CN109206505A - 变异型细胞色素蛋白及其应用 - Google Patents
变异型细胞色素蛋白及其应用 Download PDFInfo
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- CN109206505A CN109206505A CN201810721942.7A CN201810721942A CN109206505A CN 109206505 A CN109206505 A CN 109206505A CN 201810721942 A CN201810721942 A CN 201810721942A CN 109206505 A CN109206505 A CN 109206505A
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Abstract
本发明涉及一种变异型细胞色素蛋白及其应用，该变异型细胞色素蛋白为在具有3个血红素结合域的细胞色素蛋白中从N末端侧数第1个第1血红素结合域和第2个第2血红素结合域发生了缺失、优选包含第1血红素结合域和第2血红素结合域的区域发生了缺失的变异型细胞色素蛋白。
Description
技术领域
本发明涉及即使在低电位下也能够进行电子传递的变异型细胞色素蛋白。本发明的变异型细胞色素蛋白能够适合用于生物传感器等，在生物化学领域、医疗领域等中有用。
背景技术
作为受电子性蛋白质，已知有细胞色素蛋白，其与氧化还原酶的催化亚基形成复合体而被用作电化学传感器。
例如，在专利文献1中公开了将来源于伯克霍尔德菌(Burkholderia)属微生物的葡萄糖脱氢酶的催化亚基(α亚基)和细胞色素C亚基(β亚基)的复合体作为生物传感器使用的技术(专利文献1)。这样的生物传感器被用于测定生物试样中的葡萄糖浓度等，但为了更高精度地进行测定，要求通过改变蛋白质等来提高测定灵敏度。
截止到目前，作为电化学传感器蛋白的突变体，主要对催化亚基的突变体进行了研究，对于细胞色素蛋白的突变体几乎未进行研究。
现有技术文献
专利文献
专利文献1：WO 2005/023111
发明内容
在使用了观测氧化还原电流的生物传感器进行的目标物质的测定中，为了降低试样中包含的还原物质的影响，期望在可观测到酶和酶复合体与电极间的电子传递的范围内降低氧化还原电位。
但是，在使用现有的天然细胞色素蛋白作为受电子性蛋白质对氧化还原电流进行观测的情况下，需要对电极施加较大的电位。例如，在专利文献1的实施例7中，对银/氯化银电极施加了+350mV的电位，在该电位下，可能会由于抗坏血酸或乙酰氨基酚之类的还原性物质被氧化而使测定值产生正偏差。
鉴于这样的问题，本发明的课题在于提供一种能够用于生物传感器的变异型受电子性亚基蛋白，该生物传感器能够通过施加低电位进行目标物质的测定。
本发明人为了解决上述课题，制作了使包含从氨基末端数第1个第1血红素结合域和第2个第2血红素结合域的区域缺失的变异型细胞色素C蛋白，并使用其进行了生物传感器的评价。其结果令人吃惊地发现，使包含第1血红素结合域和第2血红素结合域的区域缺失的变异型细胞色素C蛋白尽管仅有1个血红素结合域(第3血红素结合域)，仍能够从催化亚基接受电子，能够在比以往更低的电位下流通与目标物质相应的电流，从而完成了本发明。
即，本发明如下所述。
[1]一种变异型细胞色素蛋白，其为在具有3个血红素结合域的细胞色素蛋白中从N末端侧数第1个第1血红素结合域和第2个第2血红素结合域发生了缺失的变异型细胞色素蛋白。
[2]如[1]所述的变异型细胞色素蛋白，其中，包含上述第1血红素结合域和第2血红素结合域的区域发生了缺失。
[3]如[2]所述的变异型细胞色素蛋白，其中，包含上述第1血红素结合域和第2血红素结合域的区域相当于序列编号4的第43～195位的区域。
[4]如[1]～[3]中任一项所述的变异型细胞色素蛋白，其中，细胞色素蛋白为细胞色素C蛋白。
[5]如[4]所述的变异型细胞色素蛋白，其中，细胞色素C蛋白来源于伯克霍尔德菌属微生物。
[6]如[5]所述的变异型细胞色素蛋白，其中，细胞色素C蛋白来源于洋葱伯克霍尔德菌。
[7]如[1]～[6]中任一项所述的变异型细胞色素蛋白，其中，改变前的细胞色素蛋白与序列编号4具有60％以上的氨基酸序列同一性。
[8]如[1]～[7]中任一项所述的变异型细胞色素蛋白，其由序列编号4的氨基酸编号314～425的氨基酸序列、序列编号4的氨基酸编号330～425的氨基酸序列、或这些氨基酸序列中1个或几个氨基酸发生了置换、缺失、插入或添加的氨基酸序列(其中，氨基酸编号334～338的CXXCH基序未被改变)构成。
[9]一种DNA，其编码[1]～[8]中任一项所述的变异型细胞色素蛋白。
[10]一种重组载体，其包含[9]所述的DNA。
[11]一种转化细胞，其利用[10]所述的重组载体进行了转化。
[12]一种氧化还原酶-细胞色素复合体，其包含[1]～[8]中任一项所述的变异型细胞色素蛋白和氧化还原酶的催化亚基蛋白。
[13]如[12]所述的氧化还原酶-细胞色素复合体，其中，氧化还原酶为葡萄糖脱氢酶。
[14]一种生物传感器，其包含[12]或[13]所述的氧化还原酶-细胞色素复合体。
[15]一种测定对象物质的测定方法，其特征在于，在[14]所述的生物传感器中添加试样，施加电位并测定响应电流，基于响应电流计算出试样中包含的测定对象物质的浓度。
[16]如[15]所述的测定方法，其中，施加电位对于银/氯化银电极为0～+300mV。
根据本发明，能够构建出一种电化学生物传感器，其能够在低于以往的电位(例如对于银/氯化银电极为0～+300mV、优选为0～+150mV、更优选为0～+100mV)下工作，能够抑制由试样中的还原物质所致的干扰，对葡萄糖等测定对象物质进行准确的测定。
附图说明
图1是示出使用了包含含野生型β亚基的葡萄糖脱氢酶(GDH)复合体的葡萄糖传感器、或包含含N末端侧缺失型β亚基的GDH复合体的葡萄糖传感器的计时安培分析法的结果的图(施加电位为0mV、+100mV、+400mV)。
具体实施方式
以下详细说明本发明。
本发明的变异型细胞色素蛋白为具有3个血红素结合域的细胞色素蛋白的突变体，其是从N末端侧数第1个第1血红素结合域和第2个第2血红素结合域发生了缺失的变异型细胞色素蛋白。
细胞色素蛋白具有3个血红素结合域，在本说明书中，将从N末端侧数第1个、第2个、第3个血红素结合域分别称为第1血红素结合域、第2血红素结合域、第3血红素结合域。血红素结合域通常由CXXCH(序列编号5)(X为任意的氨基酸)表示，在本发明的变异型细胞色素蛋白中，按照第1血红素结合域和第2血红素结合域发生缺失的方式进行了改变。
此处，“按照第1血红素结合域和第2血红素结合域发生缺失的方式进行了改变”是指第1CXXCH基序和第2CXXCH基序发生了缺失。
CXXCH基序的缺失也包括包含CXXCH基序的区域的缺失，可以是包含第1血红素结合域的区域和包含第2血红素结合域的区域分别发生了缺失，也可以是包含第1血红素结合域和第2血红素结合域的区域发生了缺失。
作为细胞色素蛋白，优选细胞色素C蛋白，可以举出由序列编号4表示的洋葱伯克霍尔德菌(Burkholderia cepacia)的细胞色素C蛋白(葡萄糖脱氢酶的β亚基)。
以下以洋葱伯克霍尔德菌的细胞色素C蛋白作为代表例进行说明。
在序列编号4中，存在第1血红素结合域(氨基酸编号43～47)、第2血红素结合域(氨基酸编号191～195)、第3血红素结合域(氨基酸编号334～338)。这些第1血红素结合域和第2血红素结合域发生缺失，但也可以是包含第1血红素结合域和第2血红素结合域的区域(氨基酸编号43～195)发生了缺失。
只要可维持第3血红素结合域、发挥出作为变异型细胞色素C蛋白的功能(即只要无损于电子传递功能)，则包含第1血红素结合域和第2血红素结合域的区域(氨基酸编号43～195)只要至少该区域发生缺失即可，也可以包括序列编号4中的该区域之前和之后的氨基酸在内而发生缺失。例如可以是序列编号4的从N末端侧到氨基酸编号195发生缺失，例如可以举出由序列编号4的氨基酸编号314～425或330～425构成的变异型细胞色素C蛋白作为包含第1血红素结合域和第2血红素结合域的区域发生了缺失的变异型细胞色素C蛋白。其中，在这些序列中，在N末端侧和/或C末端侧可以添加任意的标签序列或信号序列(例如，序列编号4的氨基酸编号1～27)等。
需要说明的是，本发明的变异型细胞色素蛋白并不限于由序列编号4的氨基酸编号314～425构成的氨基酸序列或由序列编号4的氨基酸编号330～425构成的氨基酸序列本身，只要无损于电子传递功能，也可以为这些序列中第3血红素结合域(氨基酸编号334～338)以外的1个或几个氨基酸发生了置换、缺失、添加或插入的序列。此处，“1个或几个”例如为1～20个、优选为1～10个、更优选为1～8个、进一步优选为1～5个、特别优选为1～3个。另外，置换优选为保守性置换，“保守性置换”是指酸性氨基酸彼此的置换、中性氨基酸彼此的置换、碱性氨基酸彼此的置换等性质类似的氨基酸彼此的置换。另外，可以在由序列编号4的氨基酸编号314～425构成的氨基酸序列的N末端侧添加由序列编号4的氨基酸编号196～313构成的氨基酸序列的C末端侧序列的一部分，可以在由序列编号4的氨基酸编号330～425构成的氨基酸序列的N末端侧添加由序列编号4的氨基酸编号196～329构成的氨基酸序列的C末端侧序列的一部分。
需要说明的是，本发明的变异型细胞色素C蛋白只要可保持第3血红素结合域(氨基酸编号334～338)、电子传递功能不会受损，则也可以为具有下述氨基酸序列的蛋白质，该氨基酸序列与由序列编号4的氨基酸编号314～425构成的氨基酸序列或由序列编号4的330～425构成的氨基酸序列具有80％以上、优选90％以上、更优选95％以上的序列同一性。
需要说明的是，序列编号4是洋葱伯克霍尔德菌KS1株所保有的GDHβ亚基的氨基酸序列，KS1株于2000年9月25日以保藏编号FERM BP-7306保藏于通商产业省工业技术院生命工学工业技术研究所(现在的独立行政法人日本国家高级工业科学技术学院，国际专利生物保藏中心(IPOD，NITE)，〒305-8566，日本茨城县筑波市东1丁目1番3号)。
在本发明中，导入上述改变之前的细胞色素C蛋白并不限于由序列编号4表示的洋葱伯克霍尔德菌的细胞色素C蛋白，只要具有第1血红素结合域、第2血红素结合域、第3血红素结合域即可，例如可以使用序列编号4的同源体，优选可以使用与序列编号4具有60％以上、优选80％以上、更优选90％以上的同一性的序列编号4的同源体。
作为与序列编号4具有60％以上的序列同一性的细胞色素C蛋白的氨基酸序列，优选伯克霍尔德菌属的细胞色素C蛋白，可以例示洋葱伯克霍尔德菌(Burkholderiacepacia)J2315株的细胞色素蛋白(序列编号6)、新洋葱伯克霍尔德菌(Burkholderiacenocepacia)的细胞色素C蛋白(序列编号7)、多噬伯克霍尔德菌(Burkholderiamultivorans)的细胞色素C蛋白(序列编号8)、乌汶伯克霍尔德菌(Burkholderiaubonensis)的细胞色素C蛋白(序列编号9)、礁湖伯克霍尔德菌(Burkholderia stagnalis)的细胞色素C蛋白(序列编号10)、泰国伯克霍尔德菌(Burkholderia thailandensis)的细胞色素C蛋白(序列编号11)。
另外，也可以为伯克霍尔德菌属以外的细胞色素蛋白，例如可以例示青枯雷尔氏菌(Ralstonia solanacearum)的细胞色素蛋白(序列编号12)、皮氏雷尔氏菌(Ralstoniapickettii)的细胞色素蛋白(序列编号13)。但是并不限于这些，也可以为其他生物来源的细胞色素C蛋白。以下用美国国家生物技术信息中心(NCBI)的蛋白质数据库的登录号示出与序列编号4具有60％以上的序列同一性的细胞色素C蛋白的氨基酸序列的示例。这些序列中，血红素结合域存在3个，通过使从氨基末端侧数第1个第1血红素结合域和第2个第2血红素结合域缺失而得到本发明的变异型细胞色素蛋白。
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WP_059924260.1细胞色素C[乌汶伯克霍尔德菌]
WP_060048488.1细胞色素C[乌汶伯克霍尔德菌]
WP_059892434.1细胞色素C[乌汶伯克霍尔德菌]
WP_059488090.1细胞色素C[乌汶伯克霍尔德菌]
WP_059483872.1细胞色素C[乌汶伯克霍尔德菌]
WP_059865488.1细胞色素C[乌汶伯克霍尔德菌]
WP_059633519.1细胞色素C[乌汶伯克霍尔德菌]
WP_060287380.1细胞色素C[乌汶伯克霍尔德菌]
WP_059853060.1细胞色素C[乌汶伯克霍尔德菌]
WP_059968663.1细胞色素C[乌汶伯克霍尔德菌]
WP_059752176.1细胞色素C[乌汶伯克霍尔德菌]
WP_059532869.1细胞色素C[乌汶伯克霍尔德菌]
WP_059660535.1细胞色素C[乌汶伯克霍尔德菌]
WP_059590294.1细胞色素C[乌汶伯克霍尔德菌]
WP_026043943.1细胞色素C[吡咯菌素伯克霍尔德菌]
WP_060264220.1细胞色素C[乌汶伯克霍尔德菌]
WP_060019510.1细胞色素C[乌汶伯克霍尔德菌]
WP_059946330.1细胞色素C[乌汶伯克霍尔德菌]
WP_042588017.1多菌种：细胞色素C[伯克霍尔德菌]
WP_084908871.1细胞色素C[嗜中酸假单胞菌[Pseudomonas]mesoacidophila]
WP_071751088.1细胞色素C[乌汶伯克霍尔德菌]
WP_060094261.1细胞色素C[乌汶伯克霍尔德菌]
WP_059885808.1细胞色素C[乌汶伯克霍尔德菌]
WP_059878406.1细胞色素C[乌汶伯克霍尔德菌]
WP_071852856.1细胞色素C[乌汶伯克霍尔德菌]
WP_060161154.1细胞色素C[乌汶伯克霍尔德菌]
WP_059796027.1细胞色素C[乌汶伯克霍尔德菌]
WP_059835010.1细胞色素C[乌汶伯克霍尔德菌]
WP_059911385.1细胞色素C[乌汶伯克霍尔德菌]
WP_059744627.1细胞色素C[乌汶伯克霍尔德菌]
WP_045565069.1细胞色素C[乌汶伯克霍尔德菌]
WP_060276465.1细胞色素C[乌汶伯克霍尔德菌]
WP_060052180.1细胞色素C[乌汶伯克霍尔德菌]
WP_059877204.1细胞色素C[乌汶伯克霍尔德菌]
WP_059918860.1细胞色素C[乌汶伯克霍尔德菌]
WP_059674217.1细胞色素C[乌汶伯克霍尔德菌]
WP_059626644.1细胞色素C[乌汶伯克霍尔德菌]
WP_059706954.1细胞色素C[乌汶伯克霍尔德菌]
WP_059610359.1细胞色素C[乌汶伯克霍尔德菌]
WP_071751781.1细胞色素C[乌汶伯克霍尔德菌]
WP_060345076.1细胞色素C[乌汶伯克霍尔德菌]
WP_060056668.1细胞色素C[乌汶伯克霍尔德菌]
WP_059867004.1细胞色素C[乌汶伯克霍尔德菌]
WP_059846198.1细胞色素C[乌汶伯克霍尔德菌]
WP_059956839.1细胞色素C[乌汶伯克霍尔德菌]
WP_059801215.1细胞色素C[乌汶伯克霍尔德菌]
WP_060374144.1细胞色素C[乌汶伯克霍尔德菌]
WP_060228155.1细胞色素C[乌汶伯克霍尔德菌]
WP_060449283.1细胞色素C[乌汶伯克霍尔德菌]
WP_060088315.1细胞色素C[乌汶伯克霍尔德菌]
WP_060141594.1细胞色素C[乌汶伯克霍尔德菌]
WP_059965520.1细胞色素C[乌汶伯克霍尔德菌]
WP_059733305.1细胞色素C[乌汶伯克霍尔德菌]
WP_059849627.1细胞色素C[乌汶伯克霍尔德菌]
WP_060165437.1细胞色素C[乌汶伯克霍尔德菌]
WP_059712923.1细胞色素C[乌汶伯克霍尔德菌]
WP_059737856.1细胞色素C[乌汶伯克霍尔德菌]
WP_059659999.1细胞色素C[乌汶伯克霍尔德菌]
WP_059924960.1细胞色素C[乌汶伯克霍尔德菌]
WP_059777863.1细胞色素C[乌汶伯克霍尔德菌]
WP_060058770.1细胞色素C[乌汶伯克霍尔德菌]
WP_059776014.1细胞色素C[乌汶伯克霍尔德菌]
WP_060123897.1细胞色素C[乌汶伯克霍尔德菌]
WP_060003887.1细胞色素C[乌汶伯克霍尔德菌]
WP_071753220.1细胞色素C[乌汶伯克霍尔德菌]
WP_069271183.1细胞色素C[乌汶伯克霍尔德菌]
WP_060168536.1细胞色素C[乌汶伯克霍尔德菌]
WP_059937950.1细胞色素C[乌汶伯克霍尔德菌]
WP_059651010.1细胞色素C[乌汶伯克霍尔德菌]
WP_059952547.1细胞色素C[乌汶伯克霍尔德菌]
WP_059727989.1细胞色素C[乌汶伯克霍尔德菌]
WP_060328321.1细胞色素C[乌汶伯克霍尔德菌]
WP_059872364.1细胞色素C[乌汶伯克霍尔德菌]
WP_059997503.1细胞色素C[乌汶伯克霍尔德菌]
WP_071773630.1细胞色素C[乌汶伯克霍尔德菌]
WP_059828379.1细胞色素C[乌汶伯克霍尔德菌]
WP_060088886.1细胞色素C[乌汶伯克霍尔德菌]
WP_060248705.1细胞色素C[乌汶伯克霍尔德菌]
WP_036661765.1细胞色素C[潘多拉菌(pandoraea)sp.SD6-2]
WP_059573013.1细胞色素C[伯克霍尔德菌sp.TSV86]
WP_006027348.1多菌种：细胞色素C[伯克霍尔德菌]
WP_069235777.1细胞色素C[伯克霍尔德菌sp.Bp7605]
WP_059514380.1细胞色素C[伯克霍尔德菌sp.TSV85]
WP_063533919.1细胞色素C[伯克霍尔德菌sp.MSMB1589WGS]
WP_059929957.1多菌种：细胞色素C[类鼻疽菌群(pseudomallei group)]
WP_060819951.1细胞色素C[伯克霍尔德菌sp.BDU19]
WP_059646659.1多菌种：细胞色素C[类鼻疽菌群]
WP_038746875.1细胞色素C[伯克霍尔德菌sp.ABCPW 111]
WP_059669953.1细胞色素C[伯克霍尔德菌sp.MSMB1498]
WP_038801472.1细胞色素C[俄克拉荷马伯克霍尔德菌(Burkholderiaoklahomensis)]
WP_060356553.1细胞色素C[伯克霍尔德菌sp.MSMB617WGS]
WP_059582413.1多菌种：细胞色素C[类鼻疽菌群]
WP_085508044.1细胞色素C[类鼻疽伯克霍尔德菌(Burkholderiapseudomallei)]
WP_010118598.1细胞色素C[俄克拉荷马伯克霍尔德菌]
WP_059597682.1细胞色素C[伯克霍尔德菌sp.BDU6]
WP_025405690.1细胞色素C[泰国伯克霍尔德菌]
WP_038742304.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_009900297.1细胞色素C[泰国伯克霍尔德菌](序列编号11)
WP_004532924.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_058034623.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_038758421.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_038760788.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_004540659.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_085547264.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_076883397.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_059473995.1细胞色素C[伯克霍尔德菌sp.BDU5]
WP_041191039.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_043296725.1细胞色素C[泰国伯克霍尔德菌]
WP_063597172.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_066496408.1细胞色素C[伯克霍尔德菌sp.BDU8]
WP_038799895.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_004539114.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_004524074.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_017881863.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_009897184.1细胞色素C[泰国伯克霍尔德菌]
WP_004536717.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_076885712.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_066570748.1细胞色素C[伯克霍尔德菌sp.ABCPW 14]
WP_076903220.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_076936464.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_076893617.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_043299328.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_038785043.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_041220968.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_076891198.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_076883909.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_004528232.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_085539130.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_038784065.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_038760183.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_038730591.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_038765827.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_076855330.1细胞色素C[类鼻疽伯克霍尔德菌]
WP_004199672.1细胞色素C[鼻疽伯克霍尔德菌(Burkholderia mallei)]
KGC89207.1细胞色素C家族蛋白[类鼻疽伯克霍尔德菌]
EDO93432.1葡萄糖脱氢酶，β亚基[巴斯德氏类鼻疽伯克霍尔德菌(Burkholderiapseudomallei Pasteur)52237]
WP_011205494.1细胞色素C亚基II[类鼻疽伯克霍尔德菌]
KGV82938.1细胞色素C家族蛋白[类鼻疽伯克霍尔德菌MSHR4375]
CDU31054.1推定细胞色素C亚基II[类鼻疽伯克霍尔德菌]
KGV67566.1细胞色素C家族蛋白[类鼻疽伯克霍尔德菌MSHR4299]
EDK84065.1细胞色素C家族蛋白[鼻疽伯克霍尔德菌2002721280]
WP_074287454.1细胞色素C[伯克霍尔德菌sp.GAS332]
WP_035557403.1细胞色素C[伯克霍尔德菌sp.9120]
WP_084534186.1细胞色素C[迪尔沃思副伯克霍尔德菌(Paraburkholderiadilworthii)]
WP_017774215.1细胞色素C[久留里副伯克霍尔德菌(Paraburkholderiakururiensis)]
WP_030100524.1细胞色素C[伯克霍尔德菌sp.K24]
WP_028194542.1细胞色素C[真菌副伯克霍尔德菌(Paraburkholderiafungorum)]
WP_046573324.1细胞色素C[真菌副伯克霍尔德菌]
WP_042300635.1细胞色素C[久留里副伯克霍尔德菌]
WP_084166897.1细胞色素C[苏格兰副伯克霍尔德菌(Paraburkholderiacaledonica)]
SDI05126.1细胞色素C，单血红素-二血红素变体[吩嗪副伯克霍尔德菌(Paraburkholderia phenazinium)]
WP_051120977.1细胞色素C[苔藓副伯克霍尔德菌(Paraburkholderiabryophila)]
WP_073428494.1多菌种：细胞色素C[伯克霍尔德菌科(Burkholderiaceae)]
WP_074300113.1细胞色素C[吩嗪副伯克霍尔德菌]
WP_074768708.1细胞色素C[真菌副伯克霍尔德菌]
WP_075465130.1细胞色素C[青枯雷尔氏菌]
WP_039597687.1细胞色素C[雷尔氏菌(Ralstonia)sp.A12]
WP_055334967.1细胞色素C[青枯雷尔氏菌]
WP_078223437.1细胞色素C[血液病细菌(blood disease bacterium)A2-HRMARDI]
WP_013213209.1细胞色素C[青枯雷尔氏菌]
WP_063393008.1细胞色素C[甘露醇雷尔氏菌(Ralstonia mannitolilytica)]
WP_003265144.1细胞色素C，部分[青枯雷尔氏菌](序列编号12)
WP_039568931.1细胞色素C[青枯雷尔氏菌]
WP_045786290.1细胞色素C[甘露醇雷尔氏菌]
WP_064802148.1细胞色素C[危险雷尔氏菌(Ralstonia insidiosa)]
WP_021195198.1多菌种：细胞色素C[雷尔氏菌(Ralstonia)]
WP_004629446.1细胞色素C，单血红素-二血红素变体家族[皮氏雷尔氏菌]
WP_012435095.1细胞色素C[皮氏雷尔氏菌]
WP_003279246.1细胞色素C[青枯雷尔氏菌]
WP_048931829.1细胞色素C[雷尔氏菌sp.MD27]
WP_027677928.1细胞色素C[雷尔氏菌sp.UNC404CL21Col]
WP_012761509.1细胞色素C[皮氏雷尔氏菌](序列编号13)
WP_045204557.1细胞色素C[伯克霍尔德菌科细菌26]
CUV46938.1葡糖酸2-脱氢酶细胞色素C亚基[青枯雷尔氏菌]
WP_024973348.1细胞色素C[皮氏雷尔氏菌]
AKZ27209.1细胞色素C[青枯雷尔氏菌]
WP_020749403.1氧化还原酶脱氢酶(细胞色素C亚基)[青枯雷尔氏菌]
WP_024976325.1细胞色素C[皮氏雷尔氏菌]
WP_049842155.1细胞色素C[青枯雷尔氏菌]
WP_028852719.1细胞色素C[青枯雷尔氏菌]
WP_071895582.1细胞色素C[青枯雷尔氏菌]
SFP41323.1细胞色素C，单血红素-二血红素变体[雷尔氏菌sp.NFACC01]
WP_064477581.1细胞色素C[青枯雷尔氏菌]
WP_058908222.1细胞色素C[青枯雷尔氏菌]
WP_009238766.1多菌种：细胞色素C[雷尔氏菌]
CUV21878.1葡糖酸2-脱氢酶细胞色素C亚基[青枯雷尔氏菌]
WP_011000726.1细胞色素C[青枯雷尔氏菌]
WP_071507651.1细胞色素C[青枯雷尔氏菌]
WP_065857157.1细胞色素C[皮氏雷尔氏菌]
WP_019717689.1细胞色素C[青枯雷尔氏菌]
WP_071012822.1细胞色素C[青枯雷尔氏菌]
WP_020831436.1 2-酮-D-葡糖酸脱氢酶[青枯雷尔氏菌]
CUV55668.1葡糖酸2-脱氢酶细胞色素C亚基[青枯雷尔氏菌]
本发明的变异型细胞色素C蛋白也包括与如上所述的序列编号4具有60％以上的序列同一性的氨基酸序列，例如也包括在序列编号6～13中的任一氨基酸序列中按照从氨基末端侧数第1个第1血红素结合域和第2个第2血红素结合域缺失的方式进行了改变的序列。例如，可以是在序列编号6～13中的任一氨基酸序列中第1血红素结合域和第2血红素结合域缺失的序列、在序列编号6～13中的任一氨基酸序列中包含第1血红素结合域、第2血红素结合域的区域(与序列编号4的氨基酸编号43～195相当的区域)缺失的序列。
需要说明的是，作为改变基础的序列编号6～13的氨基酸序列可以是在改变部位和第3血红素结合域以外1个或几个氨基酸发生了置换、缺失、添加或插入的序列。
因此，本发明的变异型细胞色素C蛋白只要可保持电子传递功能，则也可以为具有下述序列的蛋白质，所述序列为在序列编号6～13中的任一氨基酸序列中第1血红素结合域和第2血红素结合域发生了缺失的氨基酸序列；或在序列编号6～13中的任一氨基酸序列中包含第1血红素结合域和第2血红素结合域的区域(与序列编号4的氨基酸编号43～195相当的区域)发生了缺失的氨基酸序列中，进一步在第3血红素结合域以外的位置有1个或几个氨基酸发生了置换、缺失、添加或插入的序列。此处，“1个或几个”例如为1～20个、优选为1～10个、更优选为1～8个、进一步优选为1～5个、特别优选为1～3个。另外，置换优选保守性置换，“保守性置换”是指酸性氨基酸彼此的置换、中性氨基酸彼此的置换、碱性氨基酸彼此的置换等性质类似的氨基酸彼此的置换。
<氧化还原酶-细胞色素复合体>
本发明的变异型细胞色素蛋白可以与氧化还原酶的催化亚基一起使用，其能够接受由氧化还原反应产生的电子并将该电子传递到电极。因此，包含本发明的变异型细胞色素蛋白的氧化还原酶-细胞色素复合体能够作为电化学传感器使用。电化学传感器优选为包含变异型细胞色素蛋白与葡萄糖氧化还原酶的葡萄糖传感器。
<氧化还原酶>
氧化还原酶只要是可对测定对象物质进行氧化还原的酶即可，作为催化亚基，可以举出包含吡咯并喹啉醌(PQQ)、黄素腺嘌呤二核苷酸(FAD)中的至少一者的酶的催化亚基。例如，作为包含PQQ的氧化还原酶，可以举出PQQ葡萄糖脱氢酶(PQQGDH)；作为包含FAD的氧化还原酶，可以举出具有包含FAD的α亚基的细胞色素葡萄糖脱氢酶(CyGDH)、葡萄糖氧化酶(GOD)。此外，还可以举出胆固醇氧化酶、醌血红素乙醇脱氢酶(QHEDH(PQQ Ethanol dh)、山梨糖醇脱氢酶(Sorbitol DH)、D-果糖脱氢酶(Fructose DH)、纤维二糖脱氢酶、乳酸脱氢酶。
其中优选葡萄糖脱氢酶(GDH)。
葡萄糖脱氢酶只要具有葡萄糖脱氢酶活性就没有特别限制，可以使用包括公知的葡萄糖脱氢酶在内的来源于各种生物的葡萄糖脱氢酶催化亚基。也可以与变异型细胞色素蛋白为同样的微生物来源，具体而言，例如可以使用来源于洋葱伯克霍尔德菌KS1株的具有序列编号3的氨基酸序列的α亚基蛋白。但是，只要可发挥出作为GDH的α亚基的功能，也可以为具有在序列编号3的氨基酸序列中1个或几个氨基酸残基发生了置换、缺失、插入或添加的氨基酸序列的蛋白质。另外，只要可发挥出作为GDH的α亚基的功能，也可以为具有在KS1株以外的α亚基的氨基酸序列中1个或几个氨基酸残基发生了置换、缺失、插入或添加的氨基酸序列的蛋白质。上述“1个或几个”优选为1～20个、更优选为1～10个、特别优选为1～5个。需要说明的是，已知有数量众多的使葡萄糖脱氢酶的α亚基蛋白的酶活性或底物特异性提高的变异，可以使用具有这些变异的葡萄糖脱氢酶的α亚基蛋白。
另外，葡萄糖脱氢酶复合体除了包含本发明的变异型细胞色素蛋白和催化亚基以外，还可以进一步包含γ亚基。作为γ亚基，只要可发挥出作为γ亚基的功能就没有特别限制，可以使用包括公知的γ亚基在内的来源于各种生物的γ亚基。也可以与变异型细胞色素蛋白为同样的微生物来源，具体而言，例如可以使用来源于洋葱伯克霍尔德菌KS1株的具有序列编号2的氨基酸序列的蛋白质。但是，只要可发挥出作为γ亚基的功能，也可以为具有在由序列编号2构成的氨基酸序列中1个或几个氨基酸残基发生了置换、缺失、插入或添加的氨基酸序列的蛋白质。另外，只要可发挥出作为γ亚基的功能，也可以为具有在KS1株以外的γ亚基的氨基酸序列中1个或几个氨基酸残基发生了置换、缺失、插入或添加的氨基酸序列的蛋白质。上述“1个或几个”优选为1～15个、更优选为1～10个、特别优选为1～5个。需要说明的是，发挥出作为γ亚基的功能是指在与α亚基一起形成复合体时可提高该复合体的GDH活性的功能。
<DNA>
本发明还提供编码变异型细胞色素蛋白的DNA。关于包含变异型细胞色素蛋白的DNA，其碱基序列可以基于变异型细胞色素蛋白的氨基酸序列来确定，例如，可以通过在编码野生型细胞色素蛋白的DNA中以第1血红素结合域和第2血红素结合域或包含它们的区域缺失的方式对碱基序列进行改变而得到。
例如，作为编码洋葱伯克霍尔德菌的细胞色素蛋白的DNA的具体例，可以举出编码葡萄糖脱氢酶β亚基的、包含由序列编号1的碱基编号2386～3660构成的碱基序列的DNA。另外，编码β亚基的DNA并不限于具有由序列编号1的碱基编号2386～3660构成的碱基序列的DNA，也可以为与具有该碱基序列的互补序列的DNA在严格条件下杂交、且编码可发挥出作为β亚基的功能的蛋白质的DNA。
作为编码变异型细胞色素蛋白的DNA，例如可以举出具有由序列编号5的碱基编号3372～3660或3325～3660构成的碱基序列的DNA、或与具有该碱基序列的互补序列的DNA在严格条件下杂交且编码可发挥出作为变异型细胞色素蛋白的功能的蛋白质的DNA。
具有所期望的变异的变异型细胞色素蛋白可以通过利用定点诱变法在编码细胞色素蛋白的DNA中导入所期望的氨基酸缺失、或利用PCR法仅使特定区域扩增，由此构建出变异DNA，使用适当的表达系统对其进行表达而获得。
在使用包含变异型细胞色素蛋白的氧化还原酶-细胞色素复合体的情况下，优选将编码氧化还原酶的催化亚基的DNA与编码变异型细胞色素蛋白的DNA一起使用。
编码氧化还原酶的催化亚基的DNA没有特别限制，可以根据目的适当选择来使用，例如，作为编码葡萄糖脱氢酶α亚基的DNA，可以举出包含由序列编号1的碱基编号764～2380构成的碱基序列的DNA。另外，α亚基基因可以为与具有由序列编号1的碱基序列的碱基编号764～2380构成的碱基序列的互补序列的DNA在严格条件下杂交、且编码具有GDH活性的蛋白质的DNA。
在氧化还原酶-细胞色素复合体中包含葡萄糖脱氢酶γ亚基的情况下，优选将编码氧化还原酶的γ亚基的DNA与编码变异型细胞色素蛋白的DNA和编码催化亚基的DNA一起使用。
作为编码γ亚基的DNA的具体例，可以举出包含由序列编号1的碱基编号258～761构成的碱基序列的DNA。另外，编码γ亚基的DNA也可以为与具有由序列编号1的碱基编号258～761构成的碱基序列的互补序列的DNA在严格条件下杂交、且编码可发挥出作为γ亚基的功能的蛋白质的DNA。
作为上述严格条件，可以举出具有优选80％、更优选90％以上、特别优选95％以上的同一性的DNA彼此杂交的条件，具体而言，可以举出在杂交反应后在0.1×SSC、0.1％SDS、60℃的条件下进行清洗的条件。
编码细胞色素蛋白的DNA、编码氧化还原酶的催化亚基的DNA等可以通过以洋葱伯克霍尔德菌等微生物的染色体DNA作为模板的PCR或杂交法等而获得。
序列编号1中示出包含洋葱伯克霍尔德菌KS1株的GDHγ亚基基因、α亚基基因以及β亚基基因的染色体DNA片段的碱基序列。该碱基序列中存在3个开放读码框(ORF)，自5’末端侧起，第1个ORF编码γ亚基(序列编号2)、第2个ORF编码α亚基(序列编号3)、第3个ORF编码β亚基(序列编号4)。
变异型细胞色素蛋白、活性亚基可以各自分别地进行DNA表达，也可以使用使编码变异型细胞色素蛋白的DNA和编码活性亚基的DNA包含在多顺反子中的DNA进行表达。
<载体>
作为在变异型细胞色素蛋白、活性亚基的基因的获得、变异的导入、基因的表达等中使用的载体，只要是在宿主微生物中发挥功能的载体就没有特别限制，例如可以举出在埃希氏菌属细菌中发挥功能的载体，具体而言，可以举出pTrc99A、pBR322、pUC18、pUC118、pUC19、pUC119、pACYC184、pBBR122等。基因表达中使用的启动子也可以根据宿主适当选择，例如可以举出lac、trp、tac、trc、PL、tet、PhoA等。
<转化体>
通过将上述的DNA或包含其的载体导入到宿主微生物中而得到转化体，能够表达出变异型细胞色素蛋白或氧化还原酶-细胞色素复合体。转化可以采用公知的方法，例如可以举出基于钙处理的感受态细胞法、原生质体法或电穿孔法等。
作为宿主微生物，只要是在蛋白质表达中使用的微生物就没有特别限制，可以举出大肠埃希氏菌(Escherichia coli)等埃希氏菌属细菌、枯草芽孢杆菌(Bacillussubtilis)等芽胞杆菌属细菌、酿酒酵母(Saccharomyces cerevisiae)、毕赤酵母(Pichiapastoris)等酵母、黑曲霉(Aspergillus niger)、米曲霉(Aspergillus oryzae)等丝状菌，但并不限于这些。
<生物传感器>
本发明的变异型细胞色素蛋白可用于利用氧化还原反应的电化学生物传感器。生物传感器的种类可以根据所组合的氧化还原酶的种类而用作各种用途，例如，在使用葡萄糖氧化还原酶作为氧化还原酶的情况下，可以作为葡萄糖传感器；在使用乳酸脱氢酶作为氧化还原酶的情况下，可以作为乳酸传感器。
其中优选葡萄糖传感器，在制作葡萄糖传感器的情况下，优选在酶电极的试剂层中含有本发明的变异型细胞色素蛋白和氧化还原酶的催化亚基。还可以进一步含有γ亚基。由此，利用催化亚基将试样中的葡萄糖氧化，变异型细胞色素蛋白接受此时生成的电子并传递至电极，由此流通与试样中的葡萄糖浓度相应的响应电流，根据该电流值可以计算出葡萄糖浓度。
作为葡萄糖传感器，具体而言，可以举出下述葡萄糖传感器：该葡萄糖传感器使用在金电极、铂电极、碳电极等电极表面上固定有包含本发明的变异型细胞色素蛋白和葡萄糖脱氢酶的催化亚基的复合体的酶电极作为工作电极。传感器是指对目标待测物质的浓度进行电化学测定的测定系统，优选为包含工作电极(酶电极)、反电极(铂等)和参比电极(Ag/AgCl等)这3个电极的系统。也可以是在惯用的简易血糖值系统中所使用的由工作电极和反电极构成的2电极系统。传感器优选进一步包含放入缓冲液和待测试样的恒温池、对工作电极施加电位的电源、电流计、记录仪等。传感器可以是间歇型也可以是流动型。特别是作为流动型的传感器，可以是能够连续测量血糖值的传感器。即，可以是在连续性供给的血液试样或连续性供给的透析试样、或者血液中或细胞间液中插入固定有本发明的酶的二电极系统或三电极系统来进行测量的传感器。这样的酶传感器的结构在该技术领域中是熟知的，例如记载于Biosensors-Fundamental and Applications(生物传感器基础和应用)-Anthony P.F.Turner,Isao Karube and Geroge S.Wilson,牛津大学出版社(OxfordUniversity Press)1987中。
使用本发明的葡萄糖传感器进行的葡萄糖浓度的测定例如可以如下进行。在传感器的恒温池中加入缓冲液，维持在一定温度。作为工作电极，使用固定有本发明的变异型细胞色素蛋白和葡萄糖氧化还原酶的催化亚基蛋白的酶电极，作为反电极，例如使用铂电极，作为参比电极，例如使用Ag/AgCl电极。对工作电极施加一定的电位(例如，对于银/氯化银电极为0～+300mV、优选为0～+150mV、更优选为0～+100mV)，在电流达到稳定后，向恒温池中加入包含葡萄糖的试样，测定电流的增加。根据利用标准浓度的葡萄糖溶液制作出的校正曲线，可以计算出试样中的葡萄糖浓度。
包含本发明的变异型细胞色素蛋白和GDH催化亚基的GDH复合体还可以用作葡萄糖检测试剂盒的构成要素。在葡萄糖检测试剂盒中，除了包含本发明的变异型细胞色素蛋白和GDH催化亚基的GDH复合体以外，还可以包含显色或发光试剂、稀释用缓冲液、标准物质、使用说明书等。
例如，在美国专利公开第2004/0023330A1中记载了使用洋葱伯克霍尔德菌的野生型GDH的葡萄糖传感器和葡萄糖检测试剂盒。包含本发明的变异型细胞色素蛋白的GDH也可以同样地使用。
[实施例]
接下来举出实施例进一步具体说明本发明，但本发明并不受这些实施例的任何限定。
[实施例1]变异型GDHβ亚基基因的构建
作为变异型GDHβ亚基基因的构建中使用的质粒，使用在日本特开2012-090563所记载的质粒pTrc99Aγαβ中在α亚基中导入了变异的pTrc99Aγα(QYY)β。该质粒是将下述DNA片段插入到载体pTrc99A的克隆位点而成的质粒，该DNA片段连续地包含由洋葱伯克霍尔德菌KS1株(FERM BP-7306)染色体DNA中分离出的GDHγ亚基结构基因和α亚基结构基因、β亚基结构基因，其中在α亚基结构基因中，编码第326位残基的丝氨酸、第365位残基的丝氨酸、第472位残基的丙氨酸的密码子分别被置换成编码谷氨酰胺、酪氨酸、酪氨酸的密码子。本质粒中的GDH的各结构基因由trc启动子控制。pTrc99Aγαβ保有氨苄青霉素耐性基因。
首先，通过以上述质粒DNA作为模板、以具有下述序列的低聚核苷酸作为引物的PCR对编码至GDH的γ亚基、α亚基和β亚基的信号序列为止的DNA片段进行扩增。
[正向引物]
5’-ACGTAGCCATGGCACACAACGACAACCACTC-3’(序列编号14)
[反向引物]
5’-ATCGGCCGCGCGCGCGAAGCCCGGCAA-3’(序列编号15)
关于PCR反应，以下述反应液组成进行95℃30秒、之后重复(95℃30秒、55℃1分钟、72℃2分钟)25个循环、72℃5分钟的反应，之后保持在4℃。
[反应液组成]
另外，通过以具有下述序列的低聚核苷酸作为引物的PCR对编码β亚基的第3血红素结合域、即编码序列编号4的氨基酸编号314～425或330～425的区域的DNA片段进行扩增。需要说明的是，正向引物均在5’末端侧添加有与β亚基的信号序列相当的序列。
[正向引物314～425扩增用]
5’-TTGCCGGGCTTCGCGCGCGCGGCCGATCTGCGCGGTGTCGCGCTCGCG-3’(序列编号16)
[正向引物330～425扩增用]
5’-TTGCCGGGCTTCGCGCGCGCGGCCGATTATCTCGGCAACTGCGCGACG-3’(序列编号17)
[反向引物]
5’-GTGGTGCTCGAGTGCGGCCGCGCGCAGCTTCGCGACGTCCTG-3’(序列编号18)
关于PCR反应，以下述反应液组成进行95℃30秒、之后重复(95℃30秒、55℃1分钟、72℃30秒)25个循环、72℃5分钟的反应，之后保持在4℃。
[反应液组成]
将由上述操作得到的DNA片段分别纯化后，将它们混合作为模板，通过以具有以下序列的低聚核苷酸作为引物的重叠延伸PCR(Overlap extention PCR)进行结合、扩增。需要说明的是，在反向引物中设计了组氨酸标签序列，由此在变异型β亚基的C末端添加组氨酸标签。
[正向引物]
5’-ACGTAGCCATGGCACACAACGACAACCACTC-3’(序列编号14)
[反向引物]
5’-GTACGTAAGCTTTCAGTGGTGGTGGTGGTGGTGCTCGAGTGCGGCCGC-3’序列编号19)
关于PCR反应，以下述反应液组成进行95℃30秒、之后重复(95℃30秒、55℃1分钟、72℃4分钟)25个循环、72℃5分钟的反应，之后保持在4℃。
[反应液组成]
将所得到的PCR产物纯化后，将N末端侧用NcoI消化、C末端侧用HindIII消化，与同样处理后的pTrc99A连接。利用所得到的重组载体对大肠埃希氏菌DH5α进行转化，采集在包含羧苄西林50μg/mL的LB琼脂培养基上生长的菌落。将所得到的转化体在液体的LB培养基中进行培养，提取质粒，对其插入DNA片段进行分析，结果确认到，在γ亚基、α亚基、β亚基的信号序列之后插入有编码β亚基的第314～425位残基的基因或编码该β亚基的第330～425位残基的基因。将这些质粒分别命名为pTrc99Aγα(QYY)β314-His、pTrc99Aγα(QYY)β330-His。这些质粒中的GDH的各结构基因在β亚基的C末端具有组氨酸标签，由trc启动子进行控制。另外，均保有氨苄青霉素耐性基因。
[实施例2]包含变异型GDHβ亚基的洋葱伯克霍尔德菌的GDH的表达
使用实施例1中得到的表达质粒，制造包含变异型GDHβ亚基基因的GDH。
将导入有包含变异型GDHβ亚基基因的GDH表达质粒和具有细胞色素c成熟所需的基因群的质粒pBBJMccm的大肠埃希氏菌BL21(DE3)株在3ml的LB培养基(含有羧苄西林50μg/ml和卡那霉素50μg/ml)中使用试验管于37℃振荡培养一夜。将这些培养液以1％接种在2个300ml的振荡烧瓶中，该烧瓶包含按照使每1升培养液的组成为表1所示的方式制备出的培养基50ml(含有羧苄西林50μg/ml和卡那霉素50μg/ml)，在25℃振荡培养28小时。需要说明的是，在宿主能够稳定地使细胞色素c成熟的情况下，例如在通过同源重组等将ccm基因插入到基因组中、并进行稳定表达的转化株等中，不需要pBBJMccm之类的载体。
[表1]
表1.培养基组成(每1升)
| 胰蛋白胨 | 10g |
| 酵母提取物 | 5g |
| 氯化钠 | 5g |
| 甘油 | 0.5g |
| 葡萄糖 | 0.05g |
| 乳糖一水合物 | 0.21g |
| 硫酸铵 | 0.33g |
| 磷酸二氢钾 | 0.68g |
| 磷酸氢二钠十二水合物 | 1.79g |
| 硫酸镁 | 0.025g |
从上述培养的培养液中收集菌体，在所得到的湿菌体每1g中添加5ml的BugBuster蛋白提取试剂(Merck-Millipore)并混悬，使菌体溶解。对该混悬液进行离心分离(15000r.p.m、20分钟、4℃)，除去残渣，制成粗酶样品。
另外，对于γα(QYY)β314-His，将粗酶样品在包含0.5M氯化钠、20mM咪唑的20mM磷酸钠缓冲液(pH7.0)中透析一夜后，添加到填充有用该缓冲液平衡后的Ni-NTA琼脂糖(QIAGEN)的柱中，用包含0.5M氯化钠、54mM咪唑的20mM磷酸钠缓冲液(pH7.0)清洗后，用包含0.5M氯化钠、140mM咪唑的20mM磷酸钠缓冲液(pH7.0)洗脱，得到纯化酶样品。
[实施例3]包含变异型GDHβ亚基的洋葱伯克霍尔德菌的GDH的功能分析
GDH活性的测定利用PMS/DCIP系统和钌(Ru)/MTT系统进行。前者表示通常的GDH活性，后者表示藉由β亚基向介体(mediator)进行电子传递的情况下的GDH活性。
PMS/DCIP系统中的活性测定如下进行。在20mM磷酸钾缓冲液(pH7.0)中添加酶样品、终浓度为0.6mM的甲基吩嗪硫酸甲酯(PMS)、终浓度为0.06mM的2,6-二氯酚靛酚(DCIP)、终浓度为4mM或40mM的葡萄糖，使用分光光度计测定600nm(其为DCIP来源的吸收波长)的吸光度的每1分钟的变化量。
另外，Ru/MTT系统中的活性测定如下进行。在20mM磷酸钾缓冲液(pH7.0)中添加酶样品、终浓度为2％的六氨络钌(III)氯化物、终浓度为1mM的3-(4,5-二甲基-2-噻唑基)-2,5-二苯基-2H-四氮唑溴化物(MTT)、终浓度为4mM或40mM的葡萄糖，使用分光光度计测定565nm(其为来源于由MTT生成的甲的吸收波长)的吸光度的每1分钟的变化量。
将粗酶样品的活性测定的结果示于表2。
[表2]
表中，虽然不及包含野生型β亚基的酶，但两突变体在Ru/MTT系统中均显示出了GDH活性，暗示出这些突变体藉由β亚基进行了向介体的电子传递。另外，在野生型酶的情况下，在不含β亚基的γα复合体中，以PMS/DCIP作为电子受体时，酶活性为约45U/mg(Biochimica et Biophysica Acta,1645(2),133-138.)，仅观察到包含β亚基的情况下的值(约300U/mg：Journal of Biotechnology,123(2),127-136.)的约15％的酶活性，但包含本发明的变异细胞色素c的复合体显示出与包含野生型β亚基复合体时几乎相同的活性。因此发现了，该变异β亚基从α亚基正常接受电子、并将该电子传递至PMS/DCIP。
另外，对γα(QYY)β314进行纯化，按照上述过程用另行制备的包含野生型β亚基的γα(QYY)β一起测定相对于40mM葡萄糖的酶活性。将结果示于表3。
[表3]
在纯化酶中，γα(QYY)β314在Ru/MTT系统中也显示出GDH活性，与在PMS/DCIP系统中的GDH活性之比也与包含野生型β亚基的酶复合体是同样的。由此暗示出，该突变体与包含野生型β亚基的酶复合体同样地藉由β亚基进行了向介体的电子传递。
[实施例4]葡萄糖传感器的制作
制作在金表面藉由单分子膜形成分子固定有包含变异型β亚基的GDH的酶电极。作为单分子膜形成分子，使用以下的DSH。
[化1]
具体而言，在室温下将金导线(直径0.5mm、长度6～7cm)在食人鱼洗液(200μl)中浸渍2小时，之后用丙酮清洗，浸渍在DSH的丙酮溶液(浓度20μM)中，在25℃温育24小时，使DSH的巯基与金表面结合。接着用丙酮清洗，浸渍在包含上述GDHα、β亚基复合体(浓度0.03mg/ml))的磷酸缓冲液(300μl)中，于4℃温育一夜，使酶复合体藉由DSH的官能团进行结合，得到酶电极。
[实施例5]葡萄糖浓度的测定
使用上述酶电极，通过计时安培分析法对0mM(背景)、1mM、5mM或50mM的葡萄糖水溶液进行响应电流值的测定。葡萄糖测定中，使用反电极(Pt导线)、参比电极(银/氯化银)，使向工作电极的施加电位为0V、+0.1V、+0.4V(vs.银/氯化银)，在37℃进行测定。
<结果>
以对于银氯化银电极施加400mV后的电流值的葡萄糖浓度特性为基准，对0mV和100mV的电流值进行比较。
其结果，如图1所示，在0mV的施加电位下，在比较例(虚线)中未观测到葡萄糖浓度依赖性的氧化电流，与之相对，在实施例(实线)中观察到氧化电流的葡萄糖浓度依赖性。
在100mV的施加电位下，在比较例中也观测到葡萄糖浓度依赖性的氧化电流，但以400mV的响应电流为基准的电流值输出在实施例中更高。
由以上表明，在实施例的使用截短形式(truncated)的β亚基的传感器中，能够以比使用野生型β亚基的传感器更低的氧化电位获得催化电流。
[工业实用性]
使用了本发明的变异型细胞色素蛋白的生物传感器能够进行低电位下的电流测定，能够适合用于葡萄糖传感器等生物传感器。
序列表
<110> 爱科来株式会社(Arkray, Inc.)
究极酵素国际股份有限公司(Ultizyme International Ltd.)
<120> 变异型细胞色素蛋白及其应用(Mutant cytochrome protein and usethereof)
<130> P-170534
<150> JP2017-131345
<151> 2017-07-04
<160> 19
<170> PatentIn version 3.5
<210> 1
<211> 3706
<212> DNA
<213> 洋葱伯克霍尔德菌(Burkholderia cepacia)
<220>
<221> CDS
<222> (258)..(761)
<220>
<221> CDS
<222> (764)..(2380)
<220>
<221> CDS
<222> (2386)..(3660)
<400> 1
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<210> 2
<211> 168
<212> PRT
<213> 洋葱伯克霍尔德菌(Burkholderia cepacia)
<400> 2
Met His Asn Asp Asn Thr Pro His Ser Arg Arg His Gly Asp Ala Ala
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Asn Pro Gly Thr Ala Pro Leu Asp Thr Phe Met Thr Leu Ser Glu Ser
50 55 60
Leu Thr Gly Lys Lys Gly Leu Ser Arg Val Ile Gly Glu Arg Leu Leu
65 70 75 80
Gln Ala Leu Gln Lys Gly Ser Phe Lys Thr Ala Asp Ser Leu Pro Gln
85 90 95
Leu Ala Gly Ala Leu Ala Ser Gly Ser Leu Thr Pro Glu Gln Glu Ser
100 105 110
Leu Ala Leu Thr Ile Leu Glu Ala Trp Tyr Leu Gly Ile Val Asp Asn
115 120 125
Val Val Ile Thr Tyr Glu Glu Ala Leu Met Phe Gly Val Val Ser Asp
130 135 140
Thr Leu Val Ile Arg Ser Tyr Cys Pro Asn Lys Pro Gly Phe Trp Ala
145 150 155 160
Asp Lys Pro Ile Glu Arg Gln Ala
165
<210> 3
<211> 539
<212> PRT
<213> 洋葱伯克霍尔德菌(Burkholderia cepacia)
<400> 3
Met Ala Asp Thr Asp Thr Gln Lys Ala Asp Val Val Val Val Gly Ser
1 5 10 15
Gly Val Ala Gly Ala Ile Val Ala His Gln Leu Ala Met Ala Gly Lys
20 25 30
Ala Val Ile Leu Leu Glu Ala Gly Pro Arg Met Pro Arg Trp Glu Ile
35 40 45
Val Glu Arg Phe Arg Asn Gln Pro Asp Lys Met Asp Phe Met Ala Pro
50 55 60
Tyr Pro Ser Ser Pro Trp Ala Pro His Pro Glu Tyr Gly Pro Pro Asn
65 70 75 80
Asp Tyr Leu Ile Leu Lys Gly Glu His Lys Phe Asn Ser Gln Tyr Ile
85 90 95
Arg Ala Val Gly Gly Thr Thr Trp His Trp Ala Ala Ser Ala Trp Arg
100 105 110
Phe Ile Pro Asn Asp Phe Lys Met Lys Ser Val Tyr Gly Val Gly Arg
115 120 125
Asp Trp Pro Ile Gln Tyr Asp Asp Leu Glu Pro Tyr Tyr Gln Arg Ala
130 135 140
Glu Glu Glu Leu Gly Val Trp Gly Pro Gly Pro Glu Glu Asp Leu Tyr
145 150 155 160
Ser Pro Arg Lys Gln Pro Tyr Pro Met Pro Pro Leu Pro Leu Ser Phe
165 170 175
Asn Glu Gln Thr Ile Lys Thr Ala Leu Asn Asn Tyr Asp Pro Lys Phe
180 185 190
His Val Val Thr Glu Pro Val Ala Arg Asn Ser Arg Pro Tyr Asp Gly
195 200 205
Arg Pro Thr Cys Cys Gly Asn Asn Asn Cys Met Pro Ile Cys Pro Ile
210 215 220
Gly Ala Met Tyr Asn Gly Ile Val His Val Glu Lys Ala Glu Arg Ala
225 230 235 240
Gly Ala Lys Leu Ile Glu Asn Ala Val Val Tyr Lys Leu Glu Thr Gly
245 250 255
Pro Asp Lys Arg Ile Val Ala Ala Leu Tyr Lys Asp Lys Thr Gly Ala
260 265 270
Glu His Arg Val Glu Gly Lys Tyr Phe Val Leu Ala Ala Asn Gly Ile
275 280 285
Glu Thr Pro Lys Ile Leu Leu Met Ser Ala Asn Arg Asp Phe Pro Asn
290 295 300
Gly Val Ala Asn Ser Ser Asp Met Val Gly Arg Asn Leu Met Asp His
305 310 315 320
Pro Gly Thr Gly Val Ser Phe Tyr Ala Ser Glu Lys Leu Trp Pro Gly
325 330 335
Arg Gly Pro Gln Glu Met Thr Ser Leu Ile Gly Phe Arg Asp Gly Pro
340 345 350
Phe Arg Ala Thr Glu Ala Ala Lys Lys Ile His Leu Ser Asn Leu Ser
355 360 365
Arg Ile Asp Gln Glu Thr Gln Lys Ile Phe Lys Ala Gly Lys Leu Met
370 375 380
Lys Pro Asp Glu Leu Asp Ala Gln Ile Arg Asp Arg Ser Ala Arg Tyr
385 390 395 400
Val Gln Phe Asp Cys Phe His Glu Ile Leu Pro Gln Pro Glu Asn Arg
405 410 415
Ile Val Pro Ser Lys Thr Ala Thr Asp Ala Ile Gly Ile Pro Arg Pro
420 425 430
Glu Ile Thr Tyr Ala Ile Asp Asp Tyr Val Lys Arg Gly Ala Ala His
435 440 445
Thr Arg Glu Val Tyr Ala Thr Ala Ala Lys Val Leu Gly Gly Thr Asp
450 455 460
Val Val Phe Asn Asp Glu Phe Ala Pro Asn Asn His Ile Thr Gly Ser
465 470 475 480
Thr Ile Met Gly Ala Asp Ala Arg Asp Ser Val Val Asp Lys Asp Cys
485 490 495
Arg Thr Phe Asp His Pro Asn Leu Phe Ile Ser Ser Ser Ala Thr Met
500 505 510
Pro Thr Val Gly Thr Val Asn Val Thr Leu Thr Ile Ala Ala Leu Ala
515 520 525
Leu Arg Met Ser Asp Thr Leu Lys Lys Glu Val
530 535
<210> 4
<211> 425
<212> PRT
<213> 洋葱伯克霍尔德菌(Burkholderia cepacia)
<400> 4
Met Arg Lys Ser Thr Leu Thr Phe Leu Ile Ala Gly Cys Leu Ala Leu
1 5 10 15
Pro Gly Phe Ala Arg Ala Ala Asp Ala Ala Asp Pro Ala Leu Val Lys
20 25 30
Arg Gly Glu Tyr Leu Ala Thr Ala Gly Asp Cys Met Ala Cys His Thr
35 40 45
Val Lys Gly Gly Lys Pro Tyr Ala Gly Gly Leu Gly Met Pro Val Pro
50 55 60
Met Leu Gly Lys Ile Tyr Thr Ser Asn Ile Thr Pro Asp Pro Asp Thr
65 70 75 80
Gly Ile Gly Lys Trp Thr Phe Glu Asp Phe Glu Arg Ala Val Arg His
85 90 95
Gly Val Ser Lys Asn Gly Asp Asn Leu Tyr Pro Ala Met Pro Tyr Val
100 105 110
Ser Tyr Ala Lys Ile Thr Asp Asp Asp Val Arg Ala Leu Tyr Ala Tyr
115 120 125
Phe Met His Gly Val Glu Pro Val Lys Gln Ala Pro Pro Lys Asn Glu
130 135 140
Ile Pro Ala Leu Leu Ser Met Arg Trp Pro Leu Lys Ile Trp Asn Trp
145 150 155 160
Leu Phe Leu Lys Asp Gly Pro Tyr Gln Pro Lys Pro Ser Gln Ser Ala
165 170 175
Glu Trp Asn Arg Gly Ala Tyr Leu Val Gln Gly Leu Ala His Cys Ser
180 185 190
Thr Cys His Thr Pro Arg Gly Ile Ala Met Gln Glu Lys Ser Leu Asp
195 200 205
Glu Thr Gly Gly Ser Phe Leu Ala Gly Ser Val Leu Ala Gly Trp Asp
210 215 220
Gly Tyr Asn Ile Thr Ser Asp Pro Asn Ala Gly Ile Gly Ser Trp Thr
225 230 235 240
Gln Gln Gln Leu Val Gln Tyr Leu Arg Thr Gly Ser Val Pro Gly Val
245 250 255
Ala Gln Ala Ala Gly Pro Met Ala Glu Ala Val Glu His Ser Phe Ser
260 265 270
Lys Met Thr Glu Ala Asp Ile Gly Ala Ile Ala Thr Tyr Val Arg Thr
275 280 285
Val Pro Ala Val Ala Asp Ser Asn Ala Lys Gln Pro Arg Ser Ser Trp
290 295 300
Gly Lys Pro Ala Glu Asp Gly Leu Lys Leu Arg Gly Val Ala Leu Ala
305 310 315 320
Ser Ser Gly Ile Asp Pro Ala Arg Leu Tyr Leu Gly Asn Cys Ala Thr
325 330 335
Cys His Gln Met Gln Gly Lys Gly Thr Pro Asp Gly Tyr Tyr Pro Ser
340 345 350
Leu Phe His Asn Ser Thr Val Gly Ala Ser Asn Pro Ser Asn Leu Val
355 360 365
Gln Val Ile Leu Asn Gly Val Gln Arg Lys Ile Gly Ser Glu Asp Ile
370 375 380
Gly Met Pro Ala Phe Arg Tyr Asp Leu Asn Asp Ala Gln Ile Ala Ala
385 390 395 400
Leu Thr Asn Tyr Val Thr Ala Gln Phe Gly Asn Pro Ala Ala Lys Val
405 410 415
Thr Glu Gln Asp Val Ala Lys Leu Arg
420 425
<210> 5
<211> 5
<212> PRT
<213> 人工序列(Artificial Sequence)
<220>
<223> 血红素结合部位(heme binding site)
<220>
<221> misc_feature
<222> (2)..(3)
<223> Xaa可以为任意天然存在的氨基酸
<400> 5
Cys Xaa Xaa Cys His
1 5
<210> 6
<211> 425
<212> PRT
<213> 洋葱伯克霍尔德菌(Burkholderia cepacia)
<400> 6
Met Arg Lys Ser Thr Leu Thr Phe Leu Leu Ala Gly Cys Leu Ala Leu
1 5 10 15
Pro Gly Leu Ala Arg Ala Ala Asp Ser Ala Asp Pro Ala Gln Val Lys
20 25 30
Arg Gly Glu Tyr Leu Ala Val Ala Gly Asp Cys Met Ala Cys His Thr
35 40 45
Ala Lys Gly Gly Lys Pro Phe Ala Gly Gly Leu Gly Met Pro Val Pro
50 55 60
Met Leu Gly Lys Ile Tyr Thr Ser Asn Ile Thr Pro Asp Pro Asp Thr
65 70 75 80
Gly Ile Gly Asn Trp Thr Phe Glu Asp Phe Glu Arg Ala Val Arg His
85 90 95
Gly Val Ser Lys Asn Gly Asp Asn Leu Tyr Pro Ala Met Pro Tyr Val
100 105 110
Ser Tyr Ala Lys Ile Asn Asp Asp Asp Val Gln Ala Leu Tyr Ala Tyr
115 120 125
Phe Met His Gly Val Glu Pro Val Lys Gln Ala Pro Pro Lys Asn Glu
130 135 140
Ile Pro Ala Leu Leu Ser Met Arg Trp Pro Leu Lys Ile Trp Asn Trp
145 150 155 160
Leu Phe Leu Lys Asp Gly Val Tyr Gln Pro Lys Pro Glu Gln Ser Ala
165 170 175
Glu Trp Asn Arg Gly Ala Tyr Leu Val Gln Gly Leu Ala His Cys Ser
180 185 190
Thr Cys His Thr Pro Arg Gly Ile Ala Met Gln Glu Lys Ser Leu Asp
195 200 205
Glu Thr Gly Gly Ser Phe Leu Ser Gly Ser Val Leu Ala Gly Trp Asp
210 215 220
Gly Tyr Asn Ile Thr Ser Asp Pro Asn Ala Gly Ile Gly Gly Trp Thr
225 230 235 240
Gln Gln Gln Leu Val Gln Tyr Leu Arg Thr Gly Ser Val Pro Gly Leu
245 250 255
Ala Gln Ala Ala Gly Pro Met Ala Glu Ala Ile Glu His Ser Phe Ser
260 265 270
Lys Met Thr Glu Ala Asp Ile Gly Ala Ile Ser Thr Tyr Ile Arg Thr
275 280 285
Val Pro Ala Val Ala Ser Gly Asp Ala Lys Gln Ser Arg Ser Ser Trp
290 295 300
Gly Lys Pro Ala Glu Asp Gly Leu Lys Leu Arg Gly Val Ala Leu Ala
305 310 315 320
Ser Ser Gly Ile Asp Pro Ala Arg Leu Tyr Leu Gly Asn Cys Ala Thr
325 330 335
Cys His Gln Met Gln Gly Lys Gly Thr Pro Asp Gly Tyr Tyr Pro Pro
340 345 350
Leu Phe His Asn Ser Thr Val Gly Ala Ser Asn Pro Thr Asn Leu Val
355 360 365
Gln Val Ile Leu Asn Gly Val Gln Arg Lys Ala Gly Ser Glu Asp Val
370 375 380
Gly Met Pro Ala Phe Arg His Glu Leu Ser Asp Ala Gln Ile Ala Ala
385 390 395 400
Leu Thr Asn Tyr Leu Thr Gly Gln Phe Gly Asn Pro Ala Ala Lys Val
405 410 415
Thr Glu Gln Asp Val Ala Lys Leu Arg
420 425
<210> 7
<211> 428
<212> PRT
<213> 新洋葱伯克霍尔德菌(Burkholderia cenocepacia)
<400> 7
Met Arg Leu Pro Gly Leu Ser Val Leu Val Val Phe Phe Leu Ile Ala
1 5 10 15
Thr Asp Pro Ala Ser Ala Ala Gly Gln Ala Ala Asp Arg Ile Ala Ala
20 25 30
Gly Lys Ala Leu Ala Val Ala Ala Asp Cys Val Ala Cys His Thr Val
35 40 45
Gln Gly Gly Gln Pro Phe Ala Gly Gly Leu Ala Met Pro Leu Pro Met
50 55 60
Gly Asp Ile Tyr Ser Thr Asn Ile Thr Pro Asp Arg Gln Thr Gly Ile
65 70 75 80
Gly Asn Tyr Ser Glu Arg Asp Phe Ala Asn Val Leu Arg Lys Gly Leu
85 90 95
Arg Arg Asp Gly Gly Asn Leu Tyr Pro Ala Met Pro Tyr Pro Ser Tyr
100 105 110
Thr Lys Phe Ser Asp Asp Asp Ile Ala Asn Leu Tyr Ala Tyr Phe Met
115 120 125
Gln Gly Val Ala Pro Val Arg Gln Pro Asn Arg Ala Pro Asp Phe Pro
130 135 140
Trp Pro Leu Thr Met Arg Trp Pro Leu Lys Ile Trp Asn Ala Leu Tyr
145 150 155 160
Leu Arg Glu Gly Ala Tyr Val Pro Lys Pro Gly Arg Asp Ala Glu Trp
165 170 175
Asn Arg Gly Ala Tyr Leu Val Gln Gly Ala Ala His Cys Gly Thr Cys
180 185 190
His Thr Pro Arg Gly Ile Gly Met Gln Glu Leu Ala Tyr Asp Glu Thr
195 200 205
Gly Val Gly Tyr Leu Ala Gly Ala Pro Leu Ala Gly Trp Gln Ala Phe
210 215 220
Asn Ile Thr His Asp Arg Asp Ala Gly Ile Gly Thr Trp Thr Thr Ala
225 230 235 240
Gln Ile Val Gln Tyr Leu Arg Thr Gly Asn Val Pro Gly Lys Ala Gln
245 250 255
Ala Ala Gly Pro Met Ala Glu Ala Val Glu His Ser Phe Ser Arg Met
260 265 270
Ser Glu Arg Asp Leu Asn Ala Ile Ala Val Tyr Leu Ser Thr Val Pro
275 280 285
Ala Ala Lys Gly Ala Asp Thr Ala Pro Arg Ser Thr Gln Gly Arg Pro
290 295 300
Thr Asp Asp Tyr Val Ala Ile Arg Ala Ala Ala Ala Ala Gly Ser Trp
305 310 315 320
Thr Pro Ala Gly Ala Ser Leu Tyr Leu Asp His Cys Ala Ser Cys His
325 330 335
Gly Met Thr Gly Ala Gly Thr Thr Asp Gly Phe Phe Pro Ser Leu Phe
340 345 350
Ala Asn Ser Ala Val Gly Thr Ala Thr Ala Ser Asn Leu Leu Gln Val
355 360 365
Val Met His Gly Ala Ser Val Asn Asn Gly Thr Thr His Tyr Phe Met
370 375 380
Pro Ala Phe Gln Thr Glu Leu Asp Asp Asp Glu Val Val Thr Leu Val
385 390 395 400
Asn Tyr Leu Ser Glu Arg Phe Gly Asn Gly Arg Ala Arg Val Ser Ala
405 410 415
Ala Glu Val Ala Lys Ile Arg Thr Ala Pro Ala His
420 425
<210> 8
<211> 425
<212> PRT
<213> 多噬伯克霍尔德菌(Burkholderia multivorans)
<400> 8
Met Arg Lys Ser Thr Leu Thr Phe Leu Ile Ala Gly Cys Leu Ala Leu
1 5 10 15
Pro Gly Leu Ala Arg Ala Ala Asp Ala Ala Asp Pro Ala Leu Val Lys
20 25 30
Arg Gly Glu Tyr Leu Ala Thr Ala Gly Asp Cys Met Ala Cys His Thr
35 40 45
Val Lys Gly Gly Lys Pro Tyr Ala Gly Gly Leu Gly Met Pro Val Pro
50 55 60
Met Leu Gly Lys Ile Tyr Thr Ser Asn Ile Thr Pro Asp Pro Asp Thr
65 70 75 80
Gly Ile Gly Lys Trp Thr Phe Glu Asp Phe Glu Arg Ala Val Arg His
85 90 95
Gly Val Ser Lys Asn Gly Asp Asn Leu Tyr Pro Ala Met Pro Tyr Val
100 105 110
Ser Tyr Ala Lys Ile Thr Asp Asp Asp Val Arg Ala Leu Tyr Ala Tyr
115 120 125
Phe Met His Gly Val Glu Pro Val Lys Gln Ala Pro Pro Lys Asn Glu
130 135 140
Ile Pro Ala Leu Leu Ser Met Arg Trp Pro Leu Lys Ile Trp Asn Trp
145 150 155 160
Leu Phe Leu Lys Asp Gly Pro Tyr Gln Pro Lys Pro Ser Gln Ser Ala
165 170 175
Glu Trp Asn Arg Gly Ala Tyr Leu Val Gln Gly Leu Ala His Cys Ser
180 185 190
Thr Cys His Thr Pro Arg Gly Ile Ala Met Gln Glu Lys Ser Leu Asp
195 200 205
Glu Thr Gly Gly Ser Phe Leu Ala Gly Ser Val Leu Ala Gly Trp Asp
210 215 220
Gly Tyr Asn Ile Thr Ser Asp Pro Asn Ala Gly Ile Gly Ser Trp Thr
225 230 235 240
Gln Gln Gln Leu Val Gln Tyr Leu Arg Thr Gly Ser Val Pro Gly Val
245 250 255
Ala Gln Ala Ala Gly Pro Met Ala Glu Ala Val Glu His Ser Phe Ser
260 265 270
Lys Met Thr Glu Ala Asp Ile Gly Ala Ile Ala Thr Tyr Val Arg Thr
275 280 285
Val Pro Ala Val Ala Asp Ser Asn Ala Lys Gln Pro Arg Ser Ser Trp
290 295 300
Gly Lys Pro Ala Glu Asp Gly Leu Lys Leu Arg Gly Val Ala Leu Ala
305 310 315 320
Ser Ser Gly Ile Asp Pro Ala Arg Leu Tyr Leu Gly Asn Cys Ala Thr
325 330 335
Cys His Gln Met Gln Gly Lys Gly Thr Pro Asp Gly Tyr Tyr Pro Ser
340 345 350
Leu Phe His Asn Ser Thr Val Gly Ala Ser Asn Pro Ser Asn Leu Val
355 360 365
Gln Val Ile Leu Asn Gly Val Gln Arg Lys Ile Gly Ser Glu Asp Ile
370 375 380
Gly Met Pro Ala Phe Arg Tyr Asp Leu Asn Asp Ala Gln Ile Ala Ala
385 390 395 400
Leu Thr Asn Tyr Leu Thr Ala Gln Phe Gly Asn Pro Ala Ala Lys Val
405 410 415
Thr Glu Gln Asp Val Ala Lys Leu Arg
420 425
<210> 9
<211> 425
<212> PRT
<213> 乌汶伯克霍尔德菌(Burkholderia ubonensis)
<400> 9
Met Leu Lys Gln Thr Leu Ser Phe Leu Leu Ala Gly Cys Leu Ala Leu
1 5 10 15
Pro Gly Leu Thr Arg Ala Ala Glu Pro Ala Asp Ala Ala Leu Val Lys
20 25 30
Arg Gly Glu Tyr Leu Ala Val Ala Gly Asp Cys Met Ala Cys His Thr
35 40 45
Thr Lys Gly Gly Lys Pro Phe Ala Gly Gly Leu Gly Met Pro Val Pro
50 55 60
Leu Leu Gly Lys Ile Tyr Thr Ser Asn Ile Thr Pro Asp Ala Asp Thr
65 70 75 80
Gly Ile Gly Asn Trp Ser Tyr Asp Asp Phe Glu Arg Ala Val Arg Arg
85 90 95
Gly Val Ser Lys Asn Gly Asp Asn Leu Tyr Pro Ala Met Pro Tyr Val
100 105 110
Ser Tyr Ala Lys Val Ser Asp Asp Asp Val Lys Ala Leu Tyr Ala Tyr
115 120 125
Phe Met His Gly Val Glu Pro Val Lys Gln Thr Pro Pro Lys Asn Glu
130 135 140
Ile Pro Ala Leu Leu Ser Met Arg Trp Pro Leu Lys Ile Trp Asn Trp
145 150 155 160
Leu Phe Leu Lys Asp Gly Ala Tyr Gln Pro Lys Pro Ser Gln Ser Val
165 170 175
Glu Trp Asn Arg Gly Ala Tyr Leu Val Gln Gly Leu Ala His Cys Ser
180 185 190
Thr Cys His Thr Pro Arg Gly Ile Ala Met Gln Glu Lys Ser Leu Asp
195 200 205
Glu Thr Gly Gly Ser Phe Leu Ser Gly Ser Leu Leu Ala Gly Trp Asp
210 215 220
Gly Tyr Asn Ile Thr Ser Asp Gln Asn Ala Gly Ile Gly Gly Trp Thr
225 230 235 240
Gln Glu Gln Leu Val Gln Tyr Leu Arg Thr Gly Ser Val Pro Gly Val
245 250 255
Ala Gln Ala Ala Gly Pro Met Ala Glu Ala Val Glu His Ser Phe Ser
260 265 270
Lys Met Thr Asp Ala Asp Ile Gly Ala Ile Ala Thr Tyr Ile Arg Thr
275 280 285
Val Pro Ala Val Ala Asp Ser Ser Ala Lys Gln Pro Arg Tyr Thr Trp
290 295 300
Gly Lys Pro Ala Glu Asp Gly Leu Arg Leu Arg Gly Val Ala Leu Ala
305 310 315 320
Ser Ser Gly Ile Asp Pro Ala Arg Leu Tyr Leu Gly Asn Cys Ala Ser
325 330 335
Cys His Gln Met Gln Gly Lys Gly Thr Pro Asp Gly Tyr Tyr Pro Ser
340 345 350
Leu Phe His Asn Ser Thr Val Gly Ala Pro Asn Pro Ser Asn Leu Val
355 360 365
Gln Val Ile Leu Asn Gly Val Gln Arg Lys Ala Gly Ser Glu Asp Val
370 375 380
Gly Met Pro Ala Phe Arg His Glu Leu Ser Asp Ala Gln Ile Ala Ala
385 390 395 400
Leu Ala Asn Tyr Leu Thr Gly Gln Phe Gly Asn Pro Ala Ala Lys Val
405 410 415
Thr Glu Gln Asp Val Ala Lys Ala Arg
420 425
<210> 10
<211> 425
<212> PRT
<213> 礁湖伯克霍尔德菌(Burkholderia stagnalis)
<400> 10
Met Leu Lys Gln Ser Leu Thr Phe Leu Leu Ala Gly Cys Leu Ala Leu
1 5 10 15
Pro Gly Leu Ala Arg Ala Ala Asp Pro Ala Asp Ala Ala Leu Val Lys
20 25 30
Arg Gly Glu Tyr Leu Ala Val Ala Gly Asp Cys Met Ala Cys His Thr
35 40 45
Val Lys Gly Gly Lys Pro Phe Ala Gly Gly Leu Gly Met Pro Val Pro
50 55 60
Met Leu Gly Lys Ile Tyr Thr Ser Asn Ile Thr Pro Asp Ala Asp Thr
65 70 75 80
Gly Ile Gly Gly Trp Thr Tyr Asp Asp Phe Glu Arg Ala Val Arg His
85 90 95
Gly Val Ser Lys Asn Gly Asp Asn Leu Tyr Pro Ala Met Pro Tyr Val
100 105 110
Ser Tyr Ala Lys Val Ser Asp Asp Asp Val Lys Ala Leu Tyr Ala Tyr
115 120 125
Phe Met His Gly Val Glu Pro Val Lys Gln Thr Pro Pro Lys Asn Glu
130 135 140
Ile Pro Ala Leu Leu Ser Met Arg Trp Pro Leu Lys Ile Trp Asn Trp
145 150 155 160
Leu Phe Leu Lys Asp Gly Ala Tyr Gln Pro Lys Pro Ala Gln Ser Ala
165 170 175
Glu Trp Asn Arg Gly Ala Tyr Leu Val Gln Gly Leu Ala His Cys Ser
180 185 190
Thr Cys His Thr Pro Arg Gly Ile Ala Met Gln Glu Lys Ser Leu Asp
195 200 205
Glu Thr Gly Gly Ser Phe Leu Ser Gly Ser Val Leu Ala Gly Trp Asp
210 215 220
Gly Tyr Asn Ile Thr Ser Asp Ala Asn Ala Gly Ile Gly Gly Trp Ser
225 230 235 240
Gln Gln Gln Leu Val Gln Tyr Leu Arg Thr Gly Ser Val Pro Gly Val
245 250 255
Ala Gln Ala Ala Gly Pro Met Ala Glu Ala Val Glu His Ser Phe Ser
260 265 270
Lys Met Thr Asp Ala Asp Ile Gly Ala Ile Ala Thr Tyr Ile Arg Thr
275 280 285
Val Pro Ala Val Ala Asp Ser Ser Val Lys Gln Pro Arg Ala Thr Trp
290 295 300
Gly Lys Pro Ala Glu Asp Gly Leu Arg Leu Arg Gly Val Ala Leu Ala
305 310 315 320
Ser Ser Gly Ile Asp Pro Ala Arg Leu Tyr Leu Gly Asn Cys Ala Ser
325 330 335
Cys His Gln Met Gln Gly Lys Gly Thr Pro Asp Gly Tyr Tyr Pro Ser
340 345 350
Leu Phe His Asn Ser Thr Val Gly Ala Pro Asn Pro Thr Asn Leu Val
355 360 365
Gln Val Ile Leu Asn Gly Val Gln Arg Lys Ala Gly Ser Glu Asp Val
370 375 380
Gly Met Pro Ala Phe Arg His Glu Leu Ser Asp Ala Gln Ile Ala Ala
385 390 395 400
Leu Ala Asn Tyr Leu Thr Gly Gln Phe Gly Asn Pro Ala Ala Lys Val
405 410 415
Thr Glu Gln Asp Val Ala Lys Leu Arg
420 425
<210> 11
<211> 447
<212> PRT
<213> 泰国伯克霍尔德菌(Burkholderia thailandensis)
<400> 11
Met Leu Lys Arg Thr Leu Ser Phe Val Leu Ala Gly Cys Leu Ala Leu
1 5 10 15
Pro Gly Leu Ser Val Ala Ala Ser Ser Ala Pro Val Ala Ala Ala Thr
20 25 30
Ser Ala Ala Ala Ala Thr Ser Ala Ser Ser Ala Pro Ser Ala Asp Ala
35 40 45
Ala Leu Val Glu Arg Gly Arg Tyr Leu Ala Val Ala Gly Asp Cys Met
50 55 60
Ala Cys His Thr Ala Lys Gly Gly Lys Pro Phe Ala Gly Gly Leu Pro
65 70 75 80
Met Arg Ala Pro Leu Leu Gly Thr Ile Tyr Thr Thr Asn Ile Thr Pro
85 90 95
Asp Lys Glu Thr Gly Ile Gly Asp Trp Thr Phe Ala Asp Phe Glu Arg
100 105 110
Ala Val Arg His Gly Val Ala Lys Asn Gly Asp Asn Leu Tyr Pro Ala
115 120 125
Met Pro Tyr Val Ser Tyr Ala Lys Val Thr Asp Asp Asp Val Lys Ala
130 135 140
Leu Tyr Ala Tyr Phe Thr His Gly Val Glu Pro Val Arg Gln Pro Pro
145 150 155 160
Arg Lys Asn Asp Ile Pro Trp Tyr Leu Ser Met Arg Trp Pro Leu Lys
165 170 175
Ile Trp Asn Leu Leu Phe Leu Lys Glu Gly Val Tyr Gln Pro Lys Pro
180 185 190
Glu Arg Ser Val Glu Trp Asn Arg Gly Ala Tyr Leu Val Gln Gly Leu
195 200 205
Ala His Cys Gly Thr Cys His Thr Pro Arg Ala Val Thr Leu Gln Glu
210 215 220
Lys Ser Leu Asp Glu Thr Gly Gly Ser Phe Leu Ala Gly Ser Val Leu
225 230 235 240
Ser Gly Trp Asp Gly Tyr Asn Ile Thr Ser Asp Thr Asn Ala Gly Ile
245 250 255
Gly Gly Trp Ser Gln Gln Gln Leu Val Gln Tyr Leu Arg Thr Gly Ser
260 265 270
Val Pro Gly Leu Ala Gln Ala Ala Gly Pro Met Ala Glu Ala Val Glu
275 280 285
His Ser Phe Ser Gln Met Thr Asp Ala Asp Ile Gly Ala Ile Ala Thr
290 295 300
Tyr Ile Arg Thr Val Pro Ala Val Ala Asp Gly Thr Ala Lys Ala Arg
305 310 315 320
Ser Ala Trp Gly Lys Pro Ala Glu Asp Gly Ile Arg Leu Arg Gly Val
325 330 335
Ala Leu Ala Ala Thr Gly Ile Asp Pro Ala Arg Leu Tyr Leu Gly Asn
340 345 350
Cys Ala Ser Cys His Gln Met Gln Gly Lys Gly Thr Pro Asp Gly Tyr
355 360 365
Tyr Pro Pro Leu Leu His Asn Ser Thr Val Gly Ala Pro Asn Pro Thr
370 375 380
Asn Leu Val Gln Val Ile Leu Asn Gly Val Ala Arg Lys Ala Gly Gly
385 390 395 400
Glu Asp Val Gly Met Pro Ala Phe Arg Arg Glu Leu Ser Asp Ala Gln
405 410 415
Ile Ala Ala Leu Ala Asn Tyr Leu Thr Ala Gln Phe Gly Asn Pro Ala
420 425 430
Ala Lys Val Ser Glu Gln Asp Val Ala Lys Leu Arg Ala Ala Gln
435 440 445
<210> 12
<211> 431
<212> PRT
<213> 青枯雷尔氏菌(Ralstonia solanacearum)
<400> 12
Met Ser Lys Ala Thr Ala Gln Ala Ala Ser Phe Leu Leu Leu Ser Leu
1 5 10 15
Leu Gly Ala Ala Ala Val Ala Arg Gly Ala Asp Pro Pro Ala Asp Thr
20 25 30
Ala Gln Ile Arg Arg Gly Glu Tyr Leu Ala Ile Ala Ala Asp Cys Val
35 40 45
Ala Cys His Thr Ala Pro Gly Gly Lys Pro Phe Ala Gly Gly Leu Pro
50 55 60
Leu Gln Ile Pro Met Leu Gly Thr Ile Tyr Ser Ser Asn Ile Thr Pro
65 70 75 80
Asp Pro Asn Thr Gly Ile Gly Lys Trp Arg Tyr Glu Asp Phe Glu Arg
85 90 95
Ala Val Arg Gln Gly Val Ser Asp Asp Gly Glu Asn Leu Tyr Pro Ala
100 105 110
Met Pro Tyr Pro Ser Tyr Ala Lys Ile Asn Asp Ala Asp Met Arg Asp
115 120 125
Leu Tyr Ala Tyr Phe Arg Tyr Gly Val Ala Ala Val Glu Ser Asp Pro
130 135 140
Pro Pro Ser Thr Ile Arg Trp Pro Leu Asn Met Arg Trp Pro Leu Lys
145 150 155 160
Leu Trp Asn Leu Ala Phe Leu Lys Ser Gly Pro Tyr Ile Pro Lys Gly
165 170 175
Val Lys Thr Pro Thr Trp Asn Arg Gly Ala Tyr Leu Val Gln Gly Leu
180 185 190
Ala His Cys Gly Thr Cys His Thr Pro Arg Gly Phe Ala Met Gln Glu
195 200 205
Lys Ala Met Asp Glu Arg Gly Gln Gly Tyr Leu Ala Gly Ser Thr Leu
210 215 220
Ala Gly Trp Ser Ala Tyr Asn Ile Thr Ser Asp Pro Ala Ser Gly Ile
225 230 235 240
Gly Ser Trp Lys Pro Glu Gln Ile Val Gln Tyr Leu Arg Thr Gly Ser
245 250 255
Val Pro Gly Val Gly Gln Ala Ala Gly Pro Met Gly Glu Ala Val Gln
260 265 270
His Ser Phe Ser Arg Met Thr Glu Thr Asp Val Arg Ala Ile Ala Glu
275 280 285
Tyr Leu Arg Thr Val Pro Ala Val Gly Thr Gly Ala Glu Arg Ala Arg
290 295 300
His Asp Trp Gly Lys Pro Ala Thr Asp Val Thr Ala Leu Arg Gly Lys
305 310 315 320
Pro Ile Glu Thr Thr Ile Asp Ala Ala Arg Leu Tyr Leu Gly Asn Cys
325 330 335
Ala Thr Cys His Gln Ala Asp Gly Arg Gly Thr Pro Asp Gly Tyr Tyr
340 345 350
Pro Pro Leu Leu His Asn Ser Thr Val Gly Ala Arg Asp Thr Arg Asn
355 360 365
Leu Val Gln Val Met Leu Asn Gly Ile Glu Arg Lys Ala Gly Asp Arg
370 375 380
His Ile Gly Met Pro Ala Phe Gly Arg Gln Leu Ser Asp Ala Gln Leu
385 390 395 400
Ala Ala Leu Ala Asn Tyr Val Thr Lys Gln Phe Gly Asp Pro Ala Thr
405 410 415
Pro Ala Leu Thr Ala Glu Ala Ile Ala Lys Arg Arg Leu Pro Gln
420 425 430
<210> 13
<211> 427
<212> PRT
<213> 皮氏雷尔氏菌(Ralstonia pickettii)
<400> 13
Met Lys Ala Ile Ala Ser Phe Ser Leu Leu Leu Leu Ala Gly Pro Leu
1 5 10 15
Ala Asn Ala Trp Ala Lys Asp Thr Pro Ala Asp Thr Ala Gln Ile Lys
20 25 30
Arg Gly Glu Tyr Leu Ala Ile Ala Ala Asp Cys Ala Ala Cys His Thr
35 40 45
Ala Pro Gly Gly Lys Pro Phe Ala Gly Gly Leu Pro Met Gln Ile Pro
50 55 60
Met Leu Gly Thr Ile Tyr Ser Ser Asn Ile Thr Pro Asp Glu Lys Thr
65 70 75 80
Gly Ile Gly Lys Trp Thr Tyr Glu Glu Phe Glu Arg Ala Val Arg Lys
85 90 95
Gly Val Asp Asp Asp Gly Asn Asn Leu Tyr Pro Ala Met Pro Tyr Pro
100 105 110
Ser Tyr Ala Lys Ile Asn Asp Ala Asp Met Arg Asp Leu Tyr Ala Tyr
115 120 125
Phe Arg Asn Gly Val Thr Ala Val Glu Asn Arg Arg Pro Arg Ser Thr
130 135 140
Ile Arg Trp Pro Leu Asn Met Arg Trp Pro Leu Lys Leu Trp Asn Leu
145 150 155 160
Leu Phe Leu Lys Ser Glu Pro Tyr Thr Pro Lys Ser Asp Lys Ser Pro
165 170 175
Thr Trp Asn Arg Gly Ala Tyr Leu Thr Gln Gly Leu Ala His Cys Gly
180 185 190
Thr Cys His Thr Pro Arg Gly Phe Ala Met Gln Glu Lys Ala Leu Asp
195 200 205
Glu Arg Gly Ser Gly Tyr Leu Ala Gly Ser Thr Leu Ser Gly Trp Thr
210 215 220
Ala Phe Asn Ile Thr Ser Asp Pro Thr Ser Gly Ile Gly Ala Trp Lys
225 230 235 240
Pro Glu Gln Leu Val Gln Tyr Leu Arg Thr Gly Ser Val Pro Gly Val
245 250 255
Ala Gln Ala Ala Gly Pro Met Gly Glu Ala Val Gln His Ser Phe Ser
260 265 270
Lys Met Thr Met Gln Asp Val Gln Ala Ile Ala Glu Tyr Leu Arg Thr
275 280 285
Val Pro Ala Val Ser Asn Asn Leu Glu Arg Ala Arg Gln Asp Trp Gly
290 295 300
Lys Pro Ala Thr Asp Val Thr Met Leu Arg Gly Lys Pro Ile Glu Thr
305 310 315 320
Thr Ile Asp Ala Ala Arg Leu Tyr Leu Gly Asn Cys Ala Ser Cys His
325 330 335
Gln Ala Asp Gly Arg Gly Thr Pro Asp Gly Tyr Tyr Pro Pro Leu Leu
340 345 350
His Asn Ser Thr Val Gly Thr Arg Asp Thr Thr Asn Leu Val Gln Val
355 360 365
Ile Leu Phe Gly Ile Glu Arg Lys Ala Gly Asp Lys His Ile Gly Met
370 375 380
Pro Ala Phe Ala Arg Gln Leu Ser Asp Glu Gln Leu Ala Ala Leu Thr
385 390 395 400
Asn Tyr Val Thr Arg Gln Phe Gly Asp Pro Ala Thr Pro Glu Val Thr
405 410 415
Ala Glu Ala Ile Ala Lys Arg Arg Ser Ala Lys
420 425
<210> 14
<211> 31
<212> DNA
<213> 人工序列(Artificial Sequence)
<220>
<223> 引物(primer)
<400> 14
acgtagccat ggcacacaac gacaaccact c 31
<210> 15
<211> 27
<212> DNA
<213> 人工序列(Artificial Sequence)
<220>
<223> 引物(primer)
<400> 15
atcggccgcg cgcgcgaagc ccggcaa 27
<210> 16
<211> 48
<212> DNA
<213> 人工序列(Artificial Sequence)
<220>
<223> 引物(primer)
<400> 16
ttgccgggct tcgcgcgcgc ggccgatctg cgcggtgtcg cgctcgcg 48
<210> 17
<211> 48
<212> DNA
<213> 人工序列(Artificial Sequence)
<220>
<223> 引物(primer)
<400> 17
ttgccgggct tcgcgcgcgc ggccgattat ctcggcaact gcgcgacg 48
<210> 18
<211> 42
<212> DNA
<213> 人工序列(Artificial Sequence)
<220>
<223> 引物(primer)
<400> 18
gtggtgctcg agtgcggccg cgcgcagctt cgcgacgtcc tg 42
<210> 19
<211> 48
<212> DNA
<213> 人工序列(Artificial Sequence)
<220>
<223> 引物(primer)
<400> 19
gtacgtaagc tttcagtggt ggtggtggtg gtgctcgagt gcggccgc 48
Claims (16)
1.一种变异型细胞色素蛋白，其为在具有3个血红素结合域的细胞色素蛋白中从N末端侧数第1个第1血红素结合域和第2个第2血红素结合域发生了缺失的变异型细胞色素蛋白。
2.如权利要求1所述的变异型细胞色素蛋白，其中，包含所述第1血红素结合域和第2血红素结合域的区域发生了缺失。
3.如权利要求2所述的变异型细胞色素蛋白，其中，包含所述第1血红素结合域和第2血红素结合域的区域相当于序列编号4的第43位～第195位的区域。
4.如权利要求1～3中任一项所述的变异型细胞色素蛋白，其中，细胞色素蛋白为细胞色素C蛋白。
5.如权利要求4所述的变异型细胞色素蛋白，其中，细胞色素C蛋白来源于伯克霍尔德菌(Burkholderia)属微生物。
6.如权利要求5所述的变异型细胞色素蛋白，其中，细胞色素C蛋白来源于(Burkholderia cepacia)。
7.如权利要求1～3中任一项所述的变异型细胞色素蛋白，其中，改变前的细胞色素蛋白与序列编号4具有60％以上的氨基酸序列同一性。
8.如权利要求1～3中任一项所述的变异型细胞色素蛋白，其由序列编号4的氨基酸编号314～425的氨基酸序列、序列编号4的氨基酸编号330～425的氨基酸序列、或在这些氨基酸序列中1个或几个氨基酸发生了置换、缺失、插入或添加的氨基酸序列构成，在该发生了置换、缺失、插入或添加的氨基酸序列中，氨基酸编号334～338的CXXCH基序未被改变。
9.一种DNA，其编码权利要求1～8中任一项所述的变异型细胞色素蛋白。
10.一种重组载体，其包含权利要求9所述的DNA。
11.一种转化细胞，其利用权利要求10所述的重组载体进行了转化。
12.一种氧化还原酶-细胞色素复合体，其包含权利要求1～8中任一项所述的变异型细胞色素蛋白和氧化还原酶的催化亚基蛋白。
13.如权利要求12所述的氧化还原酶-细胞色素复合体，其中，氧化还原酶为葡萄糖脱氢酶。
14.一种生物传感器，其包含权利要求12或13所述的氧化还原酶-细胞色素复合体。
15.一种测定对象物质的测定方法，其特征在于，在权利要求14所述的生物传感器中添加试样，施加电位并测定响应电流，基于响应电流计算出试样中包含的测定对象物质的浓度。
16.如权利要求15所述的测定方法，其中，施加电位对于银/氯化银电极为0～+300mV。
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| US20040023330A1 (en) * | 2000-10-31 | 2004-02-05 | Koji Sode | Novel glucose dehydrogenase and process for producing the dehydrogenase |
| CN102453702A (zh) * | 2010-10-27 | 2012-05-16 | 爱科来株式会社 | 变异葡萄糖脱氢酶 |
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| JP4102138B2 (ja) * | 2002-08-30 | 2008-06-18 | 広司 早出 | グルコース脱水素酵素の製造方法 |
| EP1661516B1 (en) | 2003-09-02 | 2012-08-15 | Koji Sode | Glucose sensor and glucose level measuring apparatus |
| JP5586818B2 (ja) | 2003-09-30 | 2014-09-10 | 有限会社アルティザイム・インターナショナル | グルコース脱水素酵素とシトクロームとの融合蛋白質 |
| ATE455848T1 (de) | 2005-06-20 | 2010-02-15 | Arkray Inc | Mutante glucose-dehydrogenase |
| JP6856344B2 (ja) * | 2015-10-29 | 2021-04-07 | アークレイ株式会社 | 変異グルコース脱水素酵素およびその利用 |
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| Publication number | Priority date | Publication date | Assignee | Title |
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| US20040023330A1 (en) * | 2000-10-31 | 2004-02-05 | Koji Sode | Novel glucose dehydrogenase and process for producing the dehydrogenase |
| CN102453702A (zh) * | 2010-10-27 | 2012-05-16 | 爱科来株式会社 | 变异葡萄糖脱氢酶 |
Non-Patent Citations (1)
| Title |
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| TSUYA T等: "Cloning and functional expression of glucose dehydrogenase complex of Burkholderia cepacia in Escherichia coli", 《JOURNAL OF BIOTECHNOLOGY》 * |
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| US20210155675A1 (en) | 2021-05-27 |
| EP3425054A1 (en) | 2019-01-09 |
| US10927162B2 (en) | 2021-02-23 |
| JP2019013166A (ja) | 2019-01-31 |
| US20190010215A1 (en) | 2019-01-10 |
| EP3425054B1 (en) | 2023-12-20 |
| US11505596B2 (en) | 2022-11-22 |
| CN109206505B (zh) | 2023-11-17 |
| EP3425054C0 (en) | 2023-12-20 |
| JP7339723B2 (ja) | 2023-09-06 |
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