CN101163793A - 分子光电器件及其制造方法 - Google Patents
分子光电器件及其制造方法 Download PDFInfo
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- CN101163793A CN101163793A CNA2006800134334A CN200680013433A CN101163793A CN 101163793 A CN101163793 A CN 101163793A CN A2006800134334 A CNA2006800134334 A CN A2006800134334A CN 200680013433 A CN200680013433 A CN 200680013433A CN 101163793 A CN101163793 A CN 101163793A
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Abstract
本发明公开了分离的修饰多肽,该多肽包含编码光合生物光催化单元的氨基酸序列,该光催化单元能够与固体表面共价连接,并且在连接到固体表面之后仍具有光催化活性。
Description
发明领域和发明背景
本发明涉及光催化单元(photocatalytic unit),更具体地讲,本发明涉及用光催化单元制造的固相支持体。本发明还涉及包括光催化单元的分子光电器件。
纳米科学是一门研究材料中微小粒子的科学,是现代技术中最重要的研究尖端领域之一。从基本观点出发,这些微小粒子令人瞩目,因为它们能够构建具有精确性质的材料和结构。由于这种精确控制材料性质的能力,从而为科技和商业开发带来了新的机遇,纳米颗粒已经应用于或有望应用于诸如微电子学和纳米电子学、纳米流控学(nanofluidics)、涂料油漆和生物工程学等众多领域。
十分明确的是,通过增加器件元件的组装密度,微电子技术、磁记录器件和化学传感器的未来发展将得到实现。传统上,显微器件一直用较大的物体制造,但是因为这些产品越来越小,在微米水平以下,这一工艺变得日渐困难。因此预期将使用相反的方法,基本上,主要是通过纳米尺度的材料,从分子水平起建造显微器件。
目前,所制造的分子电路的清晰度远远超过常规图形化技术(例如电子束蚀刻)。然而实际上,具有亚纳米精度的分子定位是常规的,而且对操作生物复合体例如光合复合体是至关重要的。
绿色植物、蓝细菌和光合细菌依靠分子电子复合体这一镶嵌在其膜内的反应中心捕获和利用太阳光。在产氧植物和蓝细菌中,光子捕获光能并将光能转化为化学能发生在被称为类囊体的特化膜上。类囊体位于高等植物的叶绿体中,或者由蓝细菌的细胞质膜折叠构成。类囊体由垛叠的膜圆盘(称为基粒)和非垛叠的膜圆盘(称为基质)组成。类囊体膜含有两个关键的光合组分,光系统I和光系统II,分别命名为PS I和PS II。光合作用需要PSII和PSI依次工作,用水作为电子源,用CO2作为最终电子受体。
PS I是一种跨膜多亚基蛋白质-叶绿素复合体,它介导矢量的光诱导的电子传递。PS I尺度为纳米大小,产能约58%,量子效率几乎等于1[K.Brettel,Biochim.Biophys.Acta 1997,1318 322-373],这使得反应中心在分子纳米电子学应用中成为有前景的单元。PS I介导光诱导电子从质体蓝素或细胞色素C553传递到铁氧还蛋白上。
细长聚球藻(Synechococus elongatus)和植物叶绿体中PS I的晶体结构分别解析为2.5和4.4[P.Jordan等,Nature 2001,411 909-917;A.Ben Shem等,Nature 2003,426 630-635]。蓝细菌中,复合体由12个多肽组成,其中部分结合96个集光叶绿素和22个β类胡萝卜素分子。电子传递链含有P700、A0、A1、Fx、FA和FB,分别表示叶绿素a二聚体、单体叶绿素a、2个叶绿醌和3个[4Fe-4S]铁硫中心。反应中心核心复合体由异二聚体PsaA和PsaB亚基组成,含有原初电子供体P700,该供体进行光诱导电荷分离,并通过顺序传递体A0、A1和Fx传递电子。最终受体FA和FB位于另一个亚基PsaC上。原初供体P700的氧化还原电位为+0.43V,最终受体FB的氧化还原电位为-0.53V,产生的氧化还原差为-1.0V。电荷分离横跨约5nm的蛋白质高度,表示原初供体(P700)与最终受体(FB)之间中心至中心的距离。蛋白质复合体的三聚体和单体高为9nm,直径分别为21nm和15nm。
因为PS I反应中心为色素-蛋白质复合体,负责光能向化学能的光合转化,所以这些反应中心可以在各种不同的器件中用作电子元件。
这些可能的器件包括但不限于空间成像器件(spatial imagingdevice)、太阳能电池、光计算和逻辑门(optical computing and logicgate)、光电开关、光子A/D转换器和薄膜“柔性”光致电压结构(thinfilm″flexible″photovoltaic structure)。然而,为了将这些PS I反应中心整合到分子器件中,将PS I反应中心固定在基片(substrate)上而不会变性是必不可少的。
在早期的研究工作中,所关注的是将植物PS I[I.Lee等,J.Phys.Chem.B 2000,104 2439-2443;R.Das,Nano Letters 2004,4 1079-1083]和细菌反应中心[C.Nakamura等,Applied Biochemistry andBiotechnology 2000,84-6 401-408;S.A.Trammell等,Biosensors &Bioelectronics 2004,19 1649-1655]与固体表面非共价连接,以避免自组装单层膜失活。
因此,通过加入六组氨酸尾标,使得经遗传修饰的细菌反应中心[S.A.Trammell等,Biosensors & Bioelectronics 2004,19 1649-1655]和植物PS I[Das,Nano Letters 2004,4 1079-1083]与涂敷聚合物的金属表面非共价结合。连接有组氨酸的细菌反应中心显示,可以在电化学电池的溶液中产生光电流。有组氨酸尾标的PS I则显示是定向的,但未见报道能产生光电流或光电位。另外,Das(同上)所教导的有组氨酸尾标的PS I,为了与固体表面连接,需要用肽表面活性剂固定。Lee等[J.Phys.Chem.B2000]教导用有机分子涂敷金属表面,使PS I非共价地吸附在有机层上。这种情况下,蛋白质呈数个方向。Lee等[Biosensors & Bioelectronics,1996,11-4,375-387]教导铂沉积在光合膜表面,表现为与PS I的受体侧形成直接的电接触,因为这能够催化氢析出。此外,还显示可以在分离的PS I反应中心上镀铂,因为镀铂后,遇光可产生氢。
然而,这些方法都没有教导功能性PS I反应中心与固体表面的共价连接,当然也没有教导共价连接的定向方式。未定向的PS I反应中心将相互抵消,妨碍PS I固定化器件用于光电器件(例如太阳能电池或逻辑门)。
因此,普遍认为有需要而且将极其有利的是合成能够以定向方式与固体表面结合的活性PS I反应中心。
发明概述
本发明一方面提供分离的修饰多肽,该多肽包含光合生物光催化单元的多肽的氨基酸序列,该氨基酸序列能够介导光催化单元与固体表面的共价连接,并且在光催化单元连接到固体表面时仍保持光催化单元的光催化活性。
本发明另一方面提供大量的分离多肽,该多肽包含光合生物光催化单元的多肽的氨基酸序列,该氨基酸序列能够介导光催化单元与固体表面的共价连接,并且在光催化单元连接到固体表面时仍保持光催化单元的光催化活性,还能够以相对于固体表面基本相同的方向定向。
本发明又一方面提供分离的修饰光催化单元,该光催化单元包含修饰多肽,该多肽包含光合生物光催化单元的多肽的氨基酸序列,该氨基酸序列能够介导光催化单元与固体表面的共价连接,并且在光催化单元连接到固体表面时仍保持光催化单元的光催化活性。
本发明再一方面提供一种膜制品,该膜制品包含修饰光催化单元,该光催化单元包含修饰多肽,该修饰多肽包含光合生物光催化单元的多肽的氨基酸序列,该氨基酸序列能够介导光催化单元与固体表面的共价连接,并且在光催化单元连接到固体表面时仍保持光催化单元的光催化活性。
本发明还有一方面提供分离的修饰多肽编码多核苷酸,该修饰多肽包含光合生物光催化单元的多肽的氨基酸序列,该氨基酸序列能够介导光催化单元与固体表面的共价连接,并且在光催化单元连接到固体表面时仍保持光催化单元的光催化活性。
本发明还再有一方面提供一种核酸构建体,该核酸构建体包含修饰多肽编码多核苷酸,该修饰多肽包含光合生物光催化单元的多肽的氨基酸序列,该氨基酸序列能够介导光催化单元与固体表面的共价连接,并且在光催化单元连接到固体表面时仍保持光催化单元的光催化活性。
本发明另再有一方面提供一种细胞,该细胞包含分离的修饰多肽编码多核苷酸,该修饰多肽包含光合生物光催化单元的多肽的氨基酸序列,该氨基酸序列能够介导光催化单元与固体表面的共价连接,并且在光催化单元连接到固体表面时仍保持光催化单元的光催化活性。
本发明又一方面提供一种器件,该器件包括连接有大量的修饰光催化单元的固体表面,该光催化单元包含修饰多肽,该修饰多肽包含光合生物光催化单元的多肽的氨基酸序列,该氨基酸序列能够介导光催化单元与固体表面的共价连接,并且在光催化单元连接到固体表面时仍保持光催化单元的光催化活性。
根据本发明下述优选的实施方案中的另一个特征,光合生物为绿色植物。
根据本发明所述优选的实施方案中的又一个特征,光合生物为蓝细菌。
根据本发明所述优选的实施方案中的又一个特征,光催化单元为PS-I。
根据本发明所述优选的实施方案中的又一个特征,光催化单元为集胞藻(Synechosystis sp.)PCC 6803光催化单元。
根据本发明所述优选的实施方案中的又一个特征,光催化单元的多肽的氨基酸序列包含至少一个置换突变。
根据本发明所述优选的实施方案中的又一个特征,置换突变是在光催化单元的膜外环上。
根据本发明所述优选的实施方案中的又一个特征,多肽的氨基酸序列为Psa B。
根据本发明所述优选的实施方案中的又一个特征,多肽的氨基酸序列为Psa C。
根据本发明所述优选的实施方案中的又一个特征,psa B在至少一个用以下坐标标明的位置上包含置换突变:D235C、S246C、D479C、S499C、S599C、Y634C。
根据本发明所述优选的实施方案中的又一个特征,psa C在至少一个用坐标W31C标明的位置上包含置换突变。
根据本发明所述优选的实施方案中的又一个特征,至少一个置换突变为半胱氨酸。
根据本发明所述优选的实施方案中的又一个特征,该氨基酸序列如SEQ ID NO:20、21、22、23、24、25、26、27、28和29中所示。
根据本发明所述优选的实施方案中的又一个特征,固体表面为导电材料。
根据本发明所述优选的实施方案中的又一个特征,导电材料为过渡金属元素。
根据本发明所述优选的实施方案中的又一个特征,过渡金属元素选自银、金、铜、铂、镍和钯。
根据本发明所述优选的实施方案中的又一个特征,分离的修饰多肽不包含金属离子。
根据本发明所述优选的实施方案中的又一个特征,分离的修饰多肽为单体形式或三聚体形式。
根据本发明所述优选的实施方案中的又一个特征,核酸构建体还包含顺式调节元件。
根据本发明所述优选的实施方案中的又一个特征,顺式调节元件为启动子。
根据本发明所述优选的实施方案中的又一个特征,细胞为集胞藻(Synechocystis)细胞。
根据本发明所述优选的实施方案中的又一个特征,大量的修饰光催化单元每个之间的距离介于15-25nm之间。
根据本发明所述优选的实施方案中的又一个特征,大量的修饰光催化单元相对于固体表面是定向的。
根据本发明所述优选的实施方案中的又一个特征,该器件用作光电二级管的元件。
根据本发明所述优选的实施方案中的又一个特征,该器件用作光电晶体管的元件。
根据本发明所述优选的实施方案中的又一个特征,该器件用作逻辑门的元件。
根据本发明所述优选的实施方案中的又一个特征,该器件用作太阳能电池的元件。
根据本发明所述优选的实施方案中的又一个特征,该器件用作光耦合器的元件。
根据本发明所述优选的实施方案中的又一个特征,大量的修饰光催化单元与固体表面直接连接。
根据本发明所述优选的实施方案中的又一个特征,直接连接为共价连接。
本发明通过提供____成功克服了现有已知配置的不足。
除非另有规定,否则本文所用的所有科技术语都具有本发明所属领域普通技术人员一般掌握的含义。下面将介绍合适的方法与材料,尽管类似于或等同于本文所述的方法与材料也可以用于本发明的实施或测试中。所有的出版物、专利申请书、专利和其它本文引用的参考文献都通过引用全部结合到本文中。万一发生抵触,则以本专利说明书(包括定义)为准。另外,所述材料、方法和实施例仅是示例性的,而不是限制性的。
附图简述
本文仅通过实施例并参考附图描述本发明。现在具体地参考附图的细节,须强调的是,所显示的详细资料仅仅作为实例,为了说明性地论述本发明的优选实施方案,并且提供这些详细资料是为了提供所认为的对本发明的原理和构思方面最有用最容易理解的描述。在这方面,并没有试图以比基本理解本发明所必需的更为详细的方式来说明本发明的结构细节,从附图获取的说明,使得在实践中如何使本发明的所述几种形式具体化对于本领域技术人员而言是显而易见的。
附图中:
图1A-E表示PS I的半胱氨酸突变,提供了半胱氨酸突变是在PSI外面的证据。图1A-C是在集胞藻PCC 6803的psaB中通过同源重组引入突变时所用载体的示意图。将1.8kb的psaB基因片段、卡那霉素抗性赋予基因(kanamycin resistance conferring gene(KanR))和1.1kb下游侧翼区(downstream flanking region)插入pBluescript载体中构建了质粒pZBL(图1B)。为了筛选psaB缺陷型受体细胞,通过除去1.3kb的psaB下游端并插入氯霉素抗性基因(CmR)构建了pBLΔB载体(图1C)。基因组DNA的限制位点如图1A所示。图1D为在“实心”处带有所提出的突变的PS I结构的骨架图象。箭头表示光诱导电荷传递的方向。PsaB亚基上突变成为半胱氨酸的氨基酸从左到右如下排列:D235C、S246C、D479C、S499C、S599C、Y634C。这些坐标得自PDB文件JBO1,借助RasMole软件显示。图1E为分离的野生型PS I复合体(第1泳道)和经遗传修饰的PS I复合体(第2-7泳道)外露表面半胱氨酸的免疫印迹照片。
图2A-B是定向PS I反应中心在金表面的二维空间阵列图象。图象用轻敲模式原子力显微镜(tapping-mode atomic force microscopy(AFM;面积0.3μm2))获得。图2A是硅片(silicon slide)上退火的150nm金表面图象,图2B是在含有D479C突变多肽(SEQ ID NO:20)的PS I单体溶液中温育的金基片图象。
图3A-B是金表面定向PS I反应中心的二维空间图和电位图。Au-Si表面得自突变体D479C(SEQ ID NO:20)的同一批PS I反应中心三聚体的形貌图象(图3A)和电位图象(图3B)。PS I的光诱导PS I负电位见图3B。用He-Ne激光(632.8nm,5mW/cm2)提供照明,并在图象中标出。图3B所示电位的负号是由于KPFM反馈回路所致,与CPD的实际符号相反。
图4A-B是金表面定向PSI反应中心的二维空间图和电位图。Au-Si表面得自突变体D479C的同一批PS I反应中心三聚体的形貌图象(图4A)和电位图象(图4B)。PS I的光诱导PS I的负电位见图4B。示意剖面图的各光栅的扫描方向自上而下(由光至暗)。用He-Ne激光(632.8nm,5mW/cm2)提供照明。图4B所示的电位负号是由于KPFM反馈回路所致,与CPD的实际符号相反。
图5A-B是金表面定向PS I反应中心的三维形貌图象和电位图象。图5A表示Au-Si表面PS I反应中心单体的电位图象。打开He-Ne激光照明(632.8nm,5mW/cm2(hv))时,观察到PS I的光诱导PS I负电位。示意剖面图的各光栅的扫描方向自上而下(由暗至光)。图5B表示Au-Si基片上PS I三聚体的三维形貌图。
图6A-B为PS I的光谱测定。图6A表示在分离的PS I突变体(SEQID NO:20)中闪光诱导的P700的瞬时吸收变化。使激光闪光普照D479C突变PS I复合体后,在820nm(ΔA820)下监测P700的吸收变化。图6B表示定向PS I复合体的X射线吸收光谱。自组装单层膜的PS I取向用掠射X射线荧光全反射测定法(total reflectionmeasurement of grazing x-ray fluorescence)测定。PS I通过唯一的半胱氨酸与在硅基片之上钨-碳多层中钨之间形成硫键连接。各图是按与X射线束常态25(--)、45(Δ)、60(-)和90(○)度的规定角度,60秒、42秒扫描的平均值。
图7是本发明不同示例性实施方案的光电器件的示意图。
图8是本发明不同示例性实施方案的光电二级管器件的示意图。
图9是本发明不同示例性实施方案的光电晶体管30的示意图。
图10是本发明不同示例性实施方案的光耦合器40的简图。
优选实施方案的描述
本发明涉及可以与固相支持体共价连接并保持活性的修饰光催化单元。具体地讲,本发明可以在各种不同的器件中用作电子元件。
在详细说明本发明至少一个实施方案之前,应该了解本发明在其应用以下说明书或实施例中所述详细说明时并无限制。本发明可能有其它实施方案,或者能够以各种方式实施或实现。同样,应当理解本文所用措辞和术语是为了说明目的,不应视为限制。
光合作用是通过光反应和暗反应将电磁能转化为化学能的生物过程。在产氧植物和蓝细菌中,光子捕获以及光能转化为化学能发生在被称为类囊体的特化膜上。类囊体位于高等植物的叶绿体中,或者由蓝细菌的细胞质膜折叠构成。
PS I是一种跨膜多亚基蛋白质-叶绿素复合体,它介导矢量的光诱导电子从质体蓝素或细胞色C553传递到铁氧还蛋白。PS I尺度为纳米大小,产能约58%,量子效率高,这使得反应中心在分子纳米电子学应用中成为有前景的单元。然而,为了将PS I反应中心整合到分子器件上,将PS I反应中心固定在基片上而不变性是必不可少的。另外,PS I反应中心的定向连接是绝对必要的,这样诱导的电荷才不会相互抵消。
在本申请发明人将本发明付诸实施时,发现了光催化单元的多肽可以进行遗传修饰,使之包含能与固体表面共价结合而仍然保持活性的官能团。
如下面的实施例部分所述,本发明的发明人为了确保形成硫键(PS I单元和金属表面之间),利用了可公开获取的PS I 3D结构的结构数据,使细菌膜面向细胞质一侧的膜外环中PS I的Psa B多肽和PsaC多肽的氨基酸突变成为半胱氨酸。为了促进有效的电连接,在接近P700处选出不同的突变,以保证反应中心和金电极之间紧密相邻(实施例1)。本发明的发明人还指出通过在大量的光催化单元中用半胱氨酸置换相同的氨基酸,光催化单元与固相支持体的连接将会是定向的,PS I单元可在固相支持体上形式单层膜(实施例2)。选择特定的氨基酸用半胱氨酸残基置换,可以调节光催化单元在固相支持体上的精确取向。
按上述方法修饰的PS I单元,能够在金表面形成定向单层膜,如原子力显微镜观测的一样,见图2A-B。可通过以下试验证实突变PS I单元与金表面连接后具有功能活性:通过其产生明显的光诱导电位的能力,如用开尔文探针力显微镜(Kelvin probe force microscopy,KPFM)所测[图3A-B],以及通过其传递电子的能力,如用单次循环光谱(single turnover spectroscopy)所测[图6A-B]。
Trammel等[Biosensors & Bioelectronics 2004,19 1649-1655]教导对包含六组氨酸尾标的细菌反应中心的修饰。与本发明相比截然不同的是,突变后的PS I不能与金属表面共价结合。
Das[Nano Letters 2004,41079-1083]教导对包含六组氨酸尾标的植物反应中心的修饰。与本发明相比截然不同的是,突变后的PS I也不能与金属表面共价结合,为了连接在固体表面,突变后的PS I需要用肽表面活性剂固定。
Lee等[J.Phys.Chem.B2000]教导用有机分子涂敷金属表面,并使PS I非共价吸附到有机层上。与本发明相比,PS I蛋白呈数种取向,且同样无功能活性。
Lee等[Biosensors & Bioelectronics,1996,11-4,375-387]教导将铂沉积在光合膜表面,因此与PS I的受体侧产生直接电接触,从中可以催化氢析出。这些研究都未证实光电流通过外电路的电势能。同Trammel等人和Das一样,Lee等人也未教导细菌反应中心和PS I分别与固体表面的共价连接。与Trammell等人和Das相比截然不同的是,本发明PS I修饰蛋白不需要进行修饰以包含金属离子,因为它们通过引入位于PS I膜外环的半胱氨酰残基与金属表面结合。
因此,本发明一方面提供分离的修饰多肽,该多肽包含光合生物光催化单元的多肽的氨基酸序列,该氨基酸序列能够介导光催化单元与固体表面的共价连接,并且在光催化单元连接到固体表面时仍保持光催化单元的光催化活性。
本文所用术语“光催化单元(photocatalytic unit)”是指至少一个多肽和其它小分子(例如叶绿素和色素分子)的复合体,当整合在一起时,作为功能单元使光能转化为化学能。如上所述,本发明的光催化单元存在于光合生物(即将光能转化为化学能的生物)中。光合生物的实例包括但不限于绿色植物、蓝细菌、红藻、紫色细菌和绿色细菌。
因此,根据本发明这一方面,可以使用的光催化单元的实例包括生物光催化单元,例如PS I和PS II、细菌光敏感蛋白、细菌光敏感蛋白、细菌视紫红质、光催化微生物、色素(例如原黄素和视紫红质)、有机染料和藻类。优选本发明的光催化单元为光系统I(PS I)。
PS I为蛋白质-叶绿素复合体,存在于绿色植物和蓝细菌中,是类囊体膜内光合机器的部件,呈椭圆形,尺寸约9×15纳米。PS I复合体通常包含叶绿素分子,叶绿素分子作为天线发挥作用,它吸收光子并将光子能量传递给P700,这种能量在P700上被捕获并被利用以启动光化学反应。除P700和天线叶绿素外,PSI复合体还包含许多电子受体。从P700所释放的电子通过中间受体被传递到PSI还原端的末端受体上,而且电子是跨类囊体膜传递的。
PS I多肽的实例及其源生物见下表1。
表1
源生物 | 蛋白质检索号 |
强壮前沟藻(Amphidinium carterae) | CAC34545 |
中国圆柏(Juniperus chinensis) | CAC87929 |
黎巴嫩雪松(Cedrus libani) | CAC87143 |
白鹤芋(Spathiphyllum sp.)SM328 | CAC87924 |
鳄梨(Persea americana) | CAC87920 |
美洲苏铁(Zamia pumila) | CAC87935 |
钝头瓶尔小草(Ophioglossum petiolatum) | CAC87936 |
短叶红豆杉(Taxus brevifolia) | CAC87934 |
Afrocarpus gracilior | CAC87933 |
日本五针松(Pinus parviflora) | CAC87932 |
西藏云杉(Picea spinulosa) | CAC87931 |
瓶蕨叶状枝木(Phyllocladus trichomanoides) | CAC87930 |
锯叶棕榈(Serenoa repens) | CAC87923 |
沼泽三白草(Saururus cernuus) | CAC87922 |
加州悬铃木(Platanus racemosa) | CAC87921 |
顶花板凳果(Pachysandra terminalis) | CAC87919 |
睡莲(Nymphaea sp.cv.Paul Harriot) | CAC87918 |
欧亚萍蓬草(Nuphar lutea) | CAC87917 |
莲(Nelumbo nucifera) | CAC87916 |
小鸡爪槭(Acer palmatum) | CAD23045 |
绿干柏(Cupressus arizonica) | CAC87928 |
日本柳杉(Cryptomeria japonica) | CAC87927 |
欧洲白冷杉(Abies alba) | CAC87926 |
显轴买麻藤(Gnetum gnemon) | CAC87925 |
荷花玉兰(Magnolia grandiflora) | CAC87915 |
胶皮枫香树(Liquidambar styraciflua) | CAC87914 |
野百合(Lilium brownii) | CAC87913 |
Isomeris arborea | CAC87912 |
北美水青冈(Fagus grandifolia) | CAC87911 |
Eupomatia laurina | CAC87910 |
灯笼花(Enkianthus chinensis) | CAC87909 |
黄莲(Coptis laciniata) | CAC87908 |
金粟兰(Chloranthus spicatus) | CAC87907 |
加州夏腊梅(Calycanthus occidentalis) | CAC87906 |
Austrobaileya scandens | CAC87905 |
Amborella trichopoda | CAC87904 |
菖蒲(Acorus calamus) | CAC87142 |
本发明该方面一个优选的实施方案中,PS I来源于蓝细菌,更准确地讲,PS I来源于集胞藻PCC 6803。
蓝细菌中,PS I复合体由12个多肽组成,其中部分多肽结合了96个集光叶绿素分子和22个β类胡萝卜素分子。电子传递链含有P700、A0、A1、Fx、FA和FB,分别表示叶绿素a二聚体、单体叶绿素a、2个叶绿醌和3个[4Fe-4S]铁硫中心。反应中心核心复合体由异二聚体PsaA和PsaB亚基组成,含有原初电子供体P700,该复合体进行光诱导电荷分离,并通过顺序传递体A0、A1和Fx传递电子。最终受体FA和FB位于另一亚基PsaC上。
来源于蓝细菌的PS I在结构上比来源于植物和细菌反应中心的PS I更加稳定。这是因为所有的叶绿素分子和类胡萝卜素都整合在蓝细菌的核心亚基复合体中,而在植物和其它细菌反应中心内,天线叶绿素与连接到核心亚基的叶绿素-蛋白质复合体结合。因此,与来源于其它生物(例如植物和其它细菌)的PS I不同,来源于蓝细菌的PSI在连接到固体表面时,其固定不需要肽表面活性剂[R.Das等,NanoLetters 2004,4 1079-1083]。
本文所用术语“分离的”是指从其合成的天然部位(例如光合生物)至少部分取出的修饰光催化多肽。通常,该光催化多肽未与光催化单元的其它组分(即叶绿素和色素)分开,因而光催化单元保持其功能。优选多肽基本上不含存在于其体内位置的其它物质(例如其它细胞、蛋白质、核酸等)。
如上所述,本发明这一方面的光催化单元包含修饰多肽。
本文所用术语“多肽”是指可以通过重组DNA技术合成的多肽。
在本说明书和所附权利要求书部分所用的术语“氨基酸”应理解为包括:20种天然存在的氨基酸;常常在体内翻译后经修饰的氨基酸,包括例如羟脯氨酸、磷酸丝氨酸和磷酸苏氨酸;其它不常用的氨基酸,包括但不限于2-氨基己二酸、羟赖氨酸、异赖氨素、正缬氨酸、正亮氨酸和鸟氨酸。此外,术语“氨基酸”既包括D-氨基酸又包括L-氨基酸。
下面的表2和表3列举了可用于本发明的天然存在的氨基酸(表2)和不常用的或经修饰的氨基酸(表3)。
表2
氨基酸 | 三字母缩写 | 单字母缩写 |
丙氨酸 | Ala | A |
精氨酸 | Arg | R |
天冬酰胺 | Asn | N |
天冬氨酸 | Asp | D |
半胱氨酸 | Cys | C |
谷氨酰胺 | Gln | Q |
谷氨酸 | Glu | E |
甘氨酸 | Gly | G |
组氨酸 | His | H |
异亮氨酸 | Iie | I |
亮氨酸 | Leu | L |
赖氨酸 | Lys | K |
甲硫氨酸 | Met | M |
苯丙氨酸 | Phe | F |
脯氨酸 | Pro | P |
丝氨酸 | Ser | S |
苏氨酸 | Thr | T |
色氨酸 | Trp | W |
酪氨酸 | Tyr | Y |
缬氨酸 | Val | V |
上述任一种氨基酸 | Xaa | X |
表3
不常用的氨基酸 | 代码 | 不常用的氨基酸 | 代码 |
α-氨基丁酸 | Abu | L-N-甲基丙氨酸 | Nmala |
α-氨基-α-甲基丁酸 | Mgabu | L-N-甲基精氨酸 | Nmarg |
氨基环丙烷-甲酸 | Cpro | L-N-甲基天冬酰胺 | Nmasn |
L-N-甲基天冬氨酸 | Nmasp |
氨基异丁酸 | Aib | L-N-甲基半胱氨酸 | Nmcys |
氨基降冰片烷基-甲酸 | Norb | L-N-甲基谷氨酰胺 | Nmgin |
L-N-甲基谷氨酸 | Nmglu | ||
环己基丙氨酸 | Chexa | L-N-甲基组氨酸 | Nmhis |
环戊基丙氨酸 | Cpen | L-N-甲基异亮氨酸 | Nmile |
D-丙氨酸 | Dal | L-N-甲基亮氨酸 | Nmleu |
D-精氨酸 | Darg | L-N-甲基赖氨酸 | Nmlys |
D-天冬氨酸 | Dasp | L-N-甲基甲硫氨酸 | Nmmet |
D-半胱氨酸 | Dcys | L-N-甲基正亮氨酸 | Nmnle |
D-谷氨酰胺 | Dgln | L-N-甲基正缬氨酸 | Nmnva |
D-谷氨酸 | Dglu | L-N-甲基鸟氨酸 | Nmorn |
D-组氨酸 | Dhis | L-N-甲基苯丙氨酸 | Nmphe |
D-异亮氨酸 | Dile | L-N-甲基脯氨酸 | Nmpro |
D-亮氨酸 | Dleu | L-N-甲基丝氨酸 | Nmser |
D-赖氨酸 | Dlys | L-N-甲基苏氨酸 | Nmthr |
D-甲硫氨酸 | Dmet | L-N-甲基色氨酸 | Nmtrp |
D-鸟氨酸 | Dorn | L-N-甲基酪氨酸 | Nmtyr |
D-苯丙氨酸 | Dphe | L-N-甲基缬氨酸 | Nmval |
D-脯氨酸 | Dpro | L-N-甲基乙基甘氨酸 | Nmetg |
D-丝氨酸 | Dser | L-N-甲基-丁基甘氨酸 | Nmtbug |
D-苏氨酸 | Dthr | L-正亮氨酸 | Nle |
D-色氨酸 | Dtrp | L-正缬氨酸 | Nva |
D-酪氨酸 | Dtyr | α-甲基-氨基异丁酸 | Maib |
D-缬氨酸 | Dval | α-甲基-γ-氨基丁酸 | Mgabu |
D-α-甲基丙氨酸 | Dmala | αethylcyclohexylalanine | Mchexa |
D-α-甲基精氨酸 | Dmarg | α-甲基环戊基丙氨酸 | Mcpen |
D-α-甲基天冬酰胺 | Dmasn | α-甲基-α-萘基丙氨酸 | Manap |
D-α-甲基天冬氨酸 | Dmasp | α-甲基青霉胺 | Mpen |
D-α-甲基半胱氨酸 | Dmcys | N-(4-氨基丁基)甘氨酸 | Nglu |
D-α-甲基谷氨酰胺 | Dmgln | N-(2-氨基乙基)甘氨酸 | Naeg |
D-α-甲基组氨酸 | Dmhis | N-(3-氨基丙基)甘氨酸 | Norn |
D-α-甲基异亮氨酸 | Dmile | N-氨基-α-甲基丁酸 | Nmaabu |
D-α-甲基亮氨酸 | Dmleu | α-萘基丙氨酸 | Anap |
D-α-甲基赖氨酸 | Dmlys | N-苄基甘氨酸 | Nphe |
D-α-甲基甲硫氨酸 | Dmmet | N-(2-氨基甲酰乙基)甘氨酸 | Ngln |
D-α-甲基鸟氨酸 | Dmorn | N-(氨基甲酰甲基)甘氨酸 | Nasn |
D-α-甲基苯丙氨酸 | Dmphe | N-(2-羰基乙基)甘氨酸 | Nglu |
D-α-甲基脯氨酸 | Dmpro | N-(羰基甲基)甘氨酸 | Nasp |
D-α-甲基丝氨酸 | Dmser | N-环丁基甘氨酸 | Ncbut |
D-α-甲基苏氨酸 | Dmthr | N-环庚基甘氨酸 | Nchep |
D-α-甲基色氨酸 | Dmtrp | N-环己基甘氨酸 | Nchex |
D-α-甲基酪氨酸 | Dmty | N-环癸基甘氨酸 | Ncdec |
D-α-甲基缬氨酸 | Dmval | N-环十二烷基甘氨酸 | Ncdod |
D-α-甲基丙氨酸 | Dnmala | N-环辛基甘氨酸 | Ncoct |
D-α-甲基精氨酸 | Dnmarg | N-环丙基甘氨酸 | Ncpro |
D-α-甲基天冬酰胺 | Dnmasn | N-环十一烷基甘氨酸 | Ncund |
D-α-甲基天冬氨酸 | Dnmasp | N-(2,2-二苯基乙基)甘氨酸 | Nbhm |
D-α-甲基半胱氨酸 | Dnmcys | N-(3,3-二苯基丙基)甘氨酸 | Nbhe |
D-N-甲基亮氨酸 | Dnmleu | N-(3-吲哚基乙基)甘氨酸 | Nhtrp |
D-N-甲基赖氨酸 | Dnmlys | N-甲基-γ-氨基丁酸 | Nmgabu |
N-甲基环己基丙氨酸 | Nmchexa | D-N-甲基甲硫氨酸 | Dnmmet |
D-N-甲基鸟氨酸 | Dnmorn | N-甲基环戊基丙氨酸 | Nmcpen |
N-甲基甘氨酸 | Nala | D-N-甲基苯丙氨酸 | Dnmphe |
N-甲基氨基异丁酸 | Nmaib | D-N-甲基脯氨酸 | Dnmpro |
N-(1-甲基丙基)甘氨酸 | Nile | D-N-甲基丝氨酸 | Dnmser |
N-(2-甲基丙基)甘氨酸 | Nile | D-N-甲基丝氨酸 | Dnmser |
N-(2-甲基丙基)甘氨酸 | Nleu | D-N-甲基苏氨酸 | Dnmthr |
D-N-甲基色氨酸 | Dnmtrp | N-(1-甲基乙基)甘氨酸 | Nva |
D-N-甲基酪氨酸 | Dnmtyr | N-甲基-α-萘基丙氨酸 | Nmanap |
D-N-甲基缬氨酸 | Dnmval | N-甲基青霉胺 | Nmpen |
γ-氨基丁酸 | Gabu | N-(对羟基苯基)甘氨酸 | Nhtyr |
L-叔丁基甘氨酸 | Tbug | N-(硫甲基)甘氨酸 | Ncys |
L-乙基甘氨酸 | Etg | 青霉胺 | Pen |
L-高苯丙氨酸 | Hphe | L-α-甲基丙氨酸 | Mala |
L-α-甲基精氨酸 | Marg | L-α-甲基天冬酰胺 | Masn |
L-α-甲基天冬氨酸 | Masp | L-α-甲基-叔丁基甘氨酸 | Mtbug |
L-α-甲基半胱氨酸 | Mcys | L-甲基乙基甘氨酸 | Metg |
L-αthylglutamine | Mgln | L-α-甲基谷氨酸 | Mglu |
L-α-甲基组氨酸 | Mhis | L-α-甲基高苯丙氨酸 | Mhphe |
L-α-甲基异亮氨酸 | Mile | N-(2-甲硫基乙基)甘氨酸 | Nmet |
D-N-甲基谷氨酰胺 | Dnmgln | N-(3-胍基丙基)甘氨酸 | Narg |
D-N-甲基谷氨酸 | Dnmglu | N-(1-羟基乙基)甘氨酸 | Nthr |
D-N-甲基组氨酸 | Dnmhis | N-(羟基乙基)甘氨酸 | Nser |
D-N-甲基异亮氨酸 | Dnmile | N-(咪唑基乙基)甘氨酸 | Nhis |
D-N-甲基亮氨酸 | Dnmleu | N-(3-吲哚基乙基)甘氨酸 | Nhtrp |
D-N-甲基赖氨酸 | Dnmlys | N-甲基-γ-氨基丁酸 | Nmgabu |
N-甲基环己基丙氨酸 | Nmchexa | D-N-甲基甲硫氨酸 | Dnmmet |
D-N-甲基鸟氨酸 | Dnmorn | N-甲基环戊基丙氨酸 | Nmcpen |
N-甲基甘氨酸 | Nala | D-N-甲基苯丙氨酸 | Dnmphe |
N-甲基氨基异丁酸 | Nmaib | D-N-甲基脯氨酸 | Dnmpro |
N-(1-甲基丙基)甘氨酸 | Nile | D-N-甲基丝氨酸 | Dnmser |
N-(2-甲基丙基)甘氨酸 | Nleu | D-N-甲基苏氨酸 | Dnmthr |
D-N-甲基色氨酸 | Dnmtrp | N-(1-甲基乙基)甘氨酸 | Nval |
D-N-甲基酪氨酸 | Dnmtyr | N-甲基-α-萘基丙氨酸 | Nmanap |
D-N-甲基缬氨酸 | Dnmval | N-甲基青霉胺 | Nmpen |
γ-氨基丁酸 | Gabu | N-(对羟基苯基)甘氨酸 | Nhtyr |
L-叔丁基甘氨酸 | Tbug | N-(硫甲基)甘氨酸 | Ncys |
L-乙基甘氨酸 | Etg | 青霉胺 | Pen |
L-高苯丙氨酸 | Hphe | L-α-甲基丙氨酸 | Mala |
L-α-甲基精氨酸 | Marg | L-α-甲基天冬酰胺 | Masn |
L-α-甲基天冬氨酸 | Masp | L-α-甲基-叔丁基甘氨酸 | Mtbug |
L-α-甲基半胱氨酸 | Mcys | L-甲基乙基甘氨酸 | Metg |
L-α-甲基谷氨酰胺 | Mgln | L-α-甲基谷氨酸 | Mglu |
L-αethylhistidine | Mhis | L-α-甲基高苯丙氨酸 | Mhphe |
L-αthylisoleucine | Mile | N-(2-甲硫基乙基)甘氨酸 | Nmet |
L-α-甲基亮氨酸 | Mleu | L-α-甲基赖氨酸 | Mlys |
L-α-甲基甲硫氨酸 | Mmet | L-α-甲基正亮氨酸 | Mnle |
L-α-甲基正缬氨酸 | Mnva | L-α-甲基鸟氨酸 | Morn |
L-α-甲基苯丙氨酸 | Mphe | L-α-甲基脯氨酸 | Mpro |
L-α-甲基丝氨酸 | mser | L-α-甲基苏氨酸 | Mthr |
L-αethylvaline | Mtrp | L-α-甲基酪氨酸 | Mtyr |
L-α-甲基亮氨酸 | Mval | L-N-甲基高苯丙氨酸 | Nmhphe |
N-(N-(2,2-二苯基乙基)氨基甲酰甲基-甘氨酸 | Nnbhm | N-(N-(3,3-二苯基丙基)氨基甲酰甲基(1)甘氨酸 | Nnbhe |
1-羰基-1-(2,2-二苯基乙基氨基)环丙烷 | Nmbc |
本文所用术语“修饰多肽”是指与野生型多肽相比包含修饰的多肽。通常,该修饰为氨基酸修饰。本发明这一方面包括对序列进行任何的修饰,只要多肽能够与固体表面共价连接,并保持光催化活性。修饰的实例包括缺失、插入、置换及其生物活性多肽片段。插入或缺失通常的范围约1-5个氨基酸。
根据所提出的光催化单元的3D结构,选择修饰位点。有关光催化单元3D结构的证据得自X射线晶体学研究或运用蛋白质建模软件。细长聚球藻和植物叶绿体PS I的晶体结构已分别解析至2.5和4.4[P.Jordan等,Nature 2001,411 909-917;A.Ben Shem,F.Frolow,N.Nelson,Nature 2003,426 630-635]。
所要替换的氨基酸或插入位点通常在光催化单元的外面(例如在膜外环上)。优选需要替换的氨基酸或插入位点在不引起位阻的位置上。还优选的是,突变位于光催化单元的P700附近,以确保反应中心和固体表面之间紧密相邻以促进有效的电连接。
本发明该方面一个优选的实施方案中,修饰为置换(即替换),包含能够介导与固体表面结合的官能团侧链。用于集胞藻PCC 6803PsaB上PS I突变的特别优选的坐标包括单突变D235C、S246C、D479C、S499C、S599C和Y634C或者双重突变D235C/Y634C和S246C/Y634C。PsaC中,特别优选的突变位点为W31C。另外,在光催化单元中可产生三重突变(例如PsaC//PsaB W31C//D235C/Y634C)。
优选被置换的野生型氨基酸对光催化单元的活性不是必需的。为了确定哪些氨基酸在功能上是冗余的,可以通过将光催化单元的序列与已知的同源蛋白质分子的序列相比较,使高同源性区域所进行的氨基酸序列变化的数目减至最小。
在一个实施方案中,保守氨基酸置换在一个或多个预计为非必需氨基酸残基上进行。“保守氨基酸置换”是指其中氨基酸残基用具有相似侧链的氨基酸残基替换。本领域中已定义了具有相似侧链的同族氨基酸残基。这些同族氨基酸包括具有碱性侧链的氨基酸(例如赖氨酸、精氨酸、组氨酸)、具有酸性侧链的氨基酸(例如天冬氨酸、谷氨酸)、具有不带电荷的极性侧链的氨基酸(例如甘氨酸、天冬酰胺、谷氨酰胺、丝氨酸、苏氨酸、酪氨酸、半胱氨酸)、具有非极性侧链的氨基酸(例如丙氨酸、缬氨酸、亮氨酸、异亮氨酸、脯氨酸、苯丙氨酸、甲硫氨酸、色氨酸)、具有β分支侧链的氨基酸(例如苏氨酸、缬氨酸、异亮氨酸)和具有芳族侧链的氨基酸(例如酪氨酸、苯丙氨酸、色氨酸、组氨酸)。
在另一个实施方案中,可进行非保守氨基酸的置换,因为突变优选设计成能够使蛋白质与金属定向地共价连接。通过选择可以光自养生长的突变细胞,确保PS I细胞不受损伤。
优选本文上述坐标的氨基酸用能够与金属表面结合的氨基酸(例如含巯基氨基酸例如半胱氨酸)替换。
在本发明这一方面的一个优选实施方案中,多肽序列如SEQ IDNO:20、21、22、23、24、25、26、27、28和29所示。
优选采用重组技术产生本发明的多肽,因为光催化单元通常包含不止一种多肽和其它分子(例如色素分子和叶绿素),它们一起整合成复合体。另外,这些技术更适于产生相对长的多肽(例如长20个氨基酸以上)和大量的多肽。重组技术记载于Bitter等,(1987)Methods inEnzymol.153:516-544,Studier等,(1990)Methods in Enzymol.185:60-89,Brisson等,(1984)Nature 310:511-514,Takamatsu等,(1987)EMBOJ.6:307-311,Coruzzi等,(1984)EMBO J.3:1671-1680和Brogli等,(1984)Science 224:838-843,Gurley等,(1986)Mol.Cell.Biol.6:559-565和Weissbach & Weissbach,1988,Methods for Plant Molecular Biology,Academic Press,NY,第VIII节,第421-463页。
本发明的多肽用标准技术修饰,例如定点诱变(寡核苷酸介导的诱变)和PCR介导的诱变。因此,可以使编码光催化单元多肽的多核苷酸发生突变。诱变之后,光催化单元可以在合适的细胞系统中表达。
寡核苷酸介导的诱变是一门本领域公知的技术,参见Adelman等,DNA,2:183(1983)。简单地讲,通过使编码所需突变的寡核苷酸与多核苷酸模板杂交,来改变编码光催化单元多肽的多核苷酸(例如PsaB基因),其中该模板是单链形式的质粒,含有未改变或天然光催化单元多肽的多核苷酸序列。杂交之后,使用DNA聚合酶(例如DNA聚合酶I的Klenow片段)来合成模板第二条完整的互补链,因此掺入寡核苷酸引物,这将编码所选择的光催化单元多核苷酸中的变化,因而产生异源双链体分子。
然后,使该异源双链体分子转化到合适的宿主细胞中,通常为原核生物例如大肠杆菌(E.coli)JM101。细胞生长后,将细胞接种到琼脂糖平板上,进行筛选以鉴定出含有突变DNA的细菌菌落。然后取出突变区,置入用于产生蛋白质的适当载体上,一般来讲,该类型的表达载体通常用于转化合适的宿主。
一般而言,使用长度至少为25个核苷酸的寡核苷酸。最佳的寡核苷酸含12-15个与编码突变的核苷酸模板任一端完全互补的核苷酸。这就确保了寡核苷酸充分地与单链多核苷酸模板分子杂交。寡核苷酸可容易地用本领域已知技术进行合成(例如参见Crea等,Proc.Natl.Acad.Sci.USA,75:5765(1978))。
多核苷酸模板仅可由以下载体产生:源自噬菌体M13载体的载体(市售的M13mp18和M13mp19载体适用),或者含有单链噬菌体复制起点的载体(例如参见Viera等,Meth.Enzymol.,153:3(1987))。因此,需要突变的多核苷酸必须插入这些载体的一个当中以产生单链模板。单链模板的产生记载于Sambrook等,Molecular Cloning:ALaboratory Manual,第4.21-4.41节(Cold Spring Harbor Laboratory Press,N.Y.1989)。
还可用定点诱变产生不止一个氨基酸被置换的突变体。
本文下面的实施例1将介绍根据本发明的教导,采用定点诱变使PS I突变的一个示例性方法。
PCR诱变和盒式诱变也是适于对光催化单元多肽进行修饰的技术——参见Sambrook和Russell(2001,Molecular Cloning,ALaboratory Approach,Cold Spring Harbor Press,Cold Spring Harbor,N.Y.)和Ausubel等,(2002,Current Protocols in Molecular Biology,JohnWiley & Sons,NY))。
用于表达经突变的光催化序列的合适宿主包括能够合成功能性光催化单元的任何宿主。因此,宿主必须能够将色素、叶绿素分子等掺入到这个单元中。合适宿主的实例包括但不限于绿色植物细胞培养物、绿色植物和光合细菌。在本发明这一方面的一个优选实施方案中,宿主为集胞藻细菌(Synechocystis bacteria)。
根据本发明特别优选的实施方案,通过将突变DNA插入宿主基因组进行克隆。这尤其适用于当宿主细胞为光合细菌时。该方法受到包含在载体中的与光合基因组DNA中存在的序列互补的DNA序列的影响。用这一载体转染光合细菌,导致与基因组的同源重组及光催化多肽DNA的插入。
下面的实施例1描述了利用突变psaB基因通过同源重组转化集胞藻PCC 6803的示例性方法。重要的是,野生型集胞藻细胞用克隆有突变psaB基因的质粒pZBL转化,其中野生型集胞藻细胞在BG-11平板中光激活异养生长(LAHG:避光生长,只是按光子通量密度40微摩尔m2/秒,每24小时光照10分钟),BG-11平板补充了5mM葡萄糖、10mM TES-KOH、pH 8(N-三[羟甲基]-甲基-2-氨基乙烷磺酸盐)和硫代硫酸盐(3g/L)。同源重组的方案见图1A。细胞用所得质粒转化,然后用5-20mg/ml卡那霉素筛选并分离出少许子代。在卡那霉素压力下进行转化体的筛选和分离。
或者,可以将编码本发明修饰多肽的多核苷酸与核酸表达载体连接在一起,该载体包含处于顺式调节序列(例如启动子序列)转录控制之下的多核苷酸序列,该调节序列适于在宿主细胞中指导本发明多肽的组成型、组织特异性或诱导型转录。
“分离的多核苷酸”,是指分离的并按以下形式提供的单链或双链核酸序列:RNA序列、互补多核苷酸序列(cDNA)、基因组多核苷酸序列和/复合多核苷酸序列(例如上述序列的组合)。根据本发明的教导可以使用的多核苷酸序列的实例如SEQ ID NO:30、31、32、33、34、35、36、37和38中所示。
本文所用术语“互补多核苷酸序列”是指用反转录酶或任何其它依赖于RNA的DNA聚合酶,由信使RNA反转录产生的序列。该序列随后可用依赖于DNA的DNA聚合酶在体内或体外扩增。
本文所用术语“基因组多核苷酸序列”是指从染色体获得(分离)的序列,因此表示染色体的连续部分(contiguous portion)。
本文所用术语“复合多核苷酸序列(composite polynucleotidesequence)”是指一种至少部分是互补的和至少部分是基因组的序列。复合序列可以包括一些需要编码本发明多肽的外显子序列,以及一些间插在外显子序列中的内含子序列。内含子序列可以是任何来源,包括其它基因,通常可包括保守剪接信号序列。这些内含子序列还可包括顺式作用表达调节元件。
如上所述,可以将本发明的多核苷酸序列插入表达载体(即核酸构建体)中,使重组多肽能够表达。本发明的表达载体包括使这种载体适于在原核生物、真核生物或者优选两者中复制和整合的其它序列(例如穿梭载体)。典型的克隆载体含有转录起始序列和翻译起始序列(例如启动子、增强子)及转录终止子和翻译终止子(例如聚腺苷酸化信号)。
细菌系统中,可根据所要表达的多肽的用途,便利地筛选出多种表达载体。例如,如果需要大量的多肽,则可能需要指导表达高水平蛋白质产物的载体,该载体可能与指导表达产物进入细菌周质或培养基中的疏水信号序列融合,使得蛋白质产物易于纯化。可能还需要某些用特异性切割位点改造的融合蛋白,该切割位点有助于多肽的回收。适宜这种操作的载体包括但不限于大肠杆菌表达载体的pET系[Studier等,Methods in Enzymol.185:60-89(1990)]。
在使用植物表达载体的情况下,多肽编码序列的表达可以由多种启动子驱动。例如,可以使用病毒启动子,例如CaMV的35S RNA启动子和19S RNA启动子[Brisson等,Nature 310:511-514(1984)],或者可以使用TMV的外壳蛋白启动子[Takamatsu等,EMBO J.6:307-311(1987)]。或者,可以使用植物启动子,例如RUBISCO的小亚基[Coruzzi等,EMBO J.3:1671-1680(1984)和Brogli等,Science224:838-843(1984)]或热激启动子,例如大豆hsp17.5-E或hsp17.3-B[Gurley等,Mol.Cell.Biol.6:559-565(1986)]。可以用Ti质粒、Ri质粒、植物病毒载体、直接DNA转化、显微注射、电穿孔和其它本领域技术人员熟知的技术,将这些构建体导入植物细胞。参见例如Weissbach & Weissbach [Methods for Plant Molecular Biology,Academic Press,NY,第VIII节,第421-463页(1988)]。
应当理解的是,本发明的表达构建体除了含有插入编码序列(编码多肽)转录和翻译的必需元件外,还可包括经工程改造使已表达多肽的稳定性、生产、纯化、产量或活性最优化的序列。
表达和克隆载体应当含有选择基因,亦称选择标记。这是一种编码用载体转化的宿主细胞存活或生长所必需的蛋白质的基因。这一基因的存在确保了除去载体的任何宿主细胞并不会比转化宿主在生长或繁殖上获得优势。典型的选择基因编码具有以下功能的蛋白质:(a)赋予抗生素或其它毒素(例如氨苄青霉素、新霉素、氨甲蝶呤或四环素)抗性,(b)补充营养缺陷型缺乏的养分,或(c)供给不能从复合培养基得到的关键营养物。
可以用各种方法将本发明的表达载体导入宿主细胞系统。这些方法一般参见Sambrook等,Molecular Cloning:A Laboratory Manual,Cold Springs Harbor Laboratory,New York(1989,1992);Ausubel等,Current Protocols in Molecular Biology,John Wiley and Sons,Baltimore,Md.(1989);Chang等,Somatic Gene Therapy,CRC Press,Ann Arbor,Mich.(1995);Vega等,Gene Targeting,CRC Press,Ann Arbor Mich.(1995),Vectors:A Survey of Molecular Cloning Vectors and Their Uses,Butterworths,Boston Mass.(1988)和Gilboa等[Biotechniques 4(6):504-512,1986],包括例如稳定转染或瞬时转染、脂转染、电穿孔和用重组病毒载体感染。另外,有关正负双向筛选方法参见美国专利第5,464,764号和第5,487,992号。
转化细胞在有效条件下进行培养,使其以高水平表达重组多肽。有效培养条件包括但不限于允许蛋白质产生的有效培养基、生物反应器、温度、pH和氧气等条件。有效培养基是指任何用于培养细胞以产生本发明重组多肽的培养基。此培养基通常包括具有可同化碳源、氮源和磷酸盐源及适当的盐、无机物、金属和其它营养物(例如维生素)的水溶液。本发明的细胞可以在常规的发酵生物反应器、摇瓶、试管、微量滴定板和培养皿中进行培养。可以在温度、pH和氧含量适于重组细胞的条件下进行培养。这些培养条件为本领域普通技术人员所掌握。
细胞在合适条件下培养之后,优选从培养细胞中分离出光催化单元。本文以下实施例部分的实施例2描述了从光合生物提取光催化单元的示例性方法。本发明还包括采用任何其它纯化和分离方法,只要光催化单元保持功能。所分离的光催化单元可为多聚体(例如三聚体)或单个单体。光催化单元可以是完全分离的或是部分膜制品。膜提取物的制备方法是本领域公知的。例如,Qoronfleh等[J BiomedBiotechnol.2003;2003(4):249-255]教导了通过分配相分离选择性富集膜蛋白的方法。用于制备膜提取物的各种试剂盒也是市售的,例如ProteoPrepTM膜提取试剂盒(Sigma-Aldrich公司)。
因此,本发明另一方面提供分离的修饰光催化单元,该催化单元包含本发明的修饰多肽。根据本发明的这一方面,术语“分离(的)”是指光催化单元至少部分取自其天然合成部位(例如光合生物)。优选光催化单元基本上不含存在于其体内位置的物质(例如其它细胞、蛋白质、核酸等)。
可按本文下述方法,在分离出光催化单元之后,对其活性进行测试。
在分离出本发明的修饰光催化单元之后,可以通过共价键或非共价键(静电)将其与固体表面相连接。本文所用术语“共价键”是指2个原子通过共用2个电子(其中每个原子贡献1个电子)进行连接。优选光催化单元直接与固体表面键合(即既不包含任何连接分子,也不用涂敷金属离子)。
对固体表面的选择取决于光催化单元的修饰。因此,如果修饰包含如下述的半胱氨酸置换,则固体表面优选为导电材料,例如过渡金属元素。根据本发明这一方面,可以使用的过渡金属元素的实例包括但不限于银、金、铜、铂、镍、铝和钯。
本发明实施方案的修饰光催化单元可以通过使置换残基与固体基片的亲水表面直接反应,与固体表面共价连接。例如,在优选的实施方案中,置换残基为半胱氨酸,可通过将修饰光催化单元与金表面或其它金属表面一起保温长达足以形成硫键的时间进行连接。还可包括其它连接方法。本文以下实施例部分的实施例2描述了半胱氨酸置换的光催化单元的示例性共价连接方法。
根据本发明这一方面,修饰光催化单元在连接到固体表面之后仍保持光催化活性。
本文所用术语“光催化活性”是指光能向化学能的转化。优选修饰光催化单元保持野生型光催化单元在其体内环境活性的至少20%,更优选至少30%,更优选至少40%,更优选至少50%,更优选至少60%,更优选至少70%,更优选至少80%,更优选至少90%,例如约100%。本发明还包括其活性高于野生型光催化单元在其体内环境的活性的本发明光催化单元。
为了使本发明的修饰光催化单元在连接到固相支持体上后仍具有光催化活性,光催化单元必需以定向方式连接,这样它们彼此间的电荷才不会被中和。如实施例3所述,本发明的修饰多肽能够使光催化单元结合到固相支持体上,从而产生光诱导的正电位。该诱导电位是负电荷从PS I的金一侧位移的结果。本发明的发明人认为在大量的光催化单元中用半胱氨酸置换相同的氨基酸,光催化单元与固相支持体的连接将会是定向的,并且可在固相支持体上形成单层膜。
选择特定的氨基酸用半胱氨酸残基置换,可以调节光催化单元在固相支持体上的精确取向。
所制造表面的光催化活性的测定方法包括测定所制造表面的光电压性质。光电压性质可以用例如开尔文探针力显微镜(KPFM)进行测定。如图3A和图3B的KPFM图象所示,本发明的光催化单元显示清晰的光诱导电位。具体地讲,在形貌跟踪时,产生的光诱导正电位为+0.498±0.02V,其中观察到属于PS I复合体的峰值。当关掉照明时仍观察到电位变化(图4A-B),还证实了光诱导电位的可逆性。
还可通过分析光催化复合体中的电子转递,测定光催化活性。可通过分析如单次循环光谱所测定的闪光诱导的吸收变化,测定电子传递。如图6A-B所示,与野生型重组及与突变体重组之间的电荷率的差异表明,本发明教导的半胱氨酸置换没有改变光诱导电子传递作用的方式。
现参照图7,图7是本发明不同示例性实施方案的光电器件10的示意图。器件10包括固相支持体12、大量分离的光催化单元14和支持体12的表面13,其中单元14与表面13相连接。分离的光催化单元14优选经修饰以促进单元14与表面13的共价连接,同时保持光催化活性,更多详情同上。
作为光催化单元14的组成部分,光电器件10促进光诱导的电子传递。在光11激发下,在数百皮秒到几微秒的一段时间内(视光催化单元的类型而异),电子从供体位点16,经过多个中间步骤传递到受体位点18。诱导电子传递的光频率取决于获得单元14的光合生物。例如,如果使用绿色植物或绿色细菌的光催化单元,器件10对波长约500nm至约550nm的绿色光敏感,如果使用蓝细菌的光催化单元,器件10对波长约450nm至约500nm的蓝色光敏感,如果使用红藻的光催化单元,器件10对波长约650nm至约700nm的红色光敏感,如果使用紫色细菌的光催化单元,则器件10对波长约700nm至约750nm的紫色光敏感。
光电器件10可用于微电子电路和器件以及亚微电子电路和器件领域,包括但不限于空间成像器件、太阳能电池、光学计算和逻辑门、光电开关、二极管、光子A/D转换器和薄膜“柔性”光致电压结构。
现参照图8,图8是本发明不同示例性实施方案的光电二级管器件20的示意图。本领域技术人员应当理解,为了清楚表示起见,图7中出现的几个元件在图8中被省略。光电二级管器件20包括光电器件10以及两个电接触22和24,其中两个电接触22和24分别与供体位点16和受体位点18保持电通信(electrical communication)。可以与供体位点16建立电通信,例如将导电材料连接到支持体12或表面13上。通过在本发明修饰多肽(例如PS I PsaC亚基中的W31C)与沉积的金属电极上表面之间形成硫键,使受体位点共价结合。受体侧镀铂的光催化单元与顶部电极((top electrode)通过蒸发薄的金属电极而沉积可以使金属间发生电连通(electrical connection)。在光催化单层膜上沉积的导电聚合物或镀铂的光催化单层膜可用作顶部电极。光电二级管的符号图见图8(下小图)。
使用时,用光照射光催化单元,因此激发高量子效率的有效的电荷分离,通常超过95%。接触件22和24分接通过电荷分离引起的电流。根据施加在接触件22和24之间的电压,光电二级管器件20可以用作光致电压器件,或者用作反向偏压光电二级管。
具体地讲,不存在外部电压时,当用光照射时,光电二级管器件20用作产生电流的光致电压器件。该器件可以用作例如太阳能电池中的元件。
当在接触件22和24之间施加反向偏压时,只要光电二级管器件20不被光激发光催化单元的光照射,则光电二级管器件20对从接触件24流向接触件22的电流就保持高的阻抗。在适当波长的光照射下,阻抗显著减小。这种器件可在例如光检测器中用作元件。
如果不施加偏压时,光电器件10还可以用作太阳能电池。照射光催化单元后,电荷分离态导致供体位点16和受体位点18之间产生内部电压。内部电压可以通过供体位点16和受体位点18的电接触分接。如果电路外部是关闭的,电流则通过重复的光驱动电荷分离保持在太阳能电池内。
器件10中所产生的极化电荷分离态还可用于分子晶体管。具体地讲,器件10可用作光充电电容,它作为栅极,在与电容连接的沟道中改变电荷传递体的密度。
现参照图9,图9是本发明不同示例性实施方案的光电晶体管30的示意图。光电晶体管30包括源极32、漏极34、沟道36和光应答栅极38。栅极38优选包括光电器件10。沟道36优选具有半导体性能,因此电荷传递体的密度可以改变。
无光时,沟道36不含任何自由电荷传递体,且基本上是绝缘体。暴露于光下时,器件10的光催化单元形成极化电荷分离态,由此所生成的电场从源极32和漏极34中吸引电子(或更通俗地说是电荷传递体),使得沟道36变得导电。因此,光电晶体管30用作放大器或开关器件,由光控制电流从源极32和漏极34流出。
电极可由任何导电材料(例如但不限于金)制成。极间空间决定了沟道长度。电极可以沉积在半导体表面以形成源-沟道-漏结构。栅极可以用更多详情如上的本发明实施方案中分离的光催化单元形成。光电晶体管的符号图见图9的右上小图。
正如本领域普通技术人员应该了解是,当栅极38维持断路时,光电晶体管30可工作,因为门控是通过光子撞击电极38诱导的。光电晶体管30可以用作逻辑元件,因此光电晶体管可被入射光转换至“开”的状态。另外,可能是由于具有空间位置的入射光束的漏电流强烈变化,因此光电晶体管30可用作具有大型图案形成的图象传感器的主要构件(backbone)。可以组装成数种光电晶体管,各自在更多详情如上的不同波长下工作,使图象传感器对彩色图象具敏感性。可以利用具有更多改良结构的电荷贮存能力,用于存储器相关应用中,这些改良为常规半导体领域技术人员所知。
光电二级管20和/或光电晶体管30可以集成在许多电子电路中。具体地讲,这些器件可以用作结构单元,它们可以在表面结构上组装成复合电子组合件(composite electronic assembly)。例如,两个以上的光电二级管或光电晶体管可以在表面结构上组装成逻辑门、一组逻辑门或微处理器。
现参照图10,图10是本发明不同示例性实施方案的光耦合器40的简图。光耦合器40尤其用于从一个元件传递信号到其它元件而无需在元件之间设置直接电接触,例如由于电压水平不匹配。例如,光耦合器40可以用于在以低电压水平操作的微处理器和以高电压水平操作的门控交换器件之间建立接触自由通信。
本发明优选实施方案的光耦合器40包括光发射器42和光接收器44。发射器42可以是任何光源,例如但不限于发光二极管(LED)。接收器44优选包括光电器件10,并且可以是例如光电二级管(例如光电二级管20)或光电晶体管(例如光电晶体管30)。可以选择发射器42,使得所发射的辐射因此有足够的能量引起器件10的供体位点16与受体位点18之间的电荷分离。
发射器42和接收器44保持光通信但却是电去耦的。例如,发射器42和接收器44可被使光通过但却阻止任何电流流过的透明屏障46分隔。发射器42和接收器44优选彼此相对,使得从发射器42发射的辐射射到接收器44上。
受电信号触发,发射器42发射光48,光穿过屏障46,射到接收器44上。进而,接收器44产生电信号,该信号可以通过更多详情如上的合适的电接触分接。因此,光耦合器40成功地将施加于其输入端(发射器42)的电信号传输至其输出端(接收器44),输入端与输出端之间没有任何电接触。
在本发明不同的示例性实施方案中,上述电子器件(包括但不限于光电器件、太阳能电池、光电二级管、光电晶体管、逻辑门和光耦合器)的尺寸为亚毫米范围。优选电子器件的尺寸为约0.1nm至约100μm,更优选约0.1nm至约1μm。
本文所用术语“约”是指±10%。
本发明的其它目的、优势和新的特征对于本领域普通技术人员在检验以下实施例时是显而易见的,这些实施例不是限制性的。另外,本文上述的本发明每个不同的实施方案和方面,以及所附权利要求书部分所要求保护的范围都可以在下述实施例中找到实验支持。
实施例
下面参考以下实施例并结合上述描述,以非限制性方式对本发明进行说明。
一般地讲,本文所用术语和本发明所采用的实验方法包括分子、生物化学、微生物学和重组DNA技术。这些技术在文献中已有详细介绍。参见例如“Molecular Cloning:A laboratory Manual”Sambrook等,(1989);“Current Protocols in Molecular Biology”第I-III卷,Ausubel,R.M.主编,(1994);Ausubel等,“Current Protocols in MolecularBiology”;John Wiley and Sons,Baltimore,Maryland(1989);Perbal,“APractical Guide to Molecular Cloning”,John Wiley & Sons,New York(1988);Watson等,“Recombinant DNA”,Scientific American Books,New York;Birren等(主编)“Genome Analysis:A Laboratory ManualSeries”,第1-4卷,Cold Spring Harbor Laboratory Press,New York(1998);美国专利4,666,828、4,683,202、4,801,531、5,192,659和5,272,057中所公开的方法;“Cell Biology:A Laboratory Handbook”,第I-III卷,Cellis,J.E.主编(1994);“Culture of Animal Cells-A Manualof Basic Technique”Freshney,Wiley-Liss,N.Y.(1994),第三版;“Current Protocols in Immunology”第I-III卷,Coligan J.E.主编(1994);Stites等(主编),“Basic and Clinical Immunology”(第八版),Appleton & Lange,Norwalk,CT(1994);Mishell和Shiigi(主编),“Selected Methods in Cellular Immunology”,W.H.Freeman and Co.,New York(1980);将用到的免疫测定法在专利和科技文献中有大量的记载,参见例如美国专利3,791,932、3,839,153、3,850,752、3,850,578、3,853,987、3,867,517、3,879,262、3,901,654、3,935,074、3,984,533、3,996,345、4,034,074、4,098,876、4,879,219、5,011,771和5,281,521;“Oligonucleotide Synthesis”Gait,M.J.主编(1984);“Nucleic Acid Hybridization”Hames,B.D.和Higgins S.J.主编(1985);“Transcription and Translation”Hames,B.D.和Higgins S.J.主编(1984);“Animal Cell Culture”Freshney,R.L主编(1986);“ImmobilizedCells and Enzymes”IRL Press,(1986);“A Practical Guide to MolecularCloning”Perbal,B.,(1984)和“Methods in Enzymology”第1-317卷,Academic Press;“PCR Protocols:A Guide To Methods AndApplications”,Academic Press,San Diego,CA(1990);Marshak等,“Strategies for Protein Purification and Characterization-A LaboratoryCourse Manual”CSHL Press(1996);所有这些文献的内容都通过引用全部结合到本文。还提供贯穿本文的其它一般参考文献。我们认为文献方法是本领域众所周知的,仅为了方便读者而在此提供。所有所包括的资料都通过引用结合到本文中。
实施例1
集胞藻PCC 6803 psaB突变体的合成
从蓝细菌集胞藻PCC 6803中筛选完整的PS I反应中心,以确定遗传修饰是否可以有助于将反应中心连接到固相支持体上。该PS I结构稳定性的主要原因是因为所有叶绿素分子和类胡萝卜素都整合到核心亚基复合体中,而在植物和其它细菌的反应中心内,天线叶绿素与连接到核心亚基的叶绿素-蛋白质复合体结合。
基于对PS I原子结构的了解,选择氨基酸使之修饰成为半胱氨酸,从而使PS I与金表面共价连接。因此,为了确保形成硫键,使面向细菌膜细胞质一侧膜外环(当放置在固体表面时不会形成立体障碍)上的氨基酸,突变成半胱氨酸。为了促进有效的电连接,在P700附近筛选出不同的突变,确保反应中心和金电极紧密相邻。
方法与材料
集胞藻PCC 6803的psaB通过同源重组引入突变:通过同源重组实现psaB基因的定点诱变。将1.8kb psaB基因片段和1.1kb下游侧翼区插入pBluescript II KS载体中(参见图1A-C的方案)。按先前文献所述方法[Zeng等,Biochim.Biophys.Acta 2002,1556 254-264],从pUC4K上切下1.27kb卡那霉素抗性赋予基因(KanR),将其插入侧翼起始的EcoRI位点,构建载体pZBL。采用重叠延伸聚合酶链式反应[Ho等,Gene 1989,77 51-59],构建含有突变的1400bp延伸片段,分别在限制位点Sph I-Hpa I和Sph I-Sac I,通过片段交换构建载体pZBL-D479C、pZL-S499C、pZBL-S599C、pZBL-Y634C、pZBL-D235C和pZBL-S246C。
用于产生本发明多肽的引物见下表4。
表4
SEQ IDNO: | 序列(5’→3’) | 突变 | 质粒 |
psa B | |||
SEQ IDNO:1 | GTGTATGCGGCGTGTCCCGACACTGCTGGC | D235C | PZBL-D235C |
SEQ ID | AGCAGTGTCGGGACACGCCGCATACACGCC | D235C | pZBL-D235C |
NO:2 | |||
SEQ IDNO:3 | CACATTTTTGGTTGTTCTGAAGGTGCTGGT | S246C | pZBL-S246C |
SEQ IDNO:4 | AGCACCTTCAGAACAACCAAAAATGTGGCC | S246C | pZBL-S246C |
SEQ IDNO:5 | CTCTCCAATCCTTGCAGCATTGCTTCCACC | D479C | pZBL-D479C |
SEQ IDNO:6 | GGAAGCAATGCTGCAAGGATTGGAGAGCAA | D479C | pZBL-D479C |
SEQ IDNO:7 | GATGCTATCAACTGCGGCACCAACTCTCTG | S499C | pBL-S499C |
SEQ IDNO:8 | AGAGTTGGTGCCGCAGTTGATAGCATCCAA | S499C | pZBL-S499C |
SEQ IDNO:9 | CTCGGTGTTTGGTGCGGTAACGTTGCTCAG | S599C | pZBL-S599C |
SEQ IDNO:10 | AGCAACGTTACCGCACCAAACACCGAGGTG | S599C | pZBL-S599C |
SEQ IDNO:11 | GGTTACAACCCCTGCGGTGTCAACAATCTG | Y634C | pZBL-Y634C |
SEQ IDNO:12 | ATTGTTGACACCGCAGGGGTTGTAACCATT | Y634C | pZBL-Y634C |
SEQ IDNO:13 | GTGTATGCGGCGTGTCCCGACACTGCTGGCGGTTACAACCCCTGCGGTGTCAACAATCTG | D235C/Y634C | pZBL-D235C/Y634C |
SEQ IDNO:14 | AGCAGTGTCGGGACACGCCGCATACACGCCATTGTTGACACCGCAGGGGTTGTAACCATT | D235C/Y634C | pZBL-D235C/Y634C |
SEQ IDNO:15 | CACATTTITGGTTGTTCTGAAGGTGCTGGTGGTTACAACCCCTGCGGTGTCAACAATCTG | S246C/Y634C | pZBL-S246C/Y634C |
SEQ IDNO:16 | AGCACCTTCAGAACAACCAAAAATGTGGCCATTGTTGACACCGCAGGGGTTGTAACCATT | S246C/Y634C | pZBL-S246C/Y634C |
psa C | |||
SEQ IDNO:17 | GAAATGGTGCCCTGGTGTGGTTGTAAAGCC | F31C | P61-2.4-F31C |
SEQ IDNO:18 | AGCGGCTTTACAACCACACCAGGGCACCAT | RW31C | P61-2.4-R31C |
SEQ IDNO:19His-taqC-端 | AGATCTTTAGTGGTGGTGGTGGTGGTGGTAAGCTAAACCCAT | His-taq C-term | P61-2.4-His-taq C-端 |
另外,用上述同一技术,再将W31C的单突变插入PsaC,将2个双重突变(D235C/Y634C和S246C/Y634C)插入PsaB。
为了筛选psaB缺陷型受体细胞,从psaB下游端切下1.3kb(从pZBL的SpM至EcoRI),在这些位点上插入1.3kb氯霉素抗性赋予基因(CmR),构建pBLΔB载体(图1C)。野生型集胞藻细胞用pPLΔB转化,使转化体生长在“光适应异养(light adapted heterotrophic)”条件下[Zeng等,Biochim.Biophys.Acta 2002,1556 254-264],以表达D479C突变多肽(SEQ ID NO:20)、S499C突变多肽(SEQ ID NO:21)、S599C突变多肽(SEQ ID NO:22)、Y634C突变多肽(SEQ ID NO:23)、D235C突变多肽(SEQ ID NO:24)、S246C突变多肽(SEQ ID NO:25)和W31C突变多肽(SEQ ID NO:26)。
类囊体膜和PS I复合体的分离:将集胞藻细胞在弗氏压碎器于500p.s.i.下破碎,用差速离心分离出类囊体。PS I用去污剂正十二烷基β-D-麦芽糖苷增溶,在DEAE-纤维素柱中用蔗糖梯度纯化[Nechushtai,R.,Muster,P.,Binder,A.,Liveanu,V.和Nelson,N.(1983)Proc.Natl.Acad. Sci.USA 80,1179-1183]。按先前文献所述方法[Laemmli,U.K.(1970)Nature 227,680-685;Tindall,K.R.t和Kunkel,T.A.(1988)Biochemistry 27,6008-6013],用SDS聚丙烯酰胺凝胶电泳和蛋白质印迹法对亚基组成进行分析。按先前文献所述方法[Lowry,O.H.,Rosenbrough,N.L.,Farr,A.L.和Randall,R.J.(1951)J.Biol.Chem.193,265-275],在1%SDS中增溶后,对膜内的蛋白质进行测定。根据已公开的方法[Arnon,D.I.(1949)Plant Physiol.24,1-15],测定叶绿素浓度及P700化学氧化和光氧化。详细的分离方法和分析概述参见Gong等,Journal of Biological Chemistry 2003,27819141-19150。上述分析证实,分离产物就是纯化的PS I蛋白质叶绿素复合体。
PS I中外露表面的半胱氨酸用与巯基特异性反应的生物素-马来酰亚胺探测。使生物素标记的PS I复合体解离,用十二烷基硫酸钠聚丙烯酰胺凝胶电泳进行分离。对于免疫印迹测定,按先前文献所述方法[Sun等,Methods in Enzymology,Academic Press,1998,第124-139页],将蛋白质样品从凝胶转移到硝化纤维素膜上,使之与过氧化物酶缀合的抗生物素反应,然后用增强化学发光试剂显现。
结果
如图1E所示,所有纯化的修饰PS I单元都包含外露表面的半胱氨酸。虽然未修饰的蛋白包含9个游离半胱氨酸,但是当用表面活性试剂生物素-马来酰亚胺测定时,发现它们都没有暴露在外面。
实施例2
定向单层膜的制造
方法与材料
制造:通过突变PS I的半胱氨酸与新鲜洁净的亲水金表面直接反应形成Au-硫键,来制备定向单层膜。通过在玻片或硅片上蒸发5nmCr和150nm金,制成平坦的金表面。使这些表面于350℃下真空热处理1小时。含有1mg/1ml PS I的叶绿素缓冲溶液层铺在金属表面之上,然后温育。使突变PS I或天然PS I于室温下温育2小时后,未连接的蛋白质用蒸馏水彻底洗涤几次。表面用超纯氮干燥后,用原子力显微镜(AFM)观测。
原子力显微镜(AFM):用商用AFM(配有ExtenderTM电子模块(Electronics Module)的NanoscopeIIIa MultyModeTM,VeecoInstruments)进行AU测定。悬臂共振频率为300kHz,用轻敲模式进行形貌测定。
结果
PS I突变体在退火之后在金表面自组装,在温育2小时后与金表面保持共价连接。在金表面以类似方式温育的天然PS I被冲洗掉。图2B表示通过扫描玻片上0.3μm2面积的金表面所获得的原子力显微镜图象,PS I半胱氨酸突变体D479C的单体通过硫键与玻片连接在一起。图2B清楚显示出直径15-21nm,高9nm的颗粒的单层膜。这正是预期的PS I尺寸,正如晶体学所获得的一样。在发光的颗粒状表面观察到这些颗粒。图2A表示经退火未处理的裸金表面的扫描图,经退火的金颗粒直径150nm,高5nm。
实施例3
PS I突变体制造表面的光电压性质
通过开尔文探针力显微镜(KPFM)测定单层膜中半胱氨酸突变体D479C的单个PS I三聚体复合体和单体复合体的光电压。
材料与方法
KPFM装置基于商用AFM(改良的NanoScopeIIIaMultiMode,Veeco,USA),以轻敲模式操作。用所谓的“抬高模式(liftmode)”测定静电力。基本上,测量形貌以后,使针尖从样品表面缩回到固定高度。由压力引发的针尖摆动便停止,在之前用轻敲模式测量形貌所用的相同频率下,向悬臂施加AC偏流。通过使在第一共振频率下测定静电力的锁相放大器的输出信号归零,用常规方式测定CPD[Vatel等,Journal of Applied Physics 1995,772358-2362]。用连续扫描记录AFM形貌和相应的KPFM电位,扫描速率1Hz;样品面积分两段进行512线扫描以形成二维图象。对于第一段扫描打开氦-氖激光(λ=632.8nm,5mW/cm2),对于第二段则关掉激光。
结果
如图3A和图3B所示,KPFM图象证实突变PS I复合体明显的光诱导电位。在形貌跟踪时,产生的光诱导正电位为+0.498±0.02V(参见下表1),其中观测到属于PS I复合体的峰值(图3A和图3B)。
这些结果清楚表明,所有结合到金表面的突变PS I复合体都具有功能活性,并且都朝着相同的方向。诱导电位是负电荷从PS I的金一侧位移的结果,见图1D。
照明下,所测接触电位差(CPD)的变化与简单计算的电位变化(ΔΦ)十分吻合,该电位由诱导偶极层产生,偶极层高度d,与曲面法线成θ角,可用下列公式表示:Δφ=Nqd cosθ/ε,其中N为偶极层的面积密度,q为元电荷,ε为偶极层的介电常数。对于三聚体,如果N为2.2×1011cm-2,d=5nm,ε=2,θ=90°,我们得出ΔΦ=0.41伏特。因为该层偶极的角度未知,而且还因为偶极层不是一个大的实体,故介电常数具有不确定的性质,所以可部分解答预期的+1V与计算所得出的0.41V之间的偏差。此外,由于反应中心彼此相对较远,照明下所测得的CPD比实际CPD小很多。由于长距离静电力,在针尖顶下表面的点所测得的CPD是针尖附近表面电位的加权平均值。过去曾用不同算法计算针尖静电求平均值的影响[Y.Rosenwaks等,Physical Review B 2004, 70 085320-085327]。根据这些计算结果,估计CPD(照明下)更精确的值约比该测量值大2倍。在位于巯基乙醇涂敷的金上的植物PS I中,测得相似的光诱导电位[I.Lee等,J.Phys.Chem.B2000,104 2439-2443]。在这些实验中,用松散结合在单层膜上的巯基乙醇使PS I与金隔离,仅70%的复合体呈相同方向。在照明期间,金表面功函数也增加约+0.125V。部分电荷传递到位于PS I与金属界面的光氧化P700附近,预期将导致更多的负基片。金和PS I之间的电荷传递是有效电子耦合的指标。然而,当照射未处理的金表面时,CPD的变化非常小。还可以观察到暗中PS I复合体的正电位(CPD为+0.2V)比金基片的略高(图3B)。这一电位可能是由AFM反馈激光使PS I激发而产生,该反馈激光没有完全被用在KPFM装置中的悬臂所掩蔽。这种本底激发可能引起对PS I中光CPD减去暗CPD变化的低估。
关掉照明时观察到电位的变化,证实了光诱导电位的可逆性质。结果见下表5。数值表示在光下或在暗中测得的24个独立的PS I复合体的平均电位。
表5
PS I/电位 | 暗(V) | 光(V) | 光→暗(V) | 暗→光(V) |
单体 | -0.193±0.0005 | +0.118±0.0010 | +0.311±0.0010 | - |
单体 | -0.353±0.0002 | +0.004±0.0006 | - | 0.356±0.001 |
三聚体 | +0.870±0.003 | +1.220±0.0200 | +0.358±0.0014 | - |
三聚体 | +0.717±0.003 | +1.215±0.0200 | - | +0.498±0.020 |
测量或者在光下开始,然后关掉光源(光至暗),或者在暗中开始,然后打开光源(暗至光)。
表5所示,单层膜中经照射的PS I半胱氨酸突变体的三聚体产生的CPD为+1.220±0.02V,当关掉光源则降至+0.870±0.003V(见图4B)。
当关掉光源时,基片电位没有变化。在实验中施加偏流电位时,电荷不能从PS I附近快速迁移到balk金基片上。因此,与光源打开时的差异相比,光CPD减去暗CPD(SPV)在关掉光源时较小(表5)。多个PS I复合体的平均光电位减去暗电位之差为+0.358±0.014V。当金表面暴露时,接触电位中没有变化。在同一样品中或在贮藏超过一个月的样品中,光电位可以重复再生多次。
半胱氨酸突变体D479C的PS I单体易形成自组装的定向单层膜。单层膜中,单体之间(图2B)和三聚体(图5B)之间的平均距离分别为15nm和25nm。然而小单体比三聚体充满在单层膜中的密度更高,可以通过配有高清晰度悬臂针尖的AFM测定加以判断(图2B)。然而,形貌或CPD测量的灵敏度都不足于鉴别单个单体。因此,用KPFM测量在暗中的PS I单体单层膜所获得的CPD称为连续集(continuum)(图5A)。用同一装置测定形貌时,观察到类似的图象(未显示)。然而,用光照明后,CPD增加。在暗中和光下,根据单层膜上多个位置的CPD平均测量值,得出光诱导的CPD为+0.356±0.001V(表5)。经照射的单层膜在关掉光源测量变化时,观测到较小的光诱导CPD为+0.311±0.001。
实施例4
本发明其它半胱氨酸突变体的自组装
定向PS I的自组装还从其它3个突变体中获得,其中位于暴露在膜外环上的氨基酸S499C、S599C、Y634C被修饰成为半胱氨酸(见图1D)。这些突变体的PSI单体和三聚体均形成单层膜,所产生的光诱导CPD的值,与用突变体D479C制备的单层膜所测得的值相似。因此,PS I的功能性定向单层膜取决于与位于复合体膜外环的半胱氨酸形成硫键,而不局限在特定的位置。
实施例5
PS I电荷复合(charge recombination)的动力学分析
为了表征突变对PS I复合体中电子传递的作用,用单次循环光谱测定闪光诱导的吸收变化。
材料与方法
光谱测定:用前述改良的闪光光解装置[Gong等,Journal ofBiological Chemistry 2003,278 19141-19150],于700nm和820nm下测定类囊体和PS I中P700的光氧化。样品含有25mM Tris(pH 8)、10mM抗坏血酸钠、10μM吩嗪硫酸甲酯和60μg叶绿素/ml PS I复合体。吸收变化瞬态用装配多指数衰减的Marquardt最小二乘法算法程序分析(KaleidaGraph 3.5,得自Synergy Software,Reading,PA)。
结果
P700的光诱导氧化引起700nm的吸收减小或820nm的吸收增加。从突变体D479C中分离的PS I蛋白中,闪光诱导的瞬时ΔA820(和ΔA700nm未显示)衰减显示与野生型相似的结果,逆反应半衰期为4.5毫秒,这可归因于电子传递介质吩嗪硫酸甲酯使P700+还原(图6A)。对于S499C和S599C的PS I复合体也获得相似结果。结果表明,在突变PS I中电子转交给FA/FB。如果在转交给FA/FB时有干扰,则P700+还原可能是位于FA/FB之前传递体减少的结果,复合率可能比4.5毫秒更快。实际上,突变体599C吸收衰减被分解成4.5毫秒(85%)和0.5毫秒(15%)的两个组分,表明仅有部分电子传递到Fx,这导致P700+和Fx -之间的电荷复合[K.Brettel,Biochim.Biophys.Acta 1997,1318 322-373]。野生型和突变体P700+和Fx -之间相似的电荷复合率表明,定点诱变不会改变光诱导电子传递作用的模式。这些结果与突变体能够在连续光下自养生长这一事实是相符的。
实施例6
通过X射线荧光测定方向
材料与方法
X射线吸收测定:从BioCAT机构(Advanced Photon Source,Argonne National Laboratories,Argonne,IL)的波动机光束扇区(beamline Sector)18ID-D中获得X射线吸收和荧光。通过双Si(III)晶单色仪供给光束,用谐波抑制镜获得谐波抑制。聚焦光束的尺寸为100μm×150μm,10-4 DE/E带宽的流量为1014光子/秒。通过充满N2气的离子室,来监测入射X射线束强度Io,通过多层阵列检测器监测X射线荧光。Fe K边在X射线能7000eV和7900eV之间进行扫描。为了使辐射损伤减至最小,在100K下,对PS I样品的各角度扫描60秒。
结果
用掠射X射线荧光全反射测定自组装单层膜中PS I的方向。PSI通过唯一的半胱氨酸与硅基片上钨-碳多层上的钨形成硫键连接。各图是按与X射线束常态的25、45、60和90度的规定角度,60秒、42秒扫描的平均值。如图6B所示,X射线荧光k边随相对于极化X射线束角度的变化而变化。如果PS I和铁-硫簇以硅基片平面为基准取向,这种变化则是预料当中的。3个[4Fe-4S]铁-硫簇各自形成变形立方,位于中心并沿PS I假C2对称轴的两侧[P.Jordan等,Nature 2001,411 909-917]。因此,预期极化X射线束吸收衰减的变化是使光束偏振的PS I假对称轴角度的函数。实际上,早已通过电子顺磁共振在部分定向的类囊体膜中测定了铁-硫簇的方向[R.C.Prince,Biochimicaet Biophysica Acta(BBA)Bioenergetics 1980,592 323-337]。
结论
本项工作首次证明,从蓝细菌中筛选完整的反应中心PS I,加上基于晶体结构的合理的突变设计,能够在导电金属表面制造定向单层膜。通过与经突变诱导的唯一的半胱氨酸形成硫键,使蛋白质复合体直接结合在金属电极上,确保了稳定、定向、具功能和有效的电子连接。单层膜中的干膜蛋白保持其产生约+1V光电位的能力。光电二级管的性质、纳米尺度、量子产额高和几乎60%的能量转化效率使反应中心成为分子电子学和生物工程学应用中十分引人注目的纳米技术器件。
应当了解的是,为了清楚起见在不同的实施方案内容中加以描述的本发明的某些特征,同样也可合并起来在一个实施方案中提供。相反,为简明起见在一个实施方案内容中加以描述的本发明的各种特征,同样也可单独或以任何合适的再合并方式予以提供。
尽管本发明结合其具体的实施方案加以描述,但是明显的是许多替换、修改和变化对本领域技术人员而言是显而易见的。因此,本发明包括所有落入所附权利要求书精神和大范围内的这些替换、修改和变化。本说明书所提及的所有出版物、专利、专利申请都通过引用全部结合到本说明书中,其程度与每个独立出版物、专利、专利申请具体而单独指明通过引用结合到本文中一样。另外,在本申请任何参考文献的引用或标识都不得解释为承认这样的参考文献都可作为本发明的现有技术获得。
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195 200 205
Trp Asp Asn Phe Leu Ser Thr Pro Pro His Pro Ala Gly Leu Met Pro
210 215 220
Phe Phe Thr Gly Asn Trp Gly Val Tyr Ala Ala Asp Pro Asp Thr Ala
225 230 235 240
Gly His Ile Phe Gly Thr Ser Glu Gly Ala Gly Thr Ala Ile Leu Thr
245 250 255
Phe Leu Gly Gly Phe His Pro Gln Thr Glu Ser Leu Trp Leu Thr Asp
260 265 270
Ile Ala His His His Leu Ala Ile Ala Val Ile Phe Ile Ile Ala Gly
275 280 285
His Met Tyr Arg Thr Asn Trp Gly Ile Gly His Ser Ile Lys Glu Ile
290 295 300
Leu Asn Ala His Lys Gly Pro Leu Thr Gly Ala Gly His Thr Asn Leu
305 310 315 320
Tyr Asp Thr Ile Asn Asn Ser Leu His Phe Gln Leu Gly Leu Ala Leu
325 330 335
Ala Ser Leu Gly Val Ile Thr Ser Leu Val Ala Gln His Met Tyr Ser
340 345 350
Leu Pro Ser Tyr Ala Phe Ile Ala Gln Asp His Thr Thr Gln Ala Ala
355 360 365
Leu Tyr Thr His His Gln Tyr Ile Ala Gly Phe Leu Met Val Gly Ala
370 375 380
Phe Ala His Gly Ala Ile Phe Phe Val Arg Asp Tyr Asp Pro Val Ala
385 390 395 400
Asn Lys Asp Asn Val Leu Ala Arg Met Leu Glu His Lys Glu Ala Leu
405 410 415
Ile Ser His Leu Ser Trp Val Ser Leu Phe Leu Gly Phe His Thr Leu
420 425 430
Gly Leu Tyr Val His Asn Asp Val Val Val Ala Phe Gly Thr Pro Glu
435 440 445
Lys Gln Ile Leu Ile Glu Pro Val Phe Ala Gln Trp Ile Gln Ala Thr
450 455 460
Ser Gly Lys Ala Leu Tyr Gly Phe Asp Val Leu Leu Ser Asn Pro Asp
465 470 475 480
Ser Ile Ala Ser Thr Thr Gly Ala Ala Trp Leu Pro Gly Trp Leu Asp
485 490 495
Ala Ile Asn Ser Gly Thr Asn Ser Leu Phe Leu Thr Ile Gly Pro Gly
500 505 510
Asp Phe Leu Val His His Ala Ile Ala Leu Gly Leu His Thr Thr Ala
515 520 525
Leu Ile Leu Ile Lys Gly Ala Leu Asp Ala Arg Gly Ser Lys Leu Met
530 535 540
Pro Asp Lys Lys Asp Phe Gly Tyr Ser Phe Pro Cys Asp Gly Pro Gly
545 550 555 560
Arg Gly Gly Thr Cys Asp Ile Ser Ala Trp Asp Ala Phe Tyr Leu Ala
565 570 575
Met Phe Trp Met Leu Asn Thr Leu Gly Trp Leu Thr Phe Tyr Trp His
580 585 590
Trp Lys His Leu Gly Val Trp Cys Gly Asn Val Ala Gln Phe Asn Glu
595 600 605
Asn Ser Thr Tyr Leu Met Gly Trp Phe Arg Asp Tyr Leu Trp Ala Asn
610 615 620
Ser Ala Gln Leu Ile Asn Gly Tyr Asn Pro Tyr Gly Val Asn Asn Leu
625 630 635 640
Ser Val Trp Ala Trp Met Phe Leu Phe Gly His Leu Val Trp Ala Thr
645 650 655
Gly Phe Met Phe Leu Ile Ser Trp Arg Gly Tyr Trp Gln Glu Leu Ile
660 665 670
Glu Thr Ile Val Trp Ala His Glu Arg Thr Pro Leu Ala Asn Leu Val
675 680 685
Arg Trp Lys Asp Lys Pro Val Ala Leu Ser Ile Val Gln Ala Arg Leu
690 695 700
Val Gly Leu Ala His Phe Thr Val Gly Tyr Val Leu Thr Tyr Ala Ala
705 710 715 720
Phe Leu Ile Ala Ser Thr Ala Gly Lys Phe Gly
725 730
<210>23
<211>731
<212>PRT
<213>集胞藻(Synechocystis sp.)
<220>
<221>其他特征
<223>Y634C突变多肽
<400>23
Met Ala Thr Lys Phe Pro Lys Phe Ser Gln Asp Leu Ala Gln Asp Pro
1 5 10 15
Thr Thr Arg Arg Ile Trp Tyr Gly Ile Ala Thr Ala His Asp Phe Glu
20 25 30
Thr His Asp Gly Met Thr Glu Glu Asn Leu Tyr Gln Lys Ile Phe Ala
35 40 45
Ser His Phe Gly His Ile Ala Ile Ile Phe Leu Trp Thr Ser Gly Thr
50 55 60
Leu Phe His Val Ala Trp Gln Gly Asn Phe Glu Gln Trp Ile Lys Asp
65 70 75 80
Pro Leu Asn Ile Arg Pro Ile Ala His Ala Ile Trp Asp Pro His Phe
85 90 95
Gly Glu Gly Ala Val Asn Ala Phe Thr Gln Ala Gly Ala Ser Asn Pro
100 105 110
Val Asn Ile Ala Tyr Sar Gly Val Tyr His Trp Phe Tyr Thr Ile Gly
115 120 125
Met Thr Thr Asn Gln Glu Leu Tyr Ser Gly Ala Val Phe Leu Leu Val
130 135 140
Leu Ala Ser Leu Phe Leu Phe Ala Gly Trp Leu His Leu Gln Pro Lys
145 150 155 160
Phe Arg Pro Ser Leu Ala Trp Phe Lys Asn Ala Glu Ser Arg Leu Asn
165 170 175
His His Leu Ala Gly Leu Phe Gly Val Ser Ser Leu Ala Trp Ala Gly
180 185 190
His Leu Val His Val Ala Ile Pro Glu Ala Arg Gly Gln His Val Gly
195 200 205
Trp Asp Asn Phe Leu Ser Thr Pro Pro His Pro Ala Gly Leu Met Pro
210 215 220
Phe Phe Thr Gly Asn Trp Gly Val Tyr Ala Ala Asp Pro Asp Thr Ala
225 230 235 240
Gly His Ile Phe Gly Thr Ser Glu Gly Ala Gly Thr Ala Ile Leu Thr
245 250 255
Phe Leu Gly Gly Phe His Pro Gln Thr Glu Ser Leu Trp Leu Thr Asp
260 265 270
Ile Ala His His His Leu Ala Ile Ala Val Ile Phe Ile Ile Ala Gly
275 280 285
His Met Tyr Arg Thr Asn Trp Gly Ile Gly His Ser Ile Lys Glu Ile
290 295 300
Leu Asn Ala His Lys Gly Pro Leu Thr Gly Ala Gly His Thr Asn Leu
305 310 315 320
Tyr Asp Thr Ile Asn Asn Ser Leu His Phe Gln Leu Gly Leu Ala Leu
325 330 335
Ala Ser Leu Gly Val Ile Thr Ser Leu Val Ala Gln His Met Tyr Ser
340 345 350
Leu Pro Ser Tyr Ala Phe Ile Ala Gln Asp His Thr Thr Gln Ala Ala
355 360 365
Leu Tyr Thr His His Gln Tyr Ile Ala Gly Phe Leu Met Val Gly Ala
370 375 380
Phe Ala His Gly Ala Ile Phe Phe Val Arg Asp Tyr Asp Pro Val Ala
385 390 395 400
Asn Lys Asp Asn Val Leu Ala Arg Met Leu Glu His Lys Glu Ala Leu
405 410 415
Ile Ser His Leu Ser Trp Val Ser Leu Phe Leu Gly Phe His Thr Leu
420 425 430
Gly Leu Tyr Val His Asn Asp Val Val Val Ala Phe Gly Thr Pro Glu
435 440 445
Lys Gln Ile Leu Ile Glu Pro Val Phe Ala Gln Trp Ile Gln Ala Thr
450 455 460
Ser Gly Lys Ala Leu Tyr Gly Phe Asp Val Leu Leu Ser Asn Pro Asp
465 470 475 480
Ser Ile Ala Ser Thr Thr Gly Ala Ala Trp Leu Pro Gly Trp Leu Asp
485 490 495
Ala Ile Asn Ser Gly Thr Asn Ser Leu Phe Leu Thr Ile Gly Pro Gly
500 505 510
Asp Phe Leu Val His His Ala Ile Ala Leu Gly Leu His Thr Thr Ala
515 520 525
Leu Ile Leu Ile Lys Gly Ala Leu Asp Ala Arg Gly Ser Lys Leu Met
530 535 540
Pro Asp Lys Lys Asp Phe Gly Tyr Ser Phe Pro Cys Asp Gly Pro Gly
545 550 555 560
Arg Gly Gly Thr Cys Asp Ile Ser Ala Trp Asp Ala Phe Tyr Leu Ala
565 570 575
Met Phe Trp Met Leu Asn Thr Leu Gly Trp Leu Thr Phe Tyr Trp His
580 585 590
Trp Lys His Leu Gly Val Trp Ser Gly Asn Val Ala Gln Phe Asn Glu
595 600 605
Asn Ser Thr Tyr Leu Met Gly Trp Phe Arg Asp Tyr Leu Trp Ala Asn
610 615 620
Ser Ala Gln Leu Ile Asn Gly Tyr Asn Pro Cys Gly Val Asn Asn Leu
625 630 635 640
Ser Val Trp Ala Trp Met Phe Leu Phe Gly His Leu Val Trp Ala Thr
645 650 655
Gly Phe Met Phe Leu Ile Ser Trp Arg Gly Tyr Trp Gln Glu Leu Ile
660 665 670
Glu Thr Ile Val Trp Ala His Glu Arg Thr Pro Leu Ala Asn Leu Val
675 680 685
Arg Trp Lys Asp Lys Pro Val Ala Leu Ser Ile Val Gln Ala Arg Leu
690 695 700
Val Gly Leu Ala His Phe Thr Val Gly Tyr Val Leu Thr Tyr Ala Ala
705 710 715 720
Phe Leu Ile Ala Ser Thr Ala Gly Lys Phe Gly
725 730
<210>24
<211>731
<212>PRT
<213>集胞藻(Synechocystis sp.)
<220>
<221>其他特征
<223>D235C突变多肽
<400>24
Met Ala Thr Lys Phe Pro Lys Phe Ser Gln Asp Leu Ala Gln Asp Pro
1 5 10 15
Thr Thr Arg Arg Ile Trp Tyr Gly Ile Ala Thr Ala His Asp Phe Glu
20 25 30
Thr His Asp Gly Met Thr Glu Glu Asn Leu Tyr Gln Lys Ile Phe Ala
35 40 45
Ser His Phe Gly His Ile Ala Ile Ile Phe Leu Trp Thr Ser Gly Thr
50 55 60
Leu Phe His Val Ala Trp Gln Gly Asn Phe Glu Gln Trp Ile Lys Asp
65 70 75 80
Pro Leu Asn Ile Arg Pro Ile Ala His Ala Ile Trp Asp Pro His Phe
85 90 95
Gly Glu Gly Ala Val Asn Ala Phe Thr Gln Ala Gly Ala Ser Asn Pro
100 105 110
Val Asn Ile Ala Tyr Ser Gly Val Tyr His Trp Phe Tyr Thr Ile Gly
115 120 125
Met Thr Thr Asn Gln Glu Leu Tyr Ser Gly Ala Val Phe Leu Leu Val
130 135 140
Leu Ala Ser Leu Phe Leu Phe Ala Gly Trp Leu His Leu Gln Pro Lys
145 150 155 160
Phe Arg Pro Ser Leu Ala Trp Phe Lys Asn Ala Glu Ser Arg Leu Asn
165 170 175
His His Leu Ala Gly Leu Phe Gly Val Ser Ser Leu Ala Trp Ala Gly
180 185 190
His Leu Val His Val Ala Ile Pro Glu Ala Arg Gly Gln His Val Gly
195 200 205
Trp Asp Asn Phe Leu Ser Thr Pro Pro His Pro Ala Gly Leu Met Pro
210 215 220
Phe Phe Thr Gly Asn Trp Gly Val Tyr Ala Ala Cys Pro Asp Thr Ala
225 230 235 240
Gly His Ile Phe Gly Thr Ser Glu Gly Ala Gly Thr Ala Ile Leu Thr
245 250 255
Phe Leu Gly Gly Phe His Pro Gln Thr Glu Ser Leu Trp Leu Thr Asp
260 265 270
Ile Ala His His His Leu Ala Ile Ala Val Ile Phe Ile Ile Ala Gly
275 280 285
His Met Tyr Arg Thr Asn Trp Gly Ile Gly His Ser Ile Lys Glu Ile
290 295 300
Leu Asn Ala His Lys Gly Pro Leu Thr Gly Ala Gly His Thr Asn Leu
305 310 315 320
Tyr Asp Thr Ile Asn Asn Ser Leu His Phe Gln Leu Gly Leu Ala Leu
325 330 335
Ala Ser Leu Gly Val Ile Thr Ser Leu Val Ala Gln His Met Tyr Ser
340 345 350
Leu Pro Ser Tyr Ala Phe Ile Ala Gln Asp His Thr Thr Gln Ala Ala
355 360 365
Leu Tyr Thr His His Gln Tyr Ile Ala Gly Phe Leu Met Val Gly Ala
370 375 380
Phe Ala His Gly Ala Ile Phe Phe Val Arg Asp Tyr Asp Pro Val Ala
385 390 395 400
Asn Lys Asp Asn Val Leu Ala Arg Met Leu Glu His Lys Glu Ala Leu
405 410 415
Ile Ser His Leu Ser Trp Val Ser Leu Phe Leu Gly Phe His Thr Leu
420 425 430
Gly Leu Tyr Val His Asn Asp Val Val Val Ala Phe Gly Thr Pro Glu
435 440 445
Lys Gln Ile Leu Ile Glu Pro Val Phe Ala Gln Trp Ile Gln Ala Thr
450 455 460
Ser Gly Lys Ala Leu Tyr Gly Phe Asp Val Leu Leu Ser Asn Pro Asp
465 470 475 480
Ser Ile Ala Ser Thr Thr Gly Ala Ala Trp Leu Pro Gly Trp Leu Asp
485 490 495
Ala Ile Asn Ser Gly Thr Asn Ser Leu Phe Leu Thr Ile Gly Pro Gly
500 505 510
Asp Phe Leu Val His His Ala Ile Ala Leu Gly Leu His Thr Thr Ala
515 520 525
Leu Ile Leu Ile Lys Gly Ala Leu Asp Ala Arg Gly Ser Lys Leu Met
530 535 540
Pro Asp Lys Lys Asp Phe Gly Tyr Ser Phe Pro Cys Asp Gly Pro Gly
545 550 555 560
Arg Gly Gly Thr Cys Asp Ile Ser Ala Trp Asp Ala Phe Tyr Leu Ala
565 570 575
Met Phe Trp Met Leu Asn Thr Leu Gly Trp Leu Thr Phe Tyr Trp His
580 585 590
Trp Lys His Leu Gly Val Trp Ser Gly Asn Val Ala Gln Phe Asn Glu
595 600 605
Asn Ser Thr Tyr Leu Met Gly Trp Phe Arg Asp Tyr Leu Trp Ala Asn
610 615 620
Ser Ala Gln Leu Ile Asn Gly Tyr Asn Pro Tyr Gly Val Asn Asn Leu
625 630 635 640
Ser Val Trp Ala Trp Met Phe Leu Phe Gly His Leu Val Trp Ala Thr
645 650 655
Gly Phe Met Phe Leu Ile Ser Trp Arg Gly Tyr Trp Gln Glu Leu Ile
660 665 670
Glu Thr Ile Val Trp Ala His Glu Arg Thr Pro Leu Ala Asn Leu Val
675 680 685
Arg Trp Lys Asp Lys Pro Val Ala Leu Ser Ile Val Gln Ala Arg Leu
690 695 700
Val Gly Leu Ala His Phe Thr Val Gly Tyr Val Leu Thr Tyr Ala Ala
705 710 715 720
Phe Leu Ile Ala Ser Thr Ala Gly Lys Phe Gly
725 730
<210>25
<211>731
<212>PRT
<213>集胞藻(Synechocystis sp.)
<220>
<221>其他特征
<223>S246C突变多肽
<400>25
Met Ala Thr Lys Phe Pro Lys Phe Ser Gln Asp Leu Ala Gln Asp Pro
1 5 10 15
Thr Thr Arg Arg Ile Trp Tyr Gly Ile Ala Thr Ala His Asp Phe Glu
20 25 30
Thr His Asp Gly Met Thr Glu Glu Asn Leu Tyr Gln Lys Ile Phe Ala
35 40 45
Ser His Phe Gly His Ile Ala Ile Ile Phe Leu Trp Thr Ser Gly Thr
50 55 60
Leu Phe His Val Ala Trp Gln Gly Asn Phe Glu Gln Trp Ile Lys Asp
65 70 75 80
Pro Leu Asn Ile Arg Pro Ile Ala His Ala Ile Trp Asp Pro His Phe
85 90 95
Gly Glu Gly Ala Val Asn Ala Phe Thr Gln Ala Gly Ala Ser Asn Pro
100 105 110
Val Asn Ile Ala Tyr Ser Gly Val Tyr His Trp Phe Tyr Thr Ile Gly
115 120 125
Met Thr Thr Asn Gln Glu Leu Tyr Ser Gly Ala Val Phe Leu Leu Val
130 135 140
Leu Ala Ser Leu Phe Leu Phe Ala Gly Trp Leu His Leu Gln Pro Lys
145 150 155 160
Phe Arg Pro Ser Leu Ala Trp Phe Lys Asn Ala Glu Ser Arg Leu Asn
165 170 175
His His Leu Ala Gly Leu Phe Gly Val Ser Ser Leu Ala Trp Ala Gly
180 185 190
His Leu Val His Val Ala Ile Pro Glu Ala Arg Gly Gln His Val Gly
195 200 205
Trp Asp Asn Phe Leu Ser Thr Pro Pro His Pro Ala Gly Leu Met Pro
210 215 220
Phe Phe Thr Gly Asn Trp Gly Val Tyr Ala Ala Asp Pro Asp Thr Ala
225 230 235 240
Gly His Ile Phe Gly Thr Cys Glu Gly Ala Gly Thr Ala Ile Leu Thr
245 250 255
Phe Leu Gly Gly Phe His Pro Gln Thr Glu Ser Leu Trp Leu Thr Asp
260 265 270
Ile Ala His His His Leu Ala Ile Ala Val Ile Phe Ile Ile Ala Gly
275 280 285
His Met Tyr Arg Thr Asn Trp Gly Ile Gly His Ser Ile Lys Glu Ile
290 295 300
Leu Asn Ala His Lys Gly Pro Leu Thr Gly Ala Gly His Thr Asn Leu
305 310 315 320
Tyr Asp Thr Ile Asn Asn Ser Leu His Phe Gln Leu Gly Leu Ala Leu
325 330 335
Ala Ser Leu Gly Val Ile Thr Ser Leu Val Ala Gln His Met Tyr Ser
340 345 350
Leu Pro Ser Tyr Ala Phe Ile Ala Gln Asp His Thr Thr Gln Ala Ala
355 360 365
Leu Tyr Thr His His Gln Tyr Ile Ala Gly Phe Leu Met Val Gly Ala
370 375 380
Phe Ala His Gly Ala Ile Phe Phe Val Arg Asp Tyr Asp Pro Val Ala
385 390 395 400
Asn Lys Asp Asn Val Leu Ala Arg Met Leu Glu His Lys Glu Ala Leu
405 410 415
Ile Ser His Leu Ser Trp Val Ser Leu Phe Leu Gly Phe His Thr Leu
420 425 430
Gly Leu Tyr Val His Asn Asp Val Val Val Ala Phe Gly Thr Pro Glu
435 440 445
Lys Gln Ile Leu Ile Glu Pro Val Phe Ala Gln Trp Ile Gln Ala Thr
450 455 460
Ser Gly Lys Ala Leu Tyr Gly Phe Asp Val Leu Leu Ser Asn Pro Asp
465 470 475 480
Ser Ile Ala Ser Thr Thr Gly Ala Ala Trp Leu Pro Gly Trp Leu Asp
485 490 495
Ala Ile Asn Ser Gly Thr Asn Ser Leu Phe Leu Thr Ile Gly Pro Gly
500 505 510
Asp Phe Leu Val His His Ala Ile Ala Leu Gly Leu His Thr Thr Ala
515 520 525
Leu Ile Leu Ile Lys Gly Ala Leu Asp Ala Arg Gly Ser Lys Leu Met
530 535 540
Pro Asp Lys Lys Asp Phe Gly Tyr Ser Phe Pro Cys Asp Gly Pro Gly
545 550 555 560
Arg Gly Gly Thr Cys Asp Ile Ser Ala Trp Asp Ala Phe Tyr Leu Ala
565 570 575
Met Phe Trp Met Leu Asn Thr Leu Gly Trp Leu Thr Phe Tyr Trp His
580 585 590
Trp Lys His Leu Gly Val Trp Ser Gly Asn Val Ala Gln Phe Asn Glu
595 600 605
Asn Ser Thr Tyr Leu Met Gly Trp Phe Arg Asp Tyr Leu Trp Ala Asn
610 615 620
Ser Ala Gln Leu Ile Asn Gly Tyr Asn Pro Tyr Gly Val Asn Asn Leu
625 630 635 640
Ser Val Trp Ala Trp Met Phe Leu Phe Gly His Leu Val Trp Ala Thr
645 650 655
Gly Phe Met Phe Leu Ile Ser Trp Arg Gly Tyr Trp Gln Glu Leu Ile
660 665 670
Glu Thr Ile Val Trp Ala His Glu Arg Thr Pro Leu Ala Asn Leu Val
675 680 685
Arg Trp Lys Asp Lys Pro Val Ala Leu Ser Ile Val Gln Ala Arg Leu
690 695 700
Val Gly Leu Ala His Phe Thr Val Gly Tyr Val Leu Thr Tyr Ala Ala
705 710 715 720
Phe Leu Ile Ala Ser Thr Ala Gly Lys Phe Gly
725 730
<210>26
<211>81
<212>PRT
<213>集胞藻(Synechocystis sp.)
<220>
<221>其他特征
<223>W31C突变多肽
<400>26
Met Ser His Ser Val Lys Ile Tyr Asp Thr Cys Ile Gly Cys Thr Gln
1 5 10 15
Cys Val Arg Ala Cys Pro Leu Asp Val Leu Glu Met Val Pro Cys Asp
20 25 30
Gly Cys Lys Ala Ala Gln Ile Ala Ser Ser Pro Arg Thr Glu Asp Cys
35 40 45
Val Gly Cys Lys Arg Cys Glu Thr Ala Cys Pro Thr Asp Phe Leu Ser
50 55 60
Ile Arg Val Tyr Leu Gly Ala Glu Thr Thr Arg Ser Met Gly Leu Ala
65 70 75 80
Tyr
<210>27
<211>812
<212>PRT
<213>集胞藻(Synechocystis sp.)
<220>
<221>其他特征
<223>W31C/D235C/Y634突变多肽
<400>27
Met Ser His Ser Val Lys Ile Tyr Asp Thr Cys Ile Gly Cys Thr Gln
1 5 10 15
Cys Val Arg Ala Cys Pro Leu Asp Val Leu Glu Met Val Pro Cys Asp
20 25 30
Gly Cys Lys Ala Ala Gln Ile Ala Ser Ser Pro Arg Thr Glu Asp Cys
35 40 45
Val Gly Cys Lys Arg Cys Glu Thr Ala Cys Pro Thr Asp Phe Leu Ser
50 55 60
Ile Arg Val Tyr Leu Gly Ala Glu Thr Thr Arg Ser Met Gly Leu Ala
65 70 75 80
Tyr Met Ala Thr Lys Phe Pro Lys Phe Ser Gln Asp Leu Ala Gln Asp
85 90 95
Pro Thr Thr Arg Arg Ile Trp Tyr Gly Ile Ala Thr Ala His Asp Phe
100 105 110
Glu Thr His Asp Gly Met Thr Glu Glu Asn Leu Tyr Gln Lys Ile Phe
115 120 125
Ala Ser His Phe Gly His Ile Ala Ile Ile Phe Leu Trp Thr Ser Gly
130 135 140
Thr Leu Phe His Val Ala Trp Gln Gly Asn Phe Glu Gln Trp Ile Lys
145 150 155 160
Asp Pro Leu Asn Ile Arg Pro Ile Ala His Ala Ile Trp Asp Pro His
165 170 175
Phe Gly Glu Gly Ala Val Asn Ala Phe Thr Gln Ala Gly Ala Ser Asn
180 185 190
Pro Val Asn Ile Ala Tyr Ser Gly Val Tyr His Trp Phe Tyr Thr Ile
195 200 205
Gly Met Thr Thr Asn Gln Glu Leu Tyr Ser Gly Ala Val Phe Leu Leu
210 215 220
Val Leu Ala Ser Leu Phe Leu Phe Ala Gly Trp Leu His Leu Gln Pro
225 230 235 240
Lys Phe Arg Pro Ser Leu Ala Trp Phe Lys Asn Ala Glu Ser Arg Leu
245 250 255
Asn His His Leu Ala Gly Leu Phe Gly Val Ser Ser Leu Ala Trp Ala
260 265 270
Gly His Leu Val His Val Ala Ile Pro Glu Ala Arg Gly Gln His Val
275 280 285
Gly Trp Asp Asn Phe Leu Ser Thr Pro Pro His Pro Ala Gly Leu Met
290 295 300
Pro Phe Phe Thr Gly Asn Trp Gly Val Tyr Ala Ala Cys Pro Asp Thr
305 310 315 320
Ala Gly His Ile Phe Gly Thr Ser Glu Gly Ala Gly Thr Ala Ile Leu
325 330 335
Thr Phe Leu Gly Gly Phe His Pro Gln Thr Glu Ser Leu Trp Leu Thr
340 345 350
Asp Ile Ala His His His Leu Ala Ile Ala Val Ile Phe Ile Ile Ala
355 360 365
Gly His Met Tyr Arg Thr Asn Trp Gly Ile Gly His Ser Ile Lys Glu
370 375 380
Ile Leu Asn Ala His Lys Gly Pro Leu Thr Gly Ala Gly His Thr Asn
385 390 395 400
Leu Tyr Asp Thr Ile Asn Asn Ser Leu His Phe Gln Leu Gly Leu Ala
405 410 415
Leu Ala Ser Leu Gly Val Ile Thr Ser Leu Val Ala Gln His Met Tyr
420 425 430
Ser Leu Pro Ser Tyr Ala Phe Ile Ala Gln Asp His Thr Thr Gln Ala
435 440 445
Ala Leu Tyr Thr His His Gln Tyr Ile Ala Gly Phe Leu Met Val Gly
450 455 460
Ala Phe Ala His Gly Ala Ile Phe Phe Val Arg Asp Tyr Asp Pro Val
465 470 475 480
Ala Asn Lys Asp Asn Val Leu Ala Arg Met Leu Glu His Lys Glu Ala
485 490 495
Leu Ile Ser His Leu Ser Trp Val Ser Leu Phe Leu Gly Phe His Thr
500 505 510
Leu Gly Leu Tyr Val His Asn Asp Val Val Val Ala Phe Gly Thr Pro
515 520 525
Glu Lys Gln Ile Leu Ile Glu Pro Val Phe Ala Gln Trp Ile Gln Ala
530 535 540
Thr Ser Gly Lys Ala Leu Tyr Gly Phe Asp Val Leu Leu Ser Asn Pro
545 550 555 560
Asp Ser Ile Ala Ser Thr Thr Gly Ala Ala Trp Leu Pro Gly Trp Leu
565 570 575
Asp Ala Ile Asn Ser Gly Thr Asn Ser Leu Phe Leu Thr Ile Gly Pro
580 585 590
Gly Asp Phe Leu Val His His Ala Ile Ala Leu Gly Leu His Thr Thr
595 600 605
Ala Leu Ile Leu Ile Lys Gly Ala Leu Asp Ala Arg Gly Ser Lys Leu
610 615 620
Met Pro Asp Lys Lys Asp Phe Gly Tyr Ser Phe Pro Cys Asp Gly Pro
625 630 635 640
Gly Arg Gly Gly Thr Cys Asp Ile Ser Ala Trp Asp Ala Phe Tyr Leu
645 650 655
Ala Met Phe Trp Met Leu Asn Thr Leu Gly Trp Leu Thr Phe Tyr Trp
660 665 670
His Trp Lys His Leu Gly Val Trp Ser Gly Asn Val Ala Gln Phe Asn
675 680 685
Glu Asn Ser Thr Tyr Leu Met Gly Trp Phe Arg Asp Tyr Leu Trp Ala
690 695 700
Asn Ser Ala Gln Leu Ile Asn Gly Tyr Asn Pro Cys Gly Val Asn Asn
705 710 715 720
Leu Ser Val Trp Ala Trp Met Phe Leu Phe Gly His Leu Val Trp Ala
725 730 735
Thr Gly Phe Met Phe Leu Ile Ser Trp Arg Gly Tyr Trp Gln Glu Leu
740 745 750
Ile Glu Thr Ile Val Trp Ala His Glu Arg Thr Pro Leu Ala Asn Leu
755 760 765
Val Arg Trp Lys Asp Lys Pro Val Ala Leu Ser Ile Val Gln Ala Arg
770 775 780
Leu Val Gly Leu Ala His Phe Thr Val Gly Tyr Val Leu Thr Tyr Ala
785 790 795 800
Ala Phe Leu Ile Ala Ser Thr Ala Gly Lys Phe Gly
805 810
<210>28
<211>731
<212>PRT
<213>集胞藻(Synechocystis sp.)
<220>
<221>其他特征
<223>D235C/Y634突变多肽
<400>28
Met Ala Thr Lys Phe Pro Lys Phe Ser Gln Asp Leu Ala Gln Asp Pro
1 5 10 15
Thr Thr Arg Arg Ile Trp Tyr Gly Ile Ala Thr Ala His Asp Phe Glu
20 25 30
Thr His Asp Gly Met Thr Glu Glu Asn Leu Tyr Gln Lys Ile Phe Ala
35 40 45
Ser His Phe Gly His Ile Ala Ile Ile Phe Leu Trp Thr Ser Gly Thr
50 55 60
Leu Phe His Val Ala Trp Gln Gly Asn Phe Glu Gln Trp Ile Lys Asp
65 70 75 80
Pro Leu Asn Ile Arg Pro Ile Ala His Ala Ile Trp Asp Pro His Phe
85 90 95
Gly Glu Gly Ala Val Asn Ala Phe Thr Gln Ala Gly Ala Ser Asn Pro
100 105 110
Val Asn Ile Ala Tyr Ser Gly Val Tyr His Trp Phe Tyr Thr Ile Gly
115 120 125
Met Thr Thr Asn Gln Glu Leu Tyr Ser Gly Ala Val Phe Leu Leu Val
130 135 140
Leu Ala Ser Leu Phe Leu Phe Ala Gly Trp Leu His Leu Gln Pro Lys
145 150 155 160
Phe Arg Pro Ser Leu Ala Trp Phe Lys Asn Ala Glu Ser Arg Leu Asn
165 170 175
His His Leu Ala Gly Leu Phe Gly Val Ser Ser Leu Ala Trp Ala Gly
180 185 190
His Leu Val His Val Ala Ile Pro Glu Ala Arg Gly Gln His Val Gly
195 200 205
Trp Asp Asn Phe Leu Ser Thr Pro Pro His Pro Ala Gly Leu Met Pro
210 215 220
Phe Phe Thr Gly Asn Trp Gly Val Tyr Ala Ala Cys Pro Asp Thr Ala
225 230 235 240
Gly His Ile Phe Gly Thr Ser Glu Gly Ala Gly Thr Ala Ile Leu Thr
245 250 255
Phe Leu Gly Gly Phe His Pro Gln Thr Glu Ser Leu Trp Leu Thr Asp
260 265 270
Ile Ala His His His Leu Ala Ile Ala Val Ile PheIle Ile Ala Gly
275 280 285
His Met Tyr Arg Thr Asn Trp Gly Ile Gly His Ser Ile Lys Glu Ile
290 295 300
Leu Asn Ala His Lys Gly Pro Leu Thr Gly Ala Gly His Thr Asn Leu
305 310 315 320
Tyr Asp Thr Ile Asn Asn Ser Leu His Phe Gln Leu Gly Leu Ala Leu
325 330 335
Ala Ser Leu Gly Val Ile Thr Ser Leu Val Ala Gln His Met Tyr Ser
340 345 350
Leu Pro Ser Tyr Ala Phe Ile Ala Gln Asp His Thr Thr Gln Ala Ala
355 360 365
Leu Tyr Thr His His Gln Tyr Ile Ala Gly Phe Leu Met Val Gly Ala
370 375 380
Phe Ala His Gly Ala Ile Phe Phe Val Arg Asp Tyr Asp Pro Val Ala
385 390 395 400
Asn Lys Asp Asn Val Leu Ala Arg Met Leu Glu His Lys Glu Ala Leu
405 410 415
Ile Ser His Leu Ser Trp Val Ser Leu Phe Leu Gly Phe His Thr Leu
420 425 430
Gly Leu Tyr Val His Asn Asp Val Val Val Ala Phe Gly Thr Pro Glu
435 440 445
Lys Gln Ile Leu Ile Glu Pro Val Phe Ala Gln Trp Ile Gln Ala Thr
450 455 460
Ser Gly Lys Ala Leu Tyr Gly Phe Asp Val Leu Leu Ser Asn Pro Asp
465 470 475 480
Ser Ile Ala Ser Thr Thr Gly Ala Ala Trp Leu Pro Gly Trp Leu Asp
485 490 495
Ala Ile Asn Ser Gly Thr Asn Ser Leu Phe Leu Thr Ile Gly Pro Gly
500 505 510
Asp Phe Leu Val His His Ala Ile Ala Leu Gly Leu His Thr Thr Ala
515 520 525
Leu Ile Leu Ile Lys Gly Ala Leu Asp Ala Arg Gly Ser Lys Leu Met
530 535 540
Pro Asp Lys Lys Asp Phe Gly Tyr Ser Phe Pro Cys Asp Gly Pro Gly
545 550 555 560
Arg Gly Gly Thr Cys Asp Ile Ser Ala Trp Asp Ala Phe Tyr Leu Ala
565 570 575
Met Phe Trp Met Leu Asn Thr Leu Gly Trp Leu Thr Phe Tyr Trp His
580 585 590
Trp Lys His Leu Gly Val Trp Ser Gly Asn Val Ala Gln Phe Asn Glu
595 600 605
Asn Ser Thr Tyr Leu Met Gly Trp Phe Arg Asp Tyr Leu Trp Ala Asn
610 615 620
Ser Ala Gln Leu Ile Asn Gly Tyr Asn Pro Cys Gly Val Asn Asn Leu
625 630 635 640
Ser Val Trp Ala Trp Met Phe Leu Phe Gly His Leu Val Trp Ala Thr
645 650 655
Gly Phe Met Phe Leu Ile Ser Trp Arg Gly Tyr Trp Gln Glu Leu Ile
660 665 670
Glu Thr Ile Val Trp Ala His Glu Arg Thr Pro Leu Ala Asn Leu Val
675 680 685
Arg Trp Lys Asp Lys Pro Val Ala Leu Ser Ile Val Gln Ala Arg Leu
690 695 700
Val Gly Leu Ala His Phe Thr Val Gly Tyr Val Leu Thr Tyr Ala Ala
705 710 715 720
Phe Leu Ile Ala Ser Thr Ala Gly Lys Phe Gly
725 730
<210>29
<211>731
<212>PRT
<213>集胞藻(Synechocystis sp.)
<220>
<221>其他特征
<223>S246C/Y634C突变多肽
<400>29
Met Ala Thr Lys Phe Pro Lys Phe Ser Gln Asp Leu Ala Gln Asp Pro
1 5 10 15
Thr Thr Arg Arg Ile Trp Tyr Gly Ile Ala Thr Ala His Asp Phe Glu
20 25 30
Thr His Asp Gly Met Thr Glu Glu Asn Leu Tyr Gln Lys Ile Phe Ala
35 40 45
Ser His Phe Gly His Ile Ala Ile Ile Phe Leu Trp Thr Ser Gly Thr
50 55 60
Leu Phe His Val Ala Trp Gln Gly Asn Phe Glu Gln Trp Ile Lys Asp
65 70 75 80
Pro Leu Asn Ile Arg Pro Ile Ala His Ala Ile Trp Asp Pro His Phe
85 90 95
Gly Glu Gly Ala Val Asn Ala Phe Thr Gln Ala Gly Ala Ser Asn Pro
100 105 110
Val Asn Ile Ala Tyr Ser Gly Val Tyr His Trp Phe Tyr Thr Ile Gly
115 120 125
Met Thr Thr Asn Gln Glu Leu Tyr Ser Gly Ala Val Phe Leu Leu Val
130 135 140
Leu Ala Ser Leu Phe Leu Phe Ala Gly Trp Leu His Leu Gln Pro Lys
145 150 155 160
Phe Arg Pro Ser Leu Ala Trp Phe Lys Asn Ala Glu Ser Arg Leu Asn
165 170 175
His His Leu Ala Gly Leu Phe Gly Val Ser Ser Leu Ala Trp Ala Gly
180 185 190
His Leu Val His Val Ala Ile Pro Glu Ala Arg Gly Gln His Val Gly
195 200 205
Trp Asp Asn Phe Leu Ser Thr Pro Pro His Pro Ala Gly Leu Met Pro
210 215 220
Phe Phe Thr Gly Asn Trp Gly Val Tyr Ala Ala Asp Pro Asp Thr Ala
225 230 235 240
Gly His Ile Phe Gly Thr Cys Glu Gly Ala Gly Thr Ala Ile Leu Thr
245 250 255
Phe Leu Gly Gly Phe His Pro Gln Thr Glu Ser Leu Trp Leu Thr Asp
260 265 270
Ile Ala His His His Leu Ala Ile Ala Val Ile Phe Ile Ile Ala Gly
275 280 285
His Met Tyr Arg Thr Asn Trp Gly Ile Gly His Ser Ile Lys Glu Ile
290 295 300
Leu Asn Ala His Lys Gly Pro Leu Thr Gly Ala Gly His Thr Asn Leu
305 310 315 320
Tyr Asp Thr Ile Asn Asn Ser Leu His Phe Gln Leu Gly Leu Ala Leu
325 330 335
Ala Ser Leu Gly Val Ile Thr Ser Leu Val Ala Gln His Met Tyr Ser
340 345 350
Leu Pro Ser Tyr Ala Phe Ile Ala Gln Asp His Thr Thr Gln Ala Ala
355 360 365
Leu Tyr Thr His His Gln Tyr Ile Ala Gly Phe Leu Met Val Gly Ala
370 375 380
Phe Ala His Gly Ala Ile Phe Phe Val Arg Asp Tyr Asp Pro Val Ala
385 390 395 400
Asn Lys Asp Asn Val Leu Ala Arg Met Leu Glu His Lys Glu Ala Leu
405 410 415
Ile Ser His Leu Ser Trp Val Ser Leu Phe Leu Gly Phe His Thr Leu
420 425 430
Gly Leu Tyr Val His Asn Asp Val Val Val Ala Phe Gly Thr Pro Glu
435 440 445
Lys Gln Ile Leu Ile Glu Pro Val Phe Ala Gln Trp Ile Gln Ala Thr
450 455 460
Ser Gly Lys Ala Leu Tyr Gly Phe Asp Val Leu Leu Ser Asn Pro Asp
465 470 475 480
Ser Ile Ala Ser Thr Thr Gly Ala Ala Trp Leu Pro Gly Trp Leu Asp
485 490 495
Ala Ile Asn Ser Gly Thr Asn Ser Leu Phe Leu Thr Ile Gly Pro Gly
500 505 510
Asp Phe Leu Val His His Ala Ile Ala Leu Gly Leu His Thr Thr Ala
515 520 525
Leu Ile Leu Ile Lys Gly Ala Leu Asp Ala Arg Gly Ser Lys Leu Met
530 535 540
Pro Asp Lys Lys Asp Phe Gly Tyr Ser Phe Pro Cys Asp Gly Pro Gly
545 550 555 560
Arg Gly Gly Thr Cys Asp Ile Ser Ala Trp Asp Ala Phe Tyr Leu Ala
565 570 575
Met Phe Trp Met Leu Asn Thr Leu Gly Trp Leu Thr Phe Tyr Trp His
580 585 590
Trp Lys His Leu Gly Val Trp Ser Gly Asn Val Ala Gln Phe Asn Glu
595 600 605
Asn Ser Thr Tyr Leu Met Gly Trp Phe Arg Asp Tyr Leu Trp Ala Asn
610 615 620
Ser Ala Gln Leu Ile Asn Gly Tyr Asn Pro Cys Gly Val Asn Asn Leu
625 630 635 640
Ser Val Trp Ala Trp Met Phe Leu Phe Gly His Leu Val Trp Ala Thr
645 650 655
Gly Phe Met Phe Leu Ile Ser Trp Arg Gly Tyr Trp Gln Glu Leu Ile
660 665 670
Glu Thr Ile Val Trp Ala His Glu Arg Thr Pro Leu Ala Asn Leu Val
675 680 685
Arg Trp Lys Asp Lys Pro Val Ala Leu Ser Ile Val Gln Ala Arg Leu
690 695 700
Val Gly Leu Ala His Phe Thr Val Gly Tyr Val Leu Thr Tyr Ala Ala
705 710 715 720
Phe Leu Ile Ala Ser Thr Ala Gly Lys Phe Gly
725 730
<210>30
<211>2196
<212>DNA
<213>集胞藻(Synechocystis sp.)
<220>
<221>其他特征
<223>D479C突变多核苷酸
<400>30
atggcaacta aatttcctaa atttagccag gatctcgccc aagacccgac tacacggcgt 60
atttggtacg ggattgctac ggcccacgac tttgaaaccc acgatgggat gactgaggag 120
aatctttacc aaaagatttt tgcctcccac tttggacaca tcgccatcat tttcctgtgg 180
acgtctggca cccttttcca cgttgcgtgg caaggtaatt ttgaacaatg gattaaagat 240
cctttaaaca ttcgcccgat cgcccatgcg atttgggacc cccactttgg cgagggagct 300
gtcaatgctt tcacccaagc tggggcttct aacccggtta acattgctta ttccggggtt 360
taccactggt tctacaccat tggtatgacc accaaccaag agctctattc tggtgcggtc 420
ttcctattag tgttagcttc cctgttttta tttgcaggct ggttacacct ccaaccgaag 480
ttccgtccta gcttagcctg gttcaaaaat gccgaatccc gcttaaatca ccacctggct 540
ggtttgttcg gggttagctc cttggcttgg gctggtcact tggtccacgt tgcgattccc 600
gaagcccggg gtcaacacgt tggttgggat aacttcctgt ctactcctcc ccaccccgct 660
ggtttgatgc ctttcttcac cggcaactgg ggcgtgtatg cggcggatcc cgacactgct 720
ggccacattt ttggtacttc tgaaggtgct ggtaccgcaa tcctgacctt cctgggtggt 780
ttccatcccc aaacggaatc tctctggttg acggacattg cccaccacca tttggcgatc 840
gccgtgatct tcatcattgc tggtcacatg tatcgcacca actggggcat tggccacagc 900
atcaaagaaa ttcttaatgc ccataaaggt cccctaaccg gcgcaggcca taccaacctg 960
tacgacacca tcaataactc cctccacttc caactcggct tagccttggc aagcttaggg 1020
gttattactt ccctggtggc gcagcacatg tactccctgc cctcctacgc ctttattgcc 1080
caggaccaca ccacccaggc agccctctac acccatcacc agtacattgc tggattcttg 1140
atggttggtg cttttgccca cggtgccatc ttctttgtcc gggattacga tcccgtggcc 1200
aataaagata acgtgctggc ccgcatgctt gaacacaaag aggctctgat ttcccactta 1260
agctgggtgt ccctcttctt gggcttccac acccttggtc tttatgtcca taacgatgtg 1320
gtggtggcct tcggtactcc cgaaaaacaa attctgatcg agcccgtttt tgcccaatgg 1380
attcaagcaa cttccggtaa agctctctat ggctttgatg ttttgctctc caatccttgc 1440
agcattgctt ccaccactgg agccgcttgg ttacccggct ggttggatgc tatcaacagc 1500
ggcaccaact ctctgttctt gaccattggc cctggcgact tcttggttca ccacgcgatc 1560
gccctagggt tgcacaccac tgccctgatt ctaatcaaag gtgctttgga tgcccgtggt 1620
tctaagttga tgccggacaa aaaggacttc ggttactcct tcccctgtga tggccccggc 1680
cgtggcggta cctgcgacat ctctgcttgg gatgccttct acctagccat gttctggatg 1740
ctgaacacct tgggttggtt gaccttctac tggcactgga aacacctcgg tgtttggagc 1800
ggtaacgttg ctcagttcaa cgaaaactcc acctacctga tgggttggtt ccgggattac 1860
ctctgggcga actctgctca gttaatcaat ggttacaacc cctacggtgt caacaatctg 1920
tcagtttggg cttggatgtt ccttttcgga cacctggtct gggctactgg cttcatgttc 1980
ttgatctctt ggcggggtta ctggcaagag ttgattgaaa ccatcgtttg ggcccacgag 2040
cgcactcctt tggcgaactt ggttcgttgg aaagataagc ccgttgcgtt gtccattgtt 2100
caagcccgtt tggttggttt agcccacttc accgttggtt atgtgctcac ctatgcggca 2160
ttcctaattg cttccacagc cggtaagttc ggttaa 2196
<210>31
<211>2196
<212>DNA
<213>集胞藻(Synechocystis sp.)
<220>
<221>其他特征
<223>S499C突变多核苷酸
<400>31
atggcaacta aatttcctaa atttagccag gatctcgccc aagacccgac tacacggcgt 60
atttggtacg ggattgctac ggcccacgac tttgaaaccc acgatgggat gactgaggag 120
aatctttacc aaaagatttt tgcctcccac tttggacaca tcgccatcat tttcctgtgg 180
acgtctggca cccttttcca cgttgcgtgg caaggtaatt ttgaacaatg gattaaagat 240
cctttaaaca ttcgcccgat cgcccatgcg atttgggacc cccactttgg cgagggagct 300
gtcaatgctt tcacccaagc tggggcttct aacccggtta acattgctta ttccggggtt 360
taccactggt tctacaccat tggtatgacc accaaccaag agctctattc tggtgcggtc 420
ttcctattag tgttagcttc cctgttttta tttgcaggct ggttacacct ccaaccgaag 480
ttccgtccta gcttagcctg gttcaaaaat gccgaatccc gcttaaatca ccacctggct 540
ggtttgttcg gggttagctc cttggcttgg gctggtcact tggtccacgt tgcgattccc 600
gaagcccggg gtcaacacgt tggttgggat aacttcctgt ctactcctcc ccaccccgct 660
ggtttgatgc ctttcttcac cggcaactgg ggcgtgtatg cggcggatcc cgacactgct 720
ggccacattt ttggtacttc tgaaggtgct ggtaccgcaa tcctgacctt cctgggtggt 780
ttccatcccc aaacggaatc tctctggttg acggacattg cccaccacca tttggcgatc 840
gccgtgatct tcatcattgc tggtcacatg tatcgcacca actggggcat tggccacagc 900
atcaaagaaa ttcttaatgc ccataaaggt cccctaaccg gcgcaggcca taccaacctg 960
tacgacacca tcaataactc cctccacttc caactcggct tagccttggc aagcttaggg 1020
gttattactt ccctggtggc gcagcacatg tactccctgc cctcctacgc ctttattgcc 1080
caggaccaca ccacccaggc agccctctac acccatcacc agtacattgc tggattcttg 1140
atggttggtg cttttgccca cggtgccatc ttctttgtcc gggattacga tcccgtggcc 1200
aataaagata acgtgctggc ccgcatgctt gaacacaaag aggctctgat ttcccactta 1260
agctgggtgt ccctcttctt gggcttccac acccttggtc tttatgtcca taacgatgtg 1320
gtggtggcct tcggtactcc cgaaaaacaa attctgatcg agcccgtttt tgcccaatgg 1380
attcaagcaa cttccggtaa agctctctat ggctttgatg ttttgctctc caatcctgac 1440
agcattgctt ccaccactgg agccgcttgg ttacccggct ggttggatgc tatcaactgc 1500
ggcaccaact ctctgttctt gaccattggc cctggcgact tcttggttca ccacgcgatc 1560
gccctagggt tgcacaccac tgccctgatt ctaatcaaag gtgctttgga tgcccgtggt 1620
tctaagttga tgccggacaa aaaggacttc ggttactcct tcccctgtga tggccccggc 1680
cgtggcggta cctgcgacat ctctgcttgg gatgccttct acctagccat gttctggatg 1740
ctgaacacct tgggttggtt gaccttctac tggcactgga aacacctcgg tgtttggagc 1800
ggtaacgttg ctcagttcaa cgaaaactcc acctacctga tgggttggtt ccgggattac 1860
ctctgggcga actctgctca gttaatcaat ggttacaacc cctacggtgt caacaatctg 1920
tcagtttggg cttggatgtt ccttttcgga cacctggtct gggctactgg cttcatgttc 1980
ttgatctctt ggcggggtta ctggcaagag ttgattgaaa ccatcgtttg ggcccacgag 2040
cgcactcctt tggcgaactt ggttcgttgg aaagataagc ccgttgcgtt gtccattgtt 2100
caagcccgtt tggttggttt agcccacttc accgttggtt atgtgctcac ctatgcggca 2160
ttcctaattg cttccacagc cggtaagttc ggttaa 2196
<210>32
<211>2196
<212>DNA
<213>集胞藻(Synechocystis sp.)
<220>
<221>其他特征
<223>S599C突变多核苷酸
<400>32
atggcaacta aatttcctaa atttagccag gatctcgccc aagacccgac tacacggcgt 60
atttggtacg ggattgctac ggcccacgac tttgaaaccc acgatgggat gactgaggag 120
aatctttacc aaaagatttt tgcctcccac tttggacaca tcgccatcat tttcctgtgg 180
acgtctggca cccttttcca cgttgcgtgg caaggtaatt ttgaacaatg gattaaagat 240
cctttaaaca ttcgcccgat cgcccatgcg atttgggacc cccactttgg cgagggagct 300
gtcaatgctt tcacccaagc tggggcttct aacccggtta acattgctta ttccggggtt 360
taccactggt tctacaccat tggtatgacc accaaccaag agctctattc tggtgcggtc 420
ttcctattag tgttagcttc cctgttttta tttgcaggct ggttacacct ccaaccgaag 480
ttccgtccta gcttagcctg gttcaaaaat gccgaatccc gcttaaatca ccacctggct 540
ggtttgttcg gggttagctc cttggcttgg gctggtcact tggtccacgt tgcgattccc 600
gaagcccggg gtcaacacgt tggttgggat aacttcctgt ctactcctcc ccaccccgct 660
ggtttgatgc ctttcttcac cggcaactgg ggcgtgtatg cggcggatcc cgacactgct 720
ggccacattt ttggtacttc tgaaggtgct ggtaccgcaa tcctgacctt cctgggtggt 780
ttccatcccc aaacggaatc tctctggttg acggacattg cccaccacca tttggcgatc 840
gccgtgatct tcatcattgc tggtcacatg tatcgcacca actggggcat tggccacagc 900
atcaaagaaa ttcttaatgc ccataaaggt cccctaaccg gcgcaggcca taccaacctg 960
tacgacacca tcaataactc cctccacttc caactcggct tagccttggc aagcttaggg 1020
gttattactt ccctggtggc gcagcacatg tactccctgc cctcctacgc ctttattgcc 1080
caggaccaca ccacccaggc agccctctac acccatcacc agtacattgc tggattcttg 1140
atggttggtg cttttgccca cggtgccatc ttctttgtcc gggattacga tcccgtggcc 1200
aataaagata acgtgctggc ccgcatgctt gaacacaaag aggctctgat ttcccactta 1260
agctgggtgt ccctcttctt gggcttccac acccttggtc tttatgtcca taacgatgtg 1320
gtggtggcct tcggtactcc cgaaaaacaa attctgatcg agcccgtttt tgcccaatgg 1380
attcaagcaa cttccggtaa agctctctat ggctttgatg ttttgctctc caatcctgac 1440
agcattgctt ccaccactgg agccgcttgg ttacccggct ggttggatgc tatcaacagc 1500
ggcaccaact ctctgttctt gaccattggc cctggcgact tcttggttca ccacgcgatc 1560
gccctagggt tgcacaccac tgccctgatt ctaatcaaag gtgctttgga tgcccgtggt 1620
tctaagttga tgccggacaa aaaggacttc ggttactcct tcccctgtga tggccccggc 1680
cgtggcggta cctgcgacat ctctgcttgg gatgccttct acctagccat gttctggatg 1740
ctgaacacct tgggttggtt gaccttctac tggcactgga aacacctcgg tgtttggtgc 1800
ggtaacgttg ctcagttcaa cgaaaactcc acctacctga tgggttggtt ccgggattac 1860
ctctgggcga actctgctca gttaatcaat ggttacaacc cctacggtgt caacaatctg 1920
tcagtttggg cttggatgtt ccttttcgga cacctggtct gggctactgg cttcatgttc 1980
ttgatctctt ggcggggtta ctggcaagag ttgattgaaa ccatcgtttg ggcccacgag 2040
cgcactcctt tggcgaactt ggttcgttgg aaagataagc ccgttgcgtt gtccattgtt 2100
caagcccgtt tggttggttt agcccacttc accgttggtt atgtgctcac ctatgcggca 2160
ttcctaattg cttccacagc cggtaagttc ggttaa 2196
<210>33
<211>2196
<212>DNA
<213>集胞藻(Synechocystis sp.)
<220>
<221>其他特征
<223>Y634C突变多核苷酸
<400>33
atggcaacta aatttcctaa atttagccag gatctcgccc aagacccgac tacacggcgt 60
atttggtacg ggattgctac ggcccacgac tttgaaaccc acgatgggat gactgaggag 120
aatctttacc aaaagatttt tgcctcccac tttggacaca tcgccatcat tttcctgtgg 180
acgtctggca cccttttcca cgttgcgtgg caaggtaatt ttgaacaatg gattaaagat 240
cctttaaaca ttcgcccgat cgcccatgcg atttgggacc cccactttgg cgagggagct 300
gtcaatgctt tcacccaagc tggggcttct aacccggtta acattgctta ttccggggtt 360
taccactggt tctacaccat tggtatgacc accaaccaag agctctattc tggtgcggtc 420
ttcctattag tgttagcttc cctgttttta tttgcaggct ggttacacct ccaaccgaag 480
ttccgtccta gcttagcctg gttcaaaaat gccgaatccc gcttaaatca ccacctggct 540
ggtttgttcg gggttagctc cttggcttgg gctggtcact tggtccacgt tgcgattccc 600
gaagcccggg gtcaacacgt tggttgggat aacttcctgt ctactcctcc ccaccccgct 660
ggtttgatgc ctttcttcac cggcaactgg ggcgtgtatg cggcggatcc cgacactgct 720
ggccacattt ttggtacttc tgaaggtgct ggtaccgcaa tcctgacctt cctgggtggt 780
ttccatcccc aaacggaatc tctctggttg acggacattg cccaccacca tttggcgatc 840
gccgtgatct tcatcattgc tggtcacatg tatcgcacca actggggcat tggccacagc 900
atcaaagaaa ttcttaatgc ccataaaggt cccctaaccg gcgcaggcca taccaacctg 960
tacgacacca tcaataactc cctccacttc caactcggct tagccttggc aagcttaggg 1020
gttattactt ccctggtggc gcagcacatg tactccctgc cctcctacgc ctttattgcc 1080
caggaccaca ccacccaggc agccctctac acccatcacc agtacattgc tggattcttg 1140
atggttggtg cttttgccca cggtgccatc ttctttgtcc gggattacga tcccgtggcc 1200
aataaagata acgtgctggc ccgcatgctt gaacacaaag aggctctgat ttcccactta 1260
agctgggtgt ccctcttctt gggcttccac acccttggtc tttatgtcca taacgatgtg 1320
gtggtggcct tcggtactcc cgaaaaacaa attctgatcg agcccgtttt tgcccaatgg 1380
attcaagcaa cttccggtaa agctctctat ggctttgatg ttttgctctc caatcctgac 1440
agcattgctt ccaccactgg agccgcttgg ttacccggct ggttggatgc tatcaacagc 1500
ggcaccaact ctctgttctt gaccattggc cctggcgact tcttggttca ccacgcgatc 1560
gccctagggt tgcacaccac tgccctgatt ctaatcaaag gtgctttgga tgcccgtggt 1620
tctaagttga tgccggacaa aaaggacttc ggttactcct tcccctgtga tggccccggc 1680
cgtggcggta cctgcgacat ctctgcttgg gatgccttct acctagccat gttctggatg 1740
ctgaacacct tgggttggtt gaccttctac tggcactgga aacacctcgg tgtttggagc 1800
ggtaacgttg ctcagttcaa cgaaaactcc acctacctga tgggttggtt ccgggattac 1860
ctctgggcga actctgctca gttaatcaat ggttacaacc cctgcggtgt caacaatctg 1920
tcagtttggg cttggatgtt ccttttcgga cacctggtct gggctactgg cttcatgttc 1980
ttgatctctt ggcggggtta ctggcaagag ttgattgaaa ccatcgtttg ggcccacgag 2040
cgcactcctt tggcgaactt ggttcgttgg aaagataagc ccgttgcgtt gtccattgtt 2100
caagcccgtt tggttggttt agcccacttc accgttggtt atgtgctcac ctatgcggca 2160
ttcctaattg cttccacagc cggtaagttc ggttaa 2196
<210>34
<211>2196
<212>DNA
<213>集胞藻(Synechocystis sp.)
<220>
<221>其他特征
<223>D235C突变多核苷酸
<400>34
atggcaacta aatttcctaa atttagccag gatctcgccc aagacccgac tacacggcgt 60
atttggtacg ggattgctac ggcccacgac tttgaaaccc acgatgggat gactgaggag 120
aatctttacc aaaagatttt tgcctcccac tttggacaca tcgccatcat tttcctgtgg 180
acgtctggca cccttttcca cgttgcgtgg caaggtaatt ttgaacaatg gattaaagat 240
cctttaaaca ttcgcccgat cgcccatgcg atttgggacc cccactttgg cgagggagct 300
gtcaatgctt tcacccaagc tggggcttct aacccggtta acattgctta ttccggggtt 360
taccactggt tctacaccat tggtatgacc accaaccaag agctctattc tggtgcggtc 420
ttcctattag tgttagcttc cctgttttta tttgcaggct ggttacacct ccaaccgaag 480
ttccgtccta gcttagcctg gttcaaaaat gccgaatccc gcttaaatca ccacctggct 540
ggtttgttcg gggttagctc cttggcttgg gctggtcact tggtccacgt tgcgattccc 600
gaagcccggg gtcaacacgt tggttgggat aacttcctgt ctactcctcc ccaccccgct 660
ggtttgatgc ctttcttcac cggcaactgg ggcgtgtatg cggcgtgtcc cgacactgct 720
ggccacattt ttggtacttc tgaaggtgct ggtaccgcaa tcctgacctt cctgggtggt 780
ttccatcccc aaacggaatc tctctggttg acggacattg cccaccacca tttggcgatc 840
gccgtgatct tcatcattgc tggtcacatg tatcgcacca actggggcat tggccacagc 900
atcaaagaaa ttcttaatgc ccataaaggt cccctaaccg gcgcaggcca taccaacctg 960
tacgacacca tcaataactc cctccacttc caactcggct tagccttggc aagcttaggg 1020
gttattactt ccctggtggc gcagcacatg tactccctgc cctcctacgc ctttattgcc 1080
caggaccaca ccacccaggc agccctctac acccatcacc agtacattgc tggattcttg 1140
atggttggtg cttttgccca cggtgccatc ttctttgtcc gggattacga tcccgtggcc 1200
aataaagata acgtgctggc ccgcatgctt gaacacaaag aggctctgat ttcccactta 1260
agctgggtgt ccctcttctt gggcttccac acccttggtc tttatgtcca taacgatgtg 1320
gtggtggcct tcggtactcc cgaaaaacaa attctgatcg agcccgtttt tgcccaatgg 1380
attcaagcaa cttccggtaa agctctctat ggctttgatg ttttgctctc caatcctgac 1440
agcattgctt ccaccactgg agccgcttgg ttacccggct ggttggatgc tatcaacagc 1500
ggcaccaact ctctgttctt gaccattggc cctggcgact tcttggttca ccacgcgatc 1560
gccctagggt tgcacaccac tgccctgatt ctaatcaaag gtgctttgga tgcccgtggt 1620
tctaagttga tgccggacaa aaaggacttc ggttactcct tcccctgtga tggccccggc 1680
cgtggcggta cctgcgacat ctctgcttgg gatgccttct acctagccat gttctggatg 1740
ctgaacacct tgggttggtt gaccttctac tggcactgga aacacctcgg tgtttggagc 1800
ggtaacgttg ctcagttcaa cgaaaactcc acctacctga tgggttggtt ccgggattac 1860
ctctgggcga actctgctca gttaatcaat ggttacaacc cctacggtgt caacaatctg 1920
tcagtttggg cttggatgtt ccttttcgga cacctggtct gggctactgg cttcatgttc 1980
ttgatctctt ggcggggtta ctggcaagag ttgattgaaa ccatcgtttg ggcccacgag 2040
cgcactcctt tggcgaactt ggttcgttgg aaagataagc ccgttgcgtt gtccattgtt 2100
caagcccgtt tggttggttt agcccacttc accgttggtt atgtgctcac ctatgcggca 2160
ttcctaattg cttccacagc cggtaagttc ggttaa 2196
<210>35
<211>2196
<212>DNA
<213>集胞藻(Synechocystis sp.)
<220>
<221>其他特征
<223>S246C突变多核苷酸
<400>35
atggcaacta aatttcctaa atttagccag gatctcgccc aagacccgac tacacggcgt 60
atttggtacg ggattgctac ggcccacgac tttgaaaccc acgatgggat gactgaggag 120
aatctttacc aaaagatttt tgcctcccac tttggacaca tcgccatcat tttcctgtgg 180
acgtctggca cccttttcca cgttgcgtgg caaggtaatt ttgaacaatg gattaaagat 240
cctttaaaca ttcgcccgat cgcccatgcg atttgggacc cccactttgg cgagggagct 300
gtcaatgctt tcacccaagc tggggcttct aacccggtta acattgctta ttccggggtt 360
taccactggt tctacaccat tggtatgacc accaaccaag agctctattc tggtgcggtc 420
ttcctattag tgttagcttc cctgttttta tttgcaggct ggttacacct ccaaccgaag 480
ttccgtccta gcttagcctg gttcaaaaat gccgaatccc gcttaaatca ccacctggct 540
ggtttgttcg gggttagctc cttggcttgg gctggtcact tggtccacgt tgcgattccc 600
gaagcccggg gtcaacacgt tggttgggat aacttcctgt ctactcctcc ccaccccgct 660
ggtttgatgc ctttcttcac cggcaactgg ggcgtgtatg cggcggatcc cgacactgct 720
ggccacattt ttggttgttc tgaaggtgct ggtaccgcaa tcctgacctt cctgggtggt 780
ttccatcccc aaacggaatc tctctggttg acggacattg cccaccacca tttggcgatc 840
gccgtgatct tcatcattgc tggtcacatg tatcgcacca actggggcat tggccacagc 900
atcaaagaaa ttcttaatgc ccataaaggt cccctaaccg gcgcaggcca taccaacctg 960
tacgacacca tcaataactc cctccacttc caactcggct tagccttggc aagcttaggg 1020
gttattactt ccctggtggc gcagcacatg tactccctgc cctcctacgc ctttattgcc 1080
caggaccaca ccacccaggc agccctctac acccatcacc agtacattgc tggattcttg 1140
atggttggtg cttttgccca cggtgccatc ttctttgtcc gggattacga tcccgtggcc 1200
aataaagata acgtgctggc ccgcatgctt gaacacaaag aggctctgat ttcccactta 1260
agctgggtgt ccctcttctt gggcttccac acccttggtc tttatgtcca taacgatgtg 1320
gtggtggcct tcggtactcc cgaaaaacaa attctgatcg agcccgtttt tgcccaatgg 1380
attcaagcaa cttccggtaa agctctctat ggctttgatg ttttgctctc caatcctgac 1440
agcattgctt ccaccactgg agccgcttgg ttacccggct ggttggatgc tatcaacagc 1500
ggcaccaact ctctgttctt gaccattggc cctggcgact tcttggttca ccacgcgatc 1560
gccctagggt tgcacaccac tgccctgatt ctaatcaaag gtgctttgga tgcccgtggt 1620
tctaagttga tgccggacaa aaaggacttc ggttactcct tcccctgtga tggccccggc 1680
cgtggcggta cctgcgacat ctctgcttgg gatgccttct acctagccat gttctggatg 1740
ctgaacacct tgggttggtt gaccttctac tggcactgga aacacctcgg tgtttggagc 1800
ggtaacgttg ctcagttcaa cgaaaactcc acctacctga tgggttggtt ccgggattac 1860
ctctgggcga actctgctca gttaatcaat ggttacaacc cctacggtgt caacaatctg 1920
tcagtttggg cttggatgtt ccttttcgga cacctggtct gggctactgg cttcatgttc 1980
ttgatctctt ggcggggtta ctggcaagag ttgattgaaa ccatcgtttg ggcccacgag 2040
cgcactcctt tggcgaactt ggttcgttgg aaagataagc ccgttgcgtt gtccattgtt 2100
caagcccgtt tggttggttt agcccacttc accgttggtt atgtgctcac ctatgcggca 2160
ttcctaattg cttccacagc cggtaagttc ggttaa 2196
<210>36
<211>246
<212>DNA
<213>集胞藻(Synechocystis sp.)
<220>
<221>其他特征
<223>W31C突变多核苷酸
<400>36
atgtcccata gtgtaaaaat ttacgatacc tgtattggtt gtacccaatg cgtccgggct 60
tgccccctcg atgttctaga aatggtgccc tggtgtggtt gtaaagccgc tcagatcgcc 120
tcctccccta gaaccgagga ttgtgttggc tgcaaacgtt gtgagaccgc ctgtcccaca 180
gactttttaa gtatccgagt ttatttgggt gccgaaacca cccgcagtat gggtttagct 240
tactaa 246
<210>37
<211>2196
<212>DNA
<213>集胞藻(Synechocystis sp.)
<220>
<221>其他特征
<223>D235C/Y634突变多核苷酸
<400>37
atggcaacta aatttcctaa atttagccag gatctcgccc aagacccgac tacacggcgt 60
atttggtacg ggattgctac ggcccacgac tttgaaaccc acgatgggat gactgaggag 120
aatctttacc aaaagatttt tgcctcccac tttggacaca tcgccatcat tttcctgtgg 180
acgtctggca cccttttcca cgttgcgtgg caaggtaatt ttgaacaatg gattaaagat 240
cctttaaaca ttcgcccgat cgcccatgcg atttgggacc cccactttgg cgagggagct 300
gtcaatgctt tcacccaagc tggggcttct aacccggtta acattgctta ttccggggtt 360
taccactggt tctacaccat tggtatgacc accaaccaag agctctattc tggtgcggtc 420
ttcctattag tgttagcttc cctgttttta tttgcaggct ggttacacct ccaaccgaag 480
ttccgtccta gcttagcctg gttcaaaaat gccgaatccc gcttaaatca ccacctggct 540
ggtttgttcg gggttagctc cttggcttgg gctggtcact tggtccacgt tgcgattccc 600
gaagcccggg gtcaacacgt tggttgggat aacttcctgt ctactcctcc ccaccccgct 660
ggtttgatgc ctttcttcac cggcaactgg ggcgtgtatg cggcgtgtcc cgacactgct 720
ggccacattt ttggtacttc tgaaggtgct ggtaccgcaa tcctgacctt cctgggtggt 780
ttccatcccc aaacggaatc tctctggttg acggacattg cccaccacca tttggcgatc 840
gccgtgatct tcatcattgc tggtcacatg tatcgcacca actggggcat tggccacagc 900
atcaaagaaa ttcttaatgc ccataaaggt cccctaaccg gcgcaggcca taccaacctg 960
tacgacacca tcaataactc cctccacttc caactcggct tagccttggc aagcttaggg 1020
gttattactt ccctggtggc gcagcacatg tactccctgc cctcctacgc ctttattgcc 1080
caggaccaca ccacccaggc agccctctac acccatcacc agtacattgc tggattcttg 1140
atggttggtg cttttgccca cggtgccatc ttctttgtcc gggattacga tcccgtggcc 1200
aataaagata acgtgctggc ccgcatgctt gaacacaaag aggctctgat ttcccactta 1260
agctgggtgt ccctcttctt gggcttccac acccttggtc tttatgtcca taacgatgtg 1320
gtggtggcct tcggtactcc cgaaaaacaa attctgatcg agcccgtttt tgcccaatgg 1380
attcaagcaa cttccggtaa agctctctat ggctttgatg ttttgctctc caatcctgac 1440
agcattgctt ccaccactgg agccgcttgg ttacccggct ggttggatgc tatcaacagc 1500
ggcaccaact ctctgttctt gaccattggc cctggcgact tcttggttca ccacgcgatc 1560
gccctagggt tgcacaccac tgccctgatt ctaatcaaag gtgctttgga tgcccgtggt 1620
tctaagttga tgccggacaa aaaggacttc ggttactcct tcccctgtga tggccccggc 1680
cgtggcggta cctgcgacat ctctgcttgg gatgccttct acctagccat gttctggatg 1740
ctgaacacct tgggttggtt gaccttctac tggcactgga aacacctcgg tgtttggagc 1800
ggtaacgttg ctcagttcaa cgaaaactcc acctacctga tgggttggtt ccgggattac 1860
ctctgggcga actctgctca gttaatcaat ggttacaacc cctgcggtgt caacaatctg 1920
tcagtttggg cttggatgtt ccttttcgga cacctggtct gggctactgg cttcatgttc 1980
ttgatctctt ggcggggtta ctggcaagag ttgattgaaa ccatcgtttg ggcccacgag 2040
cgcactcctt tggcgaactt ggttcgttgg aaagataagc ccgttgcgtt gtccattgtt 2100
caagcccgtt tggttggttt agcccacttc accgttggtt atgtgctcac ctatgcggca 2160
ttcctaattg cttccacagc cggtaagttc ggttaa 2196
<210>38
<211>2196
<212>DNA
<213>集胞藻(Synechocystis sp.)
<220>
<221>其他特征
<223>S246C/Y634C突变多核苷酸
<400>38
atggcaacta aatttcctaa atttagccag gatctcgccc aagacccgac tacacggcgt 60
atttggtacg ggattgctac ggcccacgac tttgaaaccc acgatgggat gactgaggag 120
aatctttacc aaaagatttt tgcctcccac tttggacaca tcgccatcat tttcctgtgg 180
acgtctggca cccttttcca cgttgcgtgg caaggtaatt ttgaacaatg gattaaagat 240
cctttaaaca ttcgcccgat cgcccatgcg atttgggacc cccactttgg cgagggagct 300
gtcaatgctt tcacccaagc tggggcttct aacccggtta acattgctta ttccggggtt 360
taccactggt tctacaccat tggtatgacc accaaccaag agctctattc tggtgcggtc 420
ttcctattag tgttagcttc cctgttttta tttgcaggct ggttacacct ccaaccgaag 480
ttccgtccta gcttagcctg gttcaaaaat gccgaatccc gcttaaatca ccacctggct 540
ggtttgttcg gggttagctc cttggcttgg gctggtcact tggtccacgt tgcgattccc 600
gaagcccggg gtcaacacgt tggttgggat aacttcctgt ctactcctcc ccaccccgct 660
ggtttgatgc ctttcttcac cggcaactgg ggcgtgtatg cggcggatcc cgacactgct 720
ggccacattt ttggttgttc tgaaggtgct ggtaccgcaa tcctgacctt cctgggtggt 780
ttccatcccc aaacggaatc tctctggttg acggacattg cccaccacca tttggcgatc 840
gccgtgatct tcatcattgc tggtcacatg tatcgcacca actggggcat tggccacagc 900
atcaaagaaa ttcttaatgc ccataaaggt cccctaaccg gcgcaggcca taccaacctg 960
tacgacacca tcaataactc cctccacttc caactcggct tagccttggc aagcttaggg 1020
gttattactt ccctggtggc gcagcacatg tactccctgc cctcctacgc ctttattgcc 1080
caggaccaca ccacccaggc agccctctac acccatcacc agtacattgc tggattcttg 1140
atggttggtg cttttgccca cggtgccatc ttctttgtcc gggattacga tcccgtggcc 1200
aataaagata acgtgctggc ccgcatgctt gaacacaaag aggctctgat ttcccactta 1260
agctgggtgt ccctcttctt gggcttccac acccttggtc tttatgtcca taacgatgtg 1320
gtggtggcct tcggtactcc cgaaaaacaa attctgatcg agcccgtttt tgcccaatgg 1380
attcaagcaa cttccggtaa agctctctat ggctttgatg ttttgctctc caatcctgac 1440
agcattgctt ccaccactgg agccgcttgg ttacccggct ggttggatgc tatcaacagc 1500
ggcaccaact ctctgttctt gaccattggc cctggcgact tcttggttca ccacgcgatc 1560
gccctagggt tgcacaccac tgccctgatt ctaatcaaag gtgctttgga tgcccgtggt 1620
tctaagttga tgccggacaa aaaggacttc ggttactcct tcccctgtga tggccccggc 1680
cgtggcggta cctgcgacat ctctgcttgg gatgccttct acctagccat gttctggatg 1740
ctgaacacct tgggttggtt gaccttctac tggcactgga aacacctcgg tgtttggagc 1800
ggtaacgttg ctcagttcaa cgaaaactcc acctacctga tgggttggtt ccgggattac 1860
ctctgggcga actctgctca gttaatcaat ggttacaacc cctgcggtgt caacaatctg 1920
tcagtttggg cttggatgtt ccttttcgga cacctggtct gggctactgg cttcatgttc 1980
ttgatctctt ggcggggtta ctggcaagag ttgattgaaa ccatcgtttg ggcccacgag 2040
cgcactcctt tggcgaactt ggttcgttgg aaagataagc ccgttgcgtt gtccattgtt 2100
caagcccgtt tggttggttt agcccacttc accgttggtt atgtgctcac ctatgcggca 2160
ttcctaattg cttccacagc cggtaagttc ggttaa 2196
Claims (37)
1.一种分离的修饰多肽,该多肽包含光合生物光催化单元的多肽的氨基酸序列,所述氨基酸序列能够介导所述光催化单元与固体表面的共价连接,并且在连接到所述固体表面时仍保持所述光催化单元的光催化活性。
2.权利要求1的分离的修饰多肽,其中所述光合生物为绿色植物。
3.权利要求1的分离的修饰多肽,其中所述光合生物为蓝细菌。
4.权利要求1的分离的修饰多肽,其中所述光催化单元为PS-I。
5.权利要求3的分离的修饰多肽,其中所述光催化单元为集胞藻PCC 6803光催化单元。
6.权利要求1的分离的修饰多肽,其中所述光催化单元中所述多肽的所述氨基酸序列包含至少一个置换突变。
7.权利要求6的分离的修饰多肽,其中所述置换突变在所述光催化单元的膜外环上。
8.权利要求1的分离的修饰多肽,其中所述多肽的所述氨基酸序列为psa B。
9.权利要求1的分离的修饰多肽,其中所述多肽的所述氨基酸序列为psa C。
10.权利要求8的分离的修饰多肽,其中所述Psa B在至少一个用以下坐标标明的位置上包含置换突变:D235C、S246C、D479C、S499C、S599C、Y634C。
11.权利要求8分离的修饰多肽,其中所述Psa C在至少一个用坐标W31C标明的位置上包含置换突变。
12.权利要求6的分离的修饰多肽,其中所述至少一个置换突变为半胱氨酸。
13.权利要求6的分离的修饰多肽,其中所述氨基酸序列如SEQID NO:20、21、22、23、24、25、26、27、28和29中所示。
14.权利要求1的分离的修饰多肽,其中所述固体表面为导电材料。
15.权利要求14的分离的修饰多肽,其中所述导电材料为过渡金属元素。
16.权利要求15的分离的修饰多肽,其中所述过渡金属元素选自银、金、铜、铂、镍和钯。
17.权利要求1的分离的修饰多肽,所述多肽不包含金属离子。
18.权利要求4的分离的修饰多肽,所述多肽为单体形式或三聚体形式。
19.大量的权利要求1的分离的多肽,所述多肽能够以相对于所述固体表面基本相同的方向定向。
20.一种分离的修饰光催化单元,其包含权利要求1的修饰多肽。
21.一种膜制品,其包含权利要求20的修饰光催化单元。
22.一种分离的多核苷酸,其编码权利要求1的修饰多肽。
23.一种核酸构建体,其包含权利要求22的多核苷酸。
24.权利要求23的核酸构建体,其还包含顺式调节元件。
25.权利要求24的核酸构建体,其中所述顺式调节元件为启动子。
26.一种细胞,其包含权利要求22的分离的多核苷酸。
27.权利要求24的细胞,所述细胞是集胞藻细胞。
28.一种器件,其包括连接有大量的权利要求20的修饰光催化单元的固体表面。
29.权利要求24的器件,其中所述大量的修饰光催化单元每个之间的距离介于15-25nm之间。
30.权利要求28的器件,其中所述大量的修饰光催化单元相对于所述固体表面是定向的。
31.权利要求28的器件,用作光电二级管的元件。
32.权利要求28的器件,用作光电晶体管的元件。
33.权利要求28的器件,用作逻辑门的元件。
34.权利要求28的器件,用作太阳能电池的元件。
35.权利要求28的器件,用作光耦合器的元件。
36.权利要求28的器件,其中所述大量的修饰光催化单元与所述固体表面直接连接。
37.权利要求36的器件,其中所述直接连接为共价连接。
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US7067322B2 (en) * | 2001-07-10 | 2006-06-27 | Wisconsin Alumni Research Foundation | Fusion protein arrays on metal substrates for surface plasmon resonance imaging |
US6864118B2 (en) * | 2002-01-28 | 2005-03-08 | Hewlett-Packard Development Company, L.P. | Electronic devices containing organic semiconductor materials |
JP3933664B2 (ja) | 2002-08-01 | 2007-06-20 | 三洋電機株式会社 | 光センサ、光センサの製造方法および駆動方法、ならびに光強度検出方法 |
US7592539B2 (en) * | 2003-11-07 | 2009-09-22 | The Trustees Of Princeton University | Solid state photosensitive devices which employ isolated photosynthetic complexes |
WO2006060017A1 (en) | 2004-12-02 | 2006-06-08 | The Trustees Of Princeton University | Solid state photosensitive devices which employ isolated photosynthetic complexes |
WO2008023373A2 (en) | 2006-08-22 | 2008-02-28 | Ramot At Tel Aviv University Ltd. | Optoelectronic device and method of fabricating the same |
WO2006090381A1 (en) | 2005-02-22 | 2006-08-31 | Ramot At Tel Aviv University Ltd. | Molecular optoelectronic device and method of fabricating the same |
WO2008018982A2 (en) | 2006-08-03 | 2008-02-14 | North Carolina State University | Self-assembled photosynthesis-inspired light harvesting material and solar cells containing the same |
-
2006
- 2006-02-22 WO PCT/IL2006/000241 patent/WO2006090381A1/en active Application Filing
- 2006-02-22 AT AT06711223T patent/ATE527359T1/de not_active IP Right Cessation
- 2006-02-22 CN CN2006800134334A patent/CN101163793B/zh not_active Expired - Fee Related
- 2006-02-22 EP EP20060711223 patent/EP1853708B1/en not_active Not-in-force
- 2006-08-22 US US11/507,628 patent/US7524929B2/en not_active Expired - Fee Related
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2009
- 2009-03-30 US US12/385,083 patent/US8212005B2/en not_active Expired - Fee Related
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US20090242879A1 (en) | 2009-10-01 |
US7524929B2 (en) | 2009-04-28 |
WO2006090381A1 (en) | 2006-08-31 |
CN101163793B (zh) | 2011-11-09 |
ATE527359T1 (de) | 2011-10-15 |
US8212005B2 (en) | 2012-07-03 |
EP1853708A1 (en) | 2007-11-14 |
EP1853708B1 (en) | 2011-10-05 |
US20080305471A1 (en) | 2008-12-11 |
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