳定性显示在表9中。
表9:多肽的温度稳定性(在15分钟的预温育后50%的残余活性)。
实施例10:最佳pH
为了测定具有SEQ ID No.1、2、5、6、7、9、11、12和15的多肽的最佳pH,如在实施例1中所描述的那样,将它们以具有C-末端6×His标签的方式进行克隆,在大肠杆菌中表达,并纯化。在初步试验中,对于每种多肽,测算出那个浓度,在所述浓度下在试验条件(TeorellStenhagen缓冲液,pH 7.5,具有0.1mg/ml BSA,在30℃下)下在3小时的试验持续时间后可以确保ZEN的完全转化。在批料中使用各自的酶浓度。所述降解批料在处于3.0、4.0、5.0、5.5、6.0、6.5、7.0、7.5、8.0、8.5、9.0、9.5、10.0、11.0和12.0的pH值下的Stenhagen缓冲液中实施制备。为了进行降解,将具有0.1mg/ml BSA和5ppm ZEN的降解批料在30℃下进行温育。在具有0.1mg/ml BSA和5ppm ZEN的处于pH 3.0、pH 7.0和pH 12.0下的Teorell-Stenhagen缓冲液中的批料用作阴性对照。在0小时、0.5小时、1小时、2小时和3小时的温育时间后进行取样。随后,将多肽在99℃下加热失活10分钟,并且将样品贮存于-20℃。在解冻后,将样品转移到HPLC小瓶中,并且如在实施例8中所描述的那样借助于HPLC-DAD来进行分析。最佳pH的测算通过测算出的降解曲线的斜率来进行,其中将在斜率为最大时所处的pH值定义为最佳pH。最佳pH显示在表10中。
表10:多肽的最佳pH。
实施例11:在pH 5.0下的pH稳定性
为了测定pH稳定性,将来自实施例10的多肽在处于pH 5.0和各自最佳pH下的Teroell-Stenhagen缓冲液中在25℃下温育1小时。在降解试验中,以与对于测定最佳pH所使用的浓度相同的各自多肽的浓度,在具有0.1mg/ml BSA和5ppm ZEN的处于各自最佳pH下的100mM Tris-HCl缓冲液中,在批料中使用这些经预温育的样品。在0小时、0.5小时、1小时、2小时和3小时的温育时间后进行取样。随后,将多肽在99℃下加热失活10分钟,并且将样品贮存于-20℃。在解冻后,将样品转移到HPLC小瓶中,并且如在实施例8中所描述的那样借助于HPLC-DAD来进行分析。将pH稳定性定义为在pH 5.0下多肽的残余活性百分比,相对于在各自最佳pH下的活性而言。对于pH 5.0的pH稳定性显示在表11中。
表11:在pH 5.0下多肽的pH稳定性。
实施例12:ZEN降解试验
示例性地对于具有SEQ ID No.1、2、5、6、7、9、11、12和15的多肽,进行ZEN至HZEN和DHZEN的降解。所述降解批料在具有0.1mg/ml BSA和5ppm ZEN的Teorell Stenhagen缓冲液(pH 7.5)中实施制备。在30℃下温育所述降解批料。在0小时、0.5小时、1小时、2小时和3小时的温育时间后进行取样。随后,将多肽在99℃下加热失活10分钟,并且将样品贮存于-20℃。在解冻后,将样品转移到HPLC小瓶中,并且如在实施例8中所描述的那样借助于HPLC-DAD来进行分析。如此地来选择多肽浓度,从而在大约3小时后达到完全的降解。降解动力学描绘在图3中,其中y轴显示了以微摩尔/升(μmol/l)表示的ZEN、HZEN和DHZEN的浓度,和x轴显示了以小时(h)表示的温育时间。
*μΜ表示微体积摩尔浓度,并且相应于单位μmοl/l。
本发明的一些实施方案如下:
1.以水解方式裂解玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物的多肽,其特征在于,所述多肽是具有选自SEQ ID No.1-15的氨基酸序列或其功能性变体的水解酶,其中在所述功能性变体与至少一个所述氨基酸序列之间的序列同一性为至少40%。
2.根据实施方案1的多肽,其特征在于,所述多肽包含至少一个保守氨基酸序列区段或其功能性变体,其中所述氨基酸序列区段的功能性变体具有至少70%,优选地至少84%,更优选地至少92%,和最优选地至少98%的序列同一性,并且所述至少一个保守氨基酸序列区段选自具有SEQ ID No.1的序列的氨基酸序列+24至+50、+52至+77、+79至+87、+89至+145、+150至+171、+177至+193、+223至+228、+230至+237、+239至+247、+249至+255、+257至+261、+263至+270、+272至+279、+297至+301、+303至+313、+24至328、+1至+328。
3.根据实施方案1或2的多肽,其特征在于,所述功能性变体具有选自下列的氨基酸修饰:各自一个或多个氨基酸的置换、缺失和插入。
4.根据实施方案1、2或3之一的多肽,其特征在于,所述多肽具有至少0.01U/mg,优选地至少0.1U/mg,特别地至少1U/mg的比活性,和/或具有最高50μΜ,优选地最高3.5μΜ,特别地最高0.5μΜ的以水解方式裂解玉米赤霉烯酮的KM值,和/或具有至少0.05s-1,优选地至少0.6s-1,特别地至少5s-1的以水解方式裂解玉米赤霉烯酮的kcat值,和/或具有至少0.00001μΜ-1s-1,优选地至少0.0001μΜ-1s-1,特别地至少0.001μΜ-1s-1的以水解方式裂解玉米赤霉烯酮的vmax值。
5.根据实施方案1至4之一的多肽,其特征在于,所述多肽包含选自SEQ ID No.2、5、6、7、9、11和15的氨基酸序列或其功能性变体,其中所述功能性变体与至少一个所述氨基酸序列具有至少40%的序列同一性,并且所述多肽在pH 5.0下的pH稳定性为至少15%,优选地50%,和特别优选地至少90%。
6.根据实施方案1至4之一的多肽,其特征在于,所述多肽包含选自SEQ ID No.1、2、5、6、7、9、11、15的氨基酸序列或其功能性变体,其中所述功能性变体与至少一个所述氨基酸序列具有至少40%的序列同一性,并且所述多肽在30℃和75℃之间,优选地38℃和55℃之间,特别优选地38℃和52℃之间的温度范围内具有最高的酶促活性。
7.根据实施方案1至4之一的多肽,其特征在于,所述多肽包含选自SEQ ID No.1、5、6、9、11、12和15的氨基酸序列或其功能性变体,其中所述功能性变体与至少一个所述氨基酸序列具有至少40%的序列同一性,并且所述多肽是温度稳定的,直至90℃,优选地75℃,和特别优选地60℃的温度。
8.根据实施方案1至7之一的多肽,其特征在于,所述多肽在至少一个选自下列的位置处具有关于SEQ ID No.1的氨基酸序列的至少一个突变:22、23、25、26、27、29、31、32、35、37、42、43、46、51、53、54、57、60、69、72、73、78、80、84、88、95、97、99、114、118、119、123、132、141、146、148、149、154、163、164、165、169、170、172、176、180、182、183、190、191、194、196、197、198、201、204、205、206、207、208、209、210、212、213、214、216、217、220、221、222、229、231、233、238、240、244、245、246、248、249、251、254、256、260、262、263、266、269、271、277、280、281、282、283、284、285、286、287、292、296、298、302、307、308、309、311、314、317、319、321、323、325和326。
9.根据实施方案8的多肽,其特征在于,所述多肽具有选自下列的关于SEQ IDNo.1的氨基酸序列的至少一个突变:D22A、S23Q、S23L、N25D、I26V、F27Y、F27H、S29P、R31A、F32Y、R35K、R35Q、V37A、V42I、V43T、F46Y、S51E、S51D、D53G、N54M、N54R、L57V、L60I、S69G、P72E、V73A、A78S、N80H、F84Y、I88L、T95S、T97A、R99K、I114M、I118V、K119R、V123I、L132V、A141S、I146V、I146L、A148G、A149V、A154P、P163T、A164T、Y165C、Y165H、V169I、L170R、A172G、A176M、A176V、Y180F、D182T、F183Y、I190V、G191S、K194T、K194E、F196Y、V197C、V197R、E198R、E198S、K201D、K201G、P204S、P204A、A205S、K206P、A207M、M208A、Q209R、L210A、L210S、ΔP212、T213V、P214A、E216T、E216G、A217I、N220H、L221M、K222R、K222Q、G229A、A231V、F233W、F233Y、F233H、A238G、H240N、H240S、D244E、R245Q、M246L、S248T、S248N、S248G、Q249R、K251N、I254V、I256L、A260M、T262D、T262G、I263T、E266D、E269H、E269N、L271V、L277E、E280A、E280L、H281R、H281Q、A282V、Q283R、D284L、D284R、I285L、I286M、R287E、R287D、R292K、R292T、Q296A、Q296E、H298V、L302S、L307Q、F308S、D309A、A311P、A314V、L317F、S319Q、S319P、S319R、S321A、S321T、T323A、P325A、A326P。
10.根据实施方案1至8之一的多肽,其特征在于,包含至少一个选自SEQ IDNo.32-69的下列氨基酸基序。
11.根据实施方案9的多肽,其特征在于,所述多肽在至少一个位置处包含至少一个保守氨基酸置换,并且所述保守氨基酸置换选自:G至A;或A至G、S;或V至I、L、A、T、S;或I至V、L、M;或L至I、M、V;或M至L、I、V;或P至A、S、N;或F至Y、W、H;或Y至F、W、H;或W至Y、F、H;或R至K、E、D;或K至R、E、D;或H至Q、N、S;或D或N、E、K、R、Q;或E至Q、D、K、R、N;或S至T、A;或T至S、V、A;或C至S、T、A;或N至D、Q、H、S;或Q至E、N、H、K、R的置换。
12.分离的多核苷酸,其具有编码多肽的核苷酸序列,其中所述多肽具有水解玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物的特性,其特征在于,所述核苷酸序列编码至少一种根据实施方案1至11之一的多肽;和/或所述核苷酸序列与至少一种选自SEQ IDNo.16-31的核苷酸序列具有序列同一性程度,其中所选择的核苷酸序列为至少40%;和/或所述核苷酸序列在中等严紧条件下与至少一种选自SEQ ID No.16-31的核苷酸序列杂交,和/或与具有至少200个核苷酸,特别是至少100个核苷酸的其部分序列杂交,和/或与上述核苷酸序列或其部分序列的互补链杂交。
13.以水解方式裂解玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物的添加剂,其特征在于,所述添加剂包含至少一种具有选自SEQ ID No.1-15的氨基酸序列或其功能性变体的多肽,其中在所述功能性变体与至少一个所述氨基酸序列之间的序列同一性为至少40%;并且任选地包含助剂。
14.根据实施方案13的添加剂,其特征在于,包含至少一种根据实施方案1至11之一的多肽。
15.根据实施方案13或14之一的添加剂,其特征在于,所述助剂选自:至少一种惰性载体,以及任选地,其他成分,例如维生素和/或矿物质和/或酶和/或其他用于使真菌毒素解毒的组分。
16.根据实施方案13、14或15之一的添加剂,其特征在于,在所述添加剂中,以最高10,000U/g,优选地最高1,000U/g,更优选地最高100U/g,和最优选地最高10U/g的浓度包含至少一种根据实施方案1至11之一的多肽。
17.根据实施方案13至16之一的添加剂,其特征在于,所述添加剂以经包囊或经包衣的形式存在。
18.根据实施方案13至17之一的添加剂用于以水解方式裂解在饲料中、在食物中或在中的玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物的用途,所述饲料特别地为用于猪、家禽和水产养殖的饲料。
19.用于以水解方式裂解玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物的方法,其特征在于,所述玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物被具有选自SEQ IDNo.1-15的氨基酸序列或其功能性变体的多肽水解,其中在所述功能性变体与至少一个所述氨基酸序列之间的序列同一性为至少40%。
20.根据实施方案19的方法,其特征在于,在根据实施方案14至17之一的添加剂中使用所述多肽。
21.根据实施方案20的方法,其特征在于,将所述多肽或添加剂与被玉米赤霉烯酮和/或被至少一种玉米赤霉烯酮衍生物污染的饲料或食物相掺混,使所述被污染的饲料或食物与湿气相接触,并且所述多肽或添加剂水解在所述被污染的饲料或食物中所包含的玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物。
22.根据实施方案19至21之一的方法,其特征在于,至少70%,优选地至少80%,特别优选地至少90%的所述玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物被水解。
序列表
<110> Erber Aktiengesellschaft
<120> 多肽,分离的其多核苷酸,以及包含多肽的添加剂、其用途及方法
<130> P05341PCT
<160> 69
<170> PatentIn version 3.5
<210> 1
<211> 328
<212> PRT
<213> 红串红球菌 (Rhodococcus erythropolis)
<400> 1
Met Ala Glu Glu Gly Thr Arg Ser Glu Ala Ala Asp Ala Ala Thr Gln
1 5 10 15
Ala Arg Gln Leu Pro Asp Ser Arg Asn Ile Phe Val Ser His Arg Phe
20 25 30
Pro Glu Arg Gln Val Asp Leu Gly Glu Val Val Met Asn Phe Ala Glu
35 40 45
Ala Gly Ser Pro Asp Asn Pro Ala Leu Leu Leu Leu Pro Glu Gln Thr
50 55 60
Gly Ser Trp Trp Ser Tyr Glu Pro Val Met Gly Leu Leu Ala Glu Asn
65 70 75 80
Phe His Val Phe Ala Val Asp Ile Arg Gly Gln Gly Arg Ser Thr Trp
85 90 95
Thr Pro Arg Arg Tyr Ser Leu Asp Asn Phe Gly Asn Asp Leu Val Arg
100 105 110
Phe Ile Ala Leu Val Ile Lys Arg Pro Val Val Val Ala Gly Asn Ser
115 120 125
Ser Gly Gly Leu Leu Ala Ala Trp Leu Ser Ala Tyr Ala Met Pro Gly
130 135 140
Gln Ile Arg Ala Ala Leu Cys Glu Asp Ala Pro Phe Phe Ala Ser Glu
145 150 155 160
Leu Val Pro Ala Tyr Gly His Ser Val Leu Gln Ala Ala Gly Pro Ala
165 170 175
Phe Glu Leu Tyr Arg Asp Phe Leu Gly Asp Gln Trp Ser Ile Gly Asp
180 185 190
Trp Lys Gly Phe Val Glu Ala Ala Lys Ala Ser Pro Ala Lys Ala Met
195 200 205
Gln Leu Phe Pro Thr Pro Asp Glu Ala Pro Gln Asn Leu Lys Glu Tyr
210 215 220
Asp Pro Glu Trp Gly Arg Ala Phe Phe Glu Gly Thr Val Ala Leu His
225 230 235 240
Cys Pro His Asp Arg Met Leu Ser Gln Val Lys Thr Pro Ile Leu Ile
245 250 255
Thr His His Ala Arg Thr Ile Asp Pro Glu Thr Gly Glu Leu Leu Gly
260 265 270
Ala Leu Ser Asp Leu Gln Ala Glu His Ala Gln Asp Ile Ile Arg Ser
275 280 285
Ala Gly Val Arg Val Asp Tyr Gln Ser His Pro Asp Ala Leu His Met
290 295 300
Met His Leu Phe Asp Pro Ala Arg Tyr Ala Glu Ile Leu Thr Ser Trp
305 310 315 320
Ser Ala Thr Leu Pro Ala Asn Asp
325
<210> 2
<211> 308
<212> PRT
<213> 紫黑链霉菌 (Streptomyces violaceusniger)
<400> 2
Met Ala Asp Pro Ala Gln Arg Asp Val Tyr Val Pro His Ala Tyr Pro
1 5 10 15
Glu Lys Gln Ala Asp Leu Gly Glu Ile Thr Met Asn Tyr Ala Glu Ala
20 25 30
Gly Glu Pro Asp Met Pro Ala Val Leu Leu Ile Pro Glu Gln Thr Gly
35 40 45
Ser Trp Trp Gly Tyr Glu Glu Ala Met Gly Leu Leu Ala Glu Asn Phe
50 55 60
His Val Tyr Ala Val Asp Leu Arg Gly Gln Gly Arg Ser Ser Trp Ala
65 70 75 80
Pro Lys Arg Tyr Ser Leu Asp Asn Phe Gly Asn Asp Leu Val Arg Phe
85 90 95
Ile Ala Leu Val Val Lys Arg Pro Val Ile Val Ala Gly Asn Ser Ser
100 105 110
Gly Gly Val Leu Ala Ala Trp Leu Ser Ala Tyr Ser Met Pro Gly Gln
115 120 125
Val Arg Gly Ala Leu Cys Glu Asp Ala Pro Phe Phe Ala Ser Glu Leu
130 135 140
Val Thr Thr Cys Gly His Ser Ile Arg Gln Ala Ala Gly Pro Met Phe
145 150 155 160
Glu Leu Phe Arg Thr Tyr Leu Gly Asp Gln Trp Ser Val Gly Asp Trp
165 170 175
Thr Gly Tyr Cys Arg Ala Ala Asp Ala Ser Ser Ser Pro Met Ala Arg
180 185 190
Tyr Phe Val Ala Asp Glu Ile Pro Gln His Met Arg Glu Tyr Asp Pro
195 200 205
Glu Trp Ala Arg Ala Phe Trp Glu Gly Thr Val Ala Leu His Cys Pro
210 215 220
His Glu Gln Leu Leu Thr Gln Val Lys Thr Pro Val Leu Leu Thr His
225 230 235 240
His Met Arg Asp Ile Asp Pro Asp Thr Gly His Leu Val Gly Ala Leu
245 250 255
Ser Asp Glu Gln Ala Ala Arg Ala Arg Leu Leu Met Glu Ser Ala Gly
260 265 270
Val Lys Val Asp Tyr Ala Ser Val Pro Asp Ala Leu His Met Met His
275 280 285
Gln Phe Asp Pro Pro Arg Tyr Val Glu Ile Phe Thr Gln Trp Ala Ala
290 295 300
Thr Leu Ala Ala
305
<210> 3
<211> 309
<212> PRT
<213> 天蓝色链霉菌 (Streptomyces coelicolor)
<400> 3
Met Val Thr Ser Pro Ala Leu Arg Asp Val His Val Pro His Ala Tyr
1 5 10 15
Pro Glu Gln Gln Val Asp Leu Gly Glu Ile Thr Met Asn Tyr Ala Glu
20 25 30
Ala Gly Asp Pro Gly Arg Pro Ala Val Leu Leu Ile Pro Glu Gln Thr
35 40 45
Gly Ser Trp Trp Ser Tyr Glu Glu Ala Met Gly Leu Leu Ala Glu His
50 55 60
Phe His Val Tyr Ala Val Asp Leu Arg Gly Gln Gly Arg Ser Ser Trp
65 70 75 80
Thr Pro Lys Arg Tyr Ser Leu Asp Asn Phe Gly Asn Asp Leu Val Arg
85 90 95
Phe Ile Ala Leu Val Val Arg Arg Pro Val Val Val Ala Gly Asn Ser
100 105 110
Ser Gly Gly Val Leu Ala Ala Trp Leu Ser Ala Tyr Ser Met Pro Gly
115 120 125
Gln Ile Arg Gly Val Leu Cys Glu Asp Pro Pro Phe Phe Ala Ser Glu
130 135 140
Leu Val Pro Ala His Gly His Ser Val Arg Gln Gly Ala Gly Pro Val
145 150 155 160
Phe Glu Leu Phe Arg Thr Tyr Leu Gly Asp Gln Trp Ser Val Gly Asp
165 170 175
Trp Glu Gly Phe Arg Ser Ala Ala Asp Ala Ser Ala Ser Pro Met Ala
180 185 190
Arg Ser Phe Val Ala Asp Thr Ile Pro Gln His Leu Lys Glu Tyr Asp
195 200 205
Pro Glu Trp Ala Arg Ala Phe Tyr Glu Gly Thr Val Gly Leu Asn Cys
210 215 220
Pro His Glu Arg Met Leu Asn Arg Val Asn Thr Pro Val Leu Leu Thr
225 230 235 240
His His Met Arg Gly Thr Asp Pro Glu Thr Gly Asn Leu Leu Gly Ala
245 250 255
Leu Ser Asp Glu Gln Ala Ala Gln Val Arg Arg Leu Met Glu Ser Ala
260 265 270
Gly Val Lys Val Asp Tyr Glu Ser Val Pro Asp Ala Ser His Met Met
275 280 285
His Gln Ser Asp Pro Ala Arg Tyr Ala Glu Ile Leu Thr Pro Trp Thr
290 295 300
Ala Ala Leu Ala Pro
305
<210> 4
<211> 309
<212> PRT
<213> 雷帕霉素链霉菌 (Streptomyces rapamycinicus)
<400> 4
Met Val Thr Ser Pro Ala Leu Arg Asp Val His Val Pro His Ala Tyr
1 5 10 15
Pro Glu Gln Gln Val Asp Leu Gly Glu Ile Thr Met Asn Tyr Ala Glu
20 25 30
Ala Gly Asp Pro Asp Arg Pro Ala Val Leu Leu Ile Pro Glu Gln Thr
35 40 45
Gly Ser Trp Trp Ser Tyr Glu Glu Ala Met Gly Leu Leu Ala Glu His
50 55 60
Phe His Val Tyr Ala Val Asp Leu Arg Gly Gln Gly Arg Ser Ser Trp
65 70 75 80
Thr Pro Lys Arg Tyr Ser Leu Asp Asn Phe Gly Asn Asp Leu Val Arg
85 90 95
Phe Ile Ala Leu Val Val Lys Arg Pro Val Val Val Ala Gly Asn Ser
100 105 110
Ser Gly Gly Val Leu Ala Ala Trp Leu Ser Ala Tyr Ser Met Pro Gly
115 120 125
Gln Leu Arg Gly Val Leu Cys Glu Asp Pro Pro Phe Phe Ala Ser Glu
130 135 140
Leu Val Pro Ala His Gly His Ser Val Arg Gln Gly Ala Gly Pro Val
145 150 155 160
Phe Glu Leu Phe Arg Thr Tyr Leu Gly Asp Gln Trp Ser Val Ser Asp
165 170 175
Trp Glu Gly Phe Cys Arg Ala Ala Gly Ala Ser Ala Ser Pro Met Ala
180 185 190
Arg Ser Phe Val Ala Asp Gly Ile Pro Gln His Leu Lys Glu Tyr Asp
195 200 205
Pro Glu Trp Ala Arg Ala Phe His Glu Gly Thr Val Gly Leu Asn Cys
210 215 220
Pro His Glu Arg Met Leu Gly Arg Val Asn Thr Pro Val Leu Leu Thr
225 230 235 240
His His Met Arg Gly Thr Asp Pro Glu Thr Gly Asn Leu Leu Gly Ala
245 250 255
Leu Ser Asp Glu Gln Ala Ala Gln Ala Arg Leu Leu Met Glu Ser Ala
260 265 270
Gly Val Arg Val Asp Tyr Glu Ser Val Pro Asp Ala Ser His Met Met
275 280 285
His Gln Ser Asp Pro Ala Arg Tyr Ala Glu Ile Phe Thr Arg Trp Ala
290 295 300
Ala Ala Leu Ala Pro
305
<210> 5
<211> 309
<212> PRT
<213> 浅青紫链霉菌 (Streptomyces lividans)
<400> 5
Met Val Thr Ser Pro Ala Leu Arg Asp Val His Val Pro His Ala Tyr
1 5 10 15
Pro Glu Gln Gln Val Asp Leu Gly Glu Ile Thr Met Asn Tyr Ala Glu
20 25 30
Ala Gly Asp Pro Gly Arg Pro Ala Val Leu Leu Ile Pro Glu Gln Thr
35 40 45
Gly Ser Trp Trp Ser Tyr Glu Glu Ala Met Gly Leu Leu Ala Glu His
50 55 60
Phe His Val Tyr Ala Val Asp Leu Arg Gly Gln Gly Arg Ser Ser Trp
65 70 75 80
Thr Pro Lys Arg Tyr Ser Leu Asp Asn Phe Gly Asn Asp Leu Val Arg
85 90 95
Phe Met Ala Leu Val Val Arg Arg Pro Val Val Val Ala Gly Asn Ser
100 105 110
Ser Gly Gly Val Leu Ala Ala Trp Leu Ser Ala Tyr Ser Met Pro Gly
115 120 125
Gln Ile Arg Gly Val Leu Cys Glu Asp Pro Pro Phe Phe Ala Ser Glu
130 135 140
Leu Val Pro Ala His Gly His Ser Val Arg Gln Gly Ala Gly Pro Val
145 150 155 160
Phe Glu Leu Phe Arg Thr Tyr Leu Gly Asp Gln Trp Ser Val Gly Asp
165 170 175
Trp Glu Gly Phe Arg Ser Ala Ala Gly Ala Ser Ala Ser Pro Met Ala
180 185 190
Arg Ser Phe Val Ala Asp Thr Ile Pro Gln His Leu Lys Glu Tyr Asp
195 200 205
Pro Glu Trp Ala Arg Ala Phe Tyr Glu Gly Thr Val Gly Leu Asn Cys
210 215 220
Pro His Glu Arg Met Leu Asn Arg Val Asn Thr Pro Val Leu Leu Thr
225 230 235 240
His His Met Arg Gly Thr Asp Pro Glu Thr Gly Asn Leu Leu Gly Ala
245 250 255
Leu Ser Asp Glu Gln Ala Ala Gln Ala Arg Arg Leu Met Glu Ser Ala
260 265 270
Gly Val Lys Val Asp Tyr Glu Ser Val Pro Asp Ala Ser His Met Met
275 280 285
His Gln Ser Asp Pro Ala Arg Tyr Ala Glu Ile Leu Thr Pro Trp Ala
290 295 300
Ala Ala Leu Ala Pro
305
<210> 6
<211> 309
<212> PRT
<213> 天蓝黄链霉菌 (Streptomyces coelicoflavus)
<400> 6
Met Val Thr Ser Pro Ala Leu Arg Asp Val His Val Pro His Ala Tyr
1 5 10 15
Pro Glu Gln Gln Val Asp Leu Gly Glu Ile Thr Met Asn Tyr Ala Glu
20 25 30
Ala Gly Asp Pro Asp Arg Pro Ala Val Leu Leu Ile Pro Glu Gln Thr
35 40 45
Gly Ser Trp Trp Ser Tyr Glu Glu Ala Met Gly Leu Leu Ser Glu His
50 55 60
Phe His Val Tyr Ala Val Asp Leu Arg Gly Gln Gly Arg Ser Ser Trp
65 70 75 80
Thr Pro Lys Arg Tyr Ser Leu Asp Asn Phe Gly Asn Asp Leu Val Arg
85 90 95
Phe Ile Ala Leu Val Val Lys Arg Pro Val Val Val Ala Gly Asn Ser
100 105 110
Ser Gly Gly Val Leu Ala Ala Trp Leu Ser Ala Tyr Ser Met Pro Gly
115 120 125
Gln Leu Arg Gly Val Leu Cys Glu Asp Pro Pro Phe Phe Ala Ser Glu
130 135 140
Leu Val Pro Ala His Gly His Ser Val Arg Gln Gly Ala Gly Pro Val
145 150 155 160
Phe Glu Leu Phe Arg Thr Tyr Leu Gly Asp Gln Trp Ser Val Gly Asp
165 170 175
Trp Glu Gly Phe Cys Arg Ala Ala Gly Ala Ser Ala Ser Pro Met Ala
180 185 190
Arg Ser Phe Val Ala Asp Gly Ile Pro Gln His Leu Gln Glu Tyr Asp
195 200 205
Pro Glu Trp Ala Arg Val Phe Tyr Glu Gly Thr Val Gly Leu Ser Cys
210 215 220
Pro His Glu Arg Met Leu Gly Gln Val Lys Thr Pro Val Leu Leu Thr
225 230 235 240
His His Met Arg Gly Ile Asp Pro Glu Thr Gly Asn Leu Leu Gly Ala
245 250 255
Leu Ser Asp Glu Gln Ala Leu Arg Ala Arg Arg Leu Met Asp Ser Ala
260 265 270
Gly Val Thr Val Asp Tyr Glu Ser Val Pro Asp Ala Ser His Met Met
275 280 285
His Gln Ser Ala Pro Ala Arg Tyr Val Glu Ile Phe Thr Arg Trp Ala
290 295 300
Ala Ala Leu Ala Pro
305
<210> 7
<211> 300
<212> PRT
<213> 锥猎蝽红球菌 (Rhodococcus triatome)
<400> 7
Met Pro His Asp Tyr Glu Glu Lys Leu Val Asp Leu Gly Glu Ile Asp
1 5 10 15
Leu Asn Tyr Ala Glu Ala Gly Ser Pro Asp Lys Pro Ala Leu Leu Leu
20 25 30
Ile Pro Ser Gln Ser Glu Ser Trp Trp Gly Tyr Glu Glu Ala Met Gly
35 40 45
Leu Leu Ala Glu Asp Tyr His Val Phe Ala Val Asp Met Arg Gly Gln
50 55 60
Gly Arg Ser Thr Trp Thr Pro Gly Arg Tyr Ser Leu Asp Asn Phe Gly
65 70 75 80
Asn Asp Leu Val Arg Phe Ile Asp Leu Val Ile Gly Arg Thr Val Ile
85 90 95
Val Ser Gly Asn Ser Ser Gly Gly Val Val Ala Ala Trp Leu Ala Ala
100 105 110
Phe Ser Leu Pro Gly Gln Val Arg Ala Ala Leu Ala Glu Asp Ala Pro
115 120 125
Phe Phe Ala Ser Glu Leu Asp Pro Lys Val Gly His Thr Ile Arg Gln
130 135 140
Ala Ala Gly His Ile Phe Val Asn Trp Arg Asp Tyr Leu Gly Asp Gln
145 150 155 160
Trp Ser Val Gly Asp Tyr Ala Gly Phe Leu Lys Ala Met Lys Ser Ser
165 170 175
Glu Val Pro Met Leu Arg Gln Val Pro Leu Pro Glu Thr Ala Pro Gln
180 185 190
Asn Leu Leu Glu Tyr Asp Pro Glu Trp Ala Arg Ala Phe Tyr Glu Gly
195 200 205
Thr Val Ala Gln Thr Cys Pro His Asp Tyr Met Leu Ser Gln Val Lys
210 215 220
Val Pro Met Leu Val Thr His His Ala Arg Met Ile Asp Glu Ala Thr
225 230 235 240
Ser Gly Leu Val Gly Ala Met Ser Asp Leu Gln Val Gln Lys Ala Ala
245 250 255
Glu Ile Ile Arg Gly Thr Gly Val Gln Val Asp Val Val Asp Leu Pro
260 265 270
Glu Ala Pro His Ile Leu His Gln Leu Ala Pro Lys Glu Tyr Val Glu
275 280 285
Ile Leu Asn Asn Trp Val Glu Lys Leu Pro Pro Val
290 295 300
<210> 8
<211> 307
<212> PRT
<213> 波罗地海海氏菌 (Hirschia baltica)
<400> 8
Met Ile Gln Asn Asn Lys Thr Ala Pro Tyr Lys Tyr Lys Glu Lys Leu
1 5 10 15
Val Asp Leu Gly Glu Ile Lys Met Asn Tyr Ile Val Ala Gly Ala Asp
20 25 30
Val Ser Pro Ala Leu Leu Leu Ile Pro Gly Gln Thr Glu Ser Trp Trp
35 40 45
Gly Phe Glu Ala Ala Ile Glu Lys Leu Glu Ser Asn Phe Gln Val Phe
50 55 60
Ala Ile Asp Leu Arg Gly Gln Gly Lys Ser Thr Gln Thr Pro Gly Arg
65 70 75 80
Tyr Ser Leu Asn Leu Met Gly Asn Asp Leu Val Arg Phe Ile Ser Leu
85 90 95
Val Ile Lys Arg Pro Val Ile Val Ser Gly Asn Ser Ser Gly Gly Leu
100 105 110
Leu Ala Ala Trp Leu Ser Ala Tyr Ala Met Pro Asn Gln Ile Arg Ala
115 120 125
Ile His Cys Glu Asp Ala Pro Phe Phe Thr Ala Glu Lys Ala Pro Leu
130 135 140
Tyr Gly His Ala Ile Gln Gln Ala Ala Gly Pro Ile Phe Ser Leu Met
145 150 155 160
Ser Lys Phe Leu Gly Asp Gln Trp Ser Ile Asn Asn Trp Glu Gly Leu
165 170 175
Lys Ala Ala Gln Ala Lys Asp Thr His Pro Ala Asn Lys Met Ile Ser
180 185 190
Gln Val Glu Gln Pro Pro Gln His Leu Lys Glu Tyr Asp Pro Glu Trp
195 200 205
Gly Arg Ala Phe Ile Glu Gly Lys Phe Asn Leu Asn Ser Pro His His
210 215 220
Thr Leu Leu Ser Asp Ile Lys Thr Pro Met Leu Tyr Thr His His Met
225 230 235 240
Arg Phe Glu Asp Pro Gln Thr Gly Leu Leu Ile Gly Ala Thr Ser Asp
245 250 255
Phe Gln Ala Ser Lys Ile Lys Glu Ile Ala Leu Lys Thr Gly Asn Ser
260 265 270
Phe Glu Leu Ile Asp Ala Pro Asp Ala Phe His Ser Met His Glu Ala
275 280 285
Asp Pro Gln Arg Phe Val Asp Ile Leu Thr Ser Trp Ile Glu Arg Leu
290 295 300
Asn Leu Gln
305
<210> 9
<211> 321
<212> PRT
<213> 巴西诺卡氏菌 (Nocardia brasiliensis)
<400> 9
Met Gly Ile Ser Glu Ala Ala Asp Arg Ala Asp Thr Phe Val Ala His
1 5 10 15
Lys Phe Glu Glu Gln Leu Val Asp Leu Gly Glu Ile Arg Met Asn Tyr
20 25 30
Val Ala Ala Gly Asp Pro Thr Ser Pro Ala Leu Leu Leu Ile Pro Ala
35 40 45
Gln Gly Glu Ser Trp Trp Gly Tyr Glu Asn Ala Ile Thr Leu Leu Ala
50 55 60
Asn Asp Phe Arg Val Phe Ala Ile Asp Leu Arg Gly Gln Gly Arg Ser
65 70 75 80
Thr Trp Thr Pro Gly Arg Tyr Asn Leu Asn Thr Trp Gly Asn Asp Val
85 90 95
Glu Arg Phe Ile Asp Leu Val Ile Gly Arg Pro Thr Leu Val Ala Gly
100 105 110
Asn Ser Ser Gly Gly Val Ile Ala Ala Trp Leu Ala Ala Tyr Ala Lys
115 120 125
Pro Gly Gln Ile Arg Gly Ala Met Leu Glu Asp Pro Pro Leu Phe Ala
130 135 140
Ser Gln Ala Ala Pro Pro Tyr Gly Pro Gly Ile Met Gln Thr Leu Gly
145 150 155 160
Pro Ile Phe Val Leu Trp Ala Lys Trp Leu Gly Pro Gln Trp Ser Val
165 170 175
Gly Asp Trp Asp Gly Met Val Ala Ala Ala Pro Arg Glu Leu Pro Glu
180 185 190
Phe Leu His Pro Gly Ile Ala Phe Leu Phe Gly Asp Gly Thr Gly Glu
195 200 205
Gly Ala Ala Ala Thr Pro Pro Gln His Leu Lys Glu Tyr Asp Pro Glu
210 215 220
Trp Ala Gln Ala Trp Ala Thr Asp Val Ala Asn Ala Gly Cys Asp His
225 230 235 240
Ala Thr Met Leu Ala Gln Asn Arg Val Pro Val Leu Leu Thr His His
245 250 255
Phe His Leu Thr Asp Pro Asp Thr Gly Gln Leu Met Gly Ala Met Thr
260 265 270
Asp Ile Gln Ala Gln Gln Ala Arg Arg Leu Leu Ala Ala Thr Gly Gln
275 280 285
Pro Val Thr Phe Thr Ala Leu Asp Ala Pro His Thr Met His Asp Pro
290 295 300
Glu Pro Glu Arg Tyr Phe Glu Val Leu Thr Glu Trp Ala Ser Ala Leu
305 310 315 320
Asp
<210> 10
<211> 319
<212> PRT
<213> 母牛分枝杆菌 (Mycobacterium vaccae)
<400> 10
Met Gly Arg Tyr Ala Gly Val Phe Gly Pro His Ala Pro Glu Ser Thr
1 5 10 15
Tyr Val Gly His Ala Tyr Pro Glu Gln Leu Phe Asp Thr Gly Glu Val
20 25 30
Arg Leu Asn Tyr Ala Val Ala Gly Asp Ala Ser Ala Ser Pro Leu Leu
35 40 45
Leu Ile Pro Gly Gln Thr Glu Ser Trp Trp Gly Tyr Glu Pro Ala Met
50 55 60
Gly Leu Leu Ala Glu His Phe His Val His Ala Val Asp Leu Arg Gly
65 70 75 80
Gln Gly Arg Ser Thr Arg Thr Pro Arg Arg Tyr Thr Leu Asp Asn Ile
85 90 95
Gly Asn Asp Leu Val Arg Phe Leu Asp Gly Val Ile Gly Arg Pro Ala
100 105 110
Phe Val Ser Gly Leu Ser Ser Gly Gly Leu Leu Ser Ala Trp Leu Ser
115 120 125
Ala Phe Ala Glu Pro Gly Gln Val Leu Ala Ala Cys Tyr Glu Asp Pro
130 135 140
Pro Phe Phe Ser Ser Glu Leu Asp Pro Val Ile Gly Pro Gly Leu Met
145 150 155 160
Ser Thr Val Gly Pro Leu Phe Ala Leu Tyr Val Lys Tyr Leu Gly Asp
165 170 175
Gln Trp Ser Ile Gly Asp Trp Asp Gly Phe Val Ala Gly Ala Pro Gln
180 185 190
Glu Leu Ala Gly Trp Gln Ala His Val Ala Leu Ala Gly Gly Thr Ala
195 200 205
Glu Pro Pro Gln His Leu Lys Glu Tyr Asp Pro Glu Trp Gly Arg Ala
210 215 220
Phe Val Gly Gly Thr Phe Thr Thr Gly Cys Pro His Gln Val Met Leu
225 230 235 240
Ser Gln Val Lys Val Pro Val Leu Phe Thr His His Phe Arg Met Leu
245 250 255
Asp Asp Glu Ser Gly Ser Leu Ile Gly Ala Ala Thr Asp Asp Gln Ala
260 265 270
Ala Arg Val Val Glu Leu Val Glu Asn Ser Gly Ala Pro Leu Thr Tyr
275 280 285
Arg Ser Phe Pro Met Met Gly His Ser Met His Ala Gln Asp Pro Ala
290 295 300
Leu Phe Ala Gly Thr Leu Val Asp Trp Phe Thr Ala Ala Arg Ser
305 310 315
<210> 11
<211> 319
<212> PRT
<213> 浅黄分枝杆菌 (Mycobacterium gilvum)
<400> 11
Met Gly Arg Tyr Ala Gly Val Phe Gly Pro His Ala Pro Glu Ala Thr
1 5 10 15
Tyr Val Glu His Gly Tyr Pro Glu Arg Leu Phe Asp Thr Gly Glu Val
20 25 30
Gln Leu Asn Tyr Val Val Ala Gly Asp Ala Ala Ala Pro Pro Leu Leu
35 40 45
Leu Ile Pro Gly Gln Ser Glu Ser Trp Trp Gly Tyr Glu Ala Ala Ile
50 55 60
Pro Leu Leu Ala Arg His Phe His Val His Ala Val Asp Leu Arg Gly
65 70 75 80
Gln Gly Arg Ser Thr Arg Thr Pro Gly Arg Tyr Thr Leu Asp Asn Val
85 90 95
Gly Asn Asp Leu Val Arg Phe Leu Asp Gly Val Ile Gly Arg Pro Ala
100 105 110
Phe Val Ser Gly Leu Ser Ser Gly Gly Leu Ala Ser Ala Trp Leu Ser
115 120 125
Ala Phe Ala Lys Pro Gly Gln Val Val Ala Ala Cys Trp Glu Asp Pro
130 135 140
Pro Phe Phe Ser Ser Glu Thr Ala Pro Ile Val Gly Pro Pro Ile Thr
145 150 155 160
Asp Ser Ile Gly Pro Leu Phe Gly Met Trp Ala Arg Tyr Leu Gly Asp
165 170 175
Gln Trp Ser Val Gly Asp Trp Asp Gly Phe Val Ala Ala Val Pro Thr
180 185 190
Glu Leu Ala Asp Trp Gln Ala His Val Ala Leu Val Val Gly Thr Ala
195 200 205
Asp Pro Pro Gln Asn Leu Arg Glu Tyr Asp Pro Glu Trp Gly Lys Ala
210 215 220
Phe Ile Thr Gly Thr Phe Ala Ala Ser Cys Pro His His Val Met Leu
225 230 235 240
Ser Lys Val Lys Val Pro Val Leu Tyr Thr His His Phe Arg Met Ile
245 250 255
Asp Glu Gly Ser Gly Gly Leu Ile Gly Ala Cys Ser Asp Ile Gln Ala
260 265 270
Gly Arg Val Thr Gln Leu Ala Lys Ser Gly Gly Arg Ser Val Thr Tyr
275 280 285
Arg Ser Phe Pro Met Met Ala His Ser Met His Gly Gln Asp Pro Ala
290 295 300
Leu Phe Ser Glu Thr Leu Val Glu Trp Phe Ser Arg Phe Thr Gly
305 310 315
<210> 12
<211> 322
<212> PRT
<213> Gordonia effusa
<400> 12
Met Pro Lys Ser Glu Ala Ala Asp Arg Ala Asp Ser Phe Val Ser His
1 5 10 15
Asp Phe Lys Glu Asn Ile Val Asp Leu Gly Glu Ile Arg Met Asn Tyr
20 25 30
Val Val Gln Gly Asn Lys Lys Ser Pro Ala Leu Leu Leu Ile Pro Ala
35 40 45
Gln Gly Glu Ser Trp Trp Gly Tyr Glu Ala Ala Ile Pro Leu Leu Ala
50 55 60
Lys His Phe Gln Val Phe Ala Ile Asp Leu Arg Gly Gln Gly Arg Thr
65 70 75 80
Thr Trp Thr Pro Gly Arg Tyr Thr Leu Asp Ile Phe Gly Asn Asp Val
85 90 95
Val Arg Phe Ile Asp Leu Val Ile Gly Arg Glu Thr Leu Ile Ala Gly
100 105 110
Asn Ser Ser Gly Gly Leu Ile Gly Ala Trp Leu Ala Ala Phe Ala Lys
115 120 125
Pro Gly Gln Val Arg Ala Val Met Leu Glu Asp Pro Pro Leu Phe Ala
130 135 140
Ser Glu Ile Arg Pro Pro Tyr Gly Pro Gly Ile Trp Gln Gly Leu Gly
145 150 155 160
Pro Met Phe Ala Ala Trp Ala Lys Trp Leu Gly Pro Gln Trp Ser Ile
165 170 175
Gly Asp Trp Asp Gly Met Val Lys Ala Leu Pro Asp Glu Leu Pro Glu
180 185 190
Asp Leu Leu Pro Gly Ile Gly Phe Met Leu Gly Asp Gly Glu Ser Asp
195 200 205
Gly Ala Ala Pro Thr Pro Pro Gln His Leu Lys Glu Tyr Asp Pro Glu
210 215 220
Trp Gly Ala Ser Trp Ala Ser Gly Phe Ala Asn Thr Gly Cys Glu His
225 230 235 240
Glu Ala Val Ile Ser Gln Val Arg Val Pro Val Leu Leu Thr His His
245 250 255
Phe Arg Gln Ile Asn Glu Glu Thr Gly His Leu Met Gly Ala Leu Ser
260 265 270
Asp Leu Gln Ala Ala Gln Val Arg His Ile Ile Glu Glu Val Ala Gly
275 280 285
Gln Glu Val Thr Tyr Val Ser Leu Asp Ala Pro His Thr Met His Glu
290 295 300
Pro Gln Pro Glu Arg Tyr Thr Asp Val Leu Leu Asp Trp Val Lys Lys
305 310 315 320
Leu Gly
<210> 13
<211> 328
<212> PRT
<213> Togninia minima
<400> 13
Met Asn Tyr Ala Thr Ala Gly Ser Ser Asp Lys Pro Ala Leu Leu Leu
1 5 10 15
Val Pro Gly Gln Ser Glu Ser Trp Trp Gly Tyr Glu Met Ala Met Trp
20 25 30
Leu Leu Lys Asp Asp Tyr Gln Val Phe Ala Val Asp Met Arg Gly Gln
35 40 45
Gly Gln Ser Thr Trp Thr Pro Gly Arg Tyr Ser Leu Asp Thr Phe Gly
50 55 60
Asn Asp Leu Val Lys Phe Ile Asp Ile Val Ile Lys Arg Pro Val Val
65 70 75 80
Val Ser Gly Leu Ser Ser Gly Gly Val Val Ser Ala Trp Leu Ser Ala
85 90 95
Phe Ala Lys Pro Gly Gln Ile Arg Ala Ala Val Tyr Glu Asp Pro Pro
100 105 110
Leu Phe Ala Ser Gln Ser Lys Pro Ala Ile Gly Gln Ser Val Met Gln
115 120 125
Thr Val Ala Gly Pro Phe Phe Asn Leu Trp Tyr Lys Trp Leu Gly Ala
130 135 140
Gln Trp Thr Ile Gly Asp Gln Ala Gly Met Val Ala Ala Met Pro Lys
145 150 155 160
Glu Ile Pro Ala Trp Ile Leu Gln Tyr Leu Gly Asn Thr Thr Ser Gly
165 170 175
Pro Thr Gly Leu Asp Leu Thr Leu Asn Glu Tyr Asp Pro Glu Trp Gly
180 185 190
His Gly Phe Val Ser Gly Thr Val Asp Ala Thr Cys Asp His Glu Ala
195 200 205
Met Leu Thr His Val Lys Val Pro Val Leu Phe Thr His His Ser Arg
210 215 220
Ala Ile Asp Pro Tyr Thr Gly Asn Leu Ile Gly Ser Val Ser Asp Thr
225 230 235 240
Gln Val Ser Tyr Ala Gln Gly Leu Ile Thr Thr Asn Gly Asn Gln Ser
245 250 255
Phe Thr Leu Lys Asn Phe Pro Leu Ala Ser His Asp Met His Asn Ser
260 265 270
Asp Pro Ala Thr Tyr Val Ser Ala Ile Thr Thr Trp Met Ala Ser Leu
275 280 285
Gly Ile Gly Ser Ala Val Ile Pro Gly Pro Val Lys Val Ala Ser Ala
290 295 300
Ser Ala Gln Val Ser Ala Ala Ser Thr Ala Pro Pro Ser Cys Thr Ser
305 310 315 320
Thr Ser Ala Pro Ser Thr Gly His
325
<210> 14
<211> 280
<212> PRT
<213> 奇迹束丝放线菌 (Actinosynnema mirum)
<400> 14
Met Thr Val Val Asp Pro Pro Ala Pro Arg Asp Phe Pro Glu Leu Leu
1 5 10 15
Val Asp Leu Gly Glu Val Val Leu Asn His Ala Glu Ala Gly Ser Pro
20 25 30
Asp Arg Pro Ala Leu Val Pro Val Pro Glu Gln Gly Gly Ser Trp Trp
35 40 45
Ser Tyr Glu Arg Val Met Pro Leu Pro Ala Arg Asp Phe His Val Phe
50 55 60
Ala Val Asp Leu Arg Gly Arg Gly Arg Ser Thr Arg Thr Pro Arg Arg
65 70 75 80
Tyr Ser Leu Asp Asp Phe Gly Asn Asp Leu Val Arg Phe Leu Ala Leu
85 90 95
Val Val Arg Arg Pro Ala Val Val Ala Gly Asn Ser Ser Gly Gly Val
100 105 110
Leu Ala Ala Trp Ser Ser Ala Tyr Ala Met Pro Gly Gln Val Arg Ala
115 120 125
Val Leu Leu Glu Asp Pro Pro Leu Phe Ser Ser Glu Leu Thr Pro Val
130 135 140
Cys Gly Pro Gly Val Arg Gln Ala Ala Gly Pro Leu Phe Glu Leu Leu
145 150 155 160
Ser Thr His Leu Gly Asp Gln Trp Gly Gly Gly Arg Pro Gly Arg Val
165 170 175
His Gly Gly Val Pro Arg Leu Gly Leu Ala Ala Ala Ala Ala Val Arg
180 185 190
Val Ala Arg Arg Ala Ala Ala Thr Asp Ala Arg Gly Arg Pro Gly Ala
195 200 205
Ala Arg Gly Arg Pro Ala Gly Val Gly Gly Ala Ala Arg Arg Gly Arg
210 215 220
Gly Gly Arg Glu Arg Thr Gly Thr Thr Thr Val Leu Ser Gly Leu Thr
225 230 235 240
Gly Ser Arg Thr Ser Gly Thr Gly Arg Cys Arg Lys Pro Phe Arg Leu
245 250 255
Arg Gln Trp Trp Ala Gly Gly Ala Arg Gly Pro Pro Pro Pro Arg Gln
260 265 270
Ile Arg Ala Asp Val Arg Thr Arg
275 280
<210> 15
<211> 326
<212> PRT
<213> 白色库茨涅尔氏菌 (Kutzneria albida)
<400> 15
Met Ser Val Pro Val Thr Pro Ser Ala Arg Asn Val Phe Val Pro His
1 5 10 15
Ala Phe Pro Glu Lys Gln Ile Asp Leu Gly Glu Val Val Leu Asn Tyr
20 25 30
Ala Glu Ala Gly Thr Pro Asp Lys Pro Ala Leu Leu Leu Leu Pro Glu
35 40 45
Gln Thr Gly Ser Trp Trp Ser Tyr Glu Pro Ala Met Gly Leu Leu Ala
50 55 60
Glu His Phe His Val Phe Ala Val Asp Leu Arg Gly Gln Gly Arg Ser
65 70 75 80
Thr Trp Thr Pro Gly Arg Tyr Ser Leu Asp Asn Phe Gly Asn Asp Leu
85 90 95
Val Arg Phe Ile Ala Leu Ala Ile Arg Arg Pro Val Val Val Ala Gly
100 105 110
Cys Ser Ser Gly Gly Val Leu Ala Ala Trp Leu Ser Ala Tyr Ala Leu
115 120 125
Pro Gly Gln Ile Arg Gly Ala Leu Cys Glu Asp Ala Pro Leu Phe Ala
130 135 140
Ser Glu Leu Thr Pro Ala His Gly His Gly Val Arg Gln Gly Ala Gly
145 150 155 160
Pro Val Phe Glu Leu Tyr Arg Asp Tyr Leu Gly Asp Gln Trp Ser Val
165 170 175
Gly Asp Trp Ala Gly Leu Val Ala Ala Ala Gln Ala Ser Pro Ala Lys
180 185 190
Met Met Ser Leu Phe Lys Met Pro Gly Glu Pro Pro Gln Asn Leu Arg
195 200 205
Glu Tyr Asp Pro Glu Trp Ala Arg Val Phe Phe Glu Gly Thr Val Gly
210 215 220
Leu His Cys Pro His Asp Arg Met Leu Ser Gln Val Lys Thr Pro Val
225 230 235 240
Leu Ile Thr His His Ala Arg Thr Thr Asp Pro Glu Thr Gly Glu Phe
245 250 255
Leu Gly Ala Leu Ser Glu Leu Gln Ala Glu Arg Ala Gln Ala Ile Ile
260 265 270
Arg Ala Ala Gly Val Pro Val Asp Tyr Gln Ser Phe Pro Asp Ala Ala
275 280 285
His Ala Met His Thr Thr Glu Pro Ala Arg Tyr Ala Ala Val Leu Thr
290 295 300
Ala Trp Ala Ala Lys Leu Pro Pro Val Ala Asp Thr Ser Pro Ser Ala
305 310 315 320
Ala Ala Ser Ala His Val
325
<210> 16
<211> 987
<212> DNA
<213> 红串红球菌
<220>
<221> misc_feature
<223> 编码具有SEQ ID NO: 1的多肽的人工DNA序列
<400> 16
atggccgaag aaggaactag gtccgaagca gcggatgctg ccacacaagc gagacagcta 60
cccgattcgc ggaacatctt tgtctcgcac cgatttccgg aaaggcaggt cgatctcggt 120
gaagtggtga tgaacttcgc ggaggcgggc tctccggaca acccggcact gctcctcctc 180
cccgagcaga ccgggtcgtg gtggagttac gagccagtga tgggtcttct ggcagagaac 240
tttcatgtct ttgccgtcga tatccgtggg caaggtcgca gtacctggac gccacggcga 300
tacagcctgg acaacttcgg caatgatctg gtgcgtttca tcgctctggt catcaagcgc 360
cctgtcgtcg tggcagggaa ctcctcgggg gggctgctgg ccgcctggct ctcggcgtac 420
gcgatgcccg gccagatccg tgcagcattg tgtgaggacg caccgttctt tgcgtcggag 480
ttggtccccg catacggtca ctcggttctg caggcggcgg gtccggcatt cgagttgtac 540
cgggacttcc tcggggacca gtggtcgatt ggggactgga aagggttcgt tgaggcagcc 600
aaagcgtcgc cggcaaaggc tatgcaatta tttccgaccc cggatgaggc gccgcagaat 660
ctcaaggaat acgacccgga atgggggcgc gcattcttcg aagggactgt ggcactgcac 720
tgcccacacg acaggatgct ctcgcaagtc aagacaccaa ttctcatcac tcaccacgcg 780
cggacgatcg accccgagac gggcgagctg ttgggcgcgc tctccgacct tcaggcagag 840
catgcgcagg acatcattcg gtctgcgggc gttcgggtgg actatcagtc gcaccccgac 900
gcgcttcaca tgatgcatct gttcgatccc gctcgttacg cggagatctt gacatcctgg 960
tccgcaacac tgcctgcgaa cgactag 987
<210> 17
<211> 987
<212> DNA
<213> 人工序列
<220>
<223> ORF经密码子优化并因而不同于天然出现的DNA序列
<400> 17
atggcagaag aaggcacccg tagcgaagca gcagatgcag caacccaggc acgtcagctg 60
ccggatagcc gtaacatttt tgttagccat cgttttccgg aacgtcaggt tgatctgggt 120
gaagttgtta tgaattttgc agaagcaggt agtccggata atccggcatt actgctgctg 180
ccggaacaga ccggtagttg gtggtcttat gaaccggtta tgggtctgct ggcagaaaac 240
tttcatgttt ttgcagttga tattcgtggt cagggtcgta gcacctggac accgcgtcgt 300
tatagcctgg ataattttgg taatgatctg gtgcgtttta ttgccctggt tattaaacgt 360
ccggttgttg ttgcaggtaa tagcagcggt ggcctgctgg ctgcatggct gagcgcctat 420
gcaatgcctg gtcagattcg tgcagcactg tgtgaagatg caccgttttt tgcaagcgaa 480
ctggttcctg cctatggtca tagcgttctg caggcagcag gtccggcatt tgaactgtat 540
cgtgattttc tgggtgatca gtggtcaatt ggtgattgga aaggttttgt tgaagcagca 600
aaagcaagtc cggctaaagc aatgcagctg tttccgacac cggatgaagc accgcagaat 660
ctgaaagaat atgatccgga atggggtcgt gcattttttg aaggcaccgt tgcactgcat 720
tgtccgcatg atcgtatgct gagccaggtt aaaaccccga ttctgattac ccatcatgca 780
cgtaccatcg atccggaaac cggtgaactg ctgggtgcac tgagtgatct gcaggccgaa 840
catgcacagg atattattcg tagtgccggt gttcgtgttg attatcagag ccatcctgat 900
gcactgcaca tgatgcacct gtttgatccg gcacgttatg cagaaattct gaccagttgg 960
agcgcaaccc tgcctgcaaa tgattaa 987
<210> 18
<211> 927
<212> DNA
<213> 人工序列
<220>
<223> ORF经密码子优化并因而不同于天然出现的DNA序列
<220>
<221> misc_feature
<223> 编码具有SEQ ID NO: 2的多肽的人工DNA序列
<400> 18
atggcagatc cggcacagcg tgatgtttat gttccgcatg catatccgga aaaacaggca 60
gatctgggtg aaattaccat gaattatgcc gaagccggtg aacctgatat gcctgcagtt 120
ctgctgattc cggaacagac cggtagttgg tggggttatg aagaagcaat gggtctgctg 180
gcagaaaact ttcatgttta tgcagttgat ctgcgtggtc agggtcgtag cagctgggca 240
ccgaaacgtt atagcctgga taattttggt aatgatctgg tgcgttttat tgccctggtt 300
gttaaacgtc cggttattgt tgcaggtaat agcagcggtg gtgttctggc agcatggctg 360
agcgcatata gcatgcctgg tcaggttcgt ggtgcactgt gtgaagatgc accgtttttt 420
gcaagcgaac tggttaccac ctgtggtcat agcattcgtc aggcagcagg tccgatgttt 480
gaactgtttc gtacctatct gggcgatcag tggtcagttg gtgattggac cggctattgt 540
cgtgcagcag atgcaagcag cagcccgatg gcacgttatt ttgttgcaga tgaaattccg 600
cagcacatgc gtgaatatga tccggaatgg gcacgtgcat tttgggaagg caccgttgca 660
ctgcattgtc cgcatgaaca gctgctgacc caggttaaaa caccggtgct gctgacacat 720
cacatgcgcg atattgatcc tgataccggt catctggttg gtgccctgag tgatgaacag 780
gcagcccgtg cacgtctgct gatggaaagt gccggtgtta aagttgatta tgcaagcgtt 840
ccggatgcac tgcacatgat gcaccagttt gatccgcctc gttatgttga aatctttacc 900
cagtgggcag caaccctggc agcataa 927
<210> 19
<211> 930
<212> DNA
<213> 人工序列
<220>
<223> ORF经密码子优化并因而不同于天然出现的DNA序列
<220>
<221> misc_feature
<223> 编码具有SEQ ID NO: 3的多肽的人工DNA序列
<400> 19
atggttacca gtccggcact gcgtgatgtt catgttccgc atgcatatcc ggaacagcag 60
gttgatctgg gtgaaattac catgaattat gccgaagccg gtgatccggg tcgtccggca 120
gttctgctga tcccggaaca gaccggtagt tggtggtctt atgaagaagc aatgggtctg 180
ctggcagaac attttcatgt ttatgcagtt gatctgcgtg gtcagggtcg tagcagctgg 240
accccgaaac gttatagcct ggataatttt ggtaatgatc tggtgcgttt tattgccctg 300
gttgttcgtc gtccggttgt tgttgcaggt aatagcagcg gtggtgttct ggcagcatgg 360
ctgagcgcat atagcatgcc tggtcagatt cgtggtgtgc tgtgtgaaga tccgcctttt 420
tttgcaagcg aactggttcc ggcacatggt catagcgttc gtcagggtgc aggtccggtt 480
tttgaactgt ttcgtaccta tctgggcgat cagtggtcag ttggtgattg ggaaggtttt 540
cgtagcgcag cagatgcaag cgcaagcccg atggcacgta gctttgttgc agataccatt 600
ccgcagcatc tgaaagaata tgatccggaa tgggcacgtg cattttatga aggcaccgtt 660
ggtctgaatt gtccgcatga acgtatgctg aatcgtgtta atacaccggt gctgctgacc 720
catcacatgc gtggcaccga tccggaaacc ggtaatctgc tgggtgcact gagtgatgaa 780
caggcagcac aggtgcgtcg tctgatggaa agtgccggtg ttaaagttga ttatgaaagc 840
gttccggatg caagccacat gatgcaccag agcgatccgg cacgttatgc agaaattctg 900
accccgtgga ccgcagcact ggcaccgtaa 930
<210> 20
<211> 930
<212> DNA
<213> 人工序列
<220>
<223> ORF经密码子优化并因而不同于天然出现的DNA序列
<220>
<221> misc_feature
<223> 编码具有SEQ ID NO: 4的多肽的人工DNA序列
<400> 20
atggttacca gtccggcact gcgtgatgtt catgttccgc atgcatatcc ggaacagcag 60
gttgatctgg gtgaaattac catgaattat gccgaagccg gtgatcctga tcgtccggca 120
gttctgctga tcccggaaca gaccggtagt tggtggtcat atgaagaagc aatgggtctg 180
ctggcagaac attttcatgt ttatgcagtt gatctgcgtg gtcagggtcg tagcagctgg 240
accccgaaac gttatagcct ggataatttt ggtaatgatc tggtgcgttt tattgccctg 300
gttgttaaac gtccggttgt tgttgcaggt aatagcagcg gtggtgttct ggcagcatgg 360
ctgagcgcat atagcatgcc tggtcagctg cgtggtgtgc tgtgtgaaga tccgcctttt 420
tttgcaagcg aactggttcc ggcacatggt catagcgttc gtcagggtgc aggtccggtt 480
tttgaactgt ttcgtaccta tctgggcgat cagtggtcag ttagcgattg ggaaggtttt 540
tgtcgtgcag ccggtgcaag cgcaagcccg atggcacgta gctttgttgc agatggtatt 600
ccgcagcatc tgaaagaata tgatccggaa tgggcacgtg catttcatga aggcaccgtt 660
ggtctgaatt gtccgcatga acgtatgctg ggtcgtgtta atacaccggt gctgctgacc 720
catcatatgc gtggcaccga tccggaaacc ggtaatctgc tgggtgcact gagtgatgaa 780
caggcagcac aggcacgtct gctgatggaa agtgccggtg ttcgtgttga ttatgaaagc 840
gttccggatg caagccatat gatgcaccag agcgatccgg cacgttatgc agaaatcttt 900
acccgttggg cagcagccct ggcaccgtaa 930
<210> 21
<211> 930
<212> DNA
<213> 人工序列
<220>
<223> ORF经密码子优化并因而不同于天然出现的DNA序列
<220>
<221> misc_feature
<223> 编码具有SEQ ID NO: 5的多肽的人工DNA序列
<400> 21
atggttacca gtccggcact gcgtgatgtt catgttccgc atgcatatcc ggaacagcag 60
gttgatctgg gtgaaattac catgaattat gccgaagccg gtgatccggg tcgtccggca 120
gttctgctga tcccggaaca gaccggtagt tggtggtctt atgaagaagc aatgggtctg 180
ctggcagaac attttcatgt ttatgcagtt gatctgcgtg gtcagggtcg tagcagctgg 240
accccgaaac gttatagcct ggataatttt ggtaatgatc tggtgcgttt tatggcactg 300
gttgttcgtc gtccggttgt tgttgcaggt aatagcagcg gtggtgttct ggcagcatgg 360
ctgagcgcat atagcatgcc tggtcagatt cgtggtgtgc tgtgtgaaga tccgcctttt 420
tttgcaagcg aactggttcc ggcacatggt catagcgttc gtcagggtgc aggtccggtt 480
tttgaactgt ttcgtaccta tctgggcgat cagtggtcag ttggtgattg ggaaggtttt 540
cgtagcgcag ccggtgcaag cgcaagcccg atggcacgta gctttgttgc agataccatt 600
ccgcagcatc tgaaagaata tgatccggaa tgggcacgtg cattttatga aggcaccgtt 660
ggtctgaatt gtccgcatga acgtatgctg aatcgtgtta atacaccggt gctgctgacc 720
catcacatgc gtggcaccga tccggaaacc ggtaatctgc tgggtgcact gagtgatgaa 780
caggcagcac aggcacgtcg tctgatggaa agtgccggtg ttaaagttga ttatgaaagc 840
gttccggatg caagccacat gatgcaccag agcgatccgg cacgttatgc agaaattctg 900
accccgtggg cagcagccct ggcaccgtaa 930
<210> 22
<211> 930
<212> DNA
<213> 人工序列
<220>
<223> ORF经密码子优化并因而不同于天然出现的DNA序列
<220>
<221> misc_feature
<223> 编码具有SEQ ID NO: 6的多肽的人工DNA序列
<400> 22
atggttacca gtccggcact gcgtgatgtt catgttccgc atgcatatcc ggaacagcag 60
gttgatctgg gtgaaattac catgaattat gccgaagccg gtgatcctga tcgtccggca 120
gttctgctga tcccggaaca gaccggtagt tggtggtctt atgaagaagc aatgggtctg 180
ctgagcgaac attttcatgt ttatgcagtt gatctgcgtg gtcagggtcg tagcagctgg 240
accccgaaac gttatagcct ggataatttt ggtaatgatc tggtgcgttt tattgccctg 300
gttgttaaac gtccggttgt tgttgcaggt aatagcagcg gtggtgttct ggcagcatgg 360
ctgagcgcat atagcatgcc tggtcagctg cgtggtgtgc tgtgtgaaga tccgcctttt 420
tttgcaagcg aactggttcc ggcacatggt catagcgttc gtcagggtgc aggtccggtt 480
tttgaactgt ttcgtaccta tctgggcgat cagtggtcag ttggtgattg ggaaggtttt 540
tgtcgtgcag ccggtgcaag cgcaagcccg atggcacgta gctttgttgc agatggtatt 600
ccgcagcatc tgcaagaata tgatccggaa tgggcacgtg ttttttatga aggcaccgtt 660
ggtctgagct gtccgcatga acgtatgctg ggtcaggtta aaacaccggt gctgctgacc 720
catcacatgc gtggtatcga tccggaaacc ggtaatctgc tgggtgcact gagtgatgaa 780
caggccctgc gtgcacgtcg tctgatggat agtgccggtg ttaccgttga ttatgaaagc 840
gttccggatg caagccacat gatgcaccag agcgcaccgg cacgttatgt tgaaatcttt 900
acccgttggg cagcagccct ggcaccgtaa 930
<210> 23
<211> 903
<212> DNA
<213> 人工序列
<220>
<223> ORF经密码子优化并因而不同于天然出现的DNA序列
<220>
<221> misc_feature
<223> 编码具有SEQ ID NO: 7的多肽的人工DNA序列
<400> 23
atgccgcacg attatgaaga aaaactggtt gatctgggcg aaatcgatct gaattatgca 60
gaagcaggta gtccggataa accggcactg ctgctgattc cgagccagag cgaaagttgg 120
tggggctatg aagaagcaat gggtctgctg gccgaagatt atcatgtttt tgcagttgat 180
atgcgtggtc agggtcgtag cacctggaca ccgggtcgtt atagcctgga taattttggt 240
aatgatctgg tgcgctttat cgatctggtt attggtcgta ccgttattgt tagcggtaat 300
agcagcggtg gtgttgttgc agcatggctg gcagcattta gcctgcctgg tcaggttcgt 360
gcagcactgg cagaagatgc accgtttttt gcaagcgaac tggacccgaa agtgggtcat 420
accattcgtc aggcagcagg tcatattttt gttaactggc gtgattatct gggtgatcag 480
tggtcagttg gtgattatgc aggttttctg aaagcaatga aaagcagcga agttccgatg 540
ctgcgtcagg ttccgctgcc ggaaaccgca ccgcagaatc tgctggaata tgatccggaa 600
tgggcacgtg cattttatga aggcaccgtt gcacagacct gtccgcatga ttatatgctg 660
agccaggtta aagtgcctat gctggttacc catcatgcac gtatgattga tgaagcaacc 720
agcggtctgg ttggtgcaat gagcgatctg caggttcaga aagcagcaga aattattcgt 780
ggcaccggtg ttcaggttga tgttgttgat ctgccggaag caccgcatat tctgcatcag 840
ctggcaccga aagaatatgt ggaaattctg aataactggg tggaaaaact gcctccggtt 900
taa 903
<210> 24
<211> 924
<212> DNA
<213> 人工序列
<220>
<223> ORF经密码子优化并因而不同于天然出现的DNA序列
<220>
<221> misc_feature
<223> 编码具有SEQ ID NO: 8的多肽的人工DNA序列
<400> 24
atgatccaga acaataaaac cgcaccgtat aaatacaaag aaaaactggt tgatctgggc 60
gaaatcaaaa tgaactatat tgttgccggt gcagatgtta gtccggcact gctgctgatt 120
ccgggtcaga ccgaaagttg gtggggtttt gaagcagcaa ttgagaaact ggaaagcaac 180
tttcaggtgt ttgcaattga tctgcgtggt cagggtaaaa gcacccagac accgggtcgt 240
tatagcctga atctgatggg taatgatctg gttcgtttta ttagcctggt tattaaacgt 300
ccggttattg ttagcggtaa tagcagcggt ggtctgctgg cagcatggct gagcgcctat 360
gcaatgccga atcagattcg tgcaattcat tgtgaagatg caccgttttt taccgcagaa 420
aaagcaccgc tgtatggtca tgcaattcag caggcagcag gtccgatttt tagcctgatg 480
agcaaatttc tgggtgatca gtggtcaatt aacaattggg aaggtctgaa agcagcacag 540
gcaaaagata cccatccggc aaacaaaatg attagccagg ttgaacagcc tccgcagcat 600
ctgaaagaat atgatccgga atggggtcgt gcatttattg aaggcaaatt taacctgaac 660
agtccgcatc ataccctgct gagcgacatt aaaaccccga tgctgtatac ccatcacatg 720
cgttttgaag atccgcagac aggtctgctg attggtgcaa ccagcgattt tcaggcaagc 780
aaaatcaaag aaattgccct gaaaaccggc aatagcttcg aactgattga tgcaccggat 840
gcatttcata gtatgcatga agccgatccg cagcgttttg ttgatattct gaccagctgg 900
attgaacgtc tgaatctgca gtaa 924
<210> 25
<211> 966
<212> DNA
<213> 人工序列
<220>
<223> ORF经密码子优化并因而不同于天然出现的DNA序列
<220>
<221> misc_feature
<223> 编码具有SEQ ID NO: 9的多肽的人工DNA序列
<400> 25
atgggtatta gcgaagcagc agatcgtgca gatacctttg ttgcacataa atttgaagaa 60
cagctggttg atctgggtga aattcgtatg aattatgttg cagccggtga tccgaccagt 120
ccggcactgc tgctgattcc ggcacagggt gaaagttggt ggggttatga aaatgcaatt 180
accctgctgg caaatgattt tcgtgttttt gcaattgatc tgcgtggtca gggtcgtagc 240
acctggacac cgggtcgtta taatctgaat acctggggta atgatgtgga acgctttatt 300
gatctggtta ttggtcgtcc gaccctggtt gcaggtaata gcagcggtgg tgttattgca 360
gcatggctgg cagcctatgc aaaaccgggt cagattcgtg gtgcaatgct ggaagatccg 420
cctctgtttg caagccaggc agcaccgcct tatggtccgg gtattatgca gaccctgggt 480
ccgatttttg ttctgtgggc aaaatggctg ggtccgcagt ggtcagttgg tgattgggat 540
ggtatggttg cagcggcacc gcgtgaactg ccggaatttc tgcatccggg tatcgcattt 600
ctgtttggtg atggcaccgg tgaaggtgca gcagcaaccc ctccgcagca tctgaaagaa 660
tatgatccgg aatgggcaca ggcatgggca accgatgttg caaatgcagg ttgtgatcat 720
gcaaccatgc tggcacagaa tcgtgttccg gttctgctga cccatcattt tcatctgacc 780
gatccggata caggccagct gatgggtgca atgaccgata ttcaggcaca gcaggcacgt 840
cgtctgctgg cagcaaccgg tcagccggtt acctttaccg cactggatgc accgcatacc 900
atgcatgatc ctgaacctga acgttatttt gaagttctga ccgaatgggc aagtgcactg 960
gattaa 966
<210> 26
<211> 960
<212> DNA
<213> 人工序列
<220>
<223> ORF经密码子优化并因而不同于天然出现的DNA序列
<220>
<221> misc_feature
<223> 编码具有SEQ ID NO: 10的多肽的人工DNA序列
<400> 26
atgggtcgtt atgccggtgt ttttggtccg catgcaccgg aaagcaccta tgttggtcat 60
gcatatccgg aacaactgtt tgataccggt gaagttcgtc tgaattatgc agttgccggt 120
gatgcaagcg caagtccgct gctgctgatt ccgggtcaga ccgaaagttg gtggggttat 180
gaaccggcaa tgggtctgct ggcagaacat tttcatgttc atgcagttga tctgcgtggt 240
cagggtcgta gcacccgtac accgcgtcgt tataccctgg ataatattgg taatgatctg 300
gtgcgttttc tggatggtgt tattggtcgt ccggcatttg ttagcggtct gagcagcggt 360
ggtctgctga gcgcatggct gagcgccttt gcagaaccgg gtcaggttct ggcagcatgt 420
tatgaagatc cgcctttttt tagcagcgaa ctggacccgg tgattggtcc gggtctgatg 480
agcaccgttg gtccgctgtt tgcactgtat gttaaatatc tgggtgatca gtggtcaatt 540
ggtgattggg atggttttgt tgcaggcgca ccgcaagaac tggcaggttg gcaggcacat 600
gttgcactgg caggcggtac agcagaaccg cctcagcatc tgaaagaata tgatccggaa 660
tggggtcgtg catttgttgg tggcaccttt accaccggtt gtccgcatca ggttatgctg 720
agccaggtta aagttccggt tctgtttacc catcattttc gtatgctgga tgatgaaagc 780
ggtagcctga ttggtgcagc aaccgatgat caggcagcac gtgttgttga actggttgaa 840
aatagtggtg caccgctgac ctatcgtagc tttccgatga tgggtcatag tatgcatgca 900
caagatccgg cactgtttgc aggcaccctg gttgattggt ttaccgcagc acgtagctaa 960
<210> 27
<211> 960
<212> DNA
<213> 人工序列
<220>
<223> ORF经密码子优化并因而不同于天然出现的DNA序列
<220>
<221> misc_feature
<223> 编码具有SEQ ID NO: 11的多肽的人工DNA序列
<400> 27
atgggtcgtt atgccggtgt ttttggtccg catgcaccgg aagcaaccta tgttgaacat 60
ggttatccgg aacgtctgtt tgataccggt gaagtgcagc tgaattatgt tgttgccggt 120
gatgcagcag caccgcctct gctgctgatt ccgggtcaga gcgaaagttg gtggggttat 180
gaagcagcaa ttccgctgct ggcacgtcat tttcatgttc atgcagttga tctgcgtggt 240
cagggtcgta gcacccgtac accgggtcgc tataccctgg ataatgttgg taatgatctg 300
gtgcgttttc tggatggtgt tattggtcgt ccggcatttg ttagcggtct gagcagcggt 360
ggtctggcaa gcgcatggct gagcgcattt gcaaaaccgg gtcaggttgt tgcagcatgt 420
tgggaagatc cgcctttttt tagcagcgaa accgcaccga ttgttggtcc gcctattacc 480
gatagcattg gtccgctgtt tggtatgtgg gcacgttatc tgggtgatca gtggtcagtt 540
ggtgattggg atggttttgt tgccgcagtt ccgaccgaac tggcagattg gcaggcacat 600
gttgcactgg ttgttggcac cgcagatcct ccgcagaatc tgcgtgaata tgatccggaa 660
tggggtaaag catttattac cggcaccttt gcagcaagct gtccgcatca tgttatgctg 720
agcaaagtta aagttccggt tctgtatacc catcactttc gcatgattga tgaaggtagt 780
ggtggtctga ttggtgcatg tagcgatatt caggcaggtc gtgttaccca gctggcaaaa 840
tcaggtggtc gtagcgttac ctatcgtagc tttccgatga tggcacatag catgcatggt 900
caagatccgg cactgtttag cgaaaccctg gttgaatggt ttagccgttt taccggttaa 960
<210> 28
<211> 969
<212> DNA
<213> 人工序列
<220>
<223> ORF经密码子优化并因而不同于天然出现的DNA序列
<220>
<221> misc_feature
<223> 编码具有SEQ ID NO: 12的多肽的人工DNA序列
<400> 28
atgccgaaaa gcgaagcagc agatcgtgca gatagctttg ttagccatga tttcaaagaa 60
aacattgtgg atctgggcga aatccgcatg aattatgttg ttcagggcaa caaaaaaagt 120
ccggcactgc tgctgattcc ggcacagggt gaaagttggt ggggttatga agcagcaatt 180
ccgctgctgg caaaacattt tcaggttttt gcaattgatc tgcgtggtca gggtcgtacc 240
acctggacac cgggtcgtta taccctggat atttttggta atgatgtggt gcgctttatc 300
gatctggtta ttggtcgtga aaccctgatt gcaggtaata gcagcggtgg tctgattggt 360
gcatggctgg cagcatttgc aaaaccgggt caggttcgtg cagttatgct ggaagatccg 420
cctctgtttg caagcgaaat tcgtccgcct tatggtccgg gtatttggca gggtctgggt 480
ccgatgtttg cagcatgggc aaaatggctg ggtccgcagt ggtcaattgg tgattgggat 540
ggtatggtta aagcactgcc ggatgaactg ccggaagatc tgctgcctgg tattggtttt 600
atgctgggtg atggtgaaag tgatggtgca gcaccgaccc ctccgcagca tctgaaagaa 660
tatgatccgg aatggggtgc aagctgggca agcggttttg ccaataccgg ttgtgaacat 720
gaagcagtta ttagccaggt gcgtgttccg gttctgctga cccatcattt tcgtcagatt 780
aatgaagaaa ccggtcatct gatgggtgca ctgagcgatc tgcaggcagc acaggttcgt 840
catatcattg aagaagttgc aggtcaagag gttacctatg ttagcctgga tgcaccgcat 900
accatgcatg aaccgcagcc ggaacgttat accgatgttc tgctggattg ggttaaaaaa 960
ctgggttaa 969
<210> 29
<211> 987
<212> DNA
<213> 人工序列
<220>
<223> ORF经密码子优化并因而不同于天然出现的DNA序列
<220>
<221> misc_feature
<223> 编码具有SEQ ID NO: 13的多肽的人工DNA序列
<400> 29
atgaattatg caaccgcagg tagcagcgat aaaccggcac tgctgctggt tccgggtcag 60
agcgaaagtt ggtggggtta tgaaatggca atgtggctgc tgaaagatga ttatcaggtt 120
tttgcagttg atatgcgtgg tcagggtcag agtacctgga caccgggtcg ttatagcctg 180
gatacctttg gtaatgatct ggtgaaattc atcgatatcg tgattaaacg tccggttgtt 240
gttagcggtc tgagcagcgg tggtgttgtg agcgcatggc tgagcgcatt tgcaaaacct 300
ggtcagattc gtgcagcagt ttatgaagat ccgcctctgt ttgcaagcca gagcaaaccg 360
gcaattggtc agagtgttat gcagaccgtt gcaggtccgt tttttaacct gtggtataaa 420
tggctgggtg cacagtggac cattggtgat caggcaggta tggttgcagc aatgccgaaa 480
gaaattccgg catggattct gcagtatctg ggtaatacca ccagtggtcc gaccggtctg 540
gatctgacac tgaatgaata tgatccggaa tggggtcatg gttttgttag tggcaccgtt 600
gatgcaacct gtgatcatga agcaatgctg acccatgtta aagttccggt tctgtttacc 660
catcatagcc gtgcaattga tccgtatacc ggtaatctga ttggtagcgt tagcgatacc 720
caggttagct atgcacaggg tctgattacc accaatggca atcagagctt taccctgaaa 780
aactttccgc tggcaagcca tgatatgcat aattctgatc cggcaaccta tgttagcgca 840
attaccacct ggatggcaag cctgggtatt ggtagtgcag ttattccggg tccggttaaa 900
gttgcaagcg caagcgcaca ggttagcgca gcaagcaccg caccgcctag ctgtaccagc 960
accagcgcac cgagcaccgg tcattaa 987
<210> 30
<211> 843
<212> DNA
<213> 人工序列
<220>
<223> ORF经密码子优化并因而不同于天然出现的DNA序列
<220>
<221> misc_feature
<223> 编码具有SEQ ID NO: 14的多肽的人工DNA序列
<400> 30
atgaccgttg ttgatccgcc tgcaccgcgt gattttccgg aactgctggt tgatctgggt 60
gaagttgttc tgaatcatgc agaagcaggt agtccggatc gtccggcact ggttccggtg 120
ccggaacagg gtggtagttg gtggtcttat gaacgtgtta tgccgctgcc tgcacgcgat 180
tttcatgttt ttgcagttga tctgcgtggt cgtggtcgta gcacccgtac accgcgtcgt 240
tatagcctgg atgattttgg taatgatctg gttcgttttc tggccctggt tgttcgccgt 300
ccggcagttg ttgcaggtaa tagcagcggt ggtgttctgg cagcatggtc aagcgcctat 360
gcaatgcctg gtcaggttcg tgcagttctg ctggaagatc cgcctctgtt tagcagcgaa 420
ctgacaccgg tttgtggtcc gggtgttcgt caggcagcag gtccgctgtt tgaactgctg 480
agcacccatc tgggcgatca gtggggtggt ggtcgtccgg gtcgtgttca tggtggcgtt 540
ccgcgtctgg gtctggcagc cgcagcagca gttcgtgttg cacgtcgtgc agcagcaacc 600
gatgcacgtg gtcgccctgg tgcagcacgt ggacgtcctg ccggtgttgg tggtgcagct 660
cgtcgcggtc gcggtggtcg tgaacgcacc ggtacaacca ccgttctgag cggtctgacc 720
ggtagccgta ccagcggcac cggtcgttgt cgtaaaccgt ttcgtctgcg tcagtggtgg 780
gcaggcggtg cccgtggtcc tcctccgcct cgtcagattc gcgcagatgt tcgtacccgt 840
taa 843
<210> 31
<211> 981
<212> DNA
<213> 人工序列
<220>
<221> misc_feature
<223> 编码具有SEQ ID NO: 15的多肽的人工DNA序列
<400> 31
atgagcgttc cggttacccc gagcgcacgt aatgtttttg ttccgcatgc atttccagag 60
aaacaaattg atctgggtga agtggttctg aattatgcag aagcaggtac accggataaa 120
ccggcattac tgctgctgcc ggaacagacc ggtagttggt ggtcttatga accggcaatg 180
ggtctgctgg cagaacattt tcatgttttt gcagttgatc tgcgtggtca gggtcgtagc 240
acctggacac cgggtcgtta tagcctggat aattttggta atgatctggt gcgttttatt 300
gcactggcaa ttcgtcgtcc ggttgttgtt gcaggttgta gcagcggtgg tgttctggca 360
gcatggctga gcgcctatgc actgcctggt cagattcgtg gtgcactgtg tgaagatgca 420
ccgctgtttg caagcgaact gacaccggca catggtcatg gtgttcgtca gggtgcaggt 480
ccggtttttg aactgtatcg tgattatctg ggcgatcagt ggtcagttgg tgattgggca 540
ggtctggttg cagcagcaca ggcaagtccg gcaaaaatga tgagcctgtt taaaatgcct 600
ggtgaaccgc ctcagaatct gcgtgaatat gatccggaat gggcacgtgt tttttttgaa 660
ggcaccgttg gtctgcattg tccgcatgat cgtatgctga gccaggttaa aacaccggtt 720
ctgattaccc atcatgcacg taccaccgat ccggaaaccg gtgaatttct gggtgcactg 780
agcgaactgc aggcagaacg tgcacaggcc attattcgtg cagccggtgt tccggttgat 840
tatcagagct ttccggatgc agcacatgca atgcatacca cagaaccggc acgttatgca 900
gcagttctga ccgcatgggc agcaaaactg cctccggttg cagataccag cccgtcagca 960
gcagcaagcg cacatgttta a 981
<210> 32
<211> 7
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(7)
<400> 32
Ala Gly Asn Ser Ser Gly Gly
1 5
<210> 33
<211> 7
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(7)
<400> 33
Arg Thr Ile Asp Pro Glu Thr
1 5
<210> 34
<211> 7
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(7)
<400> 34
Asp Ala Leu His Met Met His
1 5
<210> 35
<211> 7
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(7)
<400> 35
Ala Gly Asp Ser Ser Gly Gly
1 5
<210> 36
<211> 7
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(7)
<400> 36
Ala Gly Asp Ser Ser Leu Gly
1 5
<210> 37
<211> 7
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(7)
<400> 37
Ala Gly Gln Ser Ser Gly Gly
1 5
<210> 38
<211> 7
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(7)
<400> 38
Ala Gly His Ser Ser Gly Gly
1 5
<210> 39
<211> 7
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(7)
<400> 39
Ala Gly Ser Ser Ser Gly Gly
1 5
<210> 40
<211> 7
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(7)
<400> 40
Ser Gly Asn Ser Ser Gly Gly
1 5
<210> 41
<211> 7
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(7)
<400> 41
Ser Gly Asp Ser Ser Gly Gly
1 5
<210> 42
<211> 7
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(7)
<400> 42
Ser Gly Gln Ser Ser Gly Gly
1 5
<210> 43
<211> 7
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(7)
<400> 43
Ser Gly His Ser Ser Gly Gly
1 5
<210> 44
<211> 7
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(7)
<400> 44
Ser Gly Ser Ser Ser Gly Gly
1 5
<210> 45
<211> 7
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(7)
<400> 45
Arg Thr Ile Asp Pro Glu Thr
1 5
<210> 46
<211> 7
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(7)
<400> 46
Arg Asp Ile Asp Pro Asp Thr
1 5
<210> 47
<211> 7
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(7)
<400> 47
Arg Gly Thr Asp Pro Glu Thr
1 5
<210> 48
<211> 7
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(7)
<400> 48
Arg Gly Ile Asp Pro Glu Thr
1 5
<210> 49
<211> 7
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(7)
<400> 49
Asp Ala Leu His Met Met His
1 5
<210> 50
<211> 7
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(7)
<400> 50
Asp Ala Ser His Met Met His
1 5
<210> 51
<211> 11
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(11)
<400> 51
Val Val Ala Gly Asn Ser Ser Gly Gly Leu Leu
1 5 10
<210> 52
<211> 11
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(11)
<400> 52
Ile Val Ala Gly Asn Ser Ser Gly Gly Val Leu
1 5 10
<210> 53
<211> 11
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(11)
<400> 53
His Ala Arg Thr Ile Asp Pro Glu Thr Gly Glu
1 5 10
<210> 54
<211> 11
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(11)
<400> 54
His Met Arg Asp Ile Asp Pro Asp Thr Gly His
1 5 10
<210> 55
<211> 11
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(11)
<400> 55
His Met Arg Gly Thr Asp Pro Glu Thr Gly Asn
1 5 10
<210> 56
<211> 11
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(11)
<400> 56
His Pro Asp Ala Leu His Met Met His Leu Phe
1 5 10
<210> 57
<211> 11
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(11)
<400> 57
Val Pro Asp Ala Leu His Met Met His Gln Phe
1 5 10
<210> 58
<211> 11
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(11)
<400> 58
Val Pro Asp Ala Ser His Met Met His Gln Ser
1 5 10
<210> 59
<211> 21
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(21)
<400> 59
Ile Lys Arg Pro Val Val Val Ala Gly Asn Ser Ser Gly Gly Leu Leu
1 5 10 15
Ala Ala Trp Leu Ser
20
<210> 60
<211> 21
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(21)
<400> 60
Val Lys Arg Pro Val Ile Val Ala Gly Asn Ser Ser Gly Gly Val Leu
1 5 10 15
Ala Ala Trp Leu Ser
20
<210> 61
<211> 21
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(21)
<400> 61
Val Arg Arg Pro Val Val Val Ala Gly Asn Ser Ser Gly Gly Val Leu
1 5 10 15
Ala Ala Trp Leu Ser
20
<210> 62
<211> 21
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(21)
<400> 62
Val Lys Arg Pro Val Val Val Ala Gly Asn Ser Ser Gly Gly Val Leu
1 5 10 15
Ala Ala Trp Leu Ser
20
<210> 63
<211> 21
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(21)
<400> 63
Ile Leu Ile Thr His His Ala Arg Thr Ile Asp Pro Glu Thr Gly Glu
1 5 10 15
Leu Leu Gly Ala Leu
20
<210> 64
<211> 21
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(21)
<400> 64
Val Leu Leu Thr His His Met Arg Asp Ile Asp Pro Asp Thr Gly His
1 5 10 15
Leu Val Gly Ala Leu
20
<210> 65
<211> 21
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(21)
<400> 65
Val Leu Leu Thr His His Met Arg Gly Thr Asp Pro Glu Thr Gly Asn
1 5 10 15
Leu Leu Gly Ala Leu
20
<210> 66
<211> 21
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(21)
<400> 66
Val Leu Leu Thr His His Pro Asp Ala Leu His Met Met His Leu Phe
1 5 10 15
Leu Leu Gly Ala Leu
20
<210> 67
<211> 21
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(21)
<400> 67
Val Asp Tyr Gln Ser His Pro Asp Ala Leu His Met Met His Leu Phe
1 5 10 15
Asp Pro Ala Arg Tyr
20
<210> 68
<211> 21
<212> PRT
<213> 人工序列
<220>
<223> 氨基酸基序
<220>
<221> PEPTIDE
<222> (1)..(21)
<400> 68
Val Asp Tyr Ala Ser Val Pro Asp Ala Leu His Met Met His Gln Phe
1 5 10 15
Asp Pro Pro Arg Tyr
20
<210> 69
<211> 21
<212> PRT
<213> 人工序列
<220>
<221> PEPTIDE
<222> (1)..(21)
<400> 69
Val Asp Tyr Glu Ser Val Pro Asp Ala Ser His Met Met His Gln Ser
1 5 10 15
Ala Pro Ala Arg Tyr
20
Claims (25)
1.以水解方式裂解玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物的多肽,所述玉米赤霉烯酮衍生物选自α-ZEL、β-ZEL、α-ZAL、β-ZAL、Z14G、Z14S和ZAN,其特征在于,所述多肽是具有SEQ ID No.9的氨基酸序列或其功能性变体的水解酶,其中在所述功能性变体与所述氨基酸序列之间的序列同一性为至少83%,
其中所述多肽包含至少一个保守氨基酸序列区段或其功能性变体,其中所述氨基酸序列区段的功能性变体具有至少84%的序列同一性,并且所述至少一个保守氨基酸序列区段选自具有SEQ ID No.1的序列的氨基酸序列+89至+145或+223至+228,并且
其中所述多肽具有至少0.01U/mg的比活性。
2.根据权利要求1的多肽,其特征在于,所述多肽包含至少一个保守氨基酸序列区段或其功能性变体,其中所述氨基酸序列区段的功能性变体具有至少92%的序列同一性,并且所述至少一个保守氨基酸序列区段为具有SEQ ID No.1的序列的氨基酸序列+223至+228。
3.根据权利要求2的多肽,其特征在于,所述氨基酸序列区段的功能性变体具有至少98%的序列同一性。
4.根据权利要求1至3之一的多肽,其特征在于,所述多肽在至少一个选自下列的位置处具有关于SEQ ID No.1的氨基酸序列的至少一个突变:22、23、25、26、27、29、31、32、35、37、42、43、46、51、53、54、57、60、69、72、73、78、80、84、88、95、97、99、114、118、119、123、132、141、146、148、149、154、163、164、165、169、170、172、176、180、182、183、190、191、194、196、197、198、201、204、205、206、207、208、209、210、212、213、214、216、217、220、221、222、229、231、233、238、240、244、245、246、248、249、251、254、256、260、262、263、266、269、271、277、280、281、282、283、284、285、286、287、292、296、298、302、307、308、309、311、314、317、319、321、323、325和326。
5.根据权利要求1至3之一的多肽,其特征在于,所述多肽具有选自下列的关于SEQ IDNo.1的氨基酸序列的至少一个突变:D22A、S23Q、S23L、N25D、I26V、F27Y、F27H、S29P、R31A、F32Y、R35K、R35Q、V37A、V42I、V43T、F46Y、S51E、S51D、D53G、N54M、N54R、L57V、L60I、S69G、P72E、V73A、A78S、N80H、F84Y、I88L、T95S、T97A、R99K、I114M、I118V、K119R、V123I、L132V、A141S、I146V、I146L、A148G、A149V、A154P、P163T、A164T、Y165C、Y165H、V169I、L170R、A172G、A176M、A176V、Y180F、D182T、F183Y、I190V、G191S、K194T、K194E、F196Y、V197C、V197R、E198R、E198S、K201D、K201G、P204S、P204A、A205S、K206P、A207M、M208A、Q209R、L210A、L210S、ΔP212、T213V、P214A、E216T、E216G、A217I、N220H、L221M、K222R、K222Q、G229A、A231V、F233W、F233Y、F233H、A238G、H240N、H240S、D244E、R245Q、M246L、S248T、S248N、S248G、Q249R、K251N、I254V、I256L、A260M、T262D、T262G、I263T、E266D、E269H、E269N、L271V、L277E、E280A、E280L、H281R、H281Q、A282V、Q283R、D284L、D284R、I285L、I286M、R287E、R287D、R292K、R292T、Q296A、Q296E、H298V、L302S、L307Q、F308S、D309A、A311P、A314V、L317F、S319Q、S319P、S319R、S321A、S321T、T323A、P325A、A326P。
6.分离的多核苷酸,其具有编码多肽的核苷酸序列,其中所述多肽具有水解玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物的特性,所述玉米赤霉烯酮衍生物选自α-ZEL、β-ZEL、α-ZAL、β-ZAL、Z14G、Z14S和ZAN,其特征在于,所述核苷酸序列编码至少一种根据权利要求1至5之一的多肽。
7.以水解方式裂解玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物从而产生用于猪、家禽或水产养殖的饲料的添加剂,所述添加剂用于添加至食品或干酒糟,所述玉米赤霉烯酮衍生物选自α-ZEL、β-ZEL、α-ZAL、β-ZAL、Z14G、Z14S和ZAN,其特征在于,所述添加剂包含具有SEQ ID No.9的氨基酸序列或其功能性变体的多肽,其中在所述功能性变体与所述氨基酸序列之间的序列同一性为至少83%。
8.根据权利要求7的添加剂,其特征在于,所述添加剂进一步包含助剂。
9.根据权利要求7或8的添加剂,其特征在于,包含至少一种根据权利要求1至5之一的多肽。
10.根据权利要求8的添加剂,其特征在于,所述助剂为至少一种惰性载体。
11.根据权利要求8的添加剂,其特征在于,所述助剂为至少一种惰性载体,以及其他成分。
12.根据权利要求11的添加剂,其特征在于,所述其他成分为维生素和/或矿物质和/或酶和/或其他用于使真菌毒素解毒的组分。
13.根据权利要求9的添加剂,其特征在于,在所述添加剂中,以最高10,000U/g的浓度包含至少一种根据权利要求1至5之一的多肽。
14.根据权利要求9的添加剂,其特征在于,在所述添加剂中,以最高1,000U/g的浓度包含至少一种根据权利要求1至5之一的多肽。
15.根据权利要求9的添加剂,其特征在于,在所述添加剂中,以最高100U/g的浓度包含至少一种根据权利要求1至5之一的多肽。
16.根据权利要求9的添加剂,其特征在于,在所述添加剂中,以最高10U/g的浓度包含至少一种根据权利要求1至5之一的多肽。
17.根据权利要求7或8的添加剂,其特征在于,所述添加剂以经包囊或经包衣的形式存在。
18.具有SEQ ID No.9的氨基酸序列或其功能性变体的多肽用于以水解方式裂解在饲料中、在食物中或在干酒糟中的玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物的用途,所述玉米赤霉烯酮衍生物选自α-ZEL、β-ZEL、α-ZAL、β-ZAL、Z14G、Z14S和ZAN,其中所述功能性变体与所述氨基酸序列之间的序列同一性为至少83%。
19.根据权利要求18的用途,其中所述饲料为用于猪、家禽和水产养殖的饲料。
20.用于以水解方式裂解玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物的方法,所述玉米赤霉烯酮衍生物选自α-ZEL、β-ZEL、α-ZAL、β-ZAL、Z14G、Z14S和ZAN,其特征在于,所述玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物被具有SEQ ID No.9的氨基酸序列或其功能性变体的多肽水解,其中在所述功能性变体与所述氨基酸序列之间的序列同一性为至少83%。
21.根据权利要求20的方法,其特征在于,在根据权利要求7至17之一的添加剂中使用所述多肽。
22.根据权利要求21的方法,其特征在于,将所述多肽或添加剂与被玉米赤霉烯酮和/或被至少一种玉米赤霉烯酮衍生物污染的饲料或食物相掺混,使所述被污染的饲料或食物与湿气相接触,并且所述多肽或添加剂水解在所述被污染的饲料或食物中所包含的玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物。
23.根据权利要求20至22之一的方法,其特征在于,至少70%的所述玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物被水解。
24.根据权利要求20至22之一的方法,其特征在于,至少80%的所述玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物被水解。
25.根据权利要求20至22之一的方法,其特征在于,至少90%的所述玉米赤霉烯酮和/或至少一种玉米赤霉烯酮衍生物被水解。
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