一种水稻新基因Os03g02470.3的CDS序列及其应用

技术领域

本发明属于基因工程技术领域，尤其涉及一种水稻新基因Os03g02470.3的CDS序列及其应用。

背景技术

水稻(Oryza sativa L.)是重要的粮食作物之一，世界上一半以上的人口以稻米为主食。水稻作物产量主要由三个因素决定，分别是单株穗数、每穗粒数以及籽粒重量，而粒重与粒型正相关，因此，粒型是一个十分重要的产量性状。水稻粒型性状主要包括粒长、粒宽、粒厚以及长宽比。水稻籽粒的最终大小在很大程度上取决于小穗壳(稻壳)细胞的增殖和扩张。油菜素甾醇(Brassinosteroids，BR)是一类植物特有的甾醇类激素，调节细胞伸长、分裂和分化，几乎对所有植物生长发育过程都起重要的调控作用。BR合成或信号传导相关基因突变往往导致粒型变化，如BR合成基因D11缺失，突变体籽粒明显变小，穗特异启动子驱动D11表达的回补转基因水稻籽粒大于野生型，产量提高，而且其他农艺性状没有改变。BR信号传导的正向调控因子如受体激酶OsBRI1、OsBAK1、蛋白激酶OsBSK3和LRR类激酶XIAO等功能缺失或抑制突变体都出现小粒表型，而BR信号传导抑制因子OsGSK2过表达水稻籽粒小而圆，其RNA干扰株系则出现粒长增加的表型。

BR被位于细胞膜的受体蛋白激酶识别后，启动一系列蛋白质磷酸化/去磷酸化反应，将信号传导至位于细胞核的转录因子，调节上千个基因的转录。在水稻中，已发现GRAS转录因子家族成员DLT、WRKY转录因子家族成员WRKY53都能够通过调节BR信号传导来调节水稻粒型发育。卵形家族蛋白(Ovate Family Protein，OFP)是植物特有的一类转录因子，最早在番茄中被发现调节其果实形状的变化，水稻基因组编码33个OFP蛋白，其中多个成员在BR信号传导中发挥重要作用。OsOFP3、OsOFP19和OsOFP22抑制BR信号传导，其过表达转基因水稻均出现株型收敛、半矮化，籽粒短而宽、厚度增加的表型，三者都通过调节小穗壳的生长而影响籽粒大小。OsOFP1和OsOFP8促进BR信号传导，其过表达转基因水稻籽粒长度增加，长宽比增加。

转录因子通过调控靶基因的时空转录进而调节细胞生命过程，因此，鉴定转录因子的靶基因是解析功能机制，利用其开展分子设计育种的一个重要基础。通过对OsOFP22过表达转基因水稻(OsOFP22-OE)和野生型(WT)的幼穗进行了转录组测序(RNA-Seq)，从全基因组水平分析OsOFP22调节的生物学过程。其中Os03g02470.3的转录水平在OsOFP22-OE和WT之间差异极显著，通过实时荧光定量PCR、蛋白质-DNA互作和转基因等方法，证实Os03g02470.3是水稻OFP转录因子家族的一个靶基因，OsOFP3、OsOFP19和OsOFP22均直接绑定Os03g02470.3基因的启动子，并抑制其活性。Os03g02470.3过表达转基因水稻籽粒长度和宽度均明显增加，长宽比增大，千粒重增大。但是目前，关于Os03g02470.3基因尚无明确的功能报道，在各种数据库中仅有“表达蛋白”的注释。

发明内容

本发明实施例的目的在于提供一种水稻新基因Os03g02470.3的CDS序列及其应用，旨在解决上述背景技术中提出的问题。

本发明实施例是这样实现的，一种水稻新基因Os03g02470.3的CDS序列，所述水稻新基因Os03g02470.3的核苷酸序列如SEQ ID No.1所示。

本发明实施例的另一目的在于，提供一种水稻新基因Os03g02470.3的CDS序列的应用，将所述的水稻新基因Os03g02470.3构建到由Ubiquitin驱动的植物双元表达载体pCambia1390-GFP上，获得组成型过表达载体Ubiquitin::Os03g02470.3-GFP，所述GFP的核苷酸序列如SEQ ID No.2所示，所述Ubiquitin::Os03g02470.3-GFP的全序列如SEQ IDNo.4所示。

进一步的技术方案，构建所述组成型过表达载体Ubiquitin::Os03g02470.3-GFP的具体步骤如下：

步骤1：根据Os03g02470.3基因的序列设计PCR扩增引物对，扩增引物为：正向引物F：5'-CACTAGGTACCTGCAGATGAAGAGTCCAGTATTCCCT-3'和反向引物R：5'-ATCCGTCGACCTGCAGCGAACTTAAGATGTATCCCAT-3'；

步骤2：以野生日本晴水稻总cDNA为模板，进行PCR获得Os03g02470.3全CDS序列；

步骤3：将步骤2中的PCR产物构建到其由Ubiquitin驱动的植物双元表达载体pCambia1390-GFP上，载体全序列如SEQ ID No.3所示，获得组成型过表达载体Ubiquitin::Os03g02470.3-GFP。

本发明实施例的另一目的在于，提供一种转基因水稻植株的构建方法，具体步骤为采用农杆菌介导的方法，将Ubiquitin::Os03g02470.3-GFP转入水稻愈伤组织中，用含诱导剂和农杆菌的AAM转化液进行转化，转化后的材料经过共培养、筛选、分化和生根，转基因苗的锻炼和移栽，筛选转基因水稻植株。

本发明实施例的另一目的在于，提供所述水稻Os03g02470.3基因在调控水稻高产性状中的应用。

进一步的技术方案，所述水稻Os03g02470.3基因在调控水稻高产性状中的应用的具体步骤为将水稻Os03g02470.3基因的CDS序列去掉终止密码子之后，构建到Ubiquitin启动子的下游，转化水稻。

本发明实施例提供的一种水稻新基因Os03g02470.3的CDS序列，首次将Ubiquitin启动子与Os03g02470.3基因的CDS序列与GFP进行融合，并将该融合基因转化到水稻中，从而改良水稻粒型性状，对于详细阐明水稻生长发育机理具有重要的理论价值，并且可以通过转基因手段，改良水稻产量性状，因此在生产实践中具有重要意义。

附图说明

图1为本发明实施例1中Ubiquitin::Os03g02470.3-GFP载体图谱。

图2为本发明利用DNA-pull down实验鉴定OsOFP3、OsOFP19和OsOFP22分别直接绑定Os03g02470.3基因的启动子。

图3为本发明利用Dual-luciferase实验鉴定OsOFP3、OsOFP19和OsOFP22分别抑制Os03g02470.3基因启动子的活性。

图4为本发明PCR检测Ubiquitin::Os03g02470.3-GFP转基因阳性株系，其中OE-1、OE-2和OE-3为转基因水稻株系。

图5为本发明转基因材料粒型表型比较，其中WT为野生型水稻‘Kitaake’，OE-1、OE-2和OE-3为Os03g02470.3过表达转基因水稻株系。

图6为本发明转基因材料粒型性状数据统计分析，其中WT为野生型水稻‘Kitaake’，OE-1、OE-2和OE-3为转基因水稻株系。

具体实施方式

为了使本发明的目的、技术方案及优点更加清楚明白，以下结合附图及实施例，对本发明进行进一步详细说明。应当理解，此处所描述的具体实施例仅仅用以解释本发明，并不用于限定本发明。

以下结合具体实施例对本发明的具体实现进行详细描述。

实施例1，Os03g02470.3基因CDS序列的获得及植物表达载体的构建

1.Os03g02470.3基因CDS序列的获得

在国家水稻数据中心中找到Os03g02470.3基因，根据其CDS序列设计PCR扩增引物，

正向引物F5'-CACTAGGTACCTGCAGATGAAGAGTCCAGTATTCCCT-3'，反向引物R5'-ATCCGTCGACCTGCAGCGAACTTAAGATGTATCCCAT-3'。以野生型“日本晴”水稻总cDNA为模板，利用引物F和R进行PCR扩增，获得Os03g02470.3基因完整的CDS序列(Seq ID No.1)。

2.植物表达载体的构建

将上述PCR产物构建到其由Ubiquitin驱动的植物双元表达载体pCambia1390-GFP上，获得组成型过表达载体Ubiquitin::Os03g02470.3-GFP，其全序列如SEQ ID No.4所示，载体图谱见图1。反应体系转化大肠杆菌DH5α，筛选阳性克隆。

实施例2，转基因水稻植株的获得

取水稻“Kitaake”成熟种子，人工或机械脱壳，挑选饱满光洁无菌斑的种子经消毒之后接种到诱导培养基上进行诱导培养。选择外观良好，生长力好的水稻愈伤组织为受体材料，采用农杆菌介导法将Ubiquitin::Os03g02470.3-GFP转入水稻愈伤组织中，用含有100μM的乙酰丁香酮和O.D.值为0.7的农杆菌的AAM转化液进行转化，将转化液浸泡过的愈伤组织置于共培养基上进行共培养，25℃暗培养3d后，置于筛选培养基上培养约30d，每10d继代一次。然后将筛出的抗性愈伤转移到分化培养基上分化约20d，每10d继代一次。将分化出绿色小苗的抗性愈伤转移到生根培养基上生根，待约7d长出发达根系后炼苗，并计算转化所获转基因苗数。炼苗7d后转移至大田生长。

其中，诱导培养基配方为：N6大量+B5微量+NB有机+铁盐+铜钴母液+2.5mg/L 2,4D+0.6g/L酸水解酪蛋白+2.878g/L脯氨酸+0.5g/L谷氨酰胺+30g/L蔗糖，以水配制，调pH至5.8～5.9后加入植物凝胶4g/L。

共培养基配方为：N6大量+B5微量+NB有机+铁盐+2.5mg/L 2,4D+0.5g/L谷氨酰胺+0.6g/L酸水解酪蛋白+10g/L葡萄糖+30g/L蔗糖，以水配制，调pH至5.2后加入植物凝胶4g/L。灭菌后，50℃左右加入AS(乙酰丁香酮)100～200μg/mL。

筛选培养基配方为：N6大量+B5微量+NB有机+铁盐+铜钴母液+2.5mg/L2,4D+0.6g/L酸水解酪蛋白+2.878g/L脯氨酸+0.5g/L谷氨酰胺+30g/L蔗糖，以水配制，调pH至5.8～5.9后加入植物凝胶4g/L。灭菌后加入35mg/L潮霉素。

分化培养基配方为：MS无机+MS-B5微量+MS有机+铁盐+MS-铜钴母液+0.05mg/LNAA+2.0mg/L Kinetin(激动素)+30g/L山梨醇+2g/L水解酪蛋白+30g/L蔗糖，以水配制，调pH至5.8～5.9后加入植物凝胶4g/L。

实施例3，转基因阳性株系的鉴定

为了检测Os03g02470.3基因是否整合至水稻基因组中，提取转基因水稻叶片DNA，在基因两端的植物表达载体上设计一段引物进行PCR反应，鉴定转基因植株，对PCR产物进行检测，结果发现，转基因水稻均扩增出目的条带(图4)，PCR产物经测序分析为正确的目的基因序列，由此确定Os03g02470.3基因已通过农杆菌介导的方法插入到水稻基因组中。

实施例4，转基因水稻表型和统计分析

将转基因和野生型水稻进行比较，从表型上可以明显的看出转基因植株带壳和脱壳籽粒均比野生型的粒长和粒宽大(图5)，通过统计数据分析结果可以明显的看出转基因水稻的长宽比和千粒重明显高于野生型(图6)，其中野生型的千粒重为25.76g，转基因植株的千粒重为28.22g。

以上所述仅为本发明的较佳实施例而已，并不用以限制本发明，凡在本发明的精神和原则之内所作的任何修改、等同替换和改进等，均应包含在本发明的保护范围之内。

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Thr Gly Gly Thr Gly Thr Gly Gly Thr Thr Gly Gly Gly Cys Gly Gly

10290 10295 10300

Thr Cys Gly Thr Thr Cys Ala Thr Thr Cys Gly Thr Thr Cys Thr Ala

10305 10310 10315 10320

Gly Ala Thr Cys Gly Gly Ala Gly Thr Ala Gly Ala Ala Thr Ala Cys

10325 10330 10335

Thr Gly Thr Thr Thr Cys Ala Ala Ala Cys Thr Ala Cys Cys Thr Gly

10340 10345 10350

Gly Thr Gly Thr Ala Thr Thr Thr Ala Thr Thr Ala Ala Thr Thr Thr

10355 10360 10365

Thr Gly Gly Ala Ala Cys Thr Gly Thr Ala Thr Gly Thr Gly Thr Gly

10370 10375 10380

Thr Gly Thr Cys Ala Thr Ala Cys Ala Thr Cys Thr Thr Cys Ala Thr

10385 10390 10395 10400

Ala Gly Thr Thr Ala Cys Gly Ala Gly Thr Thr Thr Ala Ala Gly Ala

10405 10410 10415

Thr Gly Gly Ala Thr Gly Gly Ala Ala Ala Thr Ala Thr Cys Gly Ala

10420 10425 10430

Thr Cys Thr Ala Gly Gly Ala Thr Ala Gly Gly Thr Ala Thr Ala Cys

10435 10440 10445

Ala Thr Gly Thr Thr Gly Ala Thr Gly Thr Gly Gly Gly Thr Thr Thr

10450 10455 10460

Thr Ala Cys Thr Gly Ala Thr Gly Cys Ala Thr Ala Thr Ala Cys Ala

10465 10470 10475 10480

Thr Gly Ala Thr Gly Gly Cys Ala Thr Ala Thr Gly Cys Ala Gly Cys

10485 10490 10495

Ala Thr Cys Thr Ala Thr Thr Cys Ala Thr Ala Thr Gly Cys Thr Cys

10500 10505 10510

Thr Ala Ala Cys Cys Thr Thr Gly Ala Gly Thr Ala Cys Cys Thr Ala

10515 10520 10525

Thr Cys Thr Ala Thr Thr Ala Thr Ala Ala Thr Ala Ala Ala Cys Ala

10530 10535 10540

Ala Gly Thr Ala Thr Gly Thr Thr Thr Thr Ala Thr Ala Ala Thr Thr

10545 10550 10555 10560

Ala Thr Thr Thr Thr Gly Ala Thr Cys Thr Thr Gly Ala Thr Ala Thr

10565 10570 10575

Ala Cys Thr Thr Gly Gly Ala Thr Gly Ala Thr Gly Gly Cys Ala Thr

10580 10585 10590

Ala Thr Gly Cys Ala Gly Cys Ala Gly Cys Thr Ala Thr Ala Thr Gly

10595 10600 10605

Thr Gly Gly Ala Thr Thr Thr Thr Thr Thr Thr Ala Gly Cys Cys Cys

10610 10615 10620

Thr Gly Cys Cys Thr Thr Cys Ala Thr Ala Cys Gly Cys Thr Ala Thr

10625 10630 10635 10640

Thr Thr Ala Thr Thr Thr Gly Cys Thr Thr Gly Gly Thr Ala Cys Thr

10645 10650 10655

Gly Thr Thr Thr Cys Thr Thr Thr Thr Gly Thr Cys Gly Ala Thr Gly

10660 10665 10670

Cys Thr Cys Ala Cys Cys Cys Thr Gly Thr Thr Gly Thr Thr Thr Gly

10675 10680 10685

Gly Thr Gly Thr Thr Ala Cys Thr Thr Cys Thr Gly Cys Ala Cys Thr

10690 10695 10700

Ala Gly Gly Thr Ala Cys Cys Thr Gly Cys Ala Gly Gly Thr Cys Gly

10705 10710 10715 10720

Ala Cys Gly Gly Ala Thr Cys Cys Ala Thr Gly Gly Thr Gly Ala Gly

10725 10730 10735

Cys Ala Ala Gly Gly Gly Cys Gly Ala Gly Gly Ala Gly Cys Thr Gly

10740 10745 10750

Thr Thr Cys Ala Cys Cys Gly Gly Gly Gly Thr Gly Gly Thr Gly Cys

10755 10760 10765

Cys Cys Ala Thr Cys Cys Thr Gly Gly Thr Cys Gly Ala Gly Cys Thr

10770 10775 10780

Gly Gly Ala Cys Gly Gly Cys Gly Ala Cys Gly Thr Ala Ala Ala Cys

10785 10790 10795 10800

Gly Gly Cys Cys Ala Cys Ala Ala Gly Thr Thr Cys Ala Gly Cys Gly

10805 10810 10815

Thr Gly Thr Cys Cys Gly Gly Cys Gly Ala Gly Gly Gly Cys Gly Ala

10820 10825 10830

Gly Gly Gly Cys Gly Ala Thr Gly Cys Cys Ala Cys Cys Thr Ala Cys

10835 10840 10845

Gly Gly Cys Ala Ala Gly Cys Thr Gly Ala Cys Cys Cys Thr Gly Ala

10850 10855 10860

Ala Gly Thr Thr Cys Ala Thr Cys Thr Gly Cys Ala Cys Cys Ala Cys

10865 10870 10875 10880

Cys Gly Gly Cys Ala Ala Gly Cys Thr Gly Cys Cys Cys Gly Thr Gly

10885 10890 10895

Cys Cys Cys Thr Gly Gly Cys Cys Cys Ala Cys Cys Cys Thr Cys Gly

10900 10905 10910

Thr Gly Ala Cys Cys Ala Cys Cys Thr Thr Cys Ala Cys Cys Thr Ala

10915 10920 10925

Cys Gly Gly Cys Gly Thr Gly Cys Ala Gly Thr Gly Cys Thr Thr Cys

10930 10935 10940

Ala Gly Cys Cys Gly Cys Thr Ala Cys Cys Cys Cys Gly Ala Cys Cys

10945 10950 10955 10960

Ala Cys Ala Thr Gly Ala Ala Gly Cys Ala Gly Cys Ala Cys Gly Ala

10965 10970 10975

Cys Thr Thr Cys Thr Thr Cys Ala Ala Gly Thr Cys Cys Gly Cys Cys

10980 10985 10990

Ala Thr Gly Cys Cys Cys Gly Ala Ala Gly Gly Cys Thr Ala Cys Gly

10995 11000 11005

Thr Cys Cys Ala Gly Gly Ala Gly Cys Gly Cys Ala Cys Cys Ala Thr

11010 11015 11020

Cys Thr Thr Cys Thr Thr Cys Ala Ala Gly Gly Ala Cys Gly Ala Cys

11025 11030 11035 11040

Gly Gly Cys Ala Ala Cys Thr Ala Cys Ala Ala Gly Ala Cys Cys Cys

11045 11050 11055

Gly Cys Gly Cys Cys Gly Ala Gly Gly Thr Gly Ala Ala Gly Thr Thr

11060 11065 11070

Cys Gly Ala Gly Gly Gly Cys Gly Ala Cys Ala Cys Cys Cys Thr Gly

11075 11080 11085

Gly Thr Gly Ala Ala Cys Cys Gly Cys Ala Thr Cys Gly Ala Gly Cys

11090 11095 11100

Thr Gly Ala Ala Gly Gly Gly Cys Ala Thr Cys Gly Ala Cys Thr Thr

11105 11110 11115 11120

Cys Ala Ala Gly Gly Ala Gly Gly Ala Cys Gly Gly Cys Ala Ala Cys

11125 11130 11135

Ala Thr Cys Cys Thr Gly Gly Gly Gly Cys Ala Cys Ala Ala Gly Cys

11140 11145 11150

Thr Gly Gly Ala Gly Thr Ala Cys Ala Ala Cys Thr Ala Cys Ala Ala

11155 11160 11165

Cys Ala Gly Cys Cys Ala Cys Ala Ala Cys Gly Thr Cys Thr Ala Thr

11170 11175 11180

Ala Thr Cys Ala Thr Gly Gly Cys Cys Gly Ala Cys Ala Ala Gly Cys

11185 11190 11195 11200

Ala Gly Ala Ala Gly Ala Ala Cys Gly Gly Cys Ala Thr Cys Ala Ala

11205 11210 11215

Gly Gly Thr Gly Ala Ala Cys Thr Thr Cys Ala Ala Gly Ala Thr Cys

11220 11225 11230

Cys Gly Cys Cys Ala Cys Ala Ala Cys Ala Thr Cys Gly Ala Gly Gly

11235 11240 11245

Ala Cys Gly Gly Gly Ala Gly Cys Gly Thr Gly Cys Ala Gly Cys Thr

11250 11255 11260

Cys Gly Cys Cys Gly Ala Cys Cys Ala Cys Thr Ala Cys Cys Ala Gly

11265 11270 11275 11280

Cys Ala Gly Ala Ala Cys Ala Cys Cys Cys Cys Cys Ala Thr Cys Gly

11285 11290 11295

Gly Cys Gly Ala Cys Gly Gly Cys Cys Cys Cys Gly Thr Gly Cys Thr

11300 11305 11310

Gly Cys Thr Gly Cys Cys Cys Gly Ala Cys Ala Ala Cys Cys Ala Cys

11315 11320 11325

Thr Ala Cys Cys Thr Gly Ala Gly Cys Ala Cys Cys Cys Ala Gly Thr

11330 11335 11340

Cys Cys Gly Cys Cys Cys Thr Gly Ala Gly Cys Ala Ala Ala Gly Ala

11345 11350 11355 11360

Cys Cys Cys Cys Ala Ala Cys Gly Ala Gly Ala Ala Gly Cys Gly Cys

11365 11370 11375

Gly Ala Thr Cys Ala Cys Ala Thr Gly Gly Thr Cys Cys Thr Gly Cys

11380 11385 11390

Thr Gly Gly Ala Gly Thr Thr Cys Gly Thr Gly Ala Cys Cys Gly Cys

11395 11400 11405

Cys Gly Cys Cys Gly Gly Gly Ala Thr Cys Ala Cys Thr Cys Ala Cys

11410 11415 11420

Gly Gly Cys Ala Thr Gly Gly Ala Cys Gly Ala Gly Cys Thr Gly Thr

11425 11430 11435 11440

Ala Cys Ala Ala Gly Thr Ala Ala Cys Cys Gly Gly Gly Ala Ala Thr

11445 11450 11455

Thr Cys Thr Ala Ala Gly Ala Gly Gly Ala Gly Thr Cys Cys Ala Cys

11460 11465 11470

Cys Ala Thr Gly Gly Thr Ala Gly Ala Thr Cys Thr Gly Ala Cys Thr

11475 11480 11485

Ala Gly Thr Gly Thr Thr Ala Ala Cys Gly Cys Thr Ala Gly Cys Cys

11490 11495 11500

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11505 11510 11515 11520

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11525 11530 11535

Gly Ala Cys Cys Ala Gly Cys Thr Cys Gly Ala Ala Thr Thr Thr Cys

11540 11545 11550

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11555 11560 11565

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11570 11575 11580

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Gly Thr Thr Thr Cys Thr Thr Ala Ala Gly Ala Thr Thr Gly Ala Ala

1 5 10 15

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20 25 30

Gly Cys Gly Ala Thr Gly Ala Thr Thr Ala Thr Cys Ala Thr Ala Thr

35 40 45

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50 55 60

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65 70 75 80

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85 90 95

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100 105 110

Ala Thr Gly Gly Gly Thr Thr Thr Thr Thr Ala Thr Gly Ala Thr Thr

115 120 125

Ala Gly Ala Gly Thr Cys Cys Cys Gly Cys Ala Ala Thr Thr Ala Thr

130 135 140

Ala Cys Ala Thr Thr Thr Ala Ala Thr Ala Cys Gly Cys Gly Ala Thr

145 150 155 160

Ala Gly Ala Ala Ala Ala Cys Ala Ala Ala Ala Thr Ala Thr Ala Gly

165 170 175

Cys Gly Cys Gly Cys Ala Ala Ala Cys Thr Ala Gly Gly Ala Thr Ala

180 185 190

Ala Ala Thr Thr Ala Thr Cys Gly Cys Gly Cys Gly Cys Gly Gly Thr

195 200 205

Gly Thr Cys Ala Thr Cys Thr Ala Thr Gly Thr Thr Ala Cys Thr Ala

210 215 220

Gly Ala Thr Cys Gly Gly Gly Ala Ala Thr Thr Ala Ala Ala Cys Thr

225 230 235 240

Ala Thr Cys Ala Gly Thr Gly Thr Thr Thr Gly Ala Cys Ala Gly Gly

245 250 255

Ala Thr Ala Thr Ala Thr Thr Gly Gly Cys Gly Gly Gly Thr Ala Ala

260 265 270

Ala Cys Cys Thr Ala Ala Gly Ala Gly Ala Ala Ala Ala Gly Ala Gly

275 280 285

Cys Gly Thr Thr Thr Ala Thr Thr Ala Gly Ala Ala Thr Ala Ala Cys

290 295 300

Gly Gly Ala Thr Ala Thr Thr Thr Ala Ala Ala Ala Gly Gly Gly Cys

305 310 315 320

Gly Thr Gly Ala Ala Ala Ala Gly Gly Thr Thr Thr Ala Thr Cys Cys

325 330 335

Gly Thr Thr Cys Gly Thr Cys Cys Ala Thr Thr Thr Gly Thr Ala Thr

340 345 350

Gly Thr Gly Cys Ala Thr Gly Cys Cys Ala Ala Cys Cys Ala Cys Ala

355 360 365

Gly Gly Gly Thr Thr Cys Cys Cys Cys Thr Cys Gly Gly Gly Ala Thr

370 375 380

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385 390 395 400

Ala Ala Cys Cys Cys Cys Thr Cys Cys Gly Cys Thr Gly Cys Thr Ala

405 410 415

Thr Ala Gly Thr Gly Cys Ala Gly Thr Cys Gly Gly Cys Thr Thr Cys

420 425 430

Thr Gly Ala Cys Gly Thr Thr Cys Ala Gly Thr Gly Cys Ala Gly Cys

435 440 445

Cys Gly Thr Cys Thr Thr Cys Thr Gly Ala Ala Ala Ala Cys Gly Ala

450 455 460

Cys Ala Thr Gly Thr Cys Gly Cys Ala Cys Ala Ala Gly Thr Cys Cys

465 470 475 480

Thr Ala Ala Gly Thr Thr Ala Cys Gly Cys Gly Ala Cys Ala Gly Gly

485 490 495

Cys Thr Gly Cys Cys Gly Cys Cys Cys Thr Gly Cys Cys Cys Thr Thr

500 505 510

Thr Thr Cys Cys Thr Gly Gly Cys Gly Thr Thr Thr Thr Cys Thr Thr

515 520 525

Gly Thr Cys Gly Cys Gly Thr Gly Thr Thr Thr Thr Ala Gly Thr Cys

530 535 540

Gly Cys Ala Thr Ala Ala Ala Gly Thr Ala Gly Ala Ala Thr Ala Cys

545 550 555 560

Thr Thr Gly Cys Gly Ala Cys Thr Ala Gly Ala Ala Cys Cys Gly Gly

565 570 575

Ala Gly Ala Cys Ala Thr Thr Ala Cys Gly Cys Cys Ala Thr Gly Ala

580 585 590

Ala Cys Ala Ala Gly Ala Gly Cys Gly Cys Cys Gly Cys Cys Gly Cys

595 600 605

Thr Gly Gly Cys Cys Thr Gly Cys Thr Gly Gly Gly Cys Thr Ala Thr

610 615 620

Gly Cys Cys Cys Gly Cys Gly Thr Cys Ala Gly Cys Ala Cys Cys Gly

625 630 635 640

Ala Cys Gly Ala Cys Cys Ala Gly Gly Ala Cys Thr Thr Gly Ala Cys

645 650 655

Cys Ala Ala Cys Cys Ala Ala Cys Gly Gly Gly Cys Cys Gly Ala Ala

660 665 670

Cys Thr Gly Cys Ala Cys Gly Cys Gly Gly Cys Cys Gly Gly Cys Thr

675 680 685

Gly Cys Ala Cys Cys Ala Ala Gly Cys Thr Gly Thr Thr Thr Thr Cys

690 695 700

Cys Gly Ala Gly Ala Ala Gly Ala Thr Cys Ala Cys Cys Gly Gly Cys

705 710 715 720

Ala Cys Cys Ala Gly Gly Cys Gly Cys Gly Ala Cys Cys Gly Cys Cys

725 730 735

Cys Gly Gly Ala Gly Cys Thr Gly Gly Cys Cys Ala Gly Gly Ala Thr

740 745 750

Gly Cys Thr Thr Gly Ala Cys Cys Ala Cys Cys Thr Ala Cys Gly Cys

755 760 765

Cys Cys Thr Gly Gly Cys Gly Ala Cys Gly Thr Thr Gly Thr Gly Ala

770 775 780

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785 790 795 800

Cys Cys Gly Cys Cys Thr Gly Gly Cys Cys Cys Gly Cys Ala Gly Cys

805 810 815

Ala Cys Cys Cys Gly Cys Gly Ala Cys Cys Thr Ala Cys Thr Gly Gly

820 825 830

Ala Cys Ala Thr Thr Gly Cys Cys Gly Ala Gly Cys Gly Cys Ala Thr

835 840 845

Cys Cys Ala Gly Gly Ala Gly Gly Cys Cys Gly Gly Cys Gly Cys Gly

850 855 860

Gly Gly Cys Cys Thr Gly Cys Gly Thr Ala Gly Cys Cys Thr Gly Gly

865 870 875 880

Cys Ala Gly Ala Gly Cys Cys Gly Thr Gly Gly Gly Cys Cys Gly Ala

885 890 895

Cys Ala Cys Cys Ala Cys Cys Ala Cys Gly Cys Cys Gly Gly Cys Cys

900 905 910

Gly Gly Cys Cys Gly Cys Ala Thr Gly Gly Thr Gly Thr Thr Gly Ala

915 920 925

Cys Cys Gly Thr Gly Thr Thr Cys Gly Cys Cys Gly Gly Cys Ala Thr

930 935 940

Thr Gly Cys Cys Gly Ala Gly Thr Thr Cys Gly Ala Gly Cys Gly Thr

945 950 955 960

Thr Cys Cys Cys Thr Ala Ala Thr Cys Ala Thr Cys Gly Ala Cys Cys

965 970 975

Gly Cys Ala Cys Cys Cys Gly Gly Ala Gly Cys Gly Gly Gly Cys Gly

980 985 990

Cys Gly Ala Gly Gly Cys Cys Gly Cys Cys Ala Ala Gly Gly Cys Cys

995 1000 1005

Cys Gly Ala Gly Gly Cys Gly Thr Gly Ala Ala Gly Thr Thr Thr Gly

1010 1015 1020

Gly Cys Cys Cys Cys Cys Gly Cys Cys Cys Thr Ala Cys Cys Cys Thr

1025 1030 1035 1040

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1045 1050 1055

Gly Cys Gly Cys Ala Cys Gly Cys Cys Cys Gly Cys Gly Ala Gly Cys

1060 1065 1070

Thr Gly Ala Thr Cys Gly Ala Cys Cys Ala Gly Gly Ala Ala Gly Gly

1075 1080 1085

Cys Cys Gly Cys Ala Cys Cys Gly Thr Gly Ala Ala Ala Gly Ala Gly

1090 1095 1100

Gly Cys Gly Gly Cys Thr Gly Cys Ala Cys Thr Gly Cys Thr Thr Gly

1105 1110 1115 1120

Gly Cys Gly Thr Gly Cys Ala Thr Cys Gly Cys Thr Cys Gly Ala Cys

1125 1130 1135

Cys Cys Thr Gly Thr Ala Cys Cys Gly Cys Gly Cys Ala Cys Thr Thr

1140 1145 1150

Gly Ala Gly Cys Gly Cys Ala Gly Cys Gly Ala Gly Gly Ala Ala Gly

1155 1160 1165

Thr Gly Ala Cys Gly Cys Cys Cys Ala Cys Cys Gly Ala Gly Gly Cys

1170 1175 1180

Cys Ala Gly Gly Cys Gly Gly Cys Gly Cys Gly Gly Thr Gly Cys Cys

1185 1190 1195 1200

Thr Thr Cys Cys Gly Thr Gly Ala Gly Gly Ala Cys Gly Cys Ala Thr

1205 1210 1215

Thr Gly Ala Cys Cys Gly Ala Gly Gly Cys Cys Gly Ala Cys Gly Cys

1220 1225 1230

Cys Cys Thr Gly Gly Cys Gly Gly Cys Cys Gly Cys Cys Gly Ala Gly

1235 1240 1245

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1250 1255 1260

Ala Ala Cys Ala Ala Gly Cys Ala Thr Gly Ala Ala Ala Cys Cys Gly

1265 1270 1275 1280

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1285 1290 1295

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1300 1305 1310

Thr Ala Cys Cys Gly Ala Ala Gly Ala Gly Ala Thr Cys Gly Ala Gly

1315 1320 1325

Gly Cys Gly Gly Ala Gly Ala Thr Gly Ala Thr Cys Gly Cys Gly Gly

1330 1335 1340

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1345 1350 1355 1360

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1365 1370 1375

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1380 1385 1390

Ala Thr Gly Ala Ala Ala Thr Cys Cys Thr Gly Gly Cys Cys Gly Gly

1395 1400 1405

Thr Thr Thr Gly Thr Cys Thr Gly Ala Thr Gly Cys Cys Ala Ala Gly

1410 1415 1420

Cys Thr Gly Gly Cys Gly Gly Cys Cys Thr Gly Gly Cys Cys Gly Gly

1425 1430 1435 1440

Cys Cys Ala Gly Cys Thr Thr Gly Gly Cys Cys Gly Cys Thr Gly Ala

1445 1450 1455

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1460 1465 1470

Cys Gly Thr Cys Thr Ala Ala Ala Ala Ala Gly Gly Thr Gly Ala Thr

1475 1480 1485

Gly Thr Gly Thr Ala Thr Thr Thr Gly Ala Gly Thr Ala Ala Ala Ala

1490 1495 1500

Cys Ala Gly Cys Thr Thr Gly Cys Gly Thr Cys Ala Thr Gly Cys Gly

1505 1510 1515 1520

Gly Thr Cys Gly Cys Thr Gly Cys Gly Thr Ala Thr Ala Thr Gly Ala

1525 1530 1535

Thr Gly Cys Gly Ala Thr Gly Ala Gly Thr Ala Ala Ala Thr Ala Ala

1540 1545 1550

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1555 1560 1565

Ala Ala Cys Gly Cys Ala Thr Gly Ala Ala Gly Gly Thr Thr Ala Thr

1570 1575 1580

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1585 1590 1595 1600

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1605 1610 1615

Ala Gly Ala Cys Gly Ala Cys Cys Ala Thr Cys Gly Cys Ala Ala Cys

1620 1625 1630

Cys Cys Ala Thr Cys Thr Ala Gly Cys Cys Cys Gly Cys Gly Cys Cys

1635 1640 1645

Cys Thr Gly Cys Ala Ala Cys Thr Cys Gly Cys Cys Gly Gly Gly Gly

1650 1655 1660

Cys Cys Gly Ala Thr Gly Thr Thr Cys Thr Gly Thr Thr Ala Gly Thr

1665 1670 1675 1680

Cys Gly Ala Thr Thr Cys Cys Gly Ala Thr Cys Cys Cys Cys Ala Gly

1685 1690 1695

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1700 1705 1710

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1715 1720 1725

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1730 1735 1740

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1745 1750 1755 1760

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1765 1770 1775

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1780 1785 1790

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1795 1800 1805

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1810 1815 1820

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1825 1830 1835 1840

Thr Thr Gly Gly Cys Thr Gly Thr Gly Thr Cys Cys Gly Cys Gly Ala

1845 1850 1855

Thr Cys Ala Ala Gly Gly Cys Ala Gly Cys Cys Gly Ala Cys Thr Thr

1860 1865 1870

Cys Gly Thr Gly Cys Thr Gly Ala Thr Thr Cys Cys Gly Gly Thr Gly

1875 1880 1885

Cys Ala Gly Cys Cys Ala Ala Gly Cys Cys Cys Thr Thr Ala Cys Gly

1890 1895 1900

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1905 1910 1915 1920

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1925 1930 1935

Gly Thr Thr Ala Ala Gly Cys Ala Gly Cys Gly Cys Ala Thr Thr Gly

1940 1945 1950

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1955 1960 1965

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1970 1975 1980

Gly Thr Cys Gly Thr Gly Thr Cys Gly Cys Gly Gly Gly Cys Gly Ala

1985 1990 1995 2000

Thr Cys Ala Ala Ala Gly Gly Cys Ala Cys Gly Cys Gly Cys Ala Thr

2005 2010 2015

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2020 2025 2030

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2035 2040 2045

Ala Cys Gly Ala Gly Cys Thr Gly Cys Cys Cys Ala Thr Thr Cys Thr

2050 2055 2060

Thr Gly Ala Gly Thr Cys Cys Cys Gly Thr Ala Thr Cys Ala Cys Gly

2065 2070 2075 2080

Cys Ala Gly Cys Gly Cys Gly Thr Gly Ala Gly Cys Thr Ala Cys Cys

2085 2090 2095

Cys Ala Gly Gly Cys Ala Cys Thr Gly Cys Cys Gly Cys Cys Gly Cys

2100 2105 2110

Cys Gly Gly Cys Ala Cys Ala Ala Cys Cys Gly Thr Thr Cys Thr Thr

2115 2120 2125

Gly Ala Ala Thr Cys Ala Gly Ala Ala Cys Cys Cys Gly Ala Gly Gly

2130 2135 2140

Gly Cys Gly Ala Cys Gly Cys Thr Gly Cys Cys Cys Gly Cys Gly Ala

2145 2150 2155 2160

Gly Gly Thr Cys Cys Ala Gly Gly Cys Gly Cys Thr Gly Gly Cys Cys

2165 2170 2175

Gly Cys Thr Gly Ala Ala Ala Thr Thr Ala Ala Ala Thr Cys Ala Ala

2180 2185 2190

Ala Ala Cys Thr Cys Ala Thr Thr Thr Gly Ala Gly Thr Thr Ala Ala

2195 2200 2205

Thr Gly Ala Gly Gly Thr Ala Ala Ala Gly Ala Gly Ala Ala Ala Ala

2210 2215 2220

Thr Gly Ala Gly Cys Ala Ala Ala Ala Gly Cys Ala Cys Ala Ala Ala

2225 2230 2235 2240

Cys Ala Cys Gly Cys Thr Ala Ala Gly Thr Gly Cys Cys Gly Gly Cys

2245 2250 2255

Cys Gly Thr Cys Cys Gly Ala Gly Cys Gly Cys Ala Cys Gly Cys Ala

2260 2265 2270

Gly Cys Ala Gly Cys Ala Ala Gly Gly Cys Thr Gly Cys Ala Ala Cys

2275 2280 2285

Gly Thr Thr Gly Gly Cys Cys Ala Gly Cys Cys Thr Gly Gly Cys Ala

2290 2295 2300

Gly Ala Cys Ala Cys Gly Cys Cys Ala Gly Cys Cys Ala Thr Gly Ala

2305 2310 2315 2320

Ala Gly Cys Gly Gly Gly Thr Cys Ala Ala Cys Thr Thr Thr Cys Ala

2325 2330 2335

Gly Thr Thr Gly Cys Cys Gly Gly Cys Gly Gly Ala Gly Gly Ala Thr

2340 2345 2350

Cys Ala Cys Ala Cys Cys Ala Ala Gly Cys Thr Gly Ala Ala Gly Ala

2355 2360 2365

Thr Gly Thr Ala Cys Gly Cys Gly Gly Thr Ala Cys Gly Cys Cys Ala

2370 2375 2380

Ala Gly Gly Cys Ala Ala Gly Ala Cys Cys Ala Thr Thr Ala Cys Cys

2385 2390 2395 2400

Gly Ala Gly Cys Thr Gly Cys Thr Ala Thr Cys Thr Gly Ala Ala Thr

2405 2410 2415

Ala Cys Ala Thr Cys Gly Cys Gly Cys Ala Gly Cys Thr Ala Cys Cys

2420 2425 2430

Ala Gly Ala Gly Thr Ala Ala Ala Thr Gly Ala Gly Cys Ala Ala Ala

2435 2440 2445

Thr Gly Ala Ala Thr Ala Ala Ala Thr Gly Ala Gly Thr Ala Gly Ala

2450 2455 2460

Thr Gly Ala Ala Thr Thr Thr Thr Ala Gly Cys Gly Gly Cys Thr Ala

2465 2470 2475 2480

Ala Ala Gly Gly Ala Gly Gly Cys Gly Gly Cys Ala Thr Gly Gly Ala

2485 2490 2495

Ala Ala Ala Thr Cys Ala Ala Gly Ala Ala Cys Ala Ala Cys Cys Ala

2500 2505 2510

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2515 2520 2525

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2530 2535 2540

Ala Gly Gly Ala Ala Cys Gly Gly Gly Cys Gly Gly Thr Thr Gly Gly

2545 2550 2555 2560

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2565 2570 2575

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2580 2585 2590

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2595 2600 2605

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2610 2615 2620

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2625 2630 2635 2640

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2645 2650 2655

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2660 2665 2670

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2675 2680 2685

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2690 2695 2700

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2705 2710 2715 2720

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2725 2730 2735

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2740 2745 2750

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2755 2760 2765

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2770 2775 2780

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2785 2790 2795 2800

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2805 2810 2815

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2820 2825 2830

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2835 2840 2845

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2850 2855 2860

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2865 2870 2875 2880

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2885 2890 2895

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2900 2905 2910

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2915 2920 2925

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2930 2935 2940

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2945 2950 2955 2960

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2965 2970 2975

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2980 2985 2990

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2995 3000 3005

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3010 3015 3020

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3025 3030 3035 3040

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3045 3050 3055

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3060 3065 3070

Thr Ala Ala Cys Cys Gly Ala Ala Thr Cys Cys Ala Thr Gly Ala Ala

3075 3080 3085

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3090 3095 3100

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3105 3110 3115 3120

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3125 3130 3135

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3140 3145 3150

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3155 3160 3165

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3170 3175 3180

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3185 3190 3195 3200

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3205 3210 3215

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3220 3225 3230

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3235 3240 3245

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3250 3255 3260

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3265 3270 3275 3280

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3285 3290 3295

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3300 3305 3310

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3315 3320 3325

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3330 3335 3340

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3345 3350 3355 3360

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3365 3370 3375

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3380 3385 3390

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3395 3400 3405

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3425 3430 3435 3440

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3475 3480 3485

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3490 3495 3500

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3505 3510 3515 3520

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3525 3530 3535

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3540 3545 3550

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