CN111684113B - 水稻绿色基因芯片与应用 - Google Patents

水稻绿色基因芯片与应用 Download PDF

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CN111684113B
CN111684113B CN201880083605.8A CN201880083605A CN111684113B CN 111684113 B CN111684113 B CN 111684113B CN 201880083605 A CN201880083605 A CN 201880083605A CN 111684113 B CN111684113 B CN 111684113B
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陆青
周发松
邱树青
雷昉
谢为博
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Wuhan Shuangluyuan Chuangxin Technology Research Institute Co ltd
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Abstract

本发明公开了水稻绿色基因芯片与应用，芯片命名为GSR40K，是基于Illumina芯片制造技术制作的SNP芯片，包含44263个位点，SNP位点来源于全世界各地的4726份栽培稻品种的重测序结果，每张芯片同时检测24个样品，可用于种质资源遗传多样性分析和亲缘关系分析、功能基因鉴定和基因定位、品种真实性鉴定、育种材料遗传背景分析。

Description

水稻绿色基因芯片与应用
技术领域
本发明涉及分子生物学、基因组学、生物信息学及植物分子育种领域，具体地，涉及水稻绿色基因芯片与应用。
背景技术
SNP标记的开发是基于DNA测序基础上的(Davey等，Genome-wide genetic markerdiscovery and genotyping using next-generation sequencing.Nat Rev Genet.12:499-510.)，自从2005年454测序仪问世以来的十多年时间，二代测序技术不断完善，基因组测序效率大大提高，测序成本大幅度下降，大量物种的全基因组序列完成，极大地推进了功能基因组研究的进展。
水稻作为单子叶植物研究的模式植物，同时也是全球最重要的粮食作物，对不同的品种进行重测序，利用重测序发现的SNP标记，构建高密度的水稻单倍型图谱(HapMap)，利用全基因组关联分析(Genome-wide Association Mapping,GWAS)对重要农艺性状进行关联分析，确定重要农艺性状相关的候选基因位点，建立起一套高效快速、成熟稳定、成本低、通量高的基因型鉴定方法(Huang等，Genome-wide association studies of14agronomic tr aits in rice landraces.Nat Genet.2010,42:961-967)，是分子生物学研究人员优先考虑的方向。
目前多个作物物种中都开发出了SNP基因芯片，水稻的Illumina InfiniumRice6K(Yu等，A whole genome SNP array(RICE6K)for genomic breeding inrice.Plant Biotech J,2014,12:28-37)、Cornell_6K_Array_Infinium_Rice(Thomson等，Large-scale deployment of a rice 6K SNP array for genetics and breedingapplications.Rice(N Y),2017,10:40)、Illu mina GoldenGate(Parida等，SNPs instress-responsive rice genes:validation,genotyping,functional relevance andpopulation structure.BMC Genomics.2012,13:426-443；)、Affyme trix GeneChip Rice44K(Zhao等，Genome-wide association mapping reveals a rich geneti carchitecture of complex traits in Oryza sativa.Nat Commun.2011,13:467)已经广泛的应用于种质资源筛选、品种真实性和纯度鉴定、育种材料遗传背景分析等工作；Ganal等(Alarge maize(Zea mays L.)SNP genotyping array:development and germplasmgenotyping,and genetic mapping to compare with the B73 reference genome.PLosOne.2011,6:e28334)利用Illumina Infinium maize SNP50基因芯片对两个玉米重组自交系群体进行分析，获得了分别包含20913和14524个标记的高密度遗传连锁图；在大豆的驯化历史研究中，Affymetrix Axiom全基因组SNP芯片NJAU 355K SoySNP分析了105个野生和262个栽培品种，得出大豆栽培品种起源于中国的中部和北部的结论(Wang等，Developmentand ap plication of a novel genome-wide SNP array reveals domesticationhistory in soybean.SciRep.2016,6:20728-20737)；为构建小麦的高密度遗传连锁图，以小麦90KInfinium iSelec t SNP芯片对4个小麦群体进行了扫描，将29692个SNP标记定位到了6倍体小麦的21条染色体上(Wen等，A high-density consensus map of commonwheat integrating four ma pping populations scanned by the 90K SNParray.Front Plant Sci.2017,doi:10.3389/fpls.2017.01389)。
发明内容
本发明的目的在于提供了水稻绿色基因芯片，所述的芯片包括SEQ ID No.1～44263所示的共44263个多核苷酸。
本发明的另一个目的在于提供了水稻绿色基因芯片的应用。
为了达到上述目的，本发明采取以下技术措施：
申请人利用4726份水稻品种重测序结果，鉴定出14,541,446个SNP位点和2,855,580个InDel位点(http://ricevarmap.ncpgr.cn/v2/)，并应用LD-KNN对其进行imputation，补缺后过滤掉缺失值比例大于30％的SNP位点，最终筛选得到44263个多核苷酸，将这些序列提交给Illumina公司，利用Infinium芯片制造技术制作SNP芯片，命名为GSR40K；所述的44263个多核苷酸为SEQ ID NO.1～44263所示。
水稻绿色基因芯片的应用，包括利用本发明提供的SNP芯片进行水稻功能基因鉴定或定位；利用本发明提供的SNP芯片进行水稻遗传多样性分析；利用本发明提供的SNP芯片进行水稻品种分析。
与现有技术相比，本发明具有以下优点：
本发明与同类型的中国种子集团有限公司的Rice6K和Rice60K以及华智水稻生物技术有限公司的56K水稻SNP芯片相比具有更好的品种代表性。Rice6K芯片的SNP位点是从520多份水稻地方品种的测序数据中鉴定筛选出来的，Rice60K芯片上的SNP位点来源于730多份水稻品种，56K水稻SNP芯片从全球89个国家和地区代表性的3024份水稻品种全基因组重测序序列信息中挖掘出来，本发明的GSR40K水稻全基因组芯片SNP位点来源于全世界各地的4726份栽培稻品种的重测序结果。
本发明与上述10中的其它芯片相比：进行了进一步优选，优异标记的比例更多，减少了籼-粳差异标记，增加了籼-籼和粳-粳标记，使得不同亚群的分型效能更加合理。
附图说明
图1为一种水稻全基因组育种芯片GSR40K所有SNP位点在全基因组上的分布示意图。
图2为一种水稻全基因组育种芯片GSR40K所有位点籼稻之间、籼粳之间以及粳稻之间多态性SNP频率分布示意图。
图3为一种水稻全基因组育种芯片GSR40K对120个水稻品种进行遗传多样性分析的结果；其中图3中a为日本晴/明恢63每100kb差异标记数全基因组分布情况；
b为日本晴/明恢63每100kb差异标记数统计；
c为9311/Balila每100kb差异标记数全基因组分布情况；
d为9311/Balila每100kb差异标记数统计。
具体实施方式
本发明所述技术方案，如未特别说明，均为本领域的常规方案；所述试剂或材料，如未特别说明，均来源于商业渠道。
实施例1：
水稻全基因组育种芯片GSR40K的获得：
本发明利用4726份水稻品种重测序结果，鉴定出14,541,446个SNP位点和2,855,580个InDel位点(http://ricevarmap.ncpgr.cn/v2/)，并应用LD-KNN对其进行imputation，补缺后过滤掉缺失值比例大于30％的SNP位点，对SNP位点的选择步骤如下：
1)应用Illumina评分系统对所有SNP位点进行评分，过滤分值小于0.6的位点。
2)过滤掉剩余SNP中第二种(按照等位基因频率降序排列)等位基因频率≤5％的位点。
3)判断剩余SNP位点是否存在两种以上基因型，并过滤掉存在两种以上基因型且第三种(按照等位基因频率降序排列)等位基因频率大于2％的位点。
4)判断剩余每个SNP位点上下游50bp中是存在第二等位基因频率大于10％的的SNP/InDe l位点，并过滤掉存在上述情况的SNP位点。
5)判断剩余SNP位点是否处于重复区间中，提取SNP上下游50bp序列，计算该段序列在水稻基因组中出现的次数(相似度≥80％)，过滤出现次数≥2的SNP位点。
6)将水稻全基因组每100kb分为一个区段，计算每区段内SNP两两间的相关系数,以相关系数(R2≥0.60)为阈值将SNP分组。
7)构建一种综合评分策略，对剩余SNP效能进行评价，具体如下：
(1)分别计算SNP在All/All_Indica/All_Japonica/Aus/IndicaI/IndicaII/Indica_Interme diate/Temperate_Japonica/Tropical_Japonica/Japonica_Intermediate/GroupVI-Aromatic/Inter mediate亚群中等位基因频率。构建打分策略Score1＝MAFAll*WAll+MAFAll_Ind*WAll_Ind+MAFAll_Jap*WAll_Jap+MAFIndI*WIndI+MAFIndII*WIndII+MAFInd_Int*WInd_Int+MAF TeJ*WTeJ+MAFTrJ*WTrJ+MAFJap_Int*WJap_Int。
按照每SNP illumina合成探针类型计算Score2＝TypeinfiniumI*0.5+TypeinfiniumII*25。
(2)按照SNP位置计算Score3：SNP位于基因间区时Score3为0；SNP位于启动子区域时，Score3为0.5；SNP位于5`-UTR或3`-UTR时，Score3为1；SNP位于基因上且为同义突变时，Score3为1；SNP位于基因上且为非同义突变时，Score3为4；SNP位于基因上且为大效应突变时，Score3为8。总分值Score＝Score1+Score2+Score3。
(3)按照每个SNP的Score值由大至小对12中每组SNP进行排序，依次选择组中分值最高SNP，进行合并，最终选择出44263个SNP位点。
所有44263个SNP位点为序列表所列的1～44263条DNA序列，将这些序列提交给Illu mina公司，利用Infinium芯片制造技术制作SNP芯片GSR40K。GSR40K所有SNP位点在全基因组上的分布如图1所示，总体上在全基因组均匀分布，同时在重要功能基因区域分布密集，能满足水稻种质资源遗传多样性分析和亲缘关系分析、满足水稻的功能基因鉴定和基因定位、满足水稻品种真实性鉴定和水稻育种材料遗传背景分析的需求。
实施例2：
利用GSRK40K水稻全基因组育种芯片检测水稻样品的方法，包括以下步骤：
1)水稻样品DNA的制备：根据检测需要利用天根生化科技(北京)有限公司的植物基因组DNA抽提试剂盒，按照试剂盒标准流程抽提不同组织、器官或个体的DNA，
2)水稻样品DNA的质量控制：用1％的琼脂糖凝胶电泳，检测基因组DNA的完整性；按照仪器使用说明，用Nanodrop2000分光光度计测定DNA的浓度，判断蛋白质和RNA的污染程度。
3)基因芯片检测：按照Illumina Infinium基因芯片检测标准流程操作。芯片扫描使用Illu mina HiScan芯片扫描仪。
4)基因芯片数据分析：Illumina HiScan扫描结果用Genome Studio软件分析基因型，并用R语言编程获得基因型比较结果。
实施例3：
GSR40K芯片标记分型效果评估
533水稻种质资源(http://ricevarmap.ncpgr.cn/v1/cultivars_information/)中随机选取亚群(IndI 98份、IndII 105份、TeJ 91份、TrJ 44份)品种，利用GSR40K芯片进行分析，比较亚群内和亚群间芯片多态性标记比例。比较数据分为七组，分别为：籼(Ind)粳(Jep)间、籼1群(IndI)和籼2(IndII)群间、温带粳稻群(TeJ)和热带粳稻群(TrJ)间、籼1群内部(IndI)、籼2群(IndII)内部、温带粳稻群(TeJ)内部、热带粳稻群(TrJ)内部。每组随机选取10份共140份数据进行比较，结果见图2。籼粳群间平均多态性标记数约22000个，籼1和籼2平均多态性标记数约14000个，温带粳稻和热带粳稻亚群间平均多态性标记数约10000个，在温带粳稻亚群内部遗传背景极为相似的情况下，多态性标记数仍然达到平均约7000个。Rice6K育种芯片包含5636个SNP标记，其籼稻与粳稻、籼稻与籼稻已经粳稻与粳稻之间多态性标记数分别平均为2600、1000和800，远低于本发明的GSR40K芯片；Rice60K育种芯片包含58290个SNP位点，位点数远高于GSR40K芯片，但其籼稻与粳稻之间、籼稻与籼稻之间以及粳稻与粳稻之间多态性标记数分别平均约为19000、10000和8000个，均低于GSR40K。
实施例4：
GSR40K芯片应用于水稻遗传背景选择：
利用实施例2所述的方法，应用GSR40K芯片对日本晴/明恢63及9311/Ballila群体的亲本日本晴、明恢63以及Ballila进行检测，确定是否可用于对上述两个杂交组合后代群体的背景选择。
分析结果显示日本晴/明恢63每1000kb平均差异标记数为56，9311/Balila每1000kb平均差异标记数为59，相当于约平均每20kb有一个可区分双亲的多态性标记，差异标记在全基因组上分布均匀，可有效满足分子育种中对背景进行精确选择的标记需求(图3)。
利用日本晴、明恢63和Ballila的测序数据，分析Rice6K和Rice60K芯片SNP位点在上述两个群体对应亲本之间的差异标记分布情况，发现Rice6K芯片SNP位点在日本晴/明恢63和9311/Ballila两个杂交组合亲本之间每1000kb的平均差异性标记数分别为7.5和8.0个，Rice60K芯片SNP位点在日本晴/明恢63和9311/Ballila两个杂交组合亲本之间每1000kb的平均差异性标记数分别为49.6和51.8个，均低于本发明的GSR40K芯片。
实施例5：
利用GSR40K芯片进行重要基因功能分析
分子设计育种的策略是将育种总体目标分解成若干个单元目标，根据每个单元设计并培育一批符合单元目标的育种材料，最后按照整体目标将各单元材料具备的基因组合在一起，获得符合总体目标的品种。解析品种中重要功能基因的存在情况尤为重要。
GSR40K芯片可解析品种抗稻瘟病、抗褐飞虱、抗白叶枯等重要农艺性状功能基因，为设计育种提供信息基础。
应用GSR40K芯片检测水稻品种IRBB-60，分析其抗稻瘟病、抗褐飞虱以及抗白叶枯基因情况，结果如下:
Figure GDA0004151989830000051
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同时对上述基因进行Sanger测序，对比芯片分析与测序的结果，发现二者对功能基因的判断结果完全一致，即IRBB-60含有白叶枯抗性基因xa5，xa13，Xa21，说明GSR40K芯片可有效进行重要功能基因等位基因的分析。
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Figure GDA0004151989830006381
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Figure GDA0004151989830012001
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Figure GDA0004151989830012011
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Figure GDA0004151989830017701
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Figure GDA0004151989830023321
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Figure GDA0004151989830024761
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Figure GDA0004151989830024771
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Figure GDA0004151989830024791
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Figure GDA0004151989830024881
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Figure GDA0004151989830024891
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Figure GDA0004151989830028661
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Figure GDA0004151989830028671
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Figure GDA0004151989830028681
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Figure GDA0004151989830028751
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Figure GDA0004151989830028761
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Figure GDA0004151989830028771
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Figure GDA0004151989830028781
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Figure GDA0004151989830028791
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Figure GDA0004151989830028841
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Figure GDA0004151989830028851
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Figure GDA0004151989830028861
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Figure GDA0004151989830028871
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Figure GDA0004151989830028881
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Figure GDA0004151989830028891
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Figure GDA0004151989830029001
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Figure GDA0004151989830029011
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Figure GDA0004151989830034651
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Figure GDA0004151989830040321
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Figure GDA0004151989830040331
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Figure GDA0004151989830045891
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Figure GDA0004151989830056881
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Figure GDA0004151989830062411
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Figure GDA0004151989830073391
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Figure GDA0004151989830084371
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Figure GDA0004151989830089821
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Figure GDA0004151989830095311
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Figure GDA0004151989830100791
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Figure GDA0004151989830106291
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Figure GDA0004151989830111741
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Figure GDA0004151989830117231
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Figure GDA0004151989830122691
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Figure GDA0004151989830128211
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Figure GDA0004151989830133671
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Claims (4)

1.一种水稻绿色基因芯片，所述的芯片包括SEQ ID No.1～44263所示的共44263个多核苷酸。
2.权利要求1所述的芯片在进行水稻功能基因鉴定或定位中的应用。
3.权利要求1所述的芯片在进行水稻遗传多样性分析中的应用。
4.权利要求1所述的芯片在进行水稻品种分析中的应用。
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CN114395639B (zh) * 2021-12-31 2023-06-09 华智生物技术有限公司 用于鉴定水稻品系纯度的snp分子标记组合及其应用

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Publication number Priority date Publication date Assignee Title
CN102747138A (zh) * 2012-03-05 2012-10-24 中国种子集团有限公司 一种水稻全基因组snp芯片及其应用
CN105008599A (zh) * 2013-02-07 2015-10-28 中国种子集团有限公司 水稻全基因组育种芯片及其应用

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CN110050092B (zh) * 2016-12-08 2023-01-03 中国种子集团有限公司 水稻全基因组育种芯片及其应用

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Publication number Priority date Publication date Assignee Title
CN102747138A (zh) * 2012-03-05 2012-10-24 中国种子集团有限公司 一种水稻全基因组snp芯片及其应用
CN105008599A (zh) * 2013-02-07 2015-10-28 中国种子集团有限公司 水稻全基因组育种芯片及其应用

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