Lingo-1结构的制作方法

专利名称：：Lingo-1结构的制作方法LINGO隱l结构相关申请本申请要求2006年2月3日提交的美国临时申请号60/765,443的权益，该申请通过引用整体并入本文。
技术领域：
本公开涉及LINGO-l多肽、LINGO-l多肽/配体复合物、LINGO-l多肽的晶体、LINGO-l多肽/配体复合物的晶体以及相关方法和软件系统。
背景技术：
：LINGO-l(含LRR和fe结构域的Hog^受体相互作用蛋白质)是p75的跨膜共受体。共受体与配体结合Nogo-66受体(NogoR)构成Nogo受体复合物，其通常位于神经元上。使得其中有LINGO-l的Nogo受体复合体失去作用能够诱导髓磷脂和神经纤维的生长。LINGO-l已经显示在已知称为少突胶质细胞的髓磷脂产生细胞中具有活性，并且，对LINGO-l的抑制也已显示能诱导髓磷脂形成。因此，人们预测LINGO-l信号传导是髓磷脂产生的负调节剂。
发明内容在一个方面，本发明涉及结晶的LINGO-l多肽。在另一个方面，本发明涉及结晶的多肽-配体复合物，其包括LINGO-l多肽和配体，所述配体是LINGO-l多肽的激动剂或拮抗剂。在又一个方面，本发明涉及一种方法，其包括使用LINGO-l多肽的三维模型来设计与LINGO-l多肽结合的物质。在另一个方面，本发明涉及一种方法，其包括使用LINGO-l多肽-配体复合物的三维模型来设计与LINGO-l多肽结合的物质。在另一个方面，本发明涉及一种方法，其包括使用包括LINGO-l多肽的复合物的三维模型来设计与LINGO-l多肽结合的物质。在另一个方面，本发明涉及一种方法，其包括通过例如用LINGO-l多肽或LINGO-l多肽-配体复合物的三维结构，进行合理的药物设计，来选择物质；将所述物质与LINGO-l多肽接触；以及，检测所述物质与LINGO-l多肽结合的能力。在一些实施方案中，所述方法还包括下迷步骤中的一种或多种获得包括LINGO-l多肽和所述物质的补充晶体复合物；测定补充晶体复合物的单位结构；通过，例如，用所述补充晶体复合物的三维结构进行合理的药物设计，来选出第二物质；将所述笫二物质与LINGO-l多肽相接触；和/或检测所述第二物质与LINGO-l多肽结合的能力。所述方法还可包括下述步骤中的一种或多种合成第二物质；检测所述第二物质抑制LINGO-l活性的能力；和/或检测所述第二物质增加体外或体内髓磷脂水平的能力。在一些实施方案中，本文公开的组合物和方法中的LINGO-l多肽包括富含亮氨酸的重复(LRR)结构域、免疫球蛋白样(Ig样)结构域和/或茎结构域或其组合(或主要由它们组成)。在其它实施方案中，LINGO-l多肽包括来自哺乳动物(例如人)或非哺乳动物来源的氨基酸序列(或主要由它们组成)；例如，LINGO-l多肽可包括SEQIDNO:1的氨基紗列或其变体(例如，其保守取代)(或由它们组成)。在其它实施方案中，LINGO-l多肽是结晶的，例如，其能将X-射线衍射为至少大约3.7A的分辨率。在一些实施方案中，结晶的LINGO-l多肽具有空间群(spacegroup)P2"J或1222。在其它实施方案中，结晶的LINGO-l多肽具有选自下述一种或多种的晶胞尺寸a=201.5A、b=149.7A、c=157.5A和a=p=y=90。、a=148.7A、b=158.6A、c-200.0A和a-p，卯0，和a-149.6A、b=157.3A、c=200.3A和/或a=p=y=90°。在又一实施方案中、结晶的LINGO-l多肽具有下述三维结构，其中包括根据表2的结构坐标，01碳原子的+/-均方根偏差不超过1.5A。在另一实施方案中，结晶的LINGO-l多肽具有下述三维结构，其包括选自SEQIDNO:1所示的LINGO-l多肽的氨基酸Aspl3、Arg20、Arg22、Arg43、Glu60、Glu62、Leu94、Leul20、Metl23、Hisl76、Tyrl42、Hisl45、Hisl85、His209、His233、Ala238、Trp346、Arg347、Asn349、Asn351、Arg352、Gln353、Phe396、Arg408、Leu420、Leu426、Phe438、Asp440、Arg446、Tyr447和/或Ile459的一个或多个原子的原子结构坐标。在其它实施方案中，使用本文公开的方法设计或篩选的物质能抑制LINGO-l活性。例如，所述物质可在配体结合位点与LINGO-l结合和/或干扰LINGO-l与配体之间的相互作用。在一些实施方案中，配体结合位点位于LRR结构域的凹面表面，例如，下述LRR结构域表面，其包括根据SEQIDNO:1的Trp346和Arg352;Hisl85、His209和His233;Aspl3;Glu60和Glu62;或Arg20、Arg22和Arg43中的一处或多处。在其它实施方案中，配体结合位点位于LRR结构域的凸面表面，例如，下述LRR结构域表面，其包括才艮据SEQIDNO:1的Tyrl42和Hisl45;Leu94、Leul20和Metl23;Hisl76;或Ala238中的一处或多处。在其它实施方案中，配体结合位点位于Ig结构域上，例如，这样的结构域，其包括才艮据SEQIDNO:1的Arg446和Tyr447;Arg408、Phe438和Asp440;Phe396;或Leu420、Leu426和Ile459中的一处或多处。在一些实施方案中，本文公开的方法包括提供包括LINGO-l多肽或LINGO-l多肽-配体复合物的组合物，和/或使所述组合物结晶，以形成包括LINGO-l多肽或LINGO-l多肽-配体复合物的结晶复合物。结晶复合物可将X-射线衍射为至少大约3.7A的分辨率。本文公开的方法可额外包括下述步骤中的一个或多个计算LINGO-l多肽的原子与该物质的原子之间的距离；测定该物质与LINGO-l多肽的相互作用；将该物质与LINGO-l多肽的相互作用与第二物质与LINGO-l多肽之间的相互作用加以比较；通过计算机建模选择该物质；合成该物质；检测该物质抑制LINGO-l活性的能力；和/或检测该物质体外或体内增加髓磷脂形成的能力。在另一个方面，本发明涉及使用本文公开的方法设计或选择的物质。在另一个方面，本发明涉及软件系统，其包括用于使计算机系统接受与LINGO-l多肽的结构相关的信息，接受与候选物质相关的信息，以及确定候选物质与LINGO-l多肽的结合特点的指令。对结合特性的确定可基于与LINGO-l多肽结构相关的信息和与候选物质相关的信息来进行。在又一方面，本发明涉及存在于计算机可读介质上的计算机程序，其具有储存于其上的多个指令，当被一个或多个处理器执行时，所述指令导致一个或多个处理器接受与包含LINGO-l多肽的复合物结构相关的信息，接受与候选物质相关的信息，以及确定候选物质与LINGO-l多肽的结合特点。对结合特性的确定可基于与LINGO-l多肽结构相关的信息和与候选物质相关的信息来进行。在另一方面，本发明涉及一种方法，其包括接受与LINGO-l多肽的结构相关的信息，以及对LINGO-l多肽与候选物质的结合特点加以建模。所述方法可通过软件系统实现。在又一方面，本发明涉及存在于计算机可读介质上的计算机程序，其具有储存于其上的多个指令，当被一个或多个处理器执行时，所述指令导致一个或多个处理器接受与LINGO-l多肽的结构相关的信息，以及对LINGO-l多肽与候选物质的结合特点加以建模。在另一个方面，本发明涉及软件系统，其中包括下述指令，用于使得计算机系统接受与LINGO-l多肽的结构相关的信息以及对LINGO-l多肽与候选物质的结合特点加以建模。在另一个方面，本发明涉及调节受试者中LINGO-l活性的方法，所述方法包括例如使用本文描述的方法(例如，通过合理的药物设计)，选择能调节LINGO-l活性的物质，将有效量的所述物质施用给受试者，使得LINGO-l活性被调节。在另一个方面，本发明涉及治疗具有与LINGO-l活性相关的病况的受试者的方法，所述方法包括例如使用本文描述的方法(例如，通过合理的药物设计)，选择能影响LINGO-l活性的物质，将治疗有效量的所述物质施用给需要其的受试者。在又一个方面，本发明涉及对与LINGO-l活性相关的病况易感的受试者进行预防性治疗的方法，所述方法包括确定所述受试者是否对与UNGO-l活性相关的病况易感，例如使用本文描述的方法(例如，通过合理的药物设计)，选择能影响LINGO-l活性的物质，以及将治疗有效量的物质施用给受试者。在一些实施方案中，本文公开的方法(例如本文公开的治疗和预防方法)中所用的物质能抑制LINGO-l活性和/或增加体内髓磷脂水平。在一些实施方案中，所述病况是脱髓鞘疾病，例如，多发性硬化。本发明的其它特征和优点将从说明书、附图和权利要求中显而易见。图1是经组氨酸标签化的LINGO-l多肽(氨基酸1-516加6个His标签)的氨基i^列(SEQIDNO:1)。用于结晶的片段被下划线示出。图2是针对精确修正过的2.7A分辨率的LINGO-l结构的2fo-fc电子密度图(也见表1)。图3是LINGO-l单体的带状图(ribbondiagram)。图4是LINGO-l单体的空间填充模型。被编号的残基指糖基化的天冬酰胺残基。图5A-5C是结构图，其说明LINGO-l晶体结构的四聚体组织。图5A是四聚体结构的顶视图；图5B是两个单体间相互作用表面的视图；图5C是四聚体结构的侧视图。图6是pSMEG质粒的图镨。图7A-1至7A-3展示了pSMEG质粒的序列(SEQIDNO:2)。图8A-8B描绘了人LINGO-l的全长氨基酸和核苷酸序列(分别是SEQIDNO:3和4)。编码序列由SEQIDNO:4的核苷酸53至1915编码。发明详述一般而言，本公开涉及LINGO-l多肽、LINGO-l多肽/配体复合物、LINGO-l多肽的晶体、LINGO-l多肽/配体复合物的晶体以及相关方法和软件系统。不希望被理论束溥，我们相信，LINGO-l多肽的晶体结构和LINGO-l多肽/配体复合物的晶体结构可用于设计或鉴定能与LINGO-l多肽相互作用的配体。能与LINGO-l多肽相互作用的配体能抑制或增加LINGO-l活性。例如，LINGO-l抑制已显示能诱导髓磷脂形成。因此，我们相信，对LINGO-l抑制剂的鉴定可用于治疗特征为髓磷脂缺乏的病症，例如多发性硬化、佩，梅病(Pelizaeus-Merzbacher)、中枢神经系统损伤、脑白质营养不良(例如Krabbe病或Canavan病)或维生素B12缺乏。图l是人LINGO-l多肽的片段(LINGO-lK549his6)的氨基餅列(SEQIDNO:1)，其包括富含亮氨酸的重复的结构域(氨基酸1-382)、免疫球蛋白样(Ig样)结构域(残基383-477)、茎区域(氨基酸478-516)以及与茎区域的C末端融合的六组氨酸标签。六组氨酸标签能有利于纯化。在全长人LINGO-l的N末端发现的33个氨基酸的信号序列在LINGO-lK549his6多肽中不存在。野生型序列还包括跨膜结构域和蛋白质C末端的短的胞质尾(Mi等人，7V^A^i/y^c/.2:221-228,2004)。图2是LINGO-lK549his6结构被解析至2.7A分辨率的电子密度图(见表1)。图3是展示了LINGO-lK549bis6多肽的结构的带状图。LINGO-lK549his6多肽晶体结构的坐标提供于表2中。为测定LINGO-l的结构，可制备人LINGO-l多肽，按照下文所述对其进行结晶。一般而言，可以按照需要制备人LINGO-l多肽。例如，在一些实施方案中，人LINGO-l多肽是从DNA质粒表达的。表达可由启动子来驱动，所述启动子例如诱导型启动子或组成型启动子，例如，巨细胞病毒启动子。人LINGO-l多肽可作为融合蛋白与合适的标签一起表达，所述标签例如多聚组氨酸(例如六组氨酸)、谷胱甘肽-S-转移酶(GST)、myc、HA、Strep或FLAG标签。标签可协助将人LINGO-l多肽与细胞分离，例如，与细菌细月包或与哺乳动物细胞系分离。例如，人LINGO-l多肽可在中国仓鼠卵巢(CHO)细胞中表达，并从中^f皮分离。例如，人LINGO-l多肽可与六组氨酸标签融合，并这样来分离将细胞提取物与镍树脂(例如，次氮基三乙酸镍(Ni-NTA)树脂)接触，以结合六组氨酸标签，然后通过用含咪唑的緩冲液洗树脂来释放多肽。此外，可在设计加入融合蛋白的蛋白酶位点(例如，在多肽与标签之间的融合位点或附近)，将融合LINGO-l多肽切割下来。人LINGO-l多肽可被放置于溶液中，以收集镨数据、NMR数据，或者用于生长晶体。例如，人LINGO-l多肽可在盐(例如，钠盐和/或铵盐)存在下结晶。晶体可通过多种方法生长，例如，沉滴或悬滴蒸汽扩散法。一般而言，结晶可在从大约4°C至大约60°C的温度(例如，从大约10°C至大约45。C，例如大约12°C、大约15°C、大约18°C、大约20°C、大约25°C、大约30。C、大约32。C、大约35°C、大约37。C)下进行。一般而言，人LINGO-l多肽可将X射线衍射至大约3.7A或更小的分辨率(例如，大约3.6A或更小、大约3.5A或更小、大约3，2A或更小、大约3.0A或更小、大约2.7A或更小、大约2.4A或更小、大约2.3入或更小、大约2JA或更小、大约2.lA或更小、大约2.0A或更小、大约1.9人或更小、大约1,8A或更小、大约1.7A或更小、大约1.6A或更小、大约1.5A或更小、或者大约1.4A或更小)。在一些实施方案中，人LINGO-l多肽的晶体可将X射线衍射至从大约1.7A至大约3.7A的分辨率(例如，人LINGO-l多肽的晶体可将X射线衍射至大约2.7A、大约3.5A或大约3.6A)。在某些实施方案中，人LINGO-l多肽的晶体属于空间群I222，具有下述晶胞参数a=148.7A、b=158.6A、c=200.0A以及a-卩，-卯。。在其它实施方案中，人LINGO-l多肽的晶体属于空间群1222,具有下述晶胞参数a=149.6A、b=157.3A、c=200.3人以及01=卩=7=卯°。在其它实施方案中，人LINGO-l多肽的晶体属于空间群P2^2，具有下述晶胞参数a=201.5A、b=149.7A、c=157.5人以及a=P=y=90°。空间群指晶体的总体对称，其包括点对称和空间对称。在某些实施方案中，人LINGO-l多肽的晶体属于空间群1222，在其不对称单位中含有两个分子的人LINGO-l多肽。在其它实施方案中，人LINGO-l多肽的晶体属于空间群P2JJ,其在不对称单位中含有四个分子的人LINGO-l多肽。不对称单位是可使用空间群的对称操作来由其产生晶体结构的最小单位。晶体通常由不对称单位上的空间群对称操作所界定的基序以及经由晶格对该基序的平移(translation)构成。可以例如通过X射线衍射获得描述晶体的结构数据，。可通过多种来源、x射线波长和检测器收集x射线衍射数据。在一些实施方案中，可使用旋转阳极和同步加速器源(例如，AdvancedLightSource(ALS),Berkeley,California或AdvancedPhotonSource(APS),Argonne，Illinois)作为X射线源。在某些实施方案中，用于产生衍射数据的X射线可具有从大约0.5A至大约1.6A的波长(例如，大约0.7A、大约0.9A、大约l.oA、大约l.iA、大约L3A、大约1.4A、大约1.5A或者大约1.6A)。在一些实施方案中，平面检测器和/或电荷耦合器件(CCD)可用作检测器。人LINGO-l多肽晶体的X射线衍射数据可用于获得复合物中原子的结构坐标。结构坐标是笛卡尔(Cartesian)坐标，其描述了三维空间中的原子相对于复合物中其它原子的位置。例如，表中列出的结构坐标是结晶人LINGO-l多肽的结构坐标。表2的结构坐标描述了人LINGO-l多肽(SEQIDNO:5)、(SEQIDNO:6)、(SEQIDNO:7)和(SEQIDNO:8)的原子互相之间的相对位置。可通过数学处理来修改结构坐标，例如通过倒数或整数加减来进行。因此，结构坐标是相对坐标。例如，描述人LINGO-l多肽中原子位置的结构坐标并不被表2中实际的x、y和z坐标特定地限制。人LINGO-l多肽的结构坐标可用于得到对多肽或多肽片段的表示(representation)(例如，二维表示或三维表示)。此类表示可用于多种应用，包括，例如，对多肽活性位点的观察、鉴定和分析。在某些实施方案中，三维表示可包括根据表2的人LINGO-l多肽结构坐标，±不超过大约1.5A(例如，不超过大约1.0A,不超过大约0.5A)的来自氨基酸a碳原子的均方根(rms)偏差。RMS偏差是与均值偏差的平方的算术均值的方根，其是表示与结构坐标的偏差或变化的方式。氨基酸的保守取代(见下文讨论)可产生具有处于指定的rms偏差内的结构坐标的分子表示。例如，由于保守氨基酸取代而相互有所不同的两个多肽分子模型具有的主链原子坐标可处于指定的rms偏差内，例如小于约1.5A(例如小于约l.O人、小于约0.5A)。多肽的主链原子包括a碳(Ca或CA)原子、羰基碳(C)原子和酰胺氮(N)原子。多种软件程序允许对一套结构坐标进行图形表示，以获得对LINGO-l多肽或多肽片段的表示。通常，此类表示应当精确反映(相对和/或绝对)结构坐标或源自结构坐标的信息，例如特征之间的距离或角度。在一些实施方案中，表示是交互式二维显示，例如交互式立体二维显示。交互式二维显示可以是，例如，可旋转以展示多肽或多肽片段的不同面的计算机显示。在一些实施方案中，表示是三维表示。例如，三维表示可以是分子结构的物理模型(例如，球和棒模型)。还例如，三维表示可以是分子结构的图形表示(例如，计算显示上呈现的图画或图)。当二维表示反映了三维信息时，例如，通过用与观察者较近的特征对与观察者较远的特征加以透视、阴影或障碍时，二维图形表示(例如图画)可对应于三维表示。在一些实施方案中，可在超过一个水平上针对表示建模。例如，当三维表示包括多肽，例如人LINGO-l多肽时，可在一个或多个不同的结构水平来表示该多肽，例如，在一级结构(氨基酸序列)、二级结构(例如a-螺旋和p-片)、三级结构(整体折叠)和四级结构(寡聚状态)上来进行。该表示可包括不同水平的细节。例如，该表示可包括蛋白质二级结构特征的相对位置，而不特别指出原子位置。更详细的该表示可例如包括原子位置。在一些实施方案中，该表示可还包括除了人LINGO-l多肽中原子的结构坐标之外的信息。例如，该表示可提供关于溶剂可达的表面的形状、模型的原子的范德华半径和溶剂(例如，水)的范德华半径。可从该表示中获得的其它特征包括，例如，静电势、大分子结构中的空隙或口袋的位置以及氢键和盐桥的位置。可通过下述方法来鉴定或设计与人LINGO-l多肽相互作用(例如结合)的物质，所述方法包括使用对多肽或多肽片段的表示。表示的示例性类型包括上文所讨论的表示。在一些实施方案中，该表示可以是类似物多肽或多肽片段的。可通过进行对候选物质与表示的计算机拟合(fitting)分析，来鉴定或设计与该表示相互作用的候选物质。通常，该物质是分子。该物质的例子包括多肽、核酸(包括DNA或RNA)、甾类和非甾类有机化合物。与多肽(例如人LINGO-l多肽)相互作用的物质能与多肽暂时或稳定地相互作用。相互作用可被本文提到的任何力介导，所述力包括，例如，氢键、静电力、疏水相互作用和范德华相互作用。如上文所述，可使用X射线晶体图来获得人LINGO-l多肽的结构坐标。但是，可使用其它技术，包括NMR技术，来获得此类结构坐标。可从光镨技术(例如，旋光色散(ORD)、圆二色镨(CD))同源性建模和计算方法(例如，可包括来自分子力学的数据的计算方法、可包括来自动力学检验的数据的计算方法)来获得其它结构信息。在一些实施方案中，可使用X射线衍射来构建人LINGO-l多肽或多肽片段的电子密度图，可使用电子密度图来获得对人LINGO-l多肽的表示(例如，二维表示、三维表示)。典型地，电子密度图的产生涉及到使用关于X射线散射相位的信息。相位信息可例如提取自衍射数据或用于完成对电子密度图构建的补充性衍射实验。用于计算来自X射线衍射数据的相位的方法包括，例如，多波长反常散射法(MAD)、多重同晶型取代法(MIR)、具反常散射的多重同晶型取代法(MIRAS)、具反常散射的单同晶型取代法(SIRAS)、倒易空间溶剂扁平化(reciprocalspacesolventflattening),分子置换(molecularreplacement)或其任何组合。这些方法通过制造天然蛋白质的同晶型(isomorphous)结构修饰(例如通过包括重原子或改变已经存在的重原子的散射强度)，然后测量天然蛋白质和每种经修饰情况的衍射幅度来产生相位信息。如果额外的重原子的位置或其^L射强度的改变是已知的，计算机程序来鉴定重原子位点的位置，所述程序例如，SHELXD(Bruker隱AXS，Madison,WI)或SHELXS(Sheldrick，InstitutAnorg.Chemie，G6ttingen,Germany)和XPREP(Bruker-AXS，Madison，WI);可使用计算机程序来加工衍射数据，所述程序例如MOSFLM、SCALA、SOLOMON和SHARP("TheCCP4Suite:ProgramsforProteinCrystallography,"爿c似O^toWogr.Sect.D,54:905-921,1997;deLaFortelleandBrigogne，Me仇五"g;附.276:472-494,1997)。4十对结晶人LINGO-l多肽的X射线散射的相位例如可通过SIRAS,使用LINGO-l多肽鉑衍生物的晶体来测定。为制造结晶LINGO-l多肽的铂衍生物，可将结晶LINGO-l多肽浸泡于含铂的溶液中。然后可在计算机程序(例如DM(CowtanandMain,J"flOj;".3^2:148-157，1993))中使用例如非晶体学对称性(NCS)平均和相位延伸，来精确修正通过SIRAS从铂衍生物获得的相位。还可通过用第二数据组进行分子置换，来进一步衍生获得的模型。例如，可用来自具有空间群P2"J的结晶LINGO-l多肽的数据组进行分子置换，对源自具有空间群1222的结晶LINGO-l多肽的模型进行精确修正。然后，可使用通过SIRAS从天然结晶多肽和铂衍生物的晶体获得的相位，来产生LINGO-l多肽的电子密度图。可使用电子密度图来获得多肽或多肽片段的表示，这通过将多肽的三维模型与电子密度图进行比对来实现。例如，可以将对应于天然结晶人LINGO-l多肽的电子密度图与对应于结晶人LINGO-l多肽复合物的铂衍生物(通过同晶型取代方法获得的)的电子密度图加以比对。比对过程产生了比较模型，其展示出计算出的电子密度图与以前已知的多肽或以前已知的复合物相比变化的程度。然后可对比较模型进行一轮或数轮(例如两轮、三轮、四轮、五轮、六轮、七轮、八轮、九轮、十轮)的精确修正，以产生与电子密度图的更好的拟合。可使用软件程序，例如CNS(Brunger等人，爿""Co^a/togr.^:905-921,1998)和REFMAC(CollaborativeComputationalProject,Number4.TheCCP4suite:ProgramsforProteinCrystallography,爿c似Oy1yto//og/%D50:760-776,1994)，来对模型加以精确修正。可例如通过Rfaet。r或Rfree值来测量比较才莫型中的拟合质量。通常，Rfact。r或Rfree的值越小就表明越好的拟合。可对比较模型中的不重合加以调整，以提供经修改的比较模型和较低的Rfact。r或Rfree值。调整可基于与人LINGO-l多肽的变化相关的信息(例如，序列变化信息、备选结构信息、重原子衍生物信息)来适当地进行。例如，在使用结晶LINGO-l多肽的重原子衍生物模型的实施方案中，调整可包括在重原子衍生物^t型上拟合天然LINGO-l多肽的大致模型。还例如，在某些实施方案中，调整可包括用具有相似结构的氨基酸替换以前已知的LINGO-l多肽中的氨基酸(保守氨基酸改变)。当对经修正的比较模型的调整满足对电子密度图的最佳拟合时，得到的模型就是被确定用来描述从其中获得X射线数据的多肽或复合物的模型。此类过程的方法被公开于，例如，Carter和Sweet，编辑，"MacromolecularCrystallography"在MethodsinEnzvmologv,第277巻，B部分，NewYork:AcademicPress,1997及其中的文章，例如,Jones和Kjeldgaard,"Electron-DensityMapInterpretation,"第173页和Kleywegt和Jones,"ModelBuildingandRefinementPractice,"第208页中。机器，例如，计算机可按程序工作，在内存中使用人LINGO-l多肽的结构坐标连同程序能够在机器相连的显示器上产生对结构坐标的图形表示。还可设计和/或利用软件系统来接受或存储结构坐标。软件系统也能够产生结构坐标的图形表示。软件系统还可能进入外部数据库，以鉴定出一种或多种可能与人LINGO-l多肽相互作用的候选物质。具有含结构数据的内存或含此类数据的软件系统的机器可协助对人LINGO-l多肽拮抗剂的人LINGO-l多肽激动剂的合理设计或选择。例如，此类机器或软件系统可协助对该物质与人LINGO-l多肽结合的能力加以评估，或者可协助通过与人LINGO-l多肽的结构或序列同源性来对相关化合物或蛋白质加以建模。在本文中使用时，激动剂指增强人LINGO-l多肽的至少一种活性的化合物，拮抗剂指抑制或抵消人LINGO-l多肽的至少一种活性的化合物。例如，化合物可通过，例如，增加神经细胞髓磷脂生产的速率，或者通过抑制人LINGO-l多肽与Nogo受体复合物的相互作用，来作为人LINGO-l多肽的拮抗剂发挥功能。机器可产生对人LINGO-l多肽或多肽片段的表示(例如，二维表示、三维表示)。软件系统例如可使得机器产生此类信息。机器可包括机器可读的存储介质，其中包括用机器可读的数据编码的数据存储材料。机器可读的数据包括人LINGO-l多肽的原子的结构坐标。机器可读的存储介质(例如，数据存储材料)包括，例如，传统的计算机硬盘、软盘、DAP磁带、CD-ROM、DVD和其它磁性、磁光、光和可适于与机器(例如计算机)一起使用的其它介质。机器还可具有工作内存，以存储用于处理机器可读的数据的指令，以及具有与工作内存和机器可读的数据存储介质相耦合的中央处理单元(CPU)，用于将机器可读的数据处理为想要的三维表示的目的。显示器可与CPU相连，使得三维表示可被使用者看到。因此，当使用编程有使用数据的指令的机器(例如，装有本文所述的种类的一种或多种程序的计算机)时，所述及其能显示本文所述的多肽、多肽片段、复合物或复合物片段中任何的图形表示(例如，二维图形表示、三维图形表示)。显示器(例如计算机显示器)可展示人LINGO-l多肽或人LINGO-l多肽片段的表示。使用者可观察所述表示，以及，使用从所述表示获得的信息，产生人LINGO-l多肽或与配体结合的多肽片段的模型。可例如通过改造已有的人LINGO-l多肽的表示来产生模型。任选地，使用者可将该物质的三维模型叠加到人LINGO-l多肽的表示上。该物质可以是人LINGO-l多肽的激动剂(例如候选激动剂)。在一些实施方案中，该物质可以是已知化合物或已知化合物的片段。在某些实施方案中，该物质可以是以前未知的化合物或以前未知的化合物的片段。其形状能补足活性位点形状的该物质可能是人们想要的。在该物质的原子和人LINGO-l多肽的原子间可以有优选的距离或距离范围。比优选距离更长的距离可能与该物质和活性位点(例如，人LINGO-l多肽的活性位点)之间的弱相互作用相关联。比优选距离更短的距离可能与能削弱该物质和多肽之间达到相互作用的斥力相关联。当原子间距离过短时可能产生空间位阻。当两个原子的位置不合理地接近时，例如，当两个原子之间分开的距离要与它们的范德华半径之和更小时，就可能产生空间位阻。如果存在空间位阻，使用者可调节该物质相对于人LINGO-l多肽的位置(例如，刚体平移或该物质的旋转)，直到空间位阻减轻。使用者可调节该物质或物质附近人LINGO-l多肽的构象，以减轻空间位阻。还可通过改造该物质的结构，例如，通过将"大块的基团"(例如芳香族环)改变为较小的基团，例如甲基或羟基，或通过将刚性基团改变为能成为不产生空间位阻的构象的柔性基团，来移除空间位阻。静电力也可能影响该物质和配体结合结构域之间的相互作用。例如，静电性质可与能削弱该物质与人LINGO-l多肽间相互作用的斥力相关联。可通过改变该物质的电荷，例如通过用中性基团替换带正电荷的基团来减轻静电斥力。可在多肽/该物质模型中评估影响各候选物质与人LINGO-l多肽间结合强度的力。它们可包括，例如，氢键、静电力、疏7jC相互作用、范德华相互作用、偶极-偶极相互作用、7T-堆集力和阳离子-7C相互作用。使用者可视觉评估这些力，例如通过观察以适于氢键的距离和角度排列的氢键供体/受体对来实现。基于这种评估，使用者可对模型加以改造，以发现人LINGO-l多肽和该物质间的更有利的相互作用。对模型的改造可包括改变多肽的三维结构，而不改变其化学结构，这例如通过改变氨基酸侧链或主链两面角的构象来实现。对模型的改造可包括按照上文所述改变该物质的位置或构象。对模型的改造还可包括改变该物质的化学结构，例如通过取代、添加或去除基团来实现。例如，如果人LINGO-l多肽上的氢键供体位于该物质上的氢键供体附近，使用者可以用氢键受体替换该物质上的氢键供体。可以调节该物质和人LINGO-l多肽的相对位置或者它们的构象，以找到特定物质与人LINGO-l多肽的最优结合几何条件。最优结合几何条件特征在于，例如，有利的氬键距离和角度、最大的静电吸引、最小的静电排斥、疏水部分从水环境的分隔、以及不存在空间碰撞。最优几何条件计算出来的能量可能是人LINGO-l多肽/物质复合物的可能的几何条件家族中最低的。可例如通过分子力学或分子动力学计算来确定最优的几何条件。可以产生针对具有不同结合的该物质的人LINGO-l多肽的一系列表示。可以针对每种表示计算出分数。分数可以描述，例如，人LINGO-l多肽和该物质间相互作用的预期强度。分数可反映上文所述的影响结合强度的因素之一。分数可以是影响超过一种因素的累积分数。可根据其分数对不同的物质加以排序。设计物质的步骤可以通过机器以自动方式进行。例如，可在机器中对人LINGO-l多肽的表示和候选物质的表示一起编程。机器可找到对于每种候选物质与活性位点的最优结合几何条件，并且计算出分数，以确定系列中的何种物质可能与人LINGO-l多肽最强烈地相互作用。可以设计和/或实现软件系统，以协助这些步骤。用于产生表示或用于进行拟合分析的软件系统(例如计算机程序)包括，例如MCSS、Ludi、QUANTA⑧(大分子X射线晶体学软件)、Insight11@(生物化合物建模和模拟软件)、Cerius^(建模和模拟软件)、CHARMm⑧(用于模拟生物学大分子的软件)、CHARMm斷用于模拟生物学大分子的软件)和来自Accdrys，Inc.(SanDiego，CA)的Modeler;SYBYL⑧(分子建才莫软件)、来自TRIPOS，Inc.(St.Louis，MO)的Unity、FleXX和LEAPFROG;AUTODOCK(ScrippsResearchInstitute,LaJolla，CA);GRID(OxfordUniversity,Oxford,UK);DOCK(UniversityofCalifornia,SanFrancisco,CA)和Flo+和Flo99(Thistlesoft,MorrisTownship,NJ)。其它有用的程序包括来自OpeneyeScientificSoftware(SantaFe，NM)的ROCS、ZAP、FRED、Vida和Szybki;来自Schrodinger，LLC(Portland，OR)的Maestro、Macromodel和Glide;MOE(ChemicalComputingGroup,Montreal,Quebec)、Allegrow(BostonDeNovo,Boston,MA)、CNS(Brunger，等人.,JctoOjW"//.Sect.D54:卯5-921,1997)和GOLD(Jones等人，丄Afo/.1^:43-53,1995)。还可使用MOLSCRIPT、RASTER3D或PYMOL(Kraulis/C，to//，946-950，1991;Bacon和Anderson,/杨/.219-220，1998;DeLano，ThePYMOLMolecularGraphicsSystem(2002)DeLanoScientific,SanCarlos,CA)，用结构坐标来观察人LINGO-l多肽的三维结构。可以例如通过篩选合适的数据库来选择物质，可通过在合适的软件系统协助下分析人LINGO-l多肽的立体构型和电荷势能来对物质加以从头设计，和/或可使用已知配体(例如神经营养因子受体p75和Nogo受体(NogoR))的特点来对物质加以设计。所述方法可用于设计或选择人LINGO-l多肽的激动剂或拮抗剂。可设计和/或实现软件系统，以协助数据库搜索和/或物质的选择和设计。一旦已经设计或鉴定了物质，其就可被获得或合成，并且针对其对于人LINGO-l多肽活性的影响对其加以评估。例如，可通过将其与人LINGO-l多肽相接触以及测量物质对于多肽活性的影响来对物质加以评估。用于评估物质的方法可包括体外或体内进行的活性测定。活性测定例如可以是基于细胞的测定。基于细胞的测定可包括监测候选物质对于髓磷脂生产的效果。针对人LINGO-l多肽抑制剂的此类测定可包括将候选抑制剂与表达人LINGO-l多肽的细胞相接触，测定对于细胞形态的影响。例如，可将神经细胞系与候选物质接触，就对于少突胶质细胞分化的影响对细胞加以监测。LINGO-l活性的减少已显示能增加细胞向成熟少突胶质细胞的分化，这由细胞突长度的增加所证实(Mi等人，7VfltA^w仍de"ce2:745-751，2005)。还可通过测量髓磷脂产生所涉及的蛋白质的水平来检验LINGO-l活性，所述蛋白质例如髓磷脂相关糖蛋白(MAG)，这是在髓鞘形成开始时表达的蛋白质；髓磷脂碱性蛋白(MBP)，这是髓磷脂的主要蛋白质组分；以及其它髓磷脂组分，包括少突胶质细胞-髓磷脂糖蛋白(OMpg)、髓磷脂少突胶质细胞糖蛋白(MOG)和环状核苷酸磷酸二酯酶(CNPase)。LINGO-l抑制剂已显示能促进对这些蛋白质各自表达的上调(Mi等人，？VM7Ve"fwde"ce5:745-751，2005)。可通过标准的蛋白质检测技术，例如通过蛋白质印迹分析或通过在培养的细胞或全组织切片中的原位杂交进行的免疫组织化学方法，来检验蛋白质水平。取决于物质对于人LINGO-l多肽的作用，物质可作为人LINGO-l多肽活性的激动剂或拮抗剂发挥作用。激动剂例如可减少髓磷脂产生的速率，或者增加人LINGO-l多肽与Nogo受体复合物的结合亲和性。相反，拮抗剂可增加髓磷脂产生的速率，或者减少人LINGO-l多肽与Nogo受体复合物的结合亲和性。可在存在Nogo受体复合物的一种或多种组分(例如p75或Nogo-66)的情况下，将物质与人LINGO-l多肽相接触，以确定物质是否能抑制人LINGO-1多肽与Nogo受体复合物结合。含有与鉴定出的物质结合的人LINGO-l多肽的晶体可生长，并通过X射线晶体学对其加以结构分析。可基于第一物质与人LINGO-1多肽的相互作用，来设计或鉴定第二物质。多种分子分析和合理的药物设计技术还公开于，例如，美国专利号5,834,228、5,939,528和5,856,116中，以及PCT申请号PCT/US98/16879(公开为WO99/09148)中。虽然已经描述了某些实施方案，但是其它实施方案也包括在本发明内。例如，虽然已经描述了涉及人LINGO-1多肽的实施方案，但本文中的描述更为一般性地针对任何LINGO-1多肽。LINGO-1多肽可以是全长的成熟多肽，包括LINGO-1多肽的任何同种型的全长氨基酸序列。同种型是初级结构有所不同的几种形式中的任何形式。人LINGO-l多肽的全长氨基酸和核苷酸序列公开于图8A-8B中。LINGO-1多肽可以是人LINGO-1多肽的片段，例如，信号序列、LRR型结构域、Ig样结构域、跨膜结构域、胞质结构域或其组合。LINGO-1多肽的片段可包括LINGO-1多肽序列(例如SEQIDNO:l)的超过40%、50%、60%、80%、90%或更多。例如，LINGO-1多肽可包括下述结构域中的一种或多种LRR型结构域(例如，SEQIDNO:l的大约1-382号氨基酸)或其片段或选定的残基(例如，SEQIDNO:1的氨基酸Aspl3、Arg20、Arg22、Arg43、Glu60、Glu62、Leu94、Leul20、Metl23、Hisl76、Tyrl42、Hisl45、Hisl85、His209、His233、Ala238、Trp346、Arg347、Asn349、Asn351、Arg352和/或Gln353中的一个或多个)；Ig样结构域(例如，SEQIDNO:1的大约383-477号氨基酸)或其片段或选定的残基(例如，SEQIDNO:l的氨基酸Phe396、Arg408、Leu420、Leu426、Phe438、Asp440、Arg446、Tyr447和/或Ile459中的一个或多个)；和/或茎区域(例如，SEQIDNO:l的大约478-516号氨基酸)或其片段或选定的残基。在其它实施方案中，LINGO-l多肽可含有本文所述的活性位点和/或结构折叠中的一种或多种。LINGO-l多肽可含有一处或多处保守氨基酸取代，所述取代产生相似的三维结构，如本文所述。例如，在一些实施方案中，LINGO-l多肽(或其片段)包括对预定序列(例如人LINGO-l氨基*歹ij(例如SEQIDNO:1))的多至5、10、20、30、50、75、100、150或200或更多处保守取代。LINGO-l多肽可具有活性位点。例如，LRR样结构域包括LINGO-l多肽的可能的活性位点。通常，活性位点包括配体结合位点或磷酸化、糖基化、烷基化、酰化或其它共价修饰位点。配体结合位点可以是Nogo受体复合物的组分或LINGO-l活性的激动剂或拮抗剂的结合位点。活性位点可包括糖基化位点。配体结合位点可包括与可能影响与配体相互作用的活性的实际结合位点接近或邻近的附加结合位点。可通过将LINGO-l多肽的结构与其它已知多肽的同源结构加以比较，来鉴定人LINGO-l多肽的候选活性位点。例如，可通过将LINGO-l多肽的结构与血小板膜糖蛋白Gplba的结构加以比较，来鉴定人LINGO-l多肽的一个或多个活性位点。LINGO-l上包括Trp346和Arg352的候选活性位点是通过其与Gplba上的配体结合位点结构的相似性鉴定出来的。位于人LINGO-l多肽表面上的腔(cavities)也代表候选的配体结合位点。人LINGO-l多肽的活性位点可包括SEQIDNO:1的氨基酸(图1)。例如，人LINGO-l多肽的活性位点可包括SEQIDNO:1氨基酸位置中示出的氨基酸Aspl3、Arg20、Arg22、Arg43、Glu60、Glu62、Leu94、Leul20、Metl23、Hisl76、Tyrl42、Hisl45、Hisl85、His209、His233、Ala238、Trp346、Arg347、Asn349、Asn351、Arg352、Gln353、Phe396、Arg408、Leu420、Leu426、Phe438、Asp440、Arg446、Tyr447和Ile459中的一处或多处。活性位点可包括位于人LINGO-l多肽特定区域的氨基酸的亚组。例如，活性位点可以位于LRR结构域的凹面，其包括SEQIDNO:1的Trp346和Arg352;His185、His209和His233;Aspl3;Glu60和Glu62;或Arg20、Arg22和Arg43。在另一实施例中，活性位点可位于LRR结构域的凸面，其包括SEQIDNO:1的Tryl42和Hisl45;Leu94、Leul20和Metl23;Hisl76;或Ala238。在另一实施例中，活性位点可位于Ig结构域，其包括SEQIDNO:1的Arg446和Tyr447;Arg408、Phe438和Asp440;Phe396;或Leu420、Leu426和Ile459。人LINGO-l多肽的氨基酸的编号可与本文所示的不同，人LINGO-l多肽的序列可含有保守氨基酸取代，其产生相同的三维结构。例如，人LINGO-l多肽的编号可不同于图1所示，并且人LINGO-l多肽的序列可含有保守氨基酸取代，但是产生与表2的坐标定义的以及图2-4和图5A-5C阐述的结构相同的结构。通过对相关氨基酸序列的视觉观察，或通过使用商业可获得的同源性软件程序(例如，MODELLAR，MSIManagementSimulations,Inc.,SanDiego,CA)，可容易地鉴定出其它同种型或类似物中对应的氨基酸和保守取代。表2所示的晶体结构坐标是从具有空间群P2,2-口的晶体测定的，但是相同的坐标+/-针对a碳原子而言不超过1.5A的均方根偏差可对应于具有不同空间群(例如1222)的结晶LINGO-l多肽。类似物是具有保守氨基酸取代的多肽。保守取代可包括用具有相似极性、立体排列或属于同一类(例如，疏水、酸性或碱性)的另一氨基酸来交换一个氨基酸，保守取代包括在对与多肽相互作用的物质的鉴定和设计方面，以及对分子置换研究和/或对同源性建模方面对于人LINGO-l多肽的三维结构具有不重要影响的取代。LINGO-l多肽的变体或突变体可用于本文7>开的方法、组合物和三维模型中。LINGO-l多肽或其片段的变体或突变体包括嵌合或融合蛋白、标记蛋白(例如放射性标记的蛋白质)、融合蛋白、突变体蛋白、具有相似(例如，高度相似)序列的蛋白质(例如，具有氨基酸取代(例如保守氨基酸取代)、缺失、插入的蛋白质)、蛋白质片段、模拟物，只要所述变体具有天然蛋白质的氨基#列的至少一部分或者具有与天然蛋白质序列高度相同的氨基酸序列的至少一部分即可。在一些实施方案中，突变体LINGO-l与SEQIDNO:l或其片段的氨基紗列具有多至5、10、20、30、50、75、100、150或200或更多个氨基酸残基的不同。在其它实施方案中，突变体LINGO-l与LINGO-l多肽序列，例如，人LINGO-l多肽(例如SEQIDNO:l)具有至少大约70%、80%、卯％、95%或更高的同一性。"嵌合蛋白"或"融合蛋白，，是编码多肽的第一氨基酸序列与第二氨基酸序列的融合，其中，所述第一和第二氨基酸序列天然状态下并不作为单个多肽链的一部分存在。在本文中使用时，术语"高度相似"(或"高度"或"充分""同源"或"相同")在本文中用于指含有足够数量的与笫二氨基酸或核苷酸序列相同或等同(例如，具有相似侧链，例如，保守氨基酸取代)的氨基酸残基或核苷酸的第一氨基酸或核苷酸序列。与本文公开的序列相似或同源(例如至少大约85。/。的序列同一性)的序列也是本申请的一部分。在一些实施方案中，序列同一性可以是大约90%、91%、92%、93%、94%、95%、96%、97%、98%、99%或更高。对两条序列间"同源性"或"序列同一性"的计算按照下文所述来进行。就最优比较的目的对序列加以比对(例如，可向第一和第二氨基酸或核^列之一或两条中都引入缺口，以进行最优比对，为了比较目的而言，非同源序列可不予考虑)。典型地，就比较目的而言被比对的参照序列的长度为参照序列长度的至少30%、优选至少40%、更优选至少50%、进一步更优选至少60%以及进一步最优选至少70%、80%、卯％、100%。然后对相应氨基酸位置或核苷酸位置上的氨基酸残基或核苷酸加以比较。当第一序列中的位置被第二序列中相应位置上相同的氨基酸残基或核苷酸所占据时，那么所述分子在该位置上就是相同的(在本文中使用时，氨基酸或核酸"同一性"与氨基酸或核酸"同源性"等同)。两条序列间的百分比同一性是序列所共享的相同位置的数量的函数，其中要考虑到为对两条序列进行最优比对需要引入的缺口的数量以及每个所述缺口的长度。对两条序列间百分比同一性的测定和对序列的比较可以使用数学算法来完成。在一个实施方案中，使用Needleman和Wunsch((1970)/Mo/.5,V/.48:444-453)算法(已被并入商业可获得的GCG软件包中的GAP程序中)来测定两条氨基酸序列间的百分比同一性，其中使用Blossum62矩阵或PAM250矩阵以及16、14、12、10、8、6或4的缺口权重以及1、2、3、4、5或6的长度权重。在另一实施方案中，使用GCG软件包中的商业可获得的GAP程序来测定两条核苷酸序列间的百分比同一性，其中使用NWSgapdna.CMP矩阵和40、50、60、3070或80的缺口权重以及1、2、3、4、5或6的长度权重。典型用于测定百分比同源性的参数是Blossum62计分矩阵，缺口惩罚为12，缺口延伸惩罚为4，框移缺口惩罚为5。还可使用E.Meyers和W.Miller((1989)CM5/OS14:11-17)的算法(其已被并入AUGN程序(版本2.0)中)来测定两条氨基酸或核苷酸序列的百分比同一性，其中使用PAM120权重残基表，缺口长度惩罚为12和缺口惩罚为4。LINGO-l多肽可来源于非哺乳动物或哺乳动物物种。哺乳动物LINGO-l多肽可来源于例如人。示例性的非人类哺乳动物包括非人灵长类(例如猴或猿)、小鼠、大鼠、山羊、母牛、公牛、猪、马、绵羊、野猪、海獭、猫和狗。示例性的非哺乳动物物种包括鸡、火鸡、坏、短吻鲟和鱼。来自数种哺乳动物和非哺乳动物物种(包括，例如，人、小鼠、大鼠、狗、黑猩猩和鸡)的LINGO-l氨基酸和核苷酸序列是本领域已知的(见，例如，Mi等人(2005)7VW7Ve"/wc/e"ce8:745-751;Carim國Todd，L.等人(2003)/7Ve"msc/.18(12):3167-3182和Okafuji等人(2005)6(1):57-62)物质可以是，例如，化学化合物(例如多肽、核酸、肽模拟物)。肽模拟物是能模拟肽识别某些生理分子(例如蛋白质和核酸)的能力的化学化合物。在一些情况下，肽模拟物包括能模拟或对抗天然亲本肽的生物学作用的非肽性结构元件。例如，易断开的肽键可被一个或多个不易断开的二肽等排物(isostere)替换。下述实施例意在说明，并非作为限制。实施例对LINGO-l结晶，和测定其结构。人LINGO-l是614个氨基酸的蛋白质，其含有信号序列、富含亮氨酸的重复样结构域、免疫球蛋白样结构域、茎结构域、跨膜结构域和短胞质尾(Mi等人，7V"tiVewwd.2:221-228，2004)。人LINGO-l的细胞外区域(包括LRR基序、Ig样结构域和茎区域，不包括信号序歹'J)在其C末端与六组氨酸标签融合(图1)。将得到的LINGO-lK549his6构建体亚克隆进pSMEG载体(SEQIDNO:2)(图6和7A-1至7A-3)的鼠巨细胞病毒(CMV)启动子之后(K549指位于全长人LINGO-l构建体的第249位氨基酸上的赖氨酸，其包括33个氨基酸的信号序列。信号序列在LINGO-lK549his6多肽中不存在)。通过测序对构建体加以验证。在5%C02和37°C下，在湿润的培养箱中生长和维持哺乳动物中国仓鼠卵巢(CHO)细胞。对Lec3.2.8.1的细胞培养和DNA转染描述于Zhong等人祝0/^j^.12^:143-150，2005)中。通过用抗his4抗体加以筛选，在Lec3.2.8.1(1042-5)中建立LINGO-lK549his6构建体的稳定克隆。大规模生产LINGO-lK549his6的条件培养基的流程以前有过描述(Zhong等人，祝ocA/肌5,V//^s.1723:143-150,2005)。为纯化LINGO-lK549his6，将表达融合蛋白的条件CHO培养基(图1)换为pH8.0的50mMTris、100mMNaCl的緩沖液，向其中加入蛋白酶抑制剂混合物(来自Roche，Nutley，NJ的完全抑制剂)。为捕获融合蛋白，在50mMTris,pH8.0，100mMNaCl中对Ni-NTA树脂加以平衡，然后加入培养基中，用于分批结合(颠倒)2小时。除去培养基之后，将Ni-NTA树脂装入柱中。将柱与AKTAExplorere(HPLC)相连，用4-5倍柱体积的50mMTris,pH8.0，100mMNaCl来洗。用12倍柱体积的、从0%至100%的50mMTris，pH8.0;100mMNaCl，300mM咪唑的梯度洗脱蛋白质。将收集的蛋白质级分渗析进50mMTris，pH8.0;500mMNaCl。为进行结晶，将蛋白质与羧肽酶A(Sigma，St.Louis,MO)—起在250C孵育14-16小时，然后渗析进50mMMES，pH6.1;50mMNaCl。渗析后，将蛋白质上样到SP琼脂糖凝胶柱(GEHealthcare)上，用4-5倍柱体积的50mMMES,pH6.1;50mMNaCl洗。用从0%至100%的50mMMES,pH6.1;750mMNaCl的梯度洗脱蛋白质。收集蛋白质峰级分，浓缩至5-8mg/ml，然后上样到用TBS平衡过的凝胶过滤Superdex200柱上。将蛋白质峰级分合并起来，浓缩至4-5mg/ml，用于结晶。纯化的蛋白质是完全糖基化的。在18。C使用悬滴蒸汽扩散法来获得LINGO-lK549his6晶体。从含4mg/mL蛋白质的蛋白质溶液和1.2-1.4M(NH4)2S04、O.IM柠檬酸钠，pH5.0的沉淀溶液来生长衍射质量晶体(diffractionqualitycrystal)。蛋白质在相同条件下以两种形式结晶，两种形式都在同样的结晶液滴中被发现1222，其每个不对称单位具有2个分子，74%的溶剂含量；以及P2J"，其每个不对称单位具有4个分子，73%的溶剂含量。为收集数据，将晶体逐渐从母液转至含2.9M丙二酸钠，pH5.2的稳定冷冻保护溶液(Holyoak等人,祝otog/c"/^9:2356-2358，2003)。该溶液(其中发现晶体在pH5-7的范围内在数天的时间内稳定)被用于晶体衍生。从钿衍生物获得允许通过具反常散射的单同晶型取代方法(SIRAS)进行结构测定的单衍生物，所述铂4汙生物是通过于pH7.0将晶体浸泡于50mMK2PtCl6和2.9M丙二酸钠中24小时来制造的。在数据收集之前，所有晶体都在氮气流下于IOOK纟皮沖冷。用从空间群1222的晶体获得的两组数据来进行相位测定3.5A天然数据库和3.6A数据组用于铂衍生物，两组都是在室内用安装于FR-ECuKa旋转阳极源(Rigaku，Japan)上的Saturn92CCD测量的。在先进光子源(APS)(Argonne，Illinois)上，用来自属于P2A2空间群晶体的SER-CAT的22-ID光束，将较高分辨率的天然数据组收集至2.7A。所有数据都^f皮整合，用HKL2000进4亍度量(scale)(Otwinowski和Minor,五"g;附o/.276:307-326,1997)。使用Pt原子在CuKa-边缘的反常差异，用SHELXD(2001,Bruker画AXS，XM，Ver.6.12;Bruker画AXS,Madison，WI)，对衍生晶体中柏(Pt)原子最初的位置加以定位。用XPREP(2001Bruker-AXS，Ver.6.12;Bruker-AXS，Madison，WI)来产生输入SAS系数。使用来自天然和衍生数据库的反常和同晶型差异，以20-3.6A的分辨率，用SHARP(deLaFortelle和Bricogne，Me^oA^:472-494，1997)来进行对重原子参数的精确修正、相位计算和密度修改。用SHARP产生的最终的3.6A的、具反常散射的单同晶型取代(SIRAS)图i普具有可供判断的质量，其揭示了不对称单位中的两个LINGO-l分子。通过两倍的非晶体学对称(NCS)平均和在DM中将相位延伸至3.5A来进一步改善SHARP相位(Cowtan和Main,^"flOyW.^[2:148-157，1993)。得到的图i普令我们可以用QUANTA(Accelrys，Inc.,SanDiego,CA)建立初始的、卯％完成的(大约855个残基)模型。然后使用该^t型，用P2,2J数据组进行分子置换，以利用更高的2.7人的分辨率数据。用PHASER(McCoy等人，j^l:458誦464，2005)来鉴定不对称单元中四个分子的清澈溶液。1222和P2JJ晶体形式共享相同的四聚体填装，其中，四聚体可通过围绕两倍轴复制二聚体来构建。用(结晶目标)-定向的工具包(COOT)(Emsley和Cowtan,^ctoO^s.D^:2126-2132，2004)和REFMAC(Murshudov等人，爿c似OjW"//ogr.Z)fl:240-255，1997)针对2.7A的数据组进行后续数轮的重构和精确4务正。最后的模型含有四个蛋白质分子(残基Al-477、B3-475、C2-477和D3-476)、39个N-乙酰葡糖胺和12个甘露糖残基以及310个水分子。B、C和D的N末端的残基1-2;B和D的C-末端的476-477和残基D32-34没有被建模进入结构，因为缺少足够的电子密度，这被假设是因为无序(disordering)造成的。用MolProbity进行的对经过最终精确修正的结构的几何分析将所有残基中的94%放进了有利的区域中，0.16%是外露的。表1中概括了针对数据收集、定相和精确修正的统计学分析。表2列出了从空间群P2^2的晶体推导出的晶体结构坐标(见表1)。具有空间群I222的天然和衍生结构揭示了高度相同的结构，它们被用于协助定相。在表2中，"#"栏向被给予坐标的每个原子指定指数。"名称"栏则表明原子的类型，"res，，栏表明原子属于何种类型的残基。"链"表示原子所述的是何种多肽。"Resr，给出了原子的残基号。例如，原子号1(表2中的第一列)是Thrl的氮(N)(根据SEQIDNO:1的序列)。在X、Y和Z栏中分别给出其x、y和z结构坐标。标题为"occ"的栏描述了被指定原子的占据率(l.OO=完全占据)，"B"栏提供了12单位的B因子(或温度因子)。标有"元素，，的栏列出了原子的元素符号。水用"HOH"表示。"NAG"和"MAN"分别表示N-乙酰葡糖胺和甘露糖残基。表l.数据收集、定相和精确修正的统计学分析数据收集天然lK2PtCl6天然2空间群12221222P2^2晶胞尺寸148.7149.6201.5b(A)158.6157.3149.7"A)200.0200.3157.5p=p=Y(度)909090源FR画ECuKaFR國ECuKaAPSID22最大分辨率(A)3.53.62.7反射(总的/独特的)251,385/29,544201,963/27,406915,151/129,631完成率(％)97.1(98.8)99.1(99.8)98.9(94.7)Rsyma(%)11.7(50.2)10.5(45.9)9.0(61.2)IM(I)16.1(3.6)15.3(3.4)20.6(1.2)定相SIRASAnom.I/o(I)(4.5A/3.6人)2.3/1.2Risob(%)43.6Ranob(%)8.6Pt位点的数量15定相力e(4.5A/3.6A)反常同晶型1.13/0.61.1/0.73晶体品质因素均值d(4.5A/3.6A)0.45/0.25模型精确修真天然2分辨率(A)50.0-2.7反射的数量122,982完成率(％)98.7(91.3)Rfacto鳥ee*(%)21.5/25.5蛋白质原子的数量15,114碳水化合物原子的数量700水的数量310与理想几何条件的r.m.s偏差键(人)角(deg)0.0091.25aRsym-HillOOi_<I(h)>I/ZI(h)i，其中<I(h)>是平均强度。括号中的数字反映了对最高分辨率外形(shell)的统计学分析。bRis。=i:IFnat(h)—FPt(h)I/ZFnat(h)，Ran。是针对Bijvoet对的阳性和阴性副本的幅度来计算的。c定相力被定义为叫FHlW〈(缺少封闭(lack-of-closure))、其中'H，代表重原子。d晶体品质因素均值是估计的相位错误的平均余弦。、free是用数据的5%计算的。LINGO-l的两种晶体结构(1222和P2JJ)揭示了与凝胶过滤数据和动力学光散射测量相一致的四聚体结构，这表明，LINGO-l作为四聚体存在于溶液中，分子量为大约240kDa。图2显示了针对经精确修正的2.7A分辨率结构的电子密度的片段。图3中显示了单体LINGO-l结构的总视图。每个单体都是由两个不同的结构域形成的残基1-382组成了N末端LRR结构域，残基383-477形成了C末端的Ig样结构域。两种组分结构域之间的相对定向赋予单体高难度结构的形状(questionmarkshape)。LRR结构域(残基l-382)采用延长的弧形，其具有在凹内面上平行的15链p-片，还具有凸外面上的高度反常的二级结构(图3)。总而言之，有12个LRR，每个23至25个残基，它们一起产生连续的右手超螺旋集合。每个LRR都从p-链开始，其具有24个残基的序列重复基序xL2xxL5xL7xxN10xL12xxLlsxxxxF20xxL23x(SEQIDNO:9)，其中，x可以是任何氨基酸；L是疏水残基，优选是L但也可以是Ile、Val、Met、Phe或Thr;N是天然较不保守的，其包括主要是Asn，但也可以是Cys、Asp、Leu或Trp;F代表疏水残基Phe或Leu。指出的位置上的共有残基构成了LRR结构域的内部。与数种其它LRR蛋白质中一样，两种LRR帽("N帽"和"C帽，，)使得LRR疏水核心的N和C末端针对溶剂而言被遮蔽(图3)。"N帽"具有两条反平行的P-链，其在底部(base)具有两个二石克键(Cys-Cys9和Cys7-Cys18)，这是与NogoR结构上非常接近的排列(He，等人，A^w/wi^:177-185，2003)。"C帽"还被两个二硫键(Cys334-Cys357和Cys336-Cys382)支持，并且，与NogoR类似，具有由一个a-螺旋和三个短310-螺旋构成的特征性基序。但与NogoR不同，LINGO-l中"C帽，，的内部基本全部填充苯丙氨酸蔟(来自Phe342、Phe350、Phe362、Phe368、Phe371、Phe380的六个芳香族环和来自Tyr379的一个芳香族环)。此外，a-螺4t之后的短环片断(残基349-353)以与NogoR中略有不同的方式整合进规范的LRR结构。这导致5个残基环的顶部离开P-片凸出进凹面空间至少7.5A，因此展示出比NogoR中同样长度的相应片断更突起的特点。凸出环的位置(图3)(对应于血小板膜糖蛋白Gplba中配体结合Ji-开关环(Huizinga等人，Sdewce122:1176-1179，2002))与表面暴露和突起侧链(Trp346、Arg347和Asn349、Asn351、Gln353和Arg352，位于环尖)的特点组合，暗示其可能是与配体相互作用的候选者。分子的静电表面(图4)揭示了在LRR结构域的所有侧面上的正电和负电补片(patch)，只除了在N末端凸侧发现的一个相对较大的疏水补片之外。基于四聚体复合物的结构，位于LRR结构域凹面上的五个氨基酸亚组被确定为候选的配体结合位点(1)Trp346和Arg352;(2)Hisl85、His209和His233;(3)Aspl3;(4)Glu60和Glu62;和(5)Arg20、Arg22和Arg43。类似地，位于LRR结构域凸面上的四个氨基酸亚组被确定为候选的配体结合位点(1)Tryl42和Hisl45;(2)Leu94、Leul20和Metl23;(8)Hisl76;和(9)Ala238。完成LRR结构域后，多肽链进入压缩的Ig样折叠(残基383-477)，其代表经典的、由一个二硫键(Cys407-Cys458)交联的两个p-片的卩-三明治。来自一个P-片的四条链(A、B、D和E)和五条链(A'、C、C'、F和G)构成了第二P-片。有一个310a-转角连接E和F链。与其它Ig样结构的比较表明，LINGO-lIg样结构域的最接近的结构的神经细胞粘附分子NCAM的Ig3模块(Soroka等人，5^""""!￡:1291-1301，2003)，计算出的两条序列的同一性为约30%。并不出人意料地，氨基酸替换的绝大多数都发生于Ig结构域表面，这改变了侧链体积和/或侧链极性，而保守氨基酸中的大多数都定位于结构的核心，并且具有等同的构象，以满足折叠。表面静电势是ABDE-表面上负和正补片的组合。相反，反向的A，CC'FG-表面则高度富含疏水残基(Ala416、Leu418、Leu420、Leu426、Leu457、Ile459、Ala461、Ala463)，它们被大量的极性和带正电荷的残基环绕。Ig结构域定向使得其远离凸LRR弧而突出，在LRR结构域的背面产生宽广的暴露给溶剂的裂缝，并且具有大约90度的急弯(图4)。裂缝的壁是由一侧的LRRC-末端凸面(重复10-12和C帽)和另一侧的Ig结构域的A，CC，FGp-片形成的。反面则相对远离，并且具有不同的性质。凸面主要由极性和带电荷残基构成，而反面的Ig面则相当平，排列有主要疏水的氨基酸(如上文所述)。在急弯的尖上发生小的结构域间相互作用，这涉及FG环上的残基(Ala463、Gly464)和LRR的C末端部分(Cys336、Arg337和Cys382)的范德华接触。需要注意的是急弯尖上的二硫键——Cys336-Cys382以及结构域间不存在接头(因为Arg383是第一P-链的第一个残基以及ABDEp-片的组分)。结构域间的此类联结限制了它们两个互相之间的定位。基于四聚体复合物的结构，位于lg结构域凹面上的五个氨基酸亚组被确定为候选的配体结合位点(1)Arg446和Tyr447;(2)Arg408、Phe438和Asp440;(3)Phe396;和(4)Leu420、Leu426和Ile459。我们预计，CHO受限细胞中LINGO-l的糖形式主要是高甘露糖形式的多糖，例如MansGlcNAc2,这与我们的质谱数据一致。结晶的糖蛋白含有八个典型的N-糖基化共有位点。其中六个位于LRR结构域中(Asn105、Asnl63、Asn225、Asn235、Asn254和Asn302),两个位于Ig样结构域中(Asn466和Asn453)。所有这些位点都位于结构域表面。可对所有这些位置上的碳7JC化合物基序给予电子密度，只除了Asn466位点。后者虽然也位于表面，但却被对称性相关LINGO-l伴侣紧密包裹，如下文所述和图5所示。因为蛋白质包裹没有给Asn466留存糖基化的空间，因此与Asn466相连的N-多糖将阻止在晶体中观察到的LINGO-l四聚化。剩下的七个位点上的寡糖在如图4的被编号的残基所示的结构上铺展，其中，#1对应于Asnl05，^2对应于Asn163，#3对应于Asn225，弁4对应于Asn235,#5对应于Asn254，糾对应于Asn302,#7对应于Asn453。在LRR结构域的四个面外，仅凸外面完全没有碳水化合物，此发现与其在自联作用中的作用一致。凹面和两个侧面(每个都具有两个N-多糖结合位点)被碳7jC化合物大大覆盖。最后的碳水化合物部分定位到Ig-面A，CC，FG上，接近C末端，并且远离分子间界面(图4)。一般而言，在糖蛋白表面上大量的糖基化的存在会限制它们与配体相互作用发生的可及性，在大多数情况下，没有被糖覆盖的表面是配体预计会结合之处(Rudd等人，/Af必293:351-366，1999)。因此，LINGO-l的凹LRR面(在弧中心具有两个糖部分)看起来不可能在容纳此类大的蛋白质伴倡(如NogoR或p75)中扮演主要角色。这与频繁使用凹面与它们的配体发生相互作用的其它含LRR受体相反(Bell等人，7>wi*/附附wto/.2^:528，2003)。因为没有证据表明与Asn相连的N-多糖直接与配体结合相关，因此也有可能是LINGO-l糖基化协助阻止非特异性蛋白质-蛋白质相互作用或者协助保持正确的骨架用于随后的配体结合。结晶、体积排阻色谱和动力学光散射都表明，LINGO-l在晶体和溶液中都采用四聚体形式。在晶体中，LINGO-l的四个独立的拷贝相互作用，构成被压紧的四聚体结构。旋转的假四倍轴将环样结构中的四个分子与中心的大洞(直径大约40A，外环直径大约95A;图5A)关联起来。这种排列将每个LRR结构域的N末端凸侧放置于邻近分子的结构域间的急弯上(图5B)，令人联想到细胞因子/受体结合模式。在后一种情况下，两个分子间的连结点是三个表面提供的N末端弯曲重复l-6形成的一个、如上文所述分别通过C末端弯曲重复11-12、"C帽"和A，CC，FGp-链形成的两个急弯表面。因此，四个LRR结构域头尾联锁成环，相应的Ig样模块从环的周界以起落架(landinggear)方式延伸(见图5C)。如果我们将这种几何条件相对细胞表面来应用的话，那么蛋白质旋转轴将以大致垂直于表面，接近膜的C末端将指向同方向，即朝向细胞膜。每个分子间界面都揭示了结合表面间的紧密几何匹配和相互作用的精确立体化学互补性。在每个格间界面下埋藏着的溶剂可达的表面区域相对较大，来自三种各組分，总共2400A2，这暗示了强烈的关联。上述证据组合相互作用的多价特点暗示，四聚体内的接触并不仅是晶体接触。表2.LINGO-lK549his6多肽的结构坐标(分别为SEOIDNO:5-8)<table>tableseeoriginaldocumentpage43</column></row><table><table>tableseeoriginaldocumentpage44</column></row><table>原子86CBASPA1347.45347.69522.8231.0078.35C原子87CGASPA1347.61548.21624.2391.0079.47C原子880D1ASPA1347■48849.45124.44310081,.370原子89OD2ASPA1347.8594739425.15110079,.74〇原子90CASPA1348.20347.85520.4581.0077.63C原子910ASPA1349.05046.98120.2751.0077.600原子92NARGA1447.3214824419.5321.0076.93N原子93CAARGA1447.09547.53018.2751.0076.17C原子94CBARGA1448.25647.74917.3001.0076.39C原子95CGARGA1448.45149.22916.9531.0077.97C原子96CDARGA1449.71449.48016.1481.0080.67C原子97NEARGA1449.92650.90915.9181.0083.36N原子98CZARGA1450.61751.42514.8991.0084.74C原子99丽lARGA1450.73952.7441477510083.73N原子100NH2ARGA1451.17950，62713，9941.0085.78N原子101CARGA1446.81146.05118.5611.0075.25C原子1020ARGA144740845.16317，9711.0075.140原子103NalaA1545.88645.82619.4931.0074.33N原子104CAALAA1545.,48144..50719.,9471.,0073.,70C原子105CBALAA1545.,54644..44621.,4521.,0073..72C原子106ca:laa1544.07244.180is-4801.0073.38c原子1070ALAA1543.24445.069IS.3261.0073.570原子108NVALA1643.80942.90119.2431.0073.08N原子109CAVALA1642.,48342.,43118.,8671..0072.,86C原子110CBVALA1642..35342.■20017..3381..0072.,76c原子111CGIVALA1641.10841.38217.0121.0072.82c原子112CG2VALA1642.30943.52716.5941.0072.51c原子113Cvala164221141.146IS-6281.0072.88c原子1140VALA1642.■97640.199IS.5501.0072710原子115NLEUA1741.11141.12620.3631.0073.25N原子116CALEUA1740,.79540..02321,.2371,.0073,.61C原子117CBLEUA1740,.95740..50322.6751.0073,.93C原子118CGLEUA1740,.83439,.52223,.83510075,.24C原子119CD1LEUA1741.87539.870248851，0077.28C原子120CD2LEUA173941939.525244401.0076.11C原子121CLEUA1739.,38439.50020.,9551.,0073..77c原子1220LEUA1738.,40840..21821■.1181.,0073..270原子123NCYSA1839.,29138.,24620.,5201.,0074.,57N原子124CACYSA1838.01537.638201281.007547C原子125CBCYSA1837.94437,50018.6081.007544C原子126SGCYSA1838.02539,.05917.7411.0076,.95S原子127CCYSA1837.82136.■26220.,7401.,0076.04C原子1280CYSA1838■,23335..26220.,1561.■0076..18〇原子129NHISA1937.,18736.,19921.,9071.,0076..81原子130CAHISA1936.95934.90822.5511.0077，38C原子131CBHISA1937.36834.95924.0181.0077.93C原子132CGHISA1938.83835.13524.2201.0079.99C原子133ND1HISA1939.41036.357244931.0081.52N原子134CE1HISA1940.71836.21024.6131.0083.13C原子135NE2HISA1941.01534.938244141，0083.73N原子136CD2HISA1939.85634,24324.17110082.59C原子137CHISA1935.51534.46722.4311.0077，21C原子1380HISA1934.60935.29422.4621.0077,010原子139NARGA2035.32333.15822.2791.0077.16N原子140CAARGA2034.00232.52122.3411.0077.64C原子141CBARGA2033.41932.63823.7571.0077.35C原子142CGARGA203412531.770247841.0078.80C原子143CDARGA2033.60232.02226.1931.0080.18C原子144NEARGA2033.97233.35426.6861.0086,15N原子145CZARGA2035.08033.62927.3821.0088.16C原子146NH1ARGA2035.95732.66827.6951.0087.59N原子147NH2ARGA2035.30834.88227.7671.0089.44N原子148CARGA2033.00033.02821.3071.0076.83C原子1490ARGA2031.81333.15721.6051.0076.960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4112.098原子3851CAARGB14113.,463原子3852CBARGB14114.,328原子3853CGARGB14114..661原子3854cdargb14115..545原子3855NEARGB14116..147原子3856CZARGB14115..630原子3857NH1ARGB14116.268原子3858NH2ARGB14114.478原子3859CARGB14113.440原子38600ARGB14114.102原子38611S1ALAB15112.652原子3862CAAiAB15112.466原子3863CBALAB15111,.022原子3864CALAB15112.886原子38650ALAB15112.674原子3866NVAXB16113.497原子3867CAVALB16113937原子3868CBVALB16115.477原子3869CGIVALB16115.97157.526-9.8561.0091.47C56.972一8.8051.0091.32058.802-10.,1231.0091.,06N59.659-9.2241.0090.79C61.058—9.8391.0090.66C59.766-7.8071.0090.60C59.770-6.8271.0090.40059.840-7.7141.00SO-54N59.993-6.4361.00SO.68C60.304-6.6651.0090.70C59.326_7.6011.0091.01C59.327_7.4371.0090.98C60.380—7.4511009098058.133_7.2991.0090.76N58.803-5.4751.0090，61c58.939-4.2741,0090.76057.644-6.0001.0090.32N56.449-5.1781.0090.06C55.■318-5.6421.0090.22C55.627-5.4411.0090.97C56，819_5.47010091.85〇54.666-5*2701.0091.65055.934_5.1931.0089.80C54.950一4.5221,0089.86056.588-5.9611.0089.53N56.,107-6.2011.■0089.■28c56.,232_4.9351.0089.34c57.■667_4.5171.■0090.■06c58.,343_5.,5621.,0091.,54c59..601-5.1141.0092.,31N60,.809_5.,3311.,0092.,89c61889—4.8921.0092.54N60.943-5.9811.0092.95N54.669-6.7551.0089.08C53.762—6.2381.0088.99054■485-7，8121.0088.79N53,.192-8..4581..0088..35C52,.760-8,.3461,.0088■.34C53.264-9.9191.0088.14C54,269-IO.5791.008822052.,195_10.4071.0088，11N52.100-ll.7891.0088.19C52.212-ll.8911008809C51.761_13;.248■0088.13C原子3870CG2VALB16115.94453.637-ll.6121.0088，25C原子3871CVALB16113.46750.754-12.3401.0088.62C原子38720VALB16113.75149.701-ll.7451.008847〇原子3873NLEUB17112.73550.793-13.4581.0088.94N原子3874CALEUB17112.■22149.■571_14.■1011.0089.,30C原子3875CBLEUB17110.,69049.,564-14.1211.,0089.,19C原子3876CGLEUB17109.94948.,860-12.9691.0089.,80C原子3877CD1LEUB17110.32649.380_11.56810089.93C原子3878CD2LEUB17108.43248.935-13.1791，0089.68C原子3879CLEUB17112.78449.391-15.5071.0089.39C原子3謂0LEUB17112.80150.335-16.2881.0089.33〇原子3881NCYSB18113.248AS-180-15.8161.0089.66N原子3882CACYSB18113.959ST.906-170711.0090.08C原子3883CBCYSB18115.46648■Oil-16.8571.0090.■26C原子3884SGCYSB18116.01749.617_16.3131.0092.53S原子3885CCYSB18113.65246.539-17.6761.0089.87C原子38860CYSB18114,56045.727-17.8741.0090.120原子3887NHISB19112.38346.298-17.9921.0089.56N原子3固CAHISB19111.95045.■018-18.■5351.0089.15C原子3889CBHISB19110.48844■763_18.1681.0089.33C原子3890CGHISB19110.22944.760-16.6911.0090.55C原子3891ND1HISB19110.13543.598-15，9521.0091.93N原子3892CE1HISB19109.90643.900-14.6851，0091.91C原子3893NE2HISB19109.85045.216-14.5751.0091.69N原子3894CD2HISB19110.04845，777-15.8141.009110C原子3895CHISB19112.14344.949-20.0491.0088.75C原子38960HISB19111.85645.911_20.7631.0088，770原子3897NARGB20112.64743.謂-20.5161.008827N原子3898CAARGB20112.,75843..481-21.,9461.,0088..00C原子3899CBARGB20111..37143,.287-22..5801..0087,.92C原子3900CGARGB20110..71641,.956-22..2171..0088..62C原子3901CDARGB20109.,29241,.852-22■.7341..0088..92C原子3902NEARGB20108,,30042.317-21,.7631,.0091,.46N原子3903CZARGB20107..54241,.515_21,.0121.,0093,.12C原子3904NH1ARGB2010"/.65440.193-21.1061.0094.06N原子3905NH2ARGB20106.66342.032-20.1581.0093.40N原子3906CARGB20113.61644.448-22.7621.0087.45C原子39070ARGB20113.25344.■831_238781.0087.■530原子3908NLYSB2111475944.■833-22.20110086■73N原子3909CALYSB21115..68045.731-22.8911.0085.86C原子3910CBLYSB21116,04846.932_22.0081.008595C原子3911CGLYSB21114,89747.568-21.2221.0086.19C原子3912CDLYSB21113.92848.363_22.0981.008689C原子3913CELYSB2111285349.028—21.2381.0087.42C<table>tableseeoriginaldocumentpage0</column></row><table>原子3958nprob27122.78048.883-16.5201.0081.29n原子3959caprob27122.81150.039-17.4131.0081.65c原子3960cbprob27121.36050.153-17.8771.■0081.54c原子3961cgprob27120.■57349.554-16.■7881.0081.08c原子3962cdprob27121.40348.412-16.2991.0081■22c原子3963cprob27123.21951.306-16.6711.0082.14c原子39640prob27122.96351.428-15.4731.0082.31o原子3965nglub28123.84752.239-17.3751.0082.58n原子3966caglub2812429453.475-16.■74410083.18c原子3967cbglub28125.44954115-17.,5271..0083.,67c原子3968cgglub28126.51953.141-18.0461■0085.■07c原子3969cdglub28126.18852.601-19.4361.0086.69c原子3970oe1glub28125.21751.800-19.55910086.47〇原子3971oe2glub28126.90152.991-20.3971.0086.32o原子3972cglub28123.13354.463-16.5951.0083.07c原子39730glub28122.04354.241-17.1351.0083.110原子3974ngi;yb29123.37255.541-15.8501.0082.94n原子3975caglyb29122.■37456.590-15.6431.,0082.,70c原子3976cglyb29121.47856.393-14.4301.0082.55c原子39770glyb29120.73657.304_14.0561.0082.75o原子3978nileb30121.53855.205-13.8261.0082.17n原子3979caileb30120.,76254..885-12..6221..0081..74c原子3980cbileb30120.,86853.,380-12,,2531.,0081.,61c原子3981cgiileb30120.28852.515-13.3741.0081.52c原子3982cd1ileb30120.57951.034-13.2261.0081.86c原子3983cg2i:leb30120，15553.086_10.9321.0081.35c原子3984cileb30121.20755.746_114351.0081.53c原子39850工:leb30122.40155.797-ll.1171.0081.46o原子3986nprob31120.24656.432-107821.0081.33n原子3987caprob31120,.55857..267-9.6231.008127c原子3988cbprob31119,19257.832-9.2171.0081.32c原子3989cgprob3111834357.707_10.4321.0081.28c原子3990cdprob3111880756.462_111031.008122c原子3991cprob31121.15256.,450-8.4761.0081.19c原子39920prob31120■83555.,270_8.3251.0081.190原子3993nthrb32122.,01557..077-7.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7原子11149〇valc450115.992-3.319-9.7791.0076.12原子11150nglnc451115.946-4.589-7.9171.0075.86原子11151cag;lnc451115.581-3.477-7.0251.0075.76原子11152CBGLNc451115.062-4.009-5.6811.0075.74原子11153CGGLNc451116.023_4.936-4.9351.0075.26原子11154CDGLNc451115.774一4.,948_3.4351.0074.39原子11155oe1g:lnc451115.415_5.981-2.8641.0073.37原子11156NE2GLNc451115.955-3.793-27911.0073.24原子11157CGLNc451116.706-2.443-6.8031.0075.82原子111580GLNc451116.688-l.693-5.8201.0075.77原子11159NASPc452117.670-2.411-7.7231.0075.77原子11160CAASPc452118.,807-l..495-76561.0075.77原子11161CBASPc452120.083-2.,246-7.2721.0075.93原子11162CGASPc452120.■006_2.,865-5.■89410076.65原子11163OD1ASPc452119.591-2.166-4.9411.0077.38原子11164OD2ASPc452120.365—4055-5.7661.007729原子11165caspc452119.022-O.795_8.9881.0075.63原子111660ASPc452119,8870.078-9.1001.0075.53原子11167NASNc453118.246-1.195-9.9961.0075.52原子11168CAASNc453118.,283_0.,563-ll.3111，0075.39原子11169CBASNc453117.,284-l.,230_12.2651.0075.52原子11170CGASNc453117.545-o..882-13.7251.0076.02原子11171OD1ASNc453117Oil0，101_14.2611.0078.12原子11172ND2ASNc453118.387-l.677-14.3691,0075.90原子11173CASNc453117.9630.921一ll.■2041.0075.14296原子11174〇ASNC453117.2151.341-10.3171-0075.040原子11175NGLYC454118.5441.710-12.1011.0074.90N原子11176CAG1jYC454118.■1953.113-12..2091..0074.,77C原子11177cg:lyc454119.3474.084-12.0831.007463c原子111780GIiYC454120，5183.696_12.1201.0074.680原子11179NTHRc455118，9865，356-11.9331.■0074.42N原子11180CATHRc455119.9296,460.8811..0074,.19C原子11181CBTHRc455119.,2937.750-12■.4481,.0074.,40C原子111820g1thrc455118.9577.553-13.8301.0074550原子11183CG2T服c455120.2498.943-12.31410074.66c原子11184CTHRc455120.4086.723_10.4591.■0073.87c原子111850THRc455119.6126.836-9.5261.0073.940原子11186NTYRc456121.7226.819-IO..3141,,0073.,32N原子11187CATYRc456122.,3457.192-9.0641.0072.86C原子11188CBTYRc456123.,4346.178-8.7131.0072.25c原子11189CGtyrc456122.,9054.834-8.2561.0071.59c原子11190CD1TYRc456122，9744.461-6.9181.0070.82c原子11191CE1TYRc4561224983.234_6..4881.0070,.25c原子11192CZTYRc456121.,9342.366_7.3981.0070.39c原子11193OHTYRc456121■.4511.163一6.9601.0070.910原子11194CE2TYRc456121.8422.706_8.73310070,.58c原子11195CD2TYRc456122.3303.936-9,.15810071,.26c原子11196CTYRc456122.9518.574-9.2471.0073.16c原子111970TYRc456123.3878.921-IO.3411.,0073.550原子11198NLEUc457122.9599.381-8.1981.0073.39N原子11199CALEUc457123.,70010,.633-8.,2461.■0073.,63c原子11200CBLEUc457122.84611,.814_7■7851..0073.■23c原子11201CGLEUc457122.11312..737-87551.0072.64c原子11202CD1LEUc457121.78814.026一8.0351-0071.11c原子11203CD2LEUc457122.91913.032—9.9931.0072.06c原子11204CLEUc457124.88010.527_7.3231.0074.22c原子112050LEUc457124.71310.140-6.1651，0074120原子11206NCYSc458126.07310.853-7.8131.0074.98N原子11207CACYSc458127.13511..181_6.8731.,0076.07C原子11208CBCYSc458128.,50510,.574-7.2071.,0076.,49c原子11209SGCYSc458129.,61811,.583-8.,1641..0081.■04s原子11210CCYSc458127.18012.677-6.7491.0075.44c原子112110cysc458126.97313397_7.7201.0075.160原子11212NILEc459127.42013.124-5.5231.0075.26N原子11213cailec459127.,39114,.529-5.,1751..0075.■02c原子11214CBILEc459126.,23214■.,—4.,2091.,0075.,21C原子11215CG1ILEc459124.92614.284_4.8131.0075.42c原子11216cd1i:lec459123.95013.787-3.8071.0075.30c原子11217CG2IIuEc459126.12916.308一3.8991.0075-48c297说明原子11218CILEC459128，69814，880-4.507l'0074.52原子11219'〇ILEC459129.01614，334-3.4571.0074.98原子11220NAIAC460129.47315.766-5.1161.0073.85原子11221CAALAC460130.71816.193-4.4931.0073.,43原子11222CBALAC460131.79516.,355-5.5121.■0073.'25原子11223CALAC460130.45217，507-3.8031.0073.37原子112240ALAC460129.90518.430-4.4161.0073.30原子11225NAIAC461130.83617.598-2.5341.0073.12原子11226CAALAC461130.51018.,781-l.7491.,0073,,04原子11227CBALAC461129.,29718.,506-O.,8891.,0073..05原子11228calac461131.659is-277_0.8891.0073.08原子112290alac461132.326is.486_0.2151.0072.86原子11230NASNC462131.89020.589-0.9321.0073.12原子11231caasnc462132.,76421.,2500.0371.0073.43原子11232CBASNC462134.23421.,284-O.4271..0073.,38原子11233CGASNC462134.,48122.,235-l.,5791..0072,.67原子11234ODlASNC462133.57922.920_2.0511,0073.18原子11235nd2asnc462135.71922.278-2.0381.007179原子11236CASNC462132.26322.6320.4541.0073.90原子11237〇ASNc462131.29023.151-0.1001.0074.02原子11238na;lac463132.93323.2221.4341.007428原子11239CAALAc46313253724.,5131.9661.0075.02原子11240cbalac463133.61125.,0372.8661.0075.13原子11241CALAc463132.18825.5570,9021.0075.74原子112420ALAc463131.27726.3491.1091.0076.32原子11243NGLYc464132.90425.564-0.2241.0076.08原子11244CAGLYc464132.■70526.,582-l.,2551..0076,.38原子11245CGLYc464131.55526.349-2.2261.0076.■74原子112460GLYc464131.08427.291-2.8761.007718原子11247NGLYc465131.10725.101-2.3431.0076.56原子11248CAGLYc465130.,05924..743-3..2961,.0076,.44原子11249CGLYc465129.95723.244-3.5051.■00"76..49原子112500GLYc465130.64222.462-2.8351.0076.,53原子11251nASNc466129.09922.835-4.4331.0076.54原子11252CAASNc466128..93421,.414一4,.7311.0076.76原子11253CBASNc466127.,86520..771-3..8431,.0076,,81原子11254CGASNc466126.,51221,.425-3,.9881,.0077.49原子11255ODlASNc466125.82721.247_4，9941.0077.99原子11256ND2ASNc466126，11122.185_2.9711.0078.87原子11257CASNc466128，67821.108-6.2021.0076.■83原子112580ASNc466128.57522.018-7.0251.0076..71原子11259nASPc467128.59319.816-6.5081.0076.■83原子11260CAASPc467128.,49419..327-7,.8711..007711原子11261CBASPc467129..88519..271一8.,4991..0077,.07原子11262CGASPC467129..85019,.303-10.0171,0078,.72原子112630D1ASPC467128.83819.764-10.6021.0080.20原子11264OD2ASPC467130.85318.874_10.6351.0080.03原子11265CASPC467127.85917.,933-7.8761.0077-20原子112660ASPC467127.87417,,222-6.8641.0077-47原子11267NSERC468127.30117.,540-9.0161.0077.09原子11268CASERC468126..68116.227-9.1381.0076.79原子11269CBSERC468125,.21316.268-8.6971.0076.76原子11270OGSERc468124.■58517.480-9.0881.0076.95原子11271CSERc468126.81515.,655-10..5391..0076..56原子112720SERc468126.92816.,394-11.,5091.,0076..68原子11273NMETc469126.84114.,329-10..6111..0076..58原子11274CAMETc469126..83313579-ll.86210076.53原子11275CBMETc469128,,23413.072-12.2221，0076.48原子11276CGMETc469129,,05714■.068-13.0091.0077.12原子11277SDMETc469128,.42814329-14.6841，0078.54原子11278CEMETc469128,.69216.101-14.9001.0077.01原子11279CMETc469125.89212.,394-ll.,7281.,0076.,44原子112800METc469125.85811.,751-10..6761..0076..46原子11281NPROc470125.09112.,127-12.,"51.,0076.,41原子11282CAPROc470124..33410..881-12.8371.0076.26原子11283CBPROc470123..34011.122_13.9851.0076.37原子11284CGPROc470123.,41312.606-14.2901007629原子11285CDPROc470124..80512.997-1393010076.35原子11286CPROc470125,2249.679-13.1491.0076.23原子112870PROc470126.2219.797-13.8701.0076.07原子11288NALAc471124.8648.541_12.5671.0076.36原子11289CAALAc471125,4367.243-12.9071.0076.,44原子11290CBALAc471126.,2846.7057681.,0076.,36原子11291CALAc471124.2736.305-13.2071.0076.51原子112920ALAc471123.2536.335-12.5121，0076.41原子11293NHISc4721244165.489-14.2481.0076.65原子11294CAHISc472123.,3074.657-14.7081.■0076.,85原子11295CBHISc472122.9565.003_16.1531,0076.,75原子11296CGHISc472122.,7156.463-16.3711.,0077.,08原子11297ND1HISc472121.5677.099_15.9481.0077.51原子11298CE1HISc472121.6338381-16.2651.0077.00原子11299NE2HISc472122.7858，601-16-8721-0076-96原子11300CD2HISc472123.4827.418_16.94910077.13原子11301CHISc472123.5693.159-14.5331.0077.00原子113020HISc472124.4442.588-15.1951.0077.19原子11303NLEUc473122.8122.532-13.6321.00"76.92原子11304CALEUc473122.9371.100_13.4061.0076.97原子11305CBLEUc473122.8880.759-ll9161.■0076.,75<table>tableseeoriginaldocumentpage300</column></row><table>说明原子113500cysd374.60655.32694.4211-0094.49原子11351nprod474.52353.45393.1351.0094.37原子11352CAPROD474.34952.44294.1881.0094.36原子11353CBPROD474.80251.15693.5031.0094.38原子11354CGPROD474."951.35992.0811.0094.40原子11355CDPROD47453852.83591.7961.0094.37原子11356CPROD475.19652.69795.4421.0094.40原子113570prod476.39752.95195.3251.0094.35原子11358NPROD57456952.61696.6351.0094.47原子11359CAPROD575.,14553.10897.8931.0094.50原子11360CBPROD573.97453.■00598.8891.0094.49原子11361CGPROD572.,74852■76798.0511.0094.45原子11362CDPROD573.,23852.02496.8531.■0094.48原子11363CPROD576.35552.31698.3891.0094.56原子113640PROD577.46552.85298.4151.0094.55原子11365NARGD676.14451.05998,7771.0094.63原子11366CAARGD677.,22850.,23299.3011.0094.72原子11367CBARGD676.,68249.046100，1081009470原子11368CGARGD677.,76748.,284100,85710094.81原子11369CDARGD677..22647.,3261018971.0094.84原子11370NEARGD678.,31846.566102.5051.0095.11原子11371CZARGD678.,33846.139103.76610095.24原子11372NH1ARGD677.32346.398104.5831.0095.17原子11373NH2ARGD67938345459104.2161.0095.16原子11374CARGD678.17749.75598.1971.0094.71原子113750ARGD679.37049.56298.4391.0094.73原子11376NCYSD777*64649.58596.9901.0094.73原子11377cacysd778..43449.,09295.8581..0094■,84原子11378CBCYSD777..52048..45094.,8091.,0094.,91原子11379SGCYSD776..19847..39395.,4931.■0095.,20原子11380CCYSD779.28650.19895.2271.0094.78原子113810CYSD778.97751，38295.3661.0094.77原子11382NGLUD880.35649.謝94.5371.0094.78原子11383CAGLUD881,,27850..76093..9171..0094.,76原子11384cbg;lud882,,74150,.33094..1271..0094..75原子11385cgg;lud883,.79751,.29993..5621.,0094..92原子11386CDGIiUD883,.87852■.64694.,3001..0095..42原子11387〇e1g;lud882.95652.99895.07210095.33原子11388OE2GLUD884.88053.36594.0961，0095.55原子11389CGLUD880.97950.98092.43410094.70原子113900GLUD881.,15450.■07491.6121.009461原子11391ncysd9so-53452.19392.1041.0094.64原子11392CACYSD9so..18552,.53690..7271..0094..,62原子11393CBCYSD978..68952,.85190..6121..0094..49说明原子11394sgcysd978.27254.60390.5111.0094.78原子11395ccysd981.03653.67690.1551.0094.60原子113960cysd981.41054.60690.8761.0094.59原子11397nserd1081.33653.58888.8591.0094.62原子11398caserd1082.10454.616881471.0094.60原子11399cbserd1083.24153.98087.3421.,0094.59原子11400〇gserd1083.,97754..96586.6371..0094.50原子11401cserd1081.21255.46587.2351.0094.67原子11402〇serd1080.15855.00986.7851.0094.64原子11403nalad1181.64956.69786.9701.0094.72原子11404caalad1180.,87857..66486.,1801.,0094■68原子11405cbalad1181.,19459.,09186.,6321.,0094.65原子11406calad1181.08857.51584.6701.0094.69原子114070alad1180.11957.46383.9091.0094.56原子11408ng:lnd1282.35457.45384.2511.0094.81原子11409cag:lnd1282..73457..30682..8411..0095..00原子11410cbg:lnd1284,.22956,.94482..7291,.0095..01原子11411cgd1284.,65956.,13281■.4881..0094.,99原子11412cdglnd1284,.65656..93180..1901..0094..90原子11413〇e1glnd1284.69858.163801981.0095.11原子11414ne2glnd1284.61656.22479.0661.0094.47原子11415cglnd1281.84256.30882.0901.0095.18原子114160glnd1281.26756.64381.0521.0095.31原子11417naspd1381.72555.09582.6321.0095.26原子11418caaspd1380..85554.,05982..0731..0095.,21原子11419cbaspd1381.,69352.,91981,,5101.,0095.,04原子11420cgaspd1382..45453,.32480.,2751..0094..95原子11421od1aspd1381.80553.64279.2561.009479原子11422od2aspd1383.7015332580.3231，0094.94原子11423caspd1379.89453.52983.1281.0095.30原子11424〇aspd1380.09153.76384.32310095.46原子11425nargd147885052.826826891.0095.25原子11426caargd1477,,93252■.16383,.6131,.0095,.13原子11427cbargd1476,.67051,.68782,.8911,.0095..01原子11428cgargd1475.76652,.78582,.3961.0094.58原子11429cdargd1474,.51052,,19881,,7921,.0094,.18原子11430neargd1473,.51553,.22881,.5041,.0093,.99原子11431czargd147343853,.90080,.3591.0093,.73原子11432nh1argd1472.49754.81880.1931,0093.04原子11433nh2argd147430153.65479.3801.0093.98原子11434cargd1478.64550.979842681.0095.■21原子114350argd1478.28749.82584,0381.,0095.■30原子11436naiad1579.6635128285.0711.,0095..18原子11437caalad1580,.45550,.26885,.7561,,0095,.11原子11438CBALAD1581.93250.63185.7131.0095.10C原子11439CALAD1579.98450.09987.1951.0095.10C原子114400AIAD1579.83151.07787.9261.0095.140原子114"NVALD1679.73348.85387.5831.0095.07N原子11442CAVALD1679.35048.52088.9511.0095.02C原子11443CBVALD1677.84448.15089.0621.0094.95C原子11444CGIVALD1677.50547.63690.45810094.94C原子11445CG2VALD1676.96149.34588.7091.0094.83C原子11446CVA!jD1680.22847-37189."11.0095.09C原子11447〇VALD16so-33746.33088.7861.0095.200原子11448NLEUD17so.87447.58490.5821.0095.12N原子11449CALEUD1781.68946.56191.2221.0095.14C原子11450CBLEUD1783.15547.00091.2821.0095.12C原子11451CGLEUD1783.,86647.23189.9451.0095.21C原子11452CD1LEUD1784.79248.43990.0351.0095.27C原子11453CD2LEUD1784.61945.97889.4731.0095.14C原子11454CLEUD1781.12446.31092.6161.0095.16C原子114550LEUD1781.68646.74793.6251.0095.26〇原子11456NCYSD1879.99145.61492.6521.0095.10N原子11457CACYSD1879.26345.35793.8871.0094.95C原子11458CBCYSD1877.76345.51493.6421.0094.90c原子11459SGCYSD1876.78245.46495.1381.0095.32s原子11460CCYSD1879.60543.97894.4701.0094.89c原子11461〇CYSD1878.78843.05294.4531.0094.780原子11462NHISD1980.82743.87594.9951.0094.87N原子11463CAHISD1981.40942.61895.4761，0094.76C原子11464CBHISD198291242.63095.2371.009479C原子11465CGHISD1983.3244193793.9841.0095.38C原子11466ND1HISD1982.92842.36192.7351.0095.86N原子11467CE1HISD198344641.56391.8191.0096.35C原子11468NE2HISD198416240.63592.4291.0096.27N原子11469CD2HISD1984.10440.84893.7841.0095.90C原子11470CHISD198116342.31096.9471.0094.61C原子114710HISD1980.95443.21797.7561.0094.530原子11472NARGD2081.23041.01797.2731.0094.39N原子11473CAARGD2081.04040.■49098.6331.0094.■16C原子11474CBARGD2082.39640.13299.2841.0094.20C原子11475CGARGD2082.32239.408100.6471，0094.34C原子11476CDARGD2081,50038.114100，6041.0094，67C原子11477NEARGD2080.95837■762101.9191.0094.90N原子11478CZARGD2079.92636.944102.1251.0094.90C原子11479NH1ARGD2079.29536.377101.0991.0094.66N原子11480NH2ARGD207951836.698103.3641.009453N原子11481CARGD2080.17441.38099.5311.0093.86C原子114820ARGD2080.67442.053100.4371.0093.850原子11483NLYSD2178.87441.39399.2431.0093.56N原子11484CALYSD2177.89942.154100..0311..0093..26C原子11485CBLYSD2177，54443.47699.3411.0093.23C原子11486CGLYSD2178.61144.55599.4951.0093.21C原子11487CDLYSD2178.49845.61998.4141.0093.06c原子11488CELYSD2179.84846.26098.1041.0092.95c原子11489NZLYSD21SO-43846.99499.2591.0092.73N原子11490CLYSD21YS.64741.333100.3561.0093.08C原子11491〇LYSD2175.63241.878100.7861.0092990原子11492NARGD2276.73240.023100.■1191.■0092.99N原子11493CAARGD2275.81239.026100,.6861.0092,.88C原子11494CBARGD2275.94139.040102,.2171.0092.94C原子11495CGARGD2275.77337.■703102,,91610093,.32c原子11496CDARGD2276.47937.755104,.2651.0094.05c原子11497NEARGD2275.93736.817105,.2461.009446N原子11498CZARGD2276.35635.565105,.4151.0094.79C原子11499NH1ARGD2277.32935.064104.6571.0094.66N原子11500NH2ARGD2275.79134.807106.3471.0094.69N原子11501CARGD2274.33639.139100.■2511.,0092.69C原子115020ARGD2273.43339.135101.■0921.,0092.,550原子11503NPHED2374.09639.22098.9411.0092.54N原子11504CAPHED2372.72639.24698.4051.0092.32C原子11505CBPHED2372.63040.10897，1411.0092.25C原子11506CGPHED2373.03841.53697，3371.0092.31C原子11507CD1PHED2372.19642.44097.9861.0092.40C原子11508CE1PHED2372.57843.76898.1631.009226C原子11509CZPHED2373.■81144.20497*6781.0092-25C原子11510CE2PHED2374.6534330897.0211.009220C原子11511CD2PHED2374.26141.98696.8521.009208C原子11512CPHED2372.22337*85098.0701.0092.14C原子115130PHED2372.87137*11797.3261.0092，270原子11514NVALD2471.06237.49598.6131.0091.90N原子11515CAVALD2470..37636..254982581.0091.■71C原子11516CBVALD2469..20135,.95299.■2351.0091.80C原子11517CGIVALD2468.64034.54599.0131.0091.67C原子11518CG2VALD2469.64036.125100，6971.0091.87C原子11519CVALD2469.84036.38096.8271.0091.61C原子115200VALD2469.7063538596.1021.0091.610原子11521NALAD2569.54537.61896.4281.0091，40N原子11522CAALAD2568,,96237,.90795.1191.0091.■20C原子11523CBALAD2567,.42637.96695..2281.,0091.,19C原子11524CALAD2569.51339.20294.5081.0091.04C原子115250ALAD257052539.■74194.9641.0090.890说明原子11526NVALD2668.82739.68593.4721.0091.00原子11527CAVALD2669.16940.92292.7651.0090.90原子11528CBVALD2668.32341.04691.4511.0091.00原子11529CGIVALD2667.8534248091.18310091.03原子11530CG2VALD26690954047890.2591.009072原子11531CVALD2669.01542.15193.6751.0090.78原子115320VALd2668.00642.27694.3771.0090.89原子11533NPROD2770.01543.06193.6631.00SO-59原子11534CAPROD2770.,06144.22394.5711.,00SO-■48原子11535CBPROD2771.,50344,71694.4231.00SO.50原子11536CGPROD2771.，744.,31793.■0351.,0090.,40原子11537CDPROD2771.,17843.■02792.■7561.0090.,38原子11538CPROD2769.07945.35794.2311.0090.35原子115390prod2768.49345.37093.1461.0090.36原子11540NGLUD2868.91146.29395.1661.0090.25原子11541CAGIaUD2868..09447..49494..9531..0090.,05原子11542CBGLUD2867,14047.72996,1331.,0090.,00原子11543cgg:lud2867.,81648.,13397,4491.,0089.,98原子11544CDGLUD2866.,85348.,77598.,4511..0090.,14原子11545OE1GLUD2865.71849.137980671.009017原子11546OE2GLUD286723548.92699.6321.0090.15原子11547CGLUD2868.97048.73094.7121.0089.95原子115480GLUD'2870.15448.74195.0641.0090.04原子11549NGLYD2968-38149.76394.1091.0089.73原子11550CAGLYD2969.,09251.,01193.8131.,0089.,46原子11551CGIiYD2970.15750.90592.7301.0089.33原子115520GLYD297123451.49992.8521.0089.25原子11553NILED3069.84950.14991.6741.0089."原子11554CAIDED3070..74549..94990.,5291■.0088.,96原子11555CBIMD3071,,04048..42290.,2871..0089..17原子11556CGIILED3071.23147.65691.6121.0089.55原子11557CD1工LiED307241248.10492.4891.0090.09原子11558CG2ILED3072.21148.201892981.008873原子11559CILED3070.12150.56189.2631.0088.72原子115600ILED3068.95350.29288.9631.0088.56原子11561NPROD3170.89351.38988.5191.0088.55原子11562CAPROD3170..39452,.01887.■2871.0088.,46原子11563CBPROD3171.,59552..83886..7921,.0088..41原子11564CGPROD3172..47153..01187,,9721.0088,.34原子11565CDPROD3172.,28551■.78588..8061,.0088,,51原子11566CPROD3169.97751.00286.2181.0088.49原子115670PROD3170.71250.04685.9431.0088■46原子11568NARGD3576.28749.09280.9731.0088.44原子11569CAARGD3577,.51648,.74680,,2551.0088.40原子11570CBARGD3578.28250.00479.8341.0088.45原子11571CGARGD3577.66350.73178.6561.0088，85原子11572CDARGD3578.60951.773780811.0089.34原子11573NEARGD3578.00852.45676.9371.0089.58原子11574CZARGD3578.17552，09475.6681.0089,54原子11575親ARGD3578.9405105175.3601.0089.66原子11576丽2ARGD3577.57752.78474.7031.0089.12原子11577CARGD3578.44147.80481.0281.0088.27原子115780argd3579.36847.23280.4491.0088.42原子11579NLEUD3678.19847.65182.3291.0088，00原子11580CALEUD3678.96046.71583.1541.008776原子11581CBLEUD3680.35647.27083.4771.0087.73原子11582CGLEUD3681.57546.34083.3691.0088.12原子11583CD1LEUD3682.85347.102837081.008834原子11584CD2LEUD3681.4604508884.2391.008842原子11585CLEUD3678.22046.40184.4461.0087.51原子115860LEUD3677.75747.29985.1431.0087.59原子11587NLEUD3778.10045.11784.7531.0087.28原子11588CALEUD3777.60044.70086.05010087.,08原子11589CBLEUD3776.14544.23185.9601.0086.98原子11590CGLEUD3775.■33944.362872591.0087.■07原子11591CD1LEUD3775.21345.81887.6901.0086.90原子11592CD2LEUD3773.96043.735871311.0087.12原子11593CLEUD3778.48743.61186.6401.0087.01原子115940LEUD3778.98142，73585.9231.0086.95原子11595NASPD387870543.69087，9481.0086.90原子11596CAASPD3879.,40742.64288.6701.0086.74原子11597CBASPD3880.89242.97288.8241.,0086.,76原子11598CGASPD3881.73341.,73989.1041..0087.,06原子11599OD1ASPD3881.25140.83689.8241.0087.29原子11600OD2ASPD3882.87841.66988.6051.0087.11原子11601CASPD3878.76842.38690.0291.0086.60原子116020ASPD3878.72843.26790.8861.0086.69原子11603NLEUD3978.25541.17390.2021.0086.47原子11604CALEUD3977.,68540.73291.,4681.,0086.,36原子11605CBLEUD3976.,20340.,38291.,3001..0086..31原子11606CGLEUD3975.,11641.,43791..5341..0086..09原子11607CD1LEUD3975.24142.64190.6081.0085.83原子11608CD2LEUD3973.75240■79291.3721.0086.08原子11609c;leud3978.46639.52591.9691.0086.50原子116100LEUD3977.88838.54692.4571.0086.45原子11611NGLYD4079.78939.60991.8401.0086.62原子11612CAGliYD4080.,68138.,53792..2541..0086..85原子11613CGLYD4080.74838.35393.7571.0087.07原子116140GLYD4080.46339.28094.5261.0087，040原子11615NLYSD4181.12537.14094.1611.0087.26N原子11616CALYSD4181.32936.76995.5681，0087.35C原子1161"7CBLYSD4182.60737.41496.1351，0087.37C原子11618CGIjYSD4183.9003701595.4271.0087，36C原子11619CDLYSD4185.12137.49096.2061.0087.55C原子11620CELYSD4186.■38837.41995.3661.0087.81C原子11621NZIiYSD4186.44738.52594.3731.0087.86N原子11622CLYSD4180.12937.07296.4661，0087.26C原子116230LYSD4180.29637.53697.5891，0087.440原子11624NASND4278.92536.80595.9711.0087.21N原子11625CAASND4277.■7093706896.74210087.26C原子11626CBASND4276.72437.91595.9281.0087.29C原子11627CGASND4277.00139.■39996.0501.0087.40c原子11628OD1ASND4276.6674001897.0581.0087.730原子11629ND2ASND4277.61339.97895.0241.0087,30N原子11630CASND4277.05135.80597.2991.0087.26C原子116310ASND4277.74734.85897.6761.0087.200原子11632NARGD4375.72135.79497*3631.0087.27N原子11633CAARGD4374.9933465897.9251.0087.40C原子11634CBARGD4374.58534,95399.3751.0087.57c原子11635CGARGD4375.61934.■584100.4271.0088.50c原子11636CDARGD4375.83133.077100.5071.0090.08c原子11637NEARGD4376.12932.645101.8701.0091.25N原子11638CZARGD4376.73931.505102.1871.0091.74C原子11639丽lARGD4377.13830.664101.2341.0091.23N原子11640丽2ARGD4376.95831.214103.4661.0091.72N原子11641CARGD4373.75834.24897.1221.0087.20C原子116420ARGD4372.85833.59697.6651.0087.320原子11643NILED4473.71234.61695.8411.0086.94N原子11644CAILED4472.,52134.,35495.0271.0086.67C原子11645CB工LED4472.■54835.,08993.6661.,0086.,66C原子11646CGIILED4472.61236.61293.9001.0086.33c原子11647CD1ILED4473.1263742392.7241.0086.76c原子11648CG2ILED4471.2813475792.8691.0086.43c原子11649CILED4472.25932.85494.8661.00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方法，其中该结构坐标是实验确定的坐标。49.权利要求48的方法，其中该结构坐标是根据表2的，a碳原子的+/-均方根偏差不超过1.5A。50.权利要求42-49的任一项的方法，其中三维模型包含原子的结构坐标，所述原子选自SEQIDNO:1所示的LINGO-1多肽的氨基酸Aspl3、Arg20、Arg22、Arg43、Glu60、Glu62、Leu94、Leul20、Metl23、Hisl76、Tyrl42、Hisl45、Hisl85、His209、His233、Ala238、Trp346、Arg347、Asn349、Asn351、Arg352、Gln353、Phe396、Arg408、Leu420、Leu426、Phe438、Asp440、Arg446、Tyr447和Ile459的原子组成的组。51.权利要求42-49的任一项的方法，还包括计算LING0-1多肽的原子和该物质的原子之间的距离。52.权利要求42-49的任一项的方法，还包括提供包含LINGO-l多肽的组合物。53.权利要求52的方法，其中该组合物包括设计的与该LINGO-l多肽结合的物质。54.权利要求42-49的任一项的方法，还包括实验确定该物质与LINGO-l多肽的相互作用。55.权利要求54的方法，还包括将该物质与LINGO-l多肽之间的相互作用与第二物质与LINGO-l多肽之间的相互作用相比较。56.—种方法，包括通过用LINGO-l多肽的三维结构进行合理药物设计来选择物质；将该物质与LINGO-l多肽相接触；和检测该物质结合该LINGO-l多肽的能力。57.的方法，其中通过计算机建模来选择该物质。58.权利要求56或57的方法，还包括合成该物质。59.权利要求幼的方法，还包括检测该物质抑制LINGO-l活性的能力。60.权利要求58的方法，还包括检测该物质体内或体外增加髓磷脂形成的能力。61.权利要求56-60的任一项的方法，还包括获得包含UNGO-l多肽和该物质的补充结晶复合物；测定补充结晶复合物的三维结构；通过用补充结晶复合物的三维结构进行合理药物设计来选择第二物质；将第二物质与LINGO-l多肽相接触；和检测第二物质结合LINGO-l多肽的能力。62.权利要求61的方法，其中，通过计算机建模来选择第二物质。63.权利要求61的方法，还包括合成笫二物质。64.权利要求61的方法，还包括检测第二物质抑制LING0-1活性的能力。65.权利要求61的方法，还包括检测第二物质体内或体外增加髓磷脂水平的能力。66.—种方法，包括使用LINGO-l多肽-配体复合物的三维结构通过进行合理药物设计来选择物质；将该物质与LINGO-l多肽相接触；和检测该物质结合LINGO-l多肽的能力。67.权利要求66的方法，其中通过计算机建模来选择该物质。68.权利要求66-67的任一项的方法，还包括合成该物质。69.权利要求68的方法，还包括检测该物质抑制LINGO-l活性的能力。70.权利要求68的方法，还包括检测该物质体内或体外增加髓磷脂形成的能力。71.权利要求66-70的任一项的方法，还包括获得包含LINGO-l多肽和该物质的补充结晶复合物；测定补充结晶复合物的三维结构；使用补充结晶复合物的三维结构通过进行合理药物设计来选择第二物质；将第二物质与LINGO-l多肽相接触；和检测第二物质结合LINGO-l多肽的能力。72.权利要求"的方法，其中通过计算机建模来选择笫二物质。73.权利要求71的方法，还包括合成第二物质。74.权利要求"的方法，还包括检测第二物质抑制LINGO-l活性的能力。75.权利要求71的方法，还包括检测第二物质体内或体外增加髓磷脂水平的能力。76.—种方法，包括提供包含LINGO-l多肽的组合物；和使该组合物结晶，形成包含LINGO-l多肽的结晶复合物，其中该结晶复合物可将X射线衍射至至少约3.7A的分辨率。77.权利要求76的方法，其中该方法包括使用悬滴蒸汽扩散法。78.—种方法，包括提供包含LINGO-l多肽-配体复合物的组合物；和使该组合物结晶，形成包含LINGO-l多肽-配体复合物的晶体，其中该晶体可将X射线衍射至至少约3.7A的分辨率。79.权利要求78的方法，其中该方法包括使用悬滴蒸汽扩散法。80.软件系统，包含对计算机系统的以下指令接受与LINGO-l多肽的结构相关的信息，接受与候选物质相关的信息，和确定候选物质与该LINGO-l多肽的结合特点，其中该确定是基于与LINGO-l多肽结构相关的信息和与候选物质相关的信息来进行。81.存在于计算机可读介质上的计算机程序，其具有储存于其上的多个指令，当被一个或多个处理器执行时该指令导致一个或多个处理器接受与包含LINGO-l多肽复合物的结构相关的信息，接受与候选物质相关的信息，和确定候选物质与LINGO-l多肽的结合特点，其中，该确定是基于与LINGO-l多肽的结构相关的信息和与候选物质相关的信息来进行。82.—种方法，包才舌接受与LINGO-l多肽的结构相关的信息；以及对该LINGO-l多肽与候选物质的结合特点加以建模，其中该方法可通过软件系统实现。83.存在于计算机可读介质上的计算机程序，其具有储存于其上的多个指令，当被一个或多个处理器执行时该指令导致一个或多个处理器接受与LINGO-l多肽的结构相关的信息，和对LINGO-l多肽与候选物质的结合特点加以建模。84.软件系统，包括对计算机系统的下述指令接受与UNGO-l多肽的结构相关的信息，和对LINGO-l多肽与候选物质的结合特点加以建模。85.调节受试者中UNGO-l活性的方法，包括使用合理的药物设计来选择能调节LINGO-l活性的物质，和将治疗有效量的该物质施用于该受试者。86.权利要求85的方法，其中该物质能体内增加髓磷脂水平。87.权利要求85或86的方法，其中合理的药物设计包括使用包含LINGO-l多肽的结晶复合物的三维结构。88.权利要求87的方法，其中结晶复合物还包含配体。89.治疗具有与LINGO-l活性相关的病况的受试者的方法，包括使用合理的药物设计来选择能影响LINGO-l活性的物质，和将治疗有效量的该物质施用于需要其的受试者。90.权利要求89的方法，其中该物质能抑制LINGO-l活性。91.权利要求89或90的方法，其中该物质能体内增加髓磷脂水平。92.权利要求89-91的任一项的方法，其中该病况是脱髓鞘疾病。93.权利要求89-91的任一项的方法，其中该病况是多发性硬化。94.预防性治疗对与LINGO-l活性相关的病况易感的受试者的方法，包括确定该受试者是否对与LINGO-l活性相关的病况易感，使用合理的药物设计来选择能影响LINGO-l活性的物质，和将治疗有效量的该物质施用于该受试者。95.权利要求94或95的方法，其中该物质能抑制LINGO-l活性。96.权利要求94-95的任一项的方法，其中该物质能体内增加髓磷脂水平。97.权利要求94-96的任一项的方法，其中该病况是脱髓鞘疾病。98.权利要求94-97的任一项的方法，其中该病况是多发性硬化。99.根据权利要求22-75的任一项设计或选择的物质在制造用于预防或治疗与LINGO-l活性相关病况的药物中的用途。100.权利要求99的用途，其中该物质能抑制LINGO-l活性。101.斥又利要求99或100的用途，其中该病况是脱髓鞘疾病。102.权利要求99-101的任一项的用途，其中该病况是多发性硬化。103.权利要求99-102的任一项的用途，其中该物质在配体结合位点与UNGO画l结合。104.权利要求103的用途，其中该配体结合位点位于LRR结构域的凹面。105.权利要求104的用途，其中该LRR结构域的凹面包含Trp346和Arg352;Hisl85、His209和His233;Aspl3;Glu60和Glu62;或Arg20、Arg22和Arg43中的一个或多个。106.权利要求103的用途，其中该配体结合位点位于LRR结构域的凸面。107.权利要求106的用途，其中该LRR结构域的凸面包含Tyrl42和Hisl45;Leu94、Leul20和Metl23;Hisl76;或Ala238中的一个或多个。108.权利要求103的用途，其中配体结合位点位于Ig结构域上。109.权利要求108的用途，其中Ig结构域包含Arg446和Try447;Arg408、Phe438和Asp440;Phe396;或Leu420、Leu426和Ile459中的一个或多个。110.根据权利要求22-75的任一项设计或选择的物质，其用于预防或治疗与LINGO-l活性相关的病况。111.权利要求110的物质，其中该物质能抑制LINGO-l活性。112.权利要求110或111的物质，其中该病况是脱髓鞘疾病。113.权利要求110-112的任一项的物质，其中该病况是多发性硬化。114.权利要求110-113的任一项的物质，其中该物质在配体结合位点与UNGO-l结合。115.权利要求114的用途，其中该配体结合位点位于LRR结构域的凹面。116.权利要求115的用途，其中LRR结构域的该凹面包含Trp346和Arg352;Hisl85、His209和His233;Aspl3;Glu60和Glu62;或Arg20、Arg22和Arg43中的一个或多个。117.权利要求114的用途，其中该配体结合位点位于LRR结构域的凸面。118.权利要求117的用途，其中LRR结构域的凸面包含Tyrl42和Hisl45;Leu94、Leul20和Metl23;Hisl76;或Ala238中的一个或多个。119.权利要求114的用途，其中该配体结合位点位于Ig结构域上。120.权利要求119的用途，其中该Ig结构域包含Arg446和Try447;Arg408、Phe438和Asp440;Phe396;或Leu420、Leu426和Ik459中的一个或多个。全文摘要本公开涉及LINGO-1多肽、LINGO-1多肽/配体复合物、LINGO-1多肽的晶体、LINGO-1多肽/配体复合物的晶体以及相关的方法和软件系统。文档编号C07K14/415GK101405297SQ200780009635公开日2009年4月8日申请日期2007年2月1日优先权日2006年2月3日发明者B·德怀尔,E·普莱斯曼,J·M·威廉,L·莫夏科,M·施塔尔,M·约翰逊,W·S·萨默斯,钟晓天申请人:惠氏公司