本发明涉及医学检验领域，具体涉及一种dnah6基因突变体及其应用。
背景技术：
精子头部异常是一种隐性遗传的导致男性不育的畸形精子症，是一种罕见的遗传疾病，其主要临床特征涉及多种头部异常亚型，包括圆头精子症、大头精子症、无头精子症等亚型。无头精子症(acephalicspermatozoa)是精子没有头部或者头部与尾部之间连接松散。早期研究中也有文献称这个病为断头精子症(decapitatedspermatozoa)、大头针状精子(pinheadsperm)。国际早期的分子发病机制研究发现sun5或tsga10基因发生了突变(1-2)。圆头精子症(globozoospermia)是指精子头部呈圆形，通常无顶体。以往分子遗传学研究发现dpy19l2基因突变会导致圆头精子症(3)。但迄今为止国际上很少报道一个患者同时患无头合并圆头精子症。
目前无头合并圆头精子症的基因突变谱尚未发现，基因型和表现型的关系也不明确。目前单基因疾病的研究开始大量采用全外显子组测序(whole-exomesequencing)及全基因组测序(whole-genomesequencing)的方法，这两种方法被成功的应用于发现稀有单基因疾病的致病基因。全外显子组测序及全基因组测序技术已被证明为降低稀有单基因疾病候选基因甚至发现其致病基因的有力、有效手段。仅通过对几个很少的个体(包括患者及正常对照)的全外显子组或基因组进行测序来筛选与疾病相关的变异，其成功率大为提升。
因此本领域对无头合并圆头精子症的研究尚不清晰，对造成该疾病的原因更不明了，因此本领域迫切需要对无头合并圆头精子症的致病机理进行研究，找到致病基因及突变位点。
技术实现要素：
本发明通过全外显子组测序的方法确定了无头合并圆头精子症的致病新基因。
根据本发明，提供一种无头合并圆头精子症的生物标记物，所述生物标记物为突变的dnah6基因或dnah6蛋白，所述生物标记物是具有选自如下的突变的dnah6基因或dnah6蛋白：
在外显子15中错义突变(c.2454a>t；p.e818d)；
在外显子48中错义突变(c.7706g>a；p.r2569h)。
本发明提供一种分离的编码dnah6突变体的核酸，其特征在于，所述核酸与seqidno:1相比，具有以下突变中的一种或两种：c.2454a>t突变、c.7706g>a突变。
其中，所述多肽为dnah6突变体，所述多肽与seqidno:2相比，具有以下突变中的一种或两种：p.e818d突变、p.r2569h突变。
其中，具有p.e818d突变的dnah6突变体是由具有c.2454a>t突变的dnah6突变基因编码，具有p.r2569h突变的dnah6突变体是由具有c.7706g>a突变的dnah6突变基因编码。
本发明还提供一种编码dnah6突变体的核酸在精子头部异常检测中的用途，其中，通过检测所述核酸在采集自受试者的生物样品中是否存在来判断所述受试者是否易患精子头部异常，其中，所述核酸与seqidno:1相比，具有以下突变中的一种或两种：c.2454a>t突变、c.7706g>a突变。
本发明还提供一种多肽在精子头部异常检测中的用途，其中，通过检测所述多肽在采集自受试者的生物样品中是否表达来判断所述受试者是否易患精子头部异常，其中，所述多肽为dnah6突变体，所述多肽与seqidno:2相比，具有以下突变中的一种或两种：p.e818d突变、p.r2569h突变。
其中，所述精子头部异常是无头精子合并圆头精子症。
本发明还提供一种检测编码dnah6突变体的核酸的方法，其中，所述方法包括使用以下至少一组引物进行pcr扩增的步骤：
seqidno:3和seqidno:4；
seqidno:5和seqidno:6。
其中，引物seqidno:3和seqidno:4用于检测与seqidno:1相比具有c.2454a>t突变的核酸，引物seqidno:5和seqidno:6用于检测与seqidno:1相比具有c.7706g>a突变的核酸。
本发明还提供一种用于检测编码dnah6突变体的核酸的引物，其中，所述引物为以下中的至少一组：
seqidno:3和seqidno:4；
seqidno:5和seqidno:6，
其中，引物seqidno:3和seqidno:4用于检测与seqidno:1相比具有c.2454a>t突变的核酸，引物seqidno:5和seqidno:6用于检测与seqidno:1相比具有c.7706g>a突变的核酸。
本发明还提供一种用于筛选易患精子头部异常的生物样品的试剂盒，其中，所述试剂盒含有特异性检测dnah6基因突变体的试剂，所述dnah6基因突变体与seqidno:1相比，具有以下突变中的一种或两种：c.2454a>t突变、c.7706g>a突变。
其中，所述精子头部异常是无头精子合并圆头精子症。
本发明还提供一种检测无头合并圆头精子症的方法，所述方法包括检测受试者的dnah6基因或dnah6蛋白中是否存在突变位点，如果有突变位点，则所述受试者被鉴定为患有无头合并圆头精子症或易患无头合并圆头精子症，所述突变位点选自如下任一种或其组合：
在外显子15中错义突变(c.2454a>t；p.e818d)；
在外显子48中错义突变(c.7706g>a；p.r2569h)。
其中，野生型dnah6基因具有seqidno:1所示序列，野生型dnah6蛋白具有seqidno:2所示序列。
具体实施方式
下面通过具体实施方式对本发明进一步详细说明。通过这些说明，本发明的特点和优点将变得更为清楚明确。
在本发明中，采用本领域通用表示法表示突变。例如，突变(c.2454a>t；p.e818d)中，c表示cdna，p表示蛋白质，dna水平的突变对应蛋白质水平的突变；
野生型dnah6基因的cdna序列如seqidno:1所示。
seqidno:2：野生型dnah6蛋白的氨基酸序列。
对于本发明说明书和权利要求书中，提及基因序列，本领域技术人员应当理解，实际包括互补双链的任意一条，或者两条。为了方便，在本说明书和权利要求书中，虽然多数情况下至给出了一条链，但实际上也公开了与之互补的另一条链。例如提及dnah6基因的cdna序列，实际上包括该序列以及其互补序列。例如，提及seqidno:1，实际包括其互补序列。本领域技术人员还可以理解，利用一条链可以检测另一条链，反之亦然。
本申请中的基因序列包括dna形式或rna形式，公开其中一种，意味着另一种也被公开。例如提及dnah6基因的cdna序列，实际也包括相应的rna序列。
实施例1确定无头合并圆头精子症的致病基因
发明人在国内收集到1个无头合并圆头精子症核心家系，患者表现为不育，无头、大头针样头或头尾部连接松散等典型的无头精子症及圆头精子疾病特征。
发明人对所有患者进行了全外显子组测序，具体步骤如下：
样品制备：采集所述患者外周血，利用试剂盒抽提外周血白细胞中的基因组dna(qiaampdnaminikit51304.qiagen,usa)，利用nanodrop2000测量dna的浓度及纯度(thermoscientific,usa)，所得的每个标本基因组dna的od260/od280均位于1.7-2.0之间，浓度不少于100ng/μl，总量不少于30μl。
然后，对上述患者样本的外显子组序列进行了测序。测序平台为illuminahiseq2000，依照illumina标准建库说明书(参见http://www.illumina.com/)进行测序，简述如下：
1)使用illumina公司试剂盒truseqdnalibraryprepkit制作dna文库。在外显子捕获过程中使用agilent公司试剂盒sureselecthumanallexonv5；
2)使用illumina公司hiseq2000测序平台对捕获的外显子区域进行平行测序读取长度为90bp，每个样本的平均测序深度最少为100；
3)使用burrows-wheeleraligner(http://bio-bwa.sourceforge.net/)将测序结果比对到人参考基因组中(hg19)。使用samtools将重复读取结果去除，通过gatk(https://www.broadinstitute.org/gatk/)进行调整和校准。然后使用samtools(http://www.samtools.sourceforge.net/)挖掘出snvs及indels，并用annovar(http://www.openbioinformatics.org/annovar/)命名。
将等位基因频率大于1％的突变位点剔除(参考数据库包括dbsnp，1000genomes，exomeaggregationconsortium(exac))。
发现该患者在基因dnah6上具有复杂杂合突变，这些位点可能是致病位点。
结果显示于表1。
因此发明人认为dnah6基因极有可能为无头合并圆头精子症的致病基因。
实施例2通过sanger法测序确认上述无头合并圆头精子症的致病基因
接着，发明人纳入无头合并圆头精子症患者及其家人，对dnah6基因为无头合并圆头精子症的致病基因进行验证。验证中利用sanger法测序验证dnah6基因的突变，其中考虑了家系内样本验证情况。
分别对1名患者、2名家系内正常人(即其中1名患者的父母，他们均未发病)基因进行检测。
具体方法步骤如下：
1.dna提取：
分别提取1名患者、2名家系内正常人的外周静脉血，按照实施例1的方法提取基因组dna和测定dna含量。
2.引物设计及pcr反应
引物设计参考人类基因组序列数据库hg19，具体见下。
a)引物序列：
b)反应体系：
pcr扩增体系：
c)pcr反应条件：
94℃变性5分钟，然后进入每个循环，即94℃变性30秒，退火(通常为55℃，根据不同引物的退火温度设定)30秒，72℃延伸1分钟，总共设定30个循环。所有循环结束，最终72℃延伸10分钟，并将pcr产物保存在4或-20℃。
3)将步骤2中获得的获自1名无头合并圆头精子症患者、2名家系内正常人(即该患者的父母，他们均未发病)的pcr扩征产物直接进行dna测序，方法同实施例1。
发明人收集到的该近亲结婚家系的无头合并圆头精子症病例在dnah6基因中检测到有意义的突变位点。同时所发现的突变位点在家族正常人中均为杂合突变，表现出家系内共分离。这些突变位点在exac数据库中的频率极低(见表1)，提示所检测到的位点很可能并非snp。因此我们认为dnah6基因为无头合并圆头精子症的致病基因。
表1：
上述说明示出并描述了本发明的一种实例，但如前所述，应当理解本发明并非局限于本文所披露的形式，不应看作是对其他实施例的排除，而可用于各种其他组合、修改和环境，并能够在本文所述发明构想范围内，通过上述教导或相关领域的技术或知识进行改动。而本领域人员所进行的改动和变化不脱离本发明的精神和范围，则都应在本发明所附加权利要求的保护范围内。
以上结合优选实施方式和范例性实例对本发明进行了详细说明。不过需要声明的是，这些具体实施方式仅是对本发明的阐述性解释，并不对本发明的保护范围构成任何限制。在不超出本发明精神和保护范围的情况下，可以对本发明技术内容及其实施方式进行各种改进、等价替换或修饰，这些均落入本发明的保护范围内。本发明的保护范围以所附权利要求为准。
序列表
<110>首都医科大学附属北京妇产医院
<120>dnah6基因突变体及其应用
<130>zf1710019c
<160>6
<170>siposequencelisting1.0
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<211>12477
<212>dna
<213>homosapiens
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ttgccaaaatttctaacagatgatgctcttctgttcagtggaatcatatctgaccttttt5100
cctggagtccaaattccagaacatgattatggtattttacaatcaacaattgtggatgtc5160
atgaatagacaaaatcttcagcctgagatgtgtatggttagaaaggtgatacagttttat5220
gaaactatgctagtaaggcatggtgttatgttagtcgggccaacaggaggcggcaagacc5280
acagtttaccgaatactagcagaaactttagggaatttacaaaaacttgggatagaaaat5340
tccttttaccaagcagttaaaacatatgttctcaaccctaaatcaattaccatgggtgaa5400
ttatatggagaggttaataacttaaccttggaatggaaagatggtttgatggcactaagt5460
gtccgagcagctgtgaatgatacttcagaagaccataaatggatcatcagtgatgggcca5520
gtagatgctctttggattgaaaacatgaatacagtgctggatgataacaagatgctttgc5580
ctggctaacagtgagaggattaaactcacacctcaaattcacatgctttttgaggtgcaa5640
gatctgcgggttgcctcccctgcaacagtcagtcgatgtggaatggtgtttgtggatcct5700
gaagaactgaaatggatgccttatgttaaaacttggatgaagggtatttctaaaaaactg5760
actgaggaaacccaagaatatatattgaatcttttccaacgttatgttgatgaaggttta5820
cattttatcaataaaaagtgcagccaagcaattccacaagtggacatcagcaaagttact5880
acactctgttgcttattggagtccttgatacttgggaaagatggagttaacttggcaatg5940
gaacaaacaaaattgaacactatactatgtcagacttttgtattctgttatttgtggtct6000
ttgggtggaaacctaactgaaaactactatgattcttttgatacatttattagaacacaa6060
tttgatgataatcctgatgccaggcttcccaattctggtgatctgtggagcattcatatg6120
gactttgacaccaaacggctggatccctgggaacgaatcatacctactttcaaatacaac6180
cgagatgttccattttttgaaatgcttgtccccacaactgacacagtgcgctatgggtat6240
ctaatggaaaaactactggcagtcaagcattccgtgttgtttactggaataactggagtg6300
ggcaagtctgtgattgcaaaaggattgctaaataaaattcaagaatcagctggctatgtc6360
cctgtttatctaaatttttctgctcaaacttcatctgcaaggacacaagagatcattgag6420
tcaaaactggagagaaaaagaaaaaatattctaggagcaccgggaaacaaacgaattgtg6480
atttttgttgatgatttaaacatgcccagactggatcgctatggctctcagcctccgatt6540
gaattacttcggcagtatcaagattttgggggattttatgacagaaacaaactgttttgg6600
aaagaaatacaggatgtaacaatcatatcggcatgtgcacctccaggcggtggccgcaac6660
cctgtgactccccgcttcatcagacacttcagcatgctgtgcctcccaatgccctcagag6720
cacagtctgaaacagatttttcaggccatcttaaatggtttcctgagtgactttccacca6780
gctgtaaagcaaactgcatcaagcattgtagaagcctcagttgagatttataacaaaatg6840
agtgttgacctcctgccaacacccgccaagtcccattatgtctttaacttgagggactta6900
tccaaatgtgtgcaaggtatcctccaatgtgatccaggaacaataagagaagaaattcag6960
atatttagactcttttgccatgagtgccaaagggtcttccatgatcgcttgattaataat7020
gaagataagcactatttccatgttattctgacagaaatggccaacaaacattttggaatt7080
gcaattgacctggaatattttttgaataagcccatcatatttggagatttcattaagttt7140
ggagcagataaagctgatcggatttatgatgacatgcctgatatagagaaaactgcaaat7200
gttctacaggactatcttgatgattataatctcacaaatcccaaagaagtaaagttggtg7260
ttcttccaggatgctatagaacatgtttcaaggattgctcggatgatacgtcaagaaaga7320
ggcaatgccctgcttgttggagtaggaggcacaggaaagcagtcactcacgagacttgca7380
gctcatatatgcggttacaaatgtttgcagattgaactcagccggggatataattatgat7440
agttttcatgaagacctgaggaagttgtacaaaatggctggtgtagaagacaagaatatg7500
gttttccttttcactgacacccagattgtagtggaggagttcctagaagatataaataac7560
atcctgaactcaggtgaagtgcctaatttatttgaaaaggatgaactggagcaggtttta7620
gcggccaccagaccaagagcaaaagaagtaggaatttctgaggggaacagagacgaggtg7680
tttcaatactttatcagcaaagtgcgtcagaagctgcacattgttctctgcatgagccca7740
gttggggaggcctttcggtcccgatgcaggatgtttccatcccttgtgaattgctgcacc7800
attgactggtttgtgcagtggcccagagaagcacttctttctgtgtcaaagacatttttc7860
tcacaagtcgatgctggaaatgaagaactgaaagaaaagcttcccttgatgtgcgtgaac7920
gttcacttgagtgtctccagcatggcagagcgctattacaatgagctgcgcaggcggtac7980
tacacgacacccacctcctacctggagcttatcaatctttacctgtctatgctgtctgaa8040
aaaaggaagcagattatttcagcacgagatcgggtgaagaatggtctcaccaagctacta8100
gaaacaaacatactagtagataaaatgaaactagatctttcagctttagagcctgtactt8160
ttagcaaaatcagaagatgttgaagccctgatggaaaaattggcagtggatcaagaaagt8220
gccgatcaggtccgtaacactgtgcaggaggatgaagcaacagcaaaagtcaaagctgaa8280
gaaacccaagcaatagctgatgatgctcaaagagatcttgacgaggcactacctgcacta8340
gatgctgccaataaagcactggattccttagataaggcagatatatctgaaatcagagtt8400
tttacaaagcccccagatttggtcatgacagtaatggaagcaatctccattcttttgaat8460
gccaagcctgattggccatcagcaaagcaacttcttggtgactctaactttctaaaaagg8520
cttttagaatatgataaggagaacataaagcctcagatattggcaaagcttcaaaagtat8580
attaataatcctgattttgtgcctgaaaaagtggagaaagtgtccaaagcatgtaaatct8640
atgtgcatgtgggtaagagctatggatttgtactctcgagtggtcaaggtcgtcgaacca8700
aaaagacaaaagctccgcgccgcacaggctgaacttgacattaccatggctaccctgaga8760
gaaaagcaagcattactaagacaagtagaagatcaaatacaggccttacaagatgaatat8820
gacaaaggtgtaaatgaaaaagaaagcctggcaaagaccatggccctgacaaaagcacgt8880
ctagtacgtgctggaaagctgacagcagcattagaagatgagcaggttcgatgggaagaa8940
agcatacagaagtttgaggaagaaatatcaaatatcactgggaacgtgttcatagcagca9000
gcttgtgtggcctactatggggctttcacagcccagtacaggcagtcacttatagagtgt9060
tggatccaggactgtcagtctctggagatcccaatcgatccttccttcagtctcattaac9120
attcttggagatccctacgagatacggcagtggaacactgatgggctgccccgtgacttg9180
atatcaacagaaaatggcattttggttactcaaggcagaagatggcctttgatgattgat9240
ccccaagatcaggcaaaccgttggataaggaacaaggaaagcaaaagtggtttaaagatc9300
attaagcttacagatagtaatttcttacgaatactcgagaattcaatccgacttggttta9360
cctgtcttactggaagagcttaaggaaaccttggatccagctctagaacccattcttttg9420
aaacaaatttttatcagtggtggccgactactcatccgtcttggagactcagacattgat9480
tatgacaaaaactttaggttctatatgacaaccaaaatgccaaatccccactatctgcct9540
gaggtatgcattaaagttaccattatcaatttcactgtaacaaaatcaggcctggaggat9600
cagttgttaagtgatgtggtgcgacttgaaaaacccaggttggaagaacaaagaattaag9660
ctcatcgtgaggatcaacactgataaaaaccagttgaaaactatcgaagagaaaatcctg9720
agaatgctctttacctctgaaggaaatattctggacaatgaagaacttattgacacactc9780
caggattcaaagatcacttctggtgccattaaaaccaggctggaagaagcagagtccact9840
gagcagatgatcaatgtggctcgtgagaagtatcgtccagtggccactcaaggctctgta9900
atgtactttgtcattgcaagcctctcagaaatagatcctatgtaccagtactcattaaaa9960
tactttaaacagttgttcaataccaccattgaaacttctgtaaagacagaaaatctacaa10020
cagcgcctggacgtactactagaacaaactctcctaactgcttatgtcaatgtttcaaga10080
ggactttttgagcaacataaactcatctacagctttatgctttgtgttgagatgatgcgt10140
cagcaaggaaccctatctgatgctgaatggaatttctttctccgaggttctgcaggattg10200
gaaaaggaacgcccacctaagcctgaagctccctggctacctactgctacatggttcgca10260
tgctgtgacttggaagaatcatttccagtttttcacggacttacccaaaatatattgtca10320
catcctatttccatacgcttaggatcttttgagacttatattaacccacagaaatgggaa10380
ggctattctaaaatgaaacacgaagataaacacatgagacaggaaaaggaggcagcacac10440
caagatccatggagtgcaggattgagttctttccataagctaattcttattaaatgttgt10500
aaagaagaaaaggtggtttttgctcttacagactttgtgatagaaaatcttggaaaacag10560
tttatagagacaccacctgtggacctgcctaccctgtatcaagacatgtcatgcaacact10620
cccctggtattcatcctaagcacaggctcagatcccatgggtgcatttcagaggtttgcc10680
agggaaagtggatattcagaacgggtgcagtcaatttcactggggcaaggacaaggacct10740
attgctgaaaaaatggtcaaggatgcaatgaaatcaggaaactgggtatttttgcaaaat10800
tgccatcttgctgtttcttggatgttggcaatggaagagctcattaaaaccttcacagat10860
ccagatagtgctatcaaggacacttttcgactttttttaagctccatgcctagtaataca10920
tttcctgttacagttcttcaaaattctgtcaaggtgaccaatgagcctccaaaaggctta10980
cgtgcaaatatcagacgagcatttactgaaatgacaccttcgttttttgaagaaaatata11040
cttggaaaaaaatggagacaaataatatttggcatttgtttcttccatgcaattattcag11100
gagagaaagaagtttggcccccttggttggaatatctgctatgaatttaatgacagtgac11160
agggaatgtgctttactgaatctcaaactctattgtaaagaaggaaagattccctgggat11220
gcactaatttacattactggtgaaattacttatggtggtagagtcacagacagctgggac11280
caaagatgccttcgtactatcttgaaaagatttttttctcctgaaacattagaagaagat11340
tataaatactctgaatcaggcatctattttgcacccatggctgacagcctacaagagttt11400
aaggactacattgaaaatctgcctttgatcgatgacccagaaatttttggaatgcatgaa11460
aatgctaatctagtcttccagtacaaagagaccagcactttaatcaacaccatacttgag11520
gttcagccaaggtcatctactggtggagagggaaaaagcaatgacgaaattgttcaagaa11580
cttgttgcttctgtccagaccagagttccagaaaaactggaaatggagggtgcttctgag11640
agcctttttgtcaaggatcttcaaggacgtctgaactccttgaccaccgttcttggacag11700
gaagtggaccggtttaacaacctgctgaagttaattcatacttctctggaaacactcaac11760
aaagccatcgctggatttgtggtgatgtctgaagaaatggaaaaagtgtataacagtttc11820
ctcaacaaccaggttcccgctctgtggtccaacacagcctacccatccctgaagccacta11880
ggatcatgggtcaaagaccttatcctgaggacctcatttgtggatctgtggctcaaaaga11940
ggacagcctaagtcctactggatctctggtttcttctttcctcaaggatttctaacagga12000
actcttcaaaatcatgctcgaaaatacaatttgcctatagatgagctgagtttcaaatac12060
agcgtaattcccacctatcgggatcaagctgcagtgatagaagctgccaagacagtgcaa12120
tttggacaagaactgcccatggacatggagttgccctctcctgaggatggtgttcttgtt12180
catgggatgttcatggatgcttctcgatgggatgataaggagatggtgatagaagatgca12240
ttgcccggacagatgaatccagtgctgcctgtggtgcattttgaaccacaacaaaactat12300
aagccaagcccaacactttaccactgcccactttataaaacaggagcccgggcaggaaca12360
ctctcaaccacaggacattcaaccaattttgtggtaaccgtcctgttaccctccaagcgg12420
tccaaagactactggattgccaagggatcagctttgctctgccagctgagcgaatga12477
<210>2
<211>4158
<212>prt
<213>homosapiens
<400>2
metthrpheargalathraspserglupheaspleuthrasnileglu
151015
glutyralagluasnseralaleuserargleuasnasnilelysala
202530
lysglnargvalsertyrvalthrserthrgluasngluseraspthr
354045
glnileleuthrphearghisilethrlysalaglnglulysthrarg
505560
lysargglnglnproilelysleugluproleuprovalleulysval
65707580
tyrglnasphislysglnproglutyrilehisgluglnasnargphe
859095
glnleumetthralaglyileilelysargprovalserilealalys
100105110
lysserphealathrserserthrglnpheleugluhisglnaspala
115120125
vallyslysmetglnilehisargprotyrvalgluvalpheserpro
130135140
serproprolysleuprohisthrglyileglylysargglyleuphe
145150155160
glythrargserseralatyrprolystyrthrphehisaspargglu
165170175
gluvalvallysalaasnileargaspproleuglnileilelysile
180185190
ilearggluasngluhisleuglypheleutyrmetileproalaval
195200205
proargserserileglutyraspthrtyrasnleulysvalvalser
210215220
tyrgluasnileasnlysasnasptyrtyrthrileserglnargala
225230235240
valthrhisiletyrasngluaspileglupheilegluileasparg
245250255
trpgluglnglutyrleutyrhisarggluleuthrlysileproile
260265270
pheserleuphearglystrplysalapheservaltrparglysasn
275280285
valargserlyslysilethrglycysglnlysserleuglnlysasn
290295300
leupheilevalasnprohisleuargproalaleuleulysileasn
305310315320
gluleucystyrhisleuserphemetglyleucystyrileglulys
325330335
cyshisthrtyrthrleuglngluphelysalaalaglnvalilearg
340345350
leualagluvalthrgluargleuglyglupheargasnglualalys
355360365
tyrvalvalargargalacysargphealaleuargalaalaglyphe
370375380
valproaspaspcysalapheglyprophegluasptyrhislysval
385390395400
glnserserglyserpheileasnthrprohisgluleuprothrtyr
405410415
glyaspserglulysmetthrtyrthrgluglnalaserlysarghis
420425430
tyrcysmetargleuthrcyspheileargleuasnasptyrleuile
435440445
gluasnthrmethisileleuthrvalasnalavalasnserleuleu
450455460
asnhisleuthrasplysleulysargthrproseralaaspvalile
465470475480
glnlystrpilethrgluglulysprogluvalproasplyslysgly
485490495
thrleumetvalglulysglnglugluaspgluserleuilepromet
500505510
pheleuthrgluleumetleuthrvalglnserleuleupheglupro
515520525
serleugluasppheleuaspglyileleuglyalavalasnhiscys
530535540
glnasnthrvalleuservalproasnleuvalproaspsertyrphe
545550555560
aspalaphethrserprotyrileasnasnlysleugluglylysthr
565570575
cysglythrglyproserleualaalavalphegluaspasplysasn
580585590
phehisthrileileserglnilelysgluthrileglnalaalaphe
595600605
gluseralaargiletyralaalathrpheglulyspheglnilephe
610615620
phelysgluasngluserleuaspleuglnalaleulysleuglnglu
625630635640
proaspileasnphephesergluglnleuglulystyrhislysgln
645650655
hislysaspalavalalaleuargprothrargasnvalglyleuleu
660665670
leuileaspthrargleuleuargglulysleuileproserproleu
675680685
argcysleugluvalleuasnphemetleuproargglnserlyslys
690695700
lysvalaspalaileilepheglualaglnaspalaglutyrlysleu
705710715720
gluphevalprothrthrthrthrglutyrvalhisserleuleuphe
725730735
leuaspgluileglngluargilegluserleugluaspgluglyasn
740745750
ilevalthrglnmettyrlysleumetgluglntyrglnvalprothr
755760765
proprogluaspphealavalphealathrmetlysproserileval
770775780
alavalargasnalaileasplysservalglyasparggluserser
785790795800
ilelysglnphecysvalhisleuglyseraspleuglugluleuasn
805810815
asngluvalasngluvallysleuglnalaglnaspproglnileleu
820825830
aspileseralaaspglnasplysileargleuileleuasnasnleu
835840845
glnservalleualaaspleuglnlysargalapheglntyrlysser
850855860
tyrglnlysasnphelysvalgluvalserlyspheglualaleuglu
865870875880
gluvalseralagluleulysleulysglnleuleutrpaspserphe
885890895
serglutrpasplysleuglnglnglutrpleulysserlyspheasp
900905910
cysleuaspprogluvalleuasnglyglnvalserlystyralalys
915920925
phevalthrglnleuglulysglyleuproproasnservalvalpro
930935940
glnleulystyrlysvalglulysmetlysglulysleuprovalile
945950955960
ileaspleuargasnprothrleulysalaarghistrpalaalaile
965970975
gluglnthrvalaspalathrleuvalaspalagluileproleuthr
980985990
leugluargleuserglnleuhisvalpheasppheglyglngluile
99510001005
glnaspileserglyglnalaserglyglualaalaleuglualaile
101010151020
leulyslysvalgluaspsertrplysthrthrgluphevalileleu
1025103010351040
prohisargaspserlysaspvalpheileleuglyglythraspasp
104510501055
ileglnvalleuleuaspaspserthrileasnvalalathrleuala
106010651070
serserargtyrleuglyproleulysthrargvalaspglutrpgln
107510801085
lysglnleualaleupheasnglnthrleugluglutrpleuthrcys
109010951100
glnargasntrpleutyrleugluserilepheasnalaproaspile
1105111011151120
glnargglnleuproalaglualalysmetpheleuglnvalasplys
112511301135
sertrplysgluilemetarglysvalasnargleuproasnalaleu
114011451150
argalaalathrglnproglyleuleugluthrpheglnasnasnasn
115511601165
alaleuleuaspglnileglnlyscysleuglualatyrleugluser
117011751180
lysargvalilepheproargphetyrpheleuserasnaspgluleu
1185119011951200
leugluileleualaglnthrargasnproglnalavalglnprohis
120512101215
leuarglyscyspheaspserileserlysleugluphealaleumet
122012251230
proproalagluglylysileproglyileaspglygluproglulys
123512401245
valtyrthrasnaspileleualametleuserprogluglygluarg
125012551260
valserleuglylysglyleulysalaargglyasnvalgluglutrp
1265127012751280
leuglylysvalgluglualametphethrserleuargargleucys
128512901295
lysalaalailealaasptyrglnglylysleuargthrasptrpval
130013051310
valalaglyhisproserglnvalileleuthrvalserglnilemet
131513201325
trpcysargaspleuthrglucysleugluthrgluhisserasnhis
133013351340
ileglnalaleulysasnpheglulysvalasnphegluargleuasn
1345135013551360
alaleualaalailevalglnglyserleuprolysleuhisargasn
136513701375
ileleuthralaleuilethrileaspvalhisalaargaspileval
138013851390
thrgluleuvalglnserlysvalgluthrvalgluserpheasptrp
139514001405
glnargglnleuargtyrtyrtrpaspileaspleuaspasncysval
141014151420
alaargmetalaleuserglntyrthrtyrglytyrglutyrleugly
1425143014351440
alacysproargleuvalilethrproleuthraspargcystyrleu
144514501455
cysleumetglyalaleuglnleuaspleuglyglyalaproalagly
146014651470
proalaglythrglylysthrgluthrthrlysaspleualalysala
147514801485
leualaileglncysvalvalpheasncysseraspglyleuasptyr
149014951500
lysmetmetglyargphepheserglyleualaglnserglyalatrp
1505151015151520
cyscyspheaspglupheasnargileaspilegluvalleuserval
152515301535
ilealaglnglnleuilethrileargasnalalysalaalalysleu
154015451550
serargphemetphegluglyarggluilelysleuvalmetthrcys
155515601565
alaalapheilethrmetasnproglytyralaglyargthrgluleu
157015751580
proaspasnleulysalaleupheargprophealametmetvalpro
1585159015951600
asntyralaleuilealagluvalileleutyrsergluglypheglu
160516101615
serserlysileleualaarglysmetthrglnmettyrlysleucys
162016251630
sergluglnleuserglnglnasphistyrasppheglymetargala
163516401645
vallysservalleuvalmetalaglyserleulysarggluasnpro
165016551660
aspleuasngluaspvalvalleuileargalaleuglnaspserasn
1665167016751680
leuprolyspheleuthraspaspalaleuleupheserglyileile
168516901695
seraspleupheproglyvalglnileprogluhisasptyrglyile
170017051710
leuglnserthrilevalaspvalmetasnargglnasnleuglnpro
171517201725
glumetcysmetvalarglysvalileglnphetyrgluthrmetleu
173017351740
valarghisglyvalmetleuvalglyprothrglyglyglylysthr
1745175017551760
thrvaltyrargileleualagluthrleuglyasnleuglnlysleu
176517701775
glyilegluasnserphetyrglnalavallysthrtyrvalleuasn
178017851790
prolysserilethrmetglygluleutyrglygluvalasnasnleu
179518001805
thrleuglutrplysaspglyleumetalaleuservalargalaala
181018151820
valasnaspthrsergluasphislystrpileileseraspglypro
1825183018351840
valaspalaleutrpilegluasnmetasnthrvalleuaspaspasn
184518501855
lysmetleucysleualaasnsergluargilelysleuthrprogln
186018651870
ilehismetleuphegluvalglnaspleuargvalalaserproala
187518801885
thrvalserargcysglymetvalphevalaspproglugluleulys
189018951900
trpmetprotyrvallysthrtrpmetlysglyileserlyslysleu
1905191019151920
thrglugluthrglnglutyrileleuasnleupheglnargtyrval
192519301935
aspgluglyleuhispheileasnlyslyscysserglnalailepro
194019451950
glnvalaspileserlysvalthrthrleucyscysleuleugluser
195519601965
leuileleuglylysaspglyvalasnleualametgluglnthrlys
197019751980
leuasnthrileleucysglnthrphevalphecystyrleutrpser
1985199019952000
leuglyglyasnleuthrgluasntyrtyraspserpheaspthrphe
200520102015
ileargthrglnpheaspaspasnproaspalaargleuproasnser
202020252030
glyaspleutrpserilehismetasppheaspthrlysargleuasp
203520402045
protrpgluargileileprothrphelystyrasnargaspvalpro
205020552060
phepheglumetleuvalprothrthraspthrvalargtyrglytyr
2065207020752080
leumetglulysleuleualavallyshisservalleuphethrgly
208520902095
ilethrglyvalglylysservalilealalysglyleuleuasnlys
210021052110
ileglngluseralaglytyrvalprovaltyrleuasnpheserala
211521202125
glnthrserseralaargthrglngluileilegluserlysleuglu
213021352140
arglysarglysasnileleuglyalaproglyasnlysargileval
2145215021552160
ilephevalaspaspleuasnmetproargleuaspargtyrglyser
216521702175
glnproproilegluleuleuargglntyrglnasppheglyglyphe
218021852190
tyraspargasnlysleuphetrplysgluileglnaspvalthrile
219522002205
ileseralacysalaproproglyglyglyargasnprovalthrpro
221022152220
argpheilearghisphesermetleucysleuprometproserglu
2225223022352240
hisserleulysglnilepheglnalaileleuasnglypheleuser
224522502255
asppheproproalavallysglnthralaserserilevalgluala
226022652270
servalgluiletyrasnlysmetservalaspleuleuprothrpro
227522802285
alalysserhistyrvalpheasnleuargaspleuserlyscysval
229022952300
glnglyileleuglncysaspproglythrileargglugluilegln
2305231023152320
ilepheargleuphecyshisglucysglnargvalphehisasparg
232523302335
leuileasnasngluasplyshistyrphehisvalileleuthrglu
234023452350
metalaasnlyshispheglyilealaileaspleuglutyrpheleu
235523602365
asnlysproileilepheglyasppheilelyspheglyalaasplys
237023752380
alaaspargiletyraspaspmetproaspileglulysthralaasn
2385239023952400
valleuglnasptyrleuaspasptyrasnleuthrasnprolysglu
240524102415
vallysleuvalphepheglnaspalailegluhisvalserargile
242024252430
alaargmetileargglngluargglyasnalaleuleuvalglyval
243524402445
glyglythrglylysglnserleuthrargleualaalahisilecys
245024552460
glytyrlyscysleuglnilegluleuserargglytyrasntyrasp
2465247024752480
serphehisgluaspleuarglysleutyrlysmetalaglyvalglu
248524902495
asplysasnmetvalpheleuphethraspthrglnilevalvalglu
250025052510
glupheleugluaspileasnasnileleuasnserglygluvalpro
251525202525
asnleupheglulysaspgluleugluglnvalleualaalathrarg
253025352540
proargalalysgluvalglyilesergluglyasnargaspgluval
2545255025552560
pheglntyrpheileserlysvalargglnlysleuhisilevalleu
256525702575
cysmetserprovalglyglualapheargserargcysargmetphe
258025852590
proserleuvalasncyscysthrileasptrpphevalglntrppro
259526002605
argglualaleuleuservalserlysthrphepheserglnvalasp
261026152620
alaglyasnglugluleulysglulysleuproleumetcysvalasn
2625263026352640
valhisleuservalsersermetalagluargtyrtyrasngluleu
264526502655
argargargtyrtyrthrthrprothrsertyrleugluleuileasn
266026652670
leutyrleusermetleuserglulysarglysglnileileserala
267526802685
argaspargvallysasnglyleuthrlysleuleugluthrasnile
269026952700
leuvalasplysmetlysleuaspleuseralaleugluprovalleu
2705271027152720
leualalyssergluaspvalglualaleumetglulysleualaval
272527302735
aspglngluseralaaspglnvalargasnthrvalglngluaspglu
274027452750
alathralalysvallysalaglugluthrglnalailealaaspasp
275527602765
alaglnargaspleuaspglualaleuproalaleuaspalaalaasn
277027752780
lysalaleuaspserleuasplysalaaspilesergluileargval
2785279027952800
phethrlysproproaspleuvalmetthrvalmetglualaileser
280528102815
ileleuleuasnalalysproasptrpproseralalysglnleuleu
282028252830
glyaspserasnpheleulysargleuleuglutyrasplysgluasn
283528402845
ilelysproglnileleualalysleuglnlystyrileasnasnpro
285028552860
aspphevalproglulysvalglulysvalserlysalacyslysser
2865287028752880
metcysmettrpvalargalametaspleutyrserargvalvallys
288528902895
valvalgluprolysargglnlysleuargalaalaglnalagluleu
290029052910
aspilethrmetalathrleuargglulysglnalaleuleuarggln
291529202925
valgluaspglnileglnalaleuglnaspglutyrasplysglyval
293029352940
asnglulysgluserleualalysthrmetalaleuthrlysalaarg
2945295029552960
leuvalargalaglylysleuthralaalaleugluaspgluglnval
296529702975
argtrpglugluserileglnlysphegluglugluileserasnile
298029852990
thrglyasnvalpheilealaalaalacysvalalatyrtyrglyala
299530003005
phethralaglntyrargglnserleuileglucystrpileglnasp
301030153020
cysglnserleugluileproileaspproserpheserleuileasn
3025303030353040
ileleuglyaspprotyrgluileargglntrpasnthraspglyleu
304530503055
proargaspleuileserthrgluasnglyileleuvalthrglngly
306030653070
argargtrpproleumetileaspproglnaspglnalaasnargtrp
307530803085
ileargasnlysgluserlysserglyleulysileilelysleuthr
309030953100
aspserasnpheleuargileleugluasnserileargleuglyleu
3105311031153120
provalleuleuglugluleulysgluthrleuaspproalaleuglu
312531303135
proileleuleulysglnilepheileserglyglyargleuleuile
314031453150
argleuglyaspseraspileasptyrasplysasnpheargphetyr
315531603165
metthrthrlysmetproasnprohistyrleuprogluvalcysile
317031753180
lysvalthrileileasnphethrvalthrlysserglyleugluasp
3185319031953200
glnleuleuseraspvalvalargleuglulysproargleugluglu
320532103215
glnargilelysleuilevalargileasnthrasplysasnglnleu
322032253230
lysthrilegluglulysileleuargmetleuphethrserglugly
323532403245
asnileleuaspasnglugluleuileaspthrleuglnaspserlys
325032553260
ilethrserglyalailelysthrargleugluglualagluserthr
3265327032753280
gluglnmetileasnvalalaargglulystyrargprovalalathr
328532903295
glnglyservalmettyrphevalilealaserleusergluileasp
330033053310
promettyrglntyrserleulystyrphelysglnleupheasnthr
331533203325
thrilegluthrservallysthrgluasnleuglnglnargleuasp
333033353340
valleuleugluglnthrleuleuthralatyrvalasnvalserarg
3345335033553360
glyleuphegluglnhislysleuiletyrserphemetleucysval
336533703375
glumetmetargglnglnglythrleuseraspalaglutrpasnphe
338033853390
pheleuargglyseralaglyleuglulysgluargproprolyspro
339534003405
glualaprotrpleuprothralathrtrpphealacyscysaspleu
341034153420
glugluserpheprovalphehisglyleuthrglnasnileleuser
3425343034353440
hisproileserileargleuglyserphegluthrtyrileasnpro
344534503455
glnlystrpgluglytyrserlysmetlyshisgluasplyshismet
346034653470
argglnglulysglualaalahisglnaspprotrpseralaglyleu
347534803485
serserphehislysleuileleuilelyscyscyslysgluglulys
349034953500
valvalphealaleuthraspphevalilegluasnleuglylysgln
3505351035153520
pheilegluthrproprovalaspleuprothrleutyrglnaspmet
352535303535
sercysasnthrproleuvalpheileleuserthrglyserasppro
354035453550
metglyalapheglnargphealaarggluserglytyrsergluarg
355535603565
valglnserileserleuglyglnglyglnglyproilealaglulys
357035753580
metvallysaspalametlysserglyasntrpvalpheleuglnasn
3585359035953600
cyshisleualavalsertrpmetleualametglugluleuilelys
360536103615
thrphethraspproaspseralailelysaspthrpheargleuphe
362036253630
leusersermetproserasnthrpheprovalthrvalleuglnasn
363536403645
servallysvalthrasngluproprolysglyleuargalaasnile
365036553660
argargalaphethrglumetthrproserphepheglugluasnile
3665367036753680
leuglylyslystrpargglnileilepheglyilecysphephehis
368536903695
alaileileglngluarglyslyspheglyproleuglytrpasnile
370037053710
cystyrglupheasnaspseraspargglucysalaleuleuasnleu
371537203725
lysleutyrcyslysgluglylysileprotrpaspalaleuiletyr
373037353740
ilethrglygluilethrtyrglyglyargvalthraspsertrpasp
3745375037553760
glnargcysleuargthrileleulysargphepheserprogluthr
376537703775
leuglugluasptyrlystyrsergluserglyiletyrphealapro
378037853790
metalaaspserleuglngluphelysasptyrilegluasnleupro
379538003805
leuileaspaspprogluilepheglymethisgluasnalaasnleu
381038153820
valpheglntyrlysgluthrserthrleuileasnthrileleuglu
3825383038353840
valglnproargserserthrglyglygluglylysserasnaspglu
384538503855
ilevalglngluleuvalalaservalglnthrargvalproglulys
386038653870
leuglumetgluglyalasergluserleuphevallysaspleugln
387538803885
glyargleuasnserleuthrthrvalleuglyglngluvalasparg
389038953900
pheasnasnleuleulysleuilehisthrserleugluthrleuasn
3905391039153920
lysalailealaglyphevalvalmetsergluglumetglulysval
392539303935
tyrasnserpheleuasnasnglnvalproalaleutrpserasnthr
394039453950
alatyrproserleulysproleuglysertrpvallysaspleuile
395539603965
leuargthrserphevalaspleutrpleulysargglyglnprolys
397039753980
sertyrtrpileserglyphephepheproglnglypheleuthrgly
3985399039954000
thrleuglnasnhisalaarglystyrasnleuproileaspgluleu
400540104015
serphelystyrservalileprothrtyrargaspglnalaalaval
402040254030
ileglualaalalysthrvalglnpheglyglngluleuprometasp
403540404045
metgluleuproserprogluaspglyvalleuvalhisglymetphe
405040554060
metaspalaserargtrpaspasplysglumetvalilegluaspala
4065407040754080
leuproglyglnmetasnprovalleuprovalvalhispheglupro
408540904095
glnglnasntyrlysproserprothrleutyrhiscysproleutyr
410041054110
lysthrglyalaargalaglythrleuserthrthrglyhisserthr
411541204125
asnphevalvalthrvalleuleuproserlysargserlysasptyr
413041354140
trpilealalysglyseralaleuleucysglnleuserglu
414541504155
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<212>dna
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<212>dna
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<212>dna
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<212>dna
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ccatccattgaaacagattttgtca25