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作物学报

作物学报 ›› 2021, Vol. 47 ›› Issue (10): 2036-2044.doi: 10.3724/SP.J.1006.2021.02082

• 研究简报 • 上一篇    下一篇

结合QTL-seq和连锁分析发掘水稻中胚轴伸长相关QTL

刘畅1(), 孟云1, 刘金栋1, 王雅美1,*(), Guoyou Ye1,2   

  1. 1中国农业科学院深圳农业基因组研究所, 广东深圳 518120
    2国际水稻研究所, 菲律宾马尼拉 DAPO Box 7777
  • 收稿日期:2020-11-30 接受日期:2021-03-23 出版日期:2021-10-12 网络出版日期:2021-04-07
  • 通讯作者: 王雅美
  • 作者简介:E-mail: woshiyonghwa@163.com
  • 基金资助:
    中国博士后科学基金项目(2020M68299);中国农业科学院科技创新工程协同创新任务(CAAS-XTCX2016001)

Combining QTL-seq and linkage analysis to identify the QTL of mesocotyl elongation in rice (Oryza sativa L.)

LIU Chang1(), MENG Yun1, LIU Jin-Dong1, WANG Ya-Mei1,*(), Guoyou Ye1,2   

  1. 1Agricultural Genomics Institute in Shenzhen, Chinese Academy of Agricultural Sciences, Shenzhen 518120, Guangdong, China
    2International Rice Research Institute, Metro Manila DAPO Box 7777, Philippines
  • Received:2020-11-30 Accepted:2021-03-23 Published:2021-10-12 Published online:2021-04-07
  • Contact: WANG Ya-Mei
  • Supported by:
    China Postdoctoral Science Foundation(2020M68299);Agricultural Science and Technology Innovation Program Cooperation and Innovation Mission of the Chinese Academy of Agricultural Sciences(CAAS-XTCX2016001)

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摘要：

中胚轴长度(mesocotyl length, ML)是影响旱直播水稻出苗和早期幼苗活力的重要性状。发掘中胚轴伸长相关位点, 解析其遗传机制, 选育长中胚轴品种是促进旱直播技术推广最为经济和有效的方式。本研究以长中胚轴品种‘Changai’和短中胚轴品种‘IR 145’为亲本构建的F2遗传分离群体为材料, 构建长池和短池并开展深度重测序(50×)。利用Δ(SNP-index)和G-value两种方法在3号染色体29.56~33.28 Mb处鉴定到1个中胚轴伸长相关位点qML3。在候选区域开发KASP标记, 对184个F2株系开展连锁分析, 将候选区间缩小到28.89~31.03 Mb。结合基因注释、连锁分析和基因表达分析结果, 推测LOC_Os03g52450、LOC_Os03g56060、LOC_Os03g58290、LOC_Os03g58300、LOC_Os03g58320、LOC_Os03g56050和LOC_Os03g57640为候选基因。这些基因分别与植物激素的调控和细胞分裂相关机制有关。本研究发掘了一个水稻中胚轴伸长相关位点, 对选育长中胚轴品种有一定帮助。

关键词: 中胚轴, 混合分组分析法, Δ(SNP-index), 连锁分析, 候选基因

Abstract:

Mesocotyl length is an important trait, which affects seedling establishment and early seedling vigor after dry direct seeding. Identifying the loci related to mesocotyl elongation, analyzing the genetic mechanism and selecting varieties with long mesocotyl is the most economic and effective way to promote the popularization of dry direct seeding technology. In this study, an F2 population constructed with ‘Changai’ (an extremely long mesocotyl variety) and ‘IR 145’ (an extremely short mesocotyl variety) was used to construct long and short DNA pool, which were re-sequenced at 50×level. Two methods, ΔSNP-index and G-value, were used to identify the quantitative trait loci (QTL) for mesocotyl elongation. A QTL named as qML3, located in the region of 29.56-33.28 Mb on chromosome 3 were detected. After linkage analysis of 184 F2 lines with the newly developed KASP markers based on this locus, the region was narrowed down to 28.89-31.03 Mb. Combining the results of gene annotation, linkage analysis, and gene expression analysis, LOC_Os03g52450, LOC_Os03g56060, LOC_Os03g58290, LOC_Os03g58300, LOC_Os03g58320, LOC_Os03g56050, and LOC_Os03g57640 were speculated to be candidate genes in this region. These genes were involved in the regulation of plant hormones as well as the mechanism of cell division. This study uncovers a locus related to rice mesocotyl elongation and is helpful for the breeding of long mesocotyl varieties.

Key words: mesocotyl, bulked segregant analysis, Δ(SNP-index), linkage analysis, candidate gene

附表1

用于连锁分析KASP标记详细信息"

Marker Chromosome position Seq C Seq X Seq Y Allele X Allele Y Changai IR 145 Long pool Short pool Sequence
OS900153_K01 1 11341599 TTTCCCTTTCACTCCAAAGAAACGA GAAGGTGACCAAGTTCATGCTTATTGTGGAAATCGCTTGACCCTCC GAAGGTCGGAGTCAACGGATTTATTGTGGAAATCGCTTGACCCTCT G A GG AA AA GG GGTGATGGCGATGCGCGGGGTCGACGGATCGGCAATGGTGAGGCGCGGGGTCAGCGGAATGACGAGGAGTGGGGGTGGCGATCAACGACGGTGAGGCGGATCCATCAGCCATATACATGTGCACACCATGTGCATGGAATCTACACCAATGTTTTCGATTTTTACTGACCTTTCCCTTTCACTCCAAAGAAACGATCTTC[G/A]GAGGGTCAAGCGATTTCCACAATAGTCTCGGTTCTAATAAGAACCGGGACTAAAGATTGTTTTTAGTCCCGGTTAAAAAAAATTTGATATTTAGTCCTGGTTCGTGATCTTTAGTCCTGATTCATCTCCTGTCAGAAGCCTGCGAGGGGGCCGAGGATCTTTAGTCCCGGTTGATTACCACTTTAGTCCCGGATGGTACT
OS900154_K01 2 8278191 TGGCACAGCCTAACAATATGATGGA GAAGGTGACCAAGTTCATGCTAATTGCTCCAGCTTCTGTGTTGTGC GAAGGTCGGAGTCAACGGATTATTGCTCCAGCTTCTGTGTTGTGA G T GG TT TT GG TAGCAAGTTAATCTGGACAATGCAGGTTAGATCTGAAATCATTTGCTCTACAGTTTCATAATGCATATAGTTCCTTATGGTATAATATCAGCTACCAACCTGCCAATATGTTCTACAATCTCATAATGCATACAATGTCTCACAATATCAGCTGATGGATGGCACAGCCTAACAATATGATGGAAATTGTGTGGTCAGAA[G/T]CACAACACAGAAGCTGGAGCAATTTTCCTTCCACTTCTACCTTGACAGAGAAGTTCTGAAGACTGATTTTCATTCTCTGATTCCACCAGCCACTATGACAGAGCATACCTTGCTTCCACTTTTTTCCTTGATGAGGTTGTAGTGTTCCCTTCCCCATATTTTTTTGTTGTTCAACTTAAGATTCTCGCCGTTAAAAAATT
OS900155_K01 2 11054736 CCAGAAGGTTCGTTACTTCCATGTCC GAAGGTGACCAAGTTCATGCTTGAATCCATTTATGTTAGGTGCCATA GAAGGTCGGAGTCAACGGATTTGAATCCATTTATGTTAGGTGCCATC A C AA CC CC AA ACCCCTTTTTAGTACAATTTAGAGCGCACTCACCACCATGTGTAAGCACATGCATGTCCTCTACATACTCTTGAAATAGAGGCATCGAAATCTTATAGAGTATGGTGGACATGCTAAATCACCTACTGAGTACATAATCATAAAATTGGTGGACACTAGTGCTAGATCTAGAATCTGAATCCATTTATGTTAGGTGCCAT[A/C]TCTTGTCGACCAGCTTCTGAATCTATGCTACAATAATTATTTTTATAAGGGACATGGAAGTAACGAACCTTCTGGAGTTGATTTTGTGAAAAATTTTGAACTCTAGAATTGGGTAAAATTCAGAACGGAGGGAGTATGTAAGAAATACGATAAAAAGTGTTAAAAACAACCAACGAGAATAGAAACAAGTCAGTTCAGAT
OS900156_K01 2 30473912 CGCAGGCACAACCCAGTTTACATA GAAGGTGACCAAGTTCATGCTCACTCCATAAAGGAAACAAAATGACTTGTA GAAGGTCGGAGTCAACGGATTTCCATAAAGGAAACAAAATGACTTGTG T C TT CC CC TT GCCAACATGCTGTCGATCCTGCATGTTAGCGGACAATGCTTACAGCAGTGATTTTATGAATGTGATTATCATGAACACTTTTGATCGTCCCGGTAAGTTTTCAGATACTTGTCCAGTTATCTACATGCAAGAAAATAATTAAGTATGTACTGATTACTCAATGTACATGGGTGTGGCGCAGGCACAACCCAGTTTACATA[T/C]ACAAGTCATTTTGTTTCCTTTATGGAGTGTTTCTACAAAACCATGAACACCAATGTTTACACAAGTACAATCGCCTACTTTCTCTCCGGTCGATGGAACAGAGATAAAAGAGATCCAAAAGCACCGAGCCAAACACCAACAAAACCGAAAAAAAAAAGGAAGAGAAAAACTTTTGAATTGAACAACGTTATACTACTTGA
OS900157_K01 5 3752390 GGCACATTACCAATGCAGGACATTT GAAGGTGACCAAGTTCATGCTACACTTTCTACAGTTGGCCTTCAGC GAAGGTCGGAGTCAACGGATTACACTTTCTACAGTTGGCCTTCAGG G C GG CC CC GG AAATCCCTACGTTTGAAACTGATTCTGCCGACAATATTAGCACCTATAGAACCATGTTCTCAATAGTGTAATATAACAATGTATAGTAGTAAAAAAAAATCAGTTTAGGCACATTACCAATGCAGGACATTTTACATGTTCAGGTGTCATTATTTAAAAAGTATTCATTTTACATATAGTAGTAGTGCTTTCGATGCATC[G/C]CTGAAGGCCAACTGTAGAAAGTGTACACTATCGTTAAAATGCCTGCTCGTGATAGTATAACTTCTAAACCTATAAAGTGATGGGGATTTTGGTGCATTCGCGTTTGGGAGGGGAGGAACCGGGATTTAAATTTGGATTTTGCAGTGAATAAAGATGTACTAATTATTGGACGAGGCATTACCAACGATCCATCCAATCTG
OS900158_K01 5 6521522 GCGTAAACCTCGGTGAATTCCTCAC GAAGGTGACCAAGTTCATGCTGACCAACAACAATATTAGCATAGCACAAAA GAAGGTCGGAGTCAACGGATTGACCAACAACAATATTAGCATAGCACAAAG T C CC TT TT CC TCTCCGCAAGGAAAGAGATACCCTTAGTGGAAGGAGGAGTGCATTTGTGCAACCTCACGAGGATAGGGTTGAAAAAGACCCGGCTCTTTGCGAGCCCCTCAACGGAGACTAGAATCCCCTTAAGGATTTGAACTCCGGGAAATACATCGGCGCGTAAACCTCGGTGAATTCCTCACTCTCTTTACCTTCTTGTTCCATTA[C/T]TTTGTGCTATGCTAATATTGTTGTTGGTCGAGCTCACCTCTTTACTTGTGGATTGAATCTAGAGCTCTTAGGTTGTTCTCTTGTTTATATTGTTGTTGTGCAGGTCAGATAGTCTAATATCAGTTCAGACTATCCGATCACATAGGACTATCCGACCTGATATTAGACAATCTGACCCTGTTTTGTTATAGCTTCCGCAT
OS900159_K01 5 14053490 TGAAATCGTGTTAGCCGACAAGTCA GAAGGTGACCAAGTTCATGCTGCATCTCCCAAAGAGCTAAATCGG GAAGGTCGGAGTCAACGGATTGCATCTCCCAAAGAGCTAAATCGA C T CC TT TT CC GGAGCTGACAGTCGGCAGCAAGACTAACCCTACAACATTTTTTGTCTTCGATGGGAAGGGTTCCTACAGCTTATTGCTTGGAAGGGATTGGATCCATGCCAATTGTTGTATTCCTTCAACCATGCATCAATGCTTGATTCAGTGGCAAGACGACAAGATTGAAATCGTGTTAGCCGACAAGTCAGTCAACGTTGCTAGTG[C/T]CGATTTAGCTCTTTGGGAGATGCCTGGTCTCGATTGCTTATCTGGAAAAGTTTGGGATGGAGATTTTCTAAAAGTGTCCAATTGTGATATACAACCAATTGAAGATGGAGAGCCCAAGCTTTTATTTTGAGGGTGTCGTGGAGGGTTCAAATGTCTACACCAAGGACACAGTAGATGATCTCAATGACAAGCAAGGACAG
OS900160_K01 5 21516100 CCATGGCAGTGGGTGTTGAGATTAT GAAGGTGACCAAGTTCATGCTAATGCGGTCATGCTAGTTTTCATCTA GAAGGTCGGAGTCAACGGATTATGCGGTCATGCTAGTTTTCATCTG T C CC TT TT CC GTCAAAGGAAAATTTGCAGATGCTATTTGCTGTTCGCAAACCGAATGAACGAGAGCGATGAGCTATAAGGCCATGCCCAATGCACCACGGTGGCCAGTGGTCTCAAGCATCCACATCGGAGCTGAGCTTCCATGGCAGTGGGTGTTGAGATTATAGGCATATAATGATTTAATCTATTCCATAAATAAATCATGACATTA[C/T]AGATGAAAACTAGCATGACCGCATTATTAGATCTATACATGTAAACTACACAATATAATATGAACAGATCAAATATACGCAATATATTGAACACGTACCGAGGTGGCGGAAAGACCGGCTGCTCGGTCGAAAATTTTCGCAGGCGTCTACGAAGGCACGCAGCACGCGAGCGAAGAGGAAGATGAACCGTCGCGGACGAA
OS900161_K01 5 24914119 AGGGCACTCACACTGCGTCACTAAT GAAGGTGACCAAGTTCATGCTGTCTAGGTGGCAGTTTTGGCTTCTT GAAGGTCGGAGTCAACGGATTGTCTAGGTGGCAGTTTTGGCTTCTC T C CC TT TT CC GAGGGGGAGAGGGGTGCCGGCGGTGGGTGGGAGAGAGAGGGGCACCAGCGGCGCAGTAGAGGAGCGGGAGCGGCGGGGAAAGGGAGAGAGAGAGAGGAGAGAGAGCGGGTGGGACCCACTGTAAGGGAAAAAGTAGACCACATGCATTGTGGAGGTTGTTGCTTACGAAGTTATTAGTCTAGGTGGCAGTTTTGGCTTCT[C/T]AAAAATTAGTGACGCAGTGTGAGTGCCCTGATAAAATCTGGCATACATGACTCAGTTGCAAGCTACTTGAAACAAGAAAATAAGAGTCGATTCTGCCTATACCGATCGTAATGACAAGACAGACAAGTCAAACACCTTGCATATAGAGGTGTACTTAGGATTTATGGTTCTAGAAAATATCATATTATATTCTAGATTAA
OS900162_K01 5 28105720 GAATTCAAATTTTCATGTTTGTGGAG GAAGGTGACCAAGTTCATGCTCATCCCTCGATATCTTCTCGTTTGG GAAGGTCGGAGTCAACGGATTCATCCCTCGATATCTTCTCGTTTGA G A GG AA AA GG TTCAGAGCCTTTTCGCGCACTTGAATGCATATGCATGAGTTAATTTGCACACTGTGGACGCAGGGGAACACAAGAACAATGGCATGCATGCATACGTTTCGTGTAGGGCAGCTGCTGGCTAGCTTCTCCCGTACAGTGTGCTCGTGCGTGTGTGCTGCTAGCTGGGAAAGTGAAATCATCCCTCGATATCTTCTCGTTTG[G/A]TGCATGTCACCTAAATAGTCAAAAAATTATTAAAAAATTATCAACATATATTAATATGAAATATATTACTCCACAAACATGAAAATTTGAATTCAATTTCTACAAGTTGTAACAAAAATAACAAATATAGTCGAATGTGTGATAACTGTTTTCAGTTTAATTTGTTATTATTGTAGCAACTTGTAGAAGTTGAATTTGGC
OS900163_K01 7 12402368 CTCAACCTGCCTACGCAACTCTTTC GAAGGTGACCAAGTTCATGCTGCAGCATGCAAGTCGAAGGTT GAAGGTCGGAGTCAACGGATTGCAGCATGCAAGTCGAAGGTC T C CC TT TT CC ACACCTGATTAGGGCGTTCGTACACATCCAACTGGAGAAGATCCCTGTTGTCTATTCCTAAAGAGGTATACCATGAAAGCAAACAGTGACAAGTGTGATCGTGAGACTGCAGGACCGGATGGGGTGCATGAAAATTACATAACAAACATGATGATGATTGAAGCATTTGGAGTGATGAGGGCAGCATGCAAGTCGAAGGT[C/T]GCATATGAAAGAGCTATGGTTATTTTGAAAGAGTTGCGTAGGCAGGTTGAGGAGGTACCACAAGATATTCAAGTTGTTGTTTGTGTTCTGTCTCTCAACCGCTCAAGACAAGGAAGGACAGTGTCTCGTGGGAATCAGGGGTAAGGGTAAGCCACCGTGCCAAATTTATTTGATTTGAGAGAGAGTACACATATCTTCTT
OS900164_K01 8 28356273 AACAGGCCGAGGCTCTTCACCTACT GAAGGTGACCAAGTTCATGCTACTCCCTAGTTCCCAAACACTCTCA GAAGGTCGGAGTCAACGGATTACTCCCTAGTTCCCAAACACTCTCT A T AA TT TT AA ATATATATATATATATATATATATATATATATATATATATATACATATATATATATATATTTGTTGGATTAATTAGTAGTAGTTGATTTATGGGGTGTTTAGGAGAGAGGGGCTAAACTTAAAAATATAAGTCTCTAGAGTAGGGACTTATATTTTTAAAAAAAGGGCTAAAGTTTACTCCCTAGTTCCCAAACACTCTC[A/T]TAGATTAGCTAGTCGATCGATCGGCTTACTGTCCAGTAGGTGAAGAGCCTCGGCCTGTTGTCCCCATACAGCTCAGGGCTGACCTGACTACAAGGAAGGAAACATGCAAATGCTTCAGATTCAGATTTAGATCCAATTATATATATGCTCAGCTCTGCTCAATACTACTACTAGTATGAATTAATACTCCTAGGAGTATA
OS900165_K01 9 19464447 ACGGGATATGTGGTGGGACCAG GAAGGTGACCAAGTTCATGCTTGTATAAATTGCGAGAATGCTCAACG GAAGGTCGGAGTCAACGGATTTGTATAAATTGCGAGAATGCTCAACT C T CC TT TT CC CGAATACACTAGGTGAAACATTTTTTTATGAAAAAAAACAAACCAAATTTCCTTAATAGATGGTTTCCAAAATATGTAGGTGATTTGTTTTATATGGATAAGTGACAGGTGGGGACACGGGATATGTGGTGGGACCAGGTGGGCACGTGATGGGGGGAGGGGGGAGCGACCGATTTTTTTCCCCTCCCCCGCGCGCCCGA[C/T]GTTGAGCATTCTCGCAATTTATACAAGTTGAATTTTCAACCTGTCTGTCTGTGGAACGCTATATTTCGTATTAATCATCTTATTAATTTTGGTCAACTTACAATAAACGAGTGGATTGTTCATCAATCCCTCTCCGGTGCACATGCTATTTAGATTCCGACGTGGACGGTGCACGTCCAGTCGTCCACCACCACCACCAC
OS900166_K01 10 180016 CGATAACCTAGGATGATGCACTGCTG GAAGGTGACCAAGTTCATGCTAAGGCTTGCTCGGCAGTTTTA GAAGGTCGGAGTCAACGGATTGTAAGGCTTGCTCGGCAGTTTTT A T TT AA AA TT CGCCTCATCATCCAATTCTATGTGGCATTTTTAGGGCAACACATCAAGTATCATATAGTACTTTGCCCTTAGTTGCTATCCTATTGGATGTATATGCCAGAGTACCATATCATTATCTTGAAATTGGGATGTCTATAAAAGTAGATGGTGTTTCTCATGTCTGACACCTATTTCCCTGGTAAGGCTTGCTCGGCAGTTTT[T/A]CAGCAGTGCATCATCCTAGGTTATCGAACTGTCGACAATTCTGAAAATAATTGATCAGCTATGGATCATGTGTTTTTCTGTGCGTGCGGTCTTGCGATAGCTTCTTTATACGTTTGATCATAACCTATATAAGATGTTAGAATGAAGGAATTCAGTATGATTAGTGACATTGCAGTTGAATTGTAATAAATAAAGATGCA
OS900167_K01 10 4232779 TCGAGGTTCGACACGTTTATAGCAA GAAGGTGACCAAGTTCATGCTGTTCTCTTCAGGATGCTTGTCGTGT GAAGGTCGGAGTCAACGGATTGTTCTCTTCAGGATGCTTGTCGTGC T C CC TT TT CC ACCCAAGAAGTCAATAAGTGGAGTTCGCCAGTCTTTCATTGTCTTGATGTCAACAATGACATGTTCGACCTCTTGAGCCCCCGAGGAGTTGTCGGTATTGGCCATGGTACCTTCGTCGTGCGACTCTTTTACAGACGGTTTTGTCAGGACATCTAAGAAAGTTCCTAGTTCAAGAGGTTCTCTTCAGGATGCTTGTCGTG[C/T]TAAGTCATCTGGTTCAATGTTATCCTTGCTATAAACGTGTCGAACCTCGATGCCGTCAAACTTCTGCTCTAGCTTCTTGACTTCAAAGAGGTACTTAGCCAACTCAGGATTGGAGCATTTGTAATCCTTATCCACTTGGTTTGCGACTAATTTTGAGTCCCCTCTCACAATCAATCGTTTAACTCCTAGTGCAACGGCTG
OS900168_K01 12 2919839 TCGTTTCGCAATCAGTAGGAGGAAA GAAGGTGACCAAGTTCATGCTGGGTGGTGTCAGATTCACTACTCATT GAAGGTCGGAGTCAACGGATTGGGTGGTGTCAGATTCACTACTCATC T C CC TT TT CC CCACCATACACTTTAAAGCCTAAAAATTAAAATAAAATTATTCTCAATGTAAAATATTCAGGTCATATTATTTGTCATTATTTTTACTTTACTTCAAAAGAAATTGATGGTTCGAGGTTAGGGAAATAACAAAAATCCACTTAACTTTTCGGAGAATCCATATGACAGTGGTGATGGGTGGTGTCAGATTCACTACTCAT[C/T]ACCACTACGTACTATTATAGAAAAGCACTTCAAGTAGTACCTACATTTTTCAAGTTGTAGCTAGATCTTTTTACGAATTTGAACTTTCCTCCTACTGATTGCGAAACGAATTAACCGGCCACCGTACAAACTTTTCATGGTACAGGCATGCATGCTTCTTGATCCTTTGCATTTCAAATCATGGTGTTTATGCCGTGTAC
OS900169_K01 12 7367534 GCTAGCACATATGACTGATGCATTACA GAAGGTGACCAAGTTCATGCTGAGTAGCTGGTGTATTTTTCAGTGATT GAAGGTCGGAGTCAACGGATTGAGTAGCTGGTGTATTTTTCAGTGATA T A TT AA AA TT TTTAATTAAATCTAACATTGAATATATTTAAATAGTATGTTTGTTTCGCATTACAAATATTATTATGTAATTTTTAATCTCGGTCAAACTTAAAAAAATAACTTAGAAAAAAATCAAAGAATTACCATAATTTGGCTAATAATTAGTTTTATTTTAGTATGGTCAAACTATTTAGAGTAGCTGGTGTATTTTTCAGTGAT[T/A]CTCTTGTAGAGTAGGATTGTAATGCATCAGTCATATGTGCTAGCTATACTATTGAATAATTGAATGTGGACCGATGTAGTTTATAGCATTGTAAGGGTTATGGTTTACGTCATATATCACATGAAGGAAATGAAGATGACGCAAACCAAGTTGATGGATTGCGTCTTAGCAAGAACGACCAAACTTATCATGGTCCATCC
OS900170_K01 12 27253671 TATAACAGCGACCACCCAGATGACC GAAGGTGACCAAGTTCATGCTATCACGTCCCCCGTTGTCA GAAGGTCGGAGTCAACGGATTATCACGTCCCCCGTTGTCG T C CC TT TT CC CAGAGGGAGGTTCGCGACCAAGGTAGCAGAACGGTGAGGGACGCCATTGGCCGAGACGGCGGAAGAAGGCAAGAGAAGGGGCACCGCGACAGGCGCCGGGATGACCGCGATGACCGTCGCGGGCGCAACAGGAGGTATAACAGCGACCACCCAGATGACCTCCGACGTGGAGGACGGCGCGACGACGACAATGATGATCG[C/T]GACAACGGGGGACGTGATGTCGCGCACGGAAGGGGCGGCGTTCGCGGCCCAGACGTCAACTTCAGGCGGGAACGCACGCACTCGCCGCGTGCCCGCGATCGCGGCGACACGGGAAGACGCGACGGGCAACGCCATCAAGCAAACTGCGGCGGCAAGGACGACGGGGCCAACGACCGCGACATGGGCAACCCACCCCCAGC
OS900129_K01 3 28890281 GCTCAGATTACTGGCTGTGTCA GAAGGTGACCAAGTTCATGCTCATCGATCGGTGTCATTCA GAAGGTCGGAGTCAACGGATTCCATCGATCGGTGTCATTCT A T TT AA AT AT AAATTGATCATTATGATTTTTTAAACAACTTTCGTATAGAAGCTTTTTACAAAAAGTATACCGTTTAGTAGTTTGAAAAACGTACGCGCGGAATACGAGTGAGAGAGGTTGGGAACAAGAAGCAACAAACATAGTACGCGTCCTATACCCCGGTGACCTCTATATATACCGTACCCCATCGCCATCGATCGGTGTCATTC[A/T]TTCTAAGCTTGACACAGCCAGTAATCTGAGCTAAATTAGTCCTAAACTCCAAGTTGATTATCCAACGAATCGTAGCCTCGAGGGGGTATCTGAATCTCCAACCACTTCAATTCAACGCGTCGCCATCGCCATCTTCGTGTAGAGGCGACCTCACCTACGCGCGCGGCGCGCCCCTGGCCATGCAGACGATAAGGTGGTCG
OS900130_K01 3 29290258 TGCATTATCCTCGAAAAACAA GAAGGTGACCAAGTTCATGCTAAAATTGTAATTGGTTGGGAAGAA GAAGGTCGGAGTCAACGGATTAAAATTGTAATTGGTTGGGAAGAG T C CC TT TC TC GAGAGTCCACATAATGTGTAAAAAATGTGCATGTTCCCTGCAAACTTAAGGTTCAACTTTACTTATGCGAAATTGCGGATAGGAAAAAAAAACTGTTTCTACGGATCGCAAGACTATGCATTATCCTCGAAAAACAATTTATACCTTGCATTGTAGCATTTAGAATTTGTTCCAACATATAACAACTTTATATTTAAAAT[C/T]TCTTCCCAACCAATTACAATTTTCCACCATTTATTTATTTTTTCATCCACTTCCTCTCTCCCATGTTTACTTACATATGTTTTTGATAACCATGTCCAATCTCAAAACTTCTTATCTTTTAAAACGGAGGTAGCAAGGCACATTCTTCTCATGATGTTAGATCGTCTTTCCTCATGTAAACAAGTACCATGGACAGTATC
OS900131_K01 3 29692468 GCACAGACATCCTCGTGTCA GAAGGTGACCAAGTTCATGCTAATCTTTAAATAGTTTCTTCTTTTCTTCTTCT GAAGGTCGGAGTCAACGGATTTCTTTAAATAGTTTCTTCTTTTCTTCTTCC T C CC TT TT CT CGCAGGTCCGGGATGCCGCCGACCATATCGTCCACAGCTTCGAGGGTTCGGCCCCTCGGCTCGCGTTCGCCCTCAACTCCGACGAGGAGGGCGATGACGGCGGTGTGGGCGACAGTGGCGACGAGGCTGGCGACCCGGGTGCATCGGAGTGAGCCCCCAGGCCCCCGCCAATCTTTAAATAGTTTCTTCTTTTCTTCTTC[C/T]GAGGCCTTCGGGCCCCCTTCTTGTATAGACTAATTTAATCTGCAATCAAATAAAGAGGAAATTTTTTGTGTCAATTCTATTTGTTGTGTGTATGACACGAGGATGTCTGTGCCGCGGTCCTTTTGTGTCTTGGCTTGAACAAGGGCTCGTGCCCAGGTCCTAGTCCTCAAATGGCTCGGGTCGGGGCTAGTGCCTGGGGA
OS900132_K01 3 30106408 CCACTAGCGAGTAGCGAAGC GAAGGTGACCAAGTTCATGCTATGGCCATGGTGAGCGTT GAAGGTCGGAGTCAACGGATTATGGCCATGGTGAGCGTC T C TT CC TC CC TTATCGAATCGCCACTGAAGTTGATGAGCTGGCTCCTCCCGTCGTCGTCGGTCCAAAGAGATAAGACGGAGACCTCACTCCACACGTCGAAATTGGCAGAGCTCAGATTGGGCGTCACCGCACAGGGGAGCGAGAGTGGAGGGAGGCATGGGGATCATGACCTTGTACTCCCACTCGAGGCGGATGGCCATGGTGAGCGT[C/T]TGCGACTGCAGCTTGTACGGATGCTTCGCTACTCGCTAGTGGCTTGAGGGTGCGGTAGCCGAGCATAACTGATGTGACCATGGCTACCACGGATGCAACCATTGCTCACATCATGGATGAACAAGAAGATATCAAGATCAAGTACTCCAAGTGCTAGAATCCGATTCGGCCACCTACTACACTGCTGACTTCAAACGGCC
OS900133_K01 3 30372501 GCTCCACCTTGAAAGCAATG GAAGGTGACCAAGTTCATGCTTTTCTAGCGTGGTGGAAGAGATA GAAGGTCGGAGTCAACGGATTTTCTAGCGTGGTGGAAGAGATT A T TT AA AT AT ACTTGTTGTCCATTTCCATCGTATGTTGTCTTCAATGTCTTGGGTTAGTGTGATGCCTGAGGTCCAGATTATCTCGAAGACCTTGAAGAAATCGGTGACGAGGACAGTTGTCAAGTTGTATCGGATGTCATCGATCCACTTGTTATCTTGTAGGGCTTCTGCGACCGTCCTTTTCTTTTTTCTAGCGTGGTGGAAGAGAT[T/A]GGGTGCCATTGCTTTCAAGGTGGAGCTGCAAGCCCATGGGGATTCCCAGAAGAGCGCCTTCCTACCATCTCCGATTGTCACTGACGTGGATGCTGCAAAGAGCTCGTTGTCCTGTTCATCACAGGGTGTTGGCATGCCGATCCATGGCTTGTCGGGGGAGGTCCATTCATACCATAGCCATCGCAGGCGAAGCGCCCTTC
OS900134_K01 3 30668336 GCCTTGGTAGCTCGGAAAA GAAGGTGACCAAGTTCATGCTAGATTCGATGTAAAGTTGAGGGAC GAAGGTCGGAGTCAACGGATTAGATTCGATGTAAAGTTGAGGGAT G A GG AA GA AA GGGACTACCCTTAGCCCGCTGTACCTGCTCTTTCCTTGCCCTGCCCTGCGCGGCCTTCTTTTCCTCCCAGCCCCAAGCTCTTCTACGCGGTCCGTTTACTTGCTTGGGATCCTCCCAGGCTCCCAGGCCATGTCTCATCTGTCCGACTGTCCGCGCCCTAATCGCCTTGGTAGCTCGGAAAAAAAATAAGTTCAGATACC[A/G]TCCCTCAACTTTACATCGAATCTGTATGATATCCCCAATATCAGATAGGGTTAATTGGATCCATGCTATTATAACTTTACCACTTTTCCCAAAAAAAAAAAACTTTACCACTTTTGATAAATGCTACTACTATTTGTTGAATTGCAAACATGCCATTATAAATCAGCTAATTTTGGAACCATGCCACTCTCTACATCCTG
OS900135_K01 3 31027370 ACTAGCCCCACACAGACCAC GAAGGTGACCAAGTTCATGCTGGATTTTATGGGCTTTTCCA GAAGGTCGGAGTCAACGGATTGGATTTTATGGGCTTTTCCG T C TT CC CT CT TACGGGGGAGATGCAGCCTGAGTCCGTCCTGATTCTAGGAGACGGTATCCCGTCCTTCCCGTTTCACGTGCACCAGCTGCGAGGGGGAACGCCCGCTGCGCGCGGGCAAAATCGCTCGCTTGCGGCCTTCTGGTCCATCATTCGTACTAGCCCCACACAGACCACAGCGGCGGCACGCTGCACGCACGCAAAATTTCCAC[T/C]GGAAAAGCCCATAAAATCCAGGAAAACGTGGTGGTTTTGCCAACCTCCCGCCCCTCGTCCAAAAAAAAGGCCCGGAAATAAAAGCGGATTGGAATCTGGTGGCTCCGGGTCCGTGCGGTGCGGGTGCGGCTCGCTCGACTTGGCGTCGTTGGAGTCCACCTGACCTCCTGTCCTCCACCGCCCGCTTCCCCTCGCCTCGG
OS900136_K01 3 31391306 GGAACGATGTGACGGAAAAA GAAGGTGACCAAGTTCATGCTGGACTTCGTCAAACTTCCAAC GAAGGTCGGAGTCAACGGATTGGACTTCGTCAAACTTCCAAA G T GG TT GT TT AGCCAAAAGGTTTATATTTAAAAATACTTTAGAAAAAGACTATTAATAAAATTAAAAATAAAGAGTCTAAAAAATAAAAATATTTTTAAGTTTGGATGTGTTTGCTTCATGCTAAAATTGGAAGTTTGGTTGAAATTGGAACGATGTGACGGAAAAATTGGAAGTTTATGCATGTAGAAAAGTTTCGATGCGATAGAAAC[T/G]TTGGAAGTTTGACGAAGTCCGTGTGCGTCACCGACATATGGGTCCCTTCTAGCGGTGCAGAAAGTGATAAGGTTCAACCAAAGTCGCAAGTCCGCAACGCGACTGTGCACACGAGGCCACTCCACTCCTCTCCTTTGCGCTCGCTCTGCTCTGCCATGGCGGAGGCGCTCGTCGCCGTCCTCCGGCTCGCGGCGTCCGCC
OS900137_K01 3 31783291 CCATGGACTATATGGGGTGTG GAAGGTGACCAAGTTCATGCTGTTTCACAAAGCAACACAGCAT GAAGGTCGGAGTCAACGGATTTTTCACAAAGCAACACAGCAC T C CC TT TC TC TGCAAGCTACTGCACTAGGAAGTATGTAATCTGGTGAATGCAAAGAGTACAAAGATTGTTTAAGCATGCACACTCATTACCTGAAGATTAGGAAAGGCAATGGATGAACAAAAGGCTCGGGAAAAGGTTGCTTGGAGCAGGACCGCATGAAATTGACTCTTTCTGTTACTTCCATAAAGGTTTCACAAAGCAACACAGCA[C/T]TAGTTGCTCACACCCCATATAGTCCATGGCTCCATGCCTTGTCTCGCCAAGATTGCCAGCTTCTGCACTGTTATCTATAACAAAAAACAGCTTCCATATTAATGTGTCGCTGCTGATTTTTCTTTTTAACATAACATAAGGATAGAAGATGTGGATGTAGAGCGAACAAGAAGCAAAAGGGACAGACCGAACAACACGTT
OS900138_K01 3 32205877 AGGAAAGATCTGGCCTTGTG GAAGGTGACCAAGTTCATGCTCGAGTGATCGGCTGTCTTT GAAGGTCGGAGTCAACGGATTCGAGTGATCGGCTGTCTTC T C TT CC TC CC CCTCGGGATTGAAGTTGATCAGGGAGAAAATGGAACCACGCTCAAGCAAACTGCCTATGCCAGAAAAGTGCTTTCTCAGTTCGGGATGCTGGAGTGTAATTCAACCTCAATTCCGATTGATCCAAGGATGCAGCTGAACAAAGATGCAGATGGTCACCCGGTGGGATGTAACTGAGTATCGGCGAGTGATCGGCTGTCTT[C/T]GATATCTGTTACACACAAGGCCAGATCTTTCCTTTGCTGTTGGAGTGGCAAGCAGATTTATGGAGCGGCCAACTCTTATGCACTTCAAAGCAGTGACACAAATTCTCAGATATCTGAAAGGTACAATTGATTGCGGCCTTGTGTATGCAGCTTGAAATGGTGATATGCAGATTACTGGGTATACAGACAGCGACCCGCTG
OS900139_K01 3 32341783 GCATTGGGTTTGTTGCTAGA GAAGGTGACCAAGTTCATGCTCCAAAAAAAGCTAATAACTCTCTCATATTT GAAGGTCGGAGTCAACGGATTCCAAAAAAAGCTAATAACTCTCTCATATTA T A TT AA TT AT GGTGATGAGGTGGAGGAGAGAAGTGAGGACAGAGATGATGAACCACCCCCCACACAAACTTCAAGAAGGATGTGAGAGTACTAAAGTCATTGATATTTGTGTATTAGAGGTGACCTATTGTATACTCTCTCCATTCCAAATTGATCTACATATTTCATAGGTACACCAAGACCAAAAAAAGCTAATAACTCTCTCATATT[T/A]TATTTACTCTAGCAACAAACCCAATGCATGCACCATCCCCACTATTTCCTAGCCAATAGCAAATTAAGATATTGCATGTGGGTTATAAATACTTGTGTGCATGGATGCATGCATCAATGTCCATTTACTCCAATGGACAAATAACGAATAGACTTAATAAATAAACACAAATATGTAGATCATTTAGGAATAACCTCAAA
OS900140_K01 3 32789243 GAGACAGCGTGAGATTGTGC GAAGGTGACCAAGTTCATGCTTCCCTTCTTTTTTTTTTTCGTTACT GAAGGTCGGAGTCAACGGATTCCCTTCTTTTTTTTTTTCGTTACC T C TT CC TC CC ATCCTTCTCGTCGTCTGGGAGATACAGAAGAAGTGGAACCAACGAATTTTTGAGCACAACGAGAGCATGACAATCACCCTCCTAGCAAAAATTAAGGAAGAAGCCAAAATTTAACCCTAGGAGGAGCTAAACCACAGTTAATTCCTAAGAGCGTATAATCTGTAAAGACGGGCTTTTCCCTTCTTTTTTTTTTTCGTTAC[C/T]AGGGGTGTATTTTTTTCCTCTCTATCTAACGAAAGATAAAGCACAATCTCACGCTGTCTCCTTTTAAAAAACACTTTCTTTACGCTGTTCATTGGCTGTAGCGGGTAGAATTCGCAAGGAATATGAATTAAATTTGGTTATAAATGAACGTACTTTTGTAATCACAACAAAGGGTCCATAGCTCAGTGGTAGAGCAATTG
OS900141_K01 3 33203208 CGCATAACCCTGATGGAGTC GAAGGTGACCAAGTTCATGCTCTCGCTGTGAAGGAGACGA GAAGGTCGGAGTCAACGGATTTCGCTGTGAAGGAGACGG T C CC TT TC TC TACCTTGCTTTCCTCGATCGGAGCGCCCAGAGTGACCAGCGTGGTGACCAGGTTGTTGAGGCGTAGCGAGAAGTCCTCGACGGACTCCCCCTCCTTGAACTCGATGGCGTCGTACTGCCGTAGAAACAGTTGCGCCTTGGACTTGCGCACTCGATCAACGCCGACTCGCATAACCCTGATGGAGTCCCACGCCTCCTTTG[C/T]CGTCTCCTTCACAGCGAGCGTCAACAGCATCTCCTGCGGCACTGCACGGAGGATGGCCTCCAGCGCCATTCTGTCTTCCTGAAAGTCTGCGTCGCCGTGCTAGACGGCTTCCCACAGACCACGGGCTTGGAGCATAACTTTCATCATCAGGGACCAGTCACCATAGTTTGTCTTCGTGAGGGTAGGATAGTTTGCTGAGC
OS900171_K01 3 14489983 GGTGACCTCGATACAAACTCGGTGA GAAGGTGACCAAGTTCATGCTCTTCCGTATGATTTTTCGTCTGTCA GAAGGTCGGAGTCAACGGATTCTTCCGTATGATTTTTCGTCTGTCG T C CT T/C CT CC ACGACGGTTGGTGGCGGCCCGGAGGAGATCGGTAGTGGCCACTAATTGGGAAGGGGAGGGAGCACGGGAAGAAAGGTGGGAGGTGGAAGAGATTAGGAGAGGGAGTGCGTGCATGAATCAGGGGATGTGATCAAGGGACCGTGATTAGGGGATCCAGATCGCTCTCTCTTAAAAGGGTGACCTCGATACAAACTCGGTGA[T/C]GACAGACGAAAAATCATACGGAAGCCTTACGTATGTTTTAAGTAGGTATAGATGTCATGTATCCTATAGATACAGATCTACTACTACGATATGTATTTAATATTTTAATATGTGACCAAATGTTATATTTTTGGGACATGTTTTGATATATCCAACTGATATGCAACTGATCTATGCAATCATTTAAAGTATGATATAAT
OS900172_K01 4 35335719 ACGGGCTGCTGGATTTACATAAGGT GAAGGTGACCAAGTTCATGCTTGCTTCAAACATATATTCATAGAGTCG GAAGGTCGGAGTCAACGGATTGCTTCAAACATATATTCATAGAGTCA G A GG AA AA GG TTTTGTAACCCTGGAAACCGCGAGACGAATTTATTAAGCCTAATTAATATGTCATTAGCACATGTAGGTTACTATAGCACTTATGGCTAATCATGGACTAATTAGGCTCAAAAGATTCGTCTCGCGATTTCCATACGAACTGTGCAATTAGTTTTTTATTTTATCATTATTTAATGCTTCAAACATATATTCATAGAGTC[G/A]ATGTGACGTTTTTCGGAAAAAAAATTAGGAACTACACAAGACCTTATGTAAATCCAGCAGCCCGTACTATTCGGGGATGACAGAAGGATGTCGATCTACACGGTTTATTAACAAGCTTTGGAGTTAAATTTCACCATTTAGCATTTTACCTTGAGCATTGGGAAAAGTAAGAAAACCCTTCGTGCAATGGAGAAATTCAT
OS900173_K01 6 29173606 AGCACAAAGGGAGTTGACATCATCA GAAGGTGACCAAGTTCATGCTGTCCTGTATCAAGTTTATTTTTAACA GAAGGTCGGAGTCAACGGATTGTCCTGTATCAAGTTTATTTTTAACG T C CC TT TT CC TAGATAGTGCATCTACACCTTTCGAATGAGGAGGAACAGCCTCTGGTCCAGCTCAAAGCTAGCGGCACTGGAGGGGTCACCCTTGAGCATCATCAGAAGGCCACCAAACGATGCATAGATCTCCCTAGAATGGAAAAGAATGCAATTCATTAAAATTCATAGCACAAAGGGAGTTGACATCATCACATGATACAGGATTA[C/T]GTTAAAAATAAACTTGATACAGGACTAACAGTAGTCTAGTACAAGCTGTGTAGCTGTAGGCCAGATTTACATATTGGCAAGGCGAAAAATTGTTCTTTGAAGGTACTTATCTTCGTATAACATGATTCACATAACAATGATACAATTAATTAAGTAAATGCACTTGAATTGAATTAACCAAGTAGCTGAAACAGTACAGA
OS900174_K01 7 23668282 AACATCTTCAGCCCTTGCAACCCTA GAAGGTGACCAAGTTCATGCTGTTCATGTTGTTGCTGGTGGGTCTA GAAGGTCGGAGTCAACGGATTGTTCATGTTGTTGCTGGTGGGTCTG T C CC TT TT CC TATTTCTTGTTCATTACATGATCAAATCAACTGGCACGCTCATACATTCGAAGGCGAGCTTTGGACCTGCACTAGAGTTAAACAGATCTCCCAGGCCTCGTGTTTTCTGTCAACAACAACACCAGCCCCAATGCCTACACCATCTTGACCAGCACCAATTGGAACATCTTCAGCCCTTGCAACCCTAGCATTCTCAGCCT[C/T]AGACCCACCAGCAACAACATGAACTTGGGCTAGATATGGGATCTCATTCTGTGGACGGAACTGAAGAAATAAAAAGATGAAACATAGTCTTAGTAAGTGGTGAAACACAAAATGTATATATACTAAAACATTCATAGTTGTGAAACATACCGGGCGAGACCCATATGTGACATAGCCAAGGTTCATCCTACAACGGTCTA
OS900175_K01 9 6324514 GCATCTCTTGACCCTATAAAATTGA GAAGGTGACCAAGTTCATGCTATGCAAGCAATGTGATTGGGAGA GAAGGTCGGAGTCAACGGATTGCAAGCAATGTGATTGGGAGG T C CC TT TT CC TAGAAACAGAAAGTTGGGGGACTCAACATACCGAGTACATCGACATAAAAACATAATCTCAATACTCCCTCCATCCCAAAATATAAGGCACAACCATTTTTCTCTCATGTCACATAATATAAGGCGTGTATGCATGTATATATTAACTAGCATCTCTTGACCCTATAAAATTGATTTGTTTTAAATCTTATACCATTAAA[C/T]CTCCCAATCACATTGCTTGCATGCATCCAAGTGATCCAACCATGAAGATGATAAAATATTTTCTTGGTCTTTGTGTTTTGAGTGTTATACCTTATATTTTAGGATGGAGGGAGTATTAAACTATACTAAAATACTATTAGGAGACCTTTAGCACCTCCTATCTTATCAACTATGACGAAAAATTTGAAGGGGGGACTCAG
OS900176_K01 9 10024501 TCGGATGTTACGTGTGTATGCAACC GAAGGTGACCAAGTTCATGCTTGGACCTCATATTCATGTTGTTTGGT GAAGGTCGGAGTCAACGGATTGGACCTCATATTCATGTTGTTTGGC T C CC TT TT CC TCTGATAGGCCACTTTCCGGATAATCCGAAATTATCAACATATGTGATATCTCTATCTATTGCATATAACATAAATTGGTTCCTTCAATGATCTAATTCATCAATTAACCTAGATTTCTTAGTCTGAGGCTTTCACCGTTTCTTGTCAGACTTGCCTAGCTGTATTTTTATATTATGGACCTCATATTCATGTTGTTTGG[C/T]ACTTAAGCATATAATTATTACGATGGATCGTTTATTGATTATTATATTATTGTTCATTGCATTACTAACTTGATATTATATTGGTTGCATACACACGTAACATCCGATAAATTTATTCCATTCCCGAACTGATTCCGCACCATTTCCGACATCCAAAATAATCCGTTCCGCATCCGCGTCCGTACACTATCCGCACCCGC
OS900177_K01 9 15799081 ACTAGCTTTCGTGCAAGGATCAACG GAAGGTGACCAAGTTCATGCTCCAAGGTACGTTATCTCCTTTAGTTG GAAGGTCGGAGTCAACGGATTCCAAGGTACGTTATCTCCTTTAGTTT G T GG TT TT GG CAGATTGTCTTGAATGATCCTGTTGAGAAGTATATGATTCCATTTCAAGAAACCCAGCCATTGCATGCTGACTTAATCTTGGAGGAGGTTGTCATGCGATTCTTCAAGGATCCTGTAGTTCCTATTGTTGATGACTGGGGAAGCTGCGTGGGTATTGTTCACCGACAGGATTGTACCAAGGTACGTTATCTCCTTTAGTT[G/T]GTACCAGCAATAATAGCGTTGATCCTTGCACGAAAGCTAGTGGAATGGTTAGCACTTGTTTTAGCGGCAGGGACCAGTCACCAGTGTTAGGATAAAAAAAATCAACCACCTTAGCTTCTCTTCCAAACAGTTCAAGTTTGGTCAAGGATTATATTACAAGGAGGAAACCAATAACAAATTTTGGTCGTTTTTCATTTTCA
OS900178_K01 10 8923315 CAGACATGGGGATGGATGCATATAGA GAAGGTGACCAAGTTCATGCTTGCAAAATAATGAAGAGATACAAGTGGAAA GAAGGTCGGAGTCAACGGATTTGCAAAATAATGAAGAGATACAAGTGGAAG T C TT CC CC TT TAACACAGAAAGTGAAAGTATATCTATCTATTTTTTTTGATCGAGTTCAAATTTTGAGATTACTGTGCTTGTCATATCAATATTTCAGGAAGATATTCAGCTTATGGCAGACATGGGGATGGATGCATATAGATTCTCAATTTCATGGTCTAGGATTTTCCCAAGTATGTAATATAATAATTCATTGGTTAATATGGGTA[T/C]TTCCACTTGTATCTCTTCATTATTTTGCATCTTTATGTTAACTTAATGGTCACACATAACCTTTGAAGTTTTTCTTCATTATGAAATGTTTTATCTGTGGATAAACAGACCTGCACTACTAACAATAATCATATCTGCACTCTTATTCAGATGGCACTGGTGAGGTCAACCAGGCAGGCATTGATCACTACAACAAGCTA
OS900179_K01 10 12020045 TTCAAGTTTTAGTTTGGACACTCTG GAAGGTGACCAAGTTCATGCTCAGCTGGTGAGTTCAAAGTGAAAAGATA GAAGGTCGGAGTCAACGGATTGCTGGTGAGTTCAAAGTGAAAAGATG T C CC TT TT CC GTCACGCATGAAGTACTATTCACATTTTATCATCTAATAACAATAAAAAATACTAATTATAAACAAATTAAATAAGACGGTGAAACGTTGGACATTGAAACCCACAACTACACTTAAAATGATACGGGGAAATAAATTGAGTAAATTGCAATTTGGACCATCTTTTATTATTCAAGTTTTAGTTTGGACACTCTGTATTT[C/T]ATCTTTTCACTTTGAACTCACCAGCTGATCAGAGTTGCGCTTAGCCATCGTTGTAGTTTTTTTTTCTCCTTTTCCTTGCTGCAGCTTTTGTTTTGCTTTTTCTCTCTTCCCCTGGTTTGGTTTGGACCGGTCTGCGGGTGAAGTAAAAAGATGACCGGTCTAATTAAACCAGGGAAAGGGAGAAAAAGAGTTTTTGTTAC
OS900180_K01 10 22817323 ATATGAATGTTGTGCCGGTGGTGAC GAAGGTGACCAAGTTCATGCTGATGACTCCCTATGTTAACCGATCCT GAAGGTCGGAGTCAACGGATTATGACTCCCTATGTTAACCGATCCC T C CC TT TT CC TTTCCATTCTCGCGAACCTTCCACACACCGAGAGCTTAGAATCTCACCCAGGGCCCCCGGCCGAACCGGCAAAGGGGGGCCTGCGCGGTCTCCCGGTGAGGAGCCTCACGCTCCGTCAGGTACACTAGCCGGTTTTGACAAGTCTTAGCTTAGATCACCGCAATAATCTTTAAGGGATGACTCCCTATGTTAACCGATCC[C/T]GCCTCAGATCCGCGCTCGTCACCACCGGCACAACATTCATATCCGGCCGATATGGCGATTTAATATATGCTACTATCTTCCTTCCAAAATATAAGGAATATAAGGAGTTATATTTTGAGATGGAGAAAGTACTAGTGATGATGAGGAAGATACACTAGTGCTTGAAGGCTTAGTTGTTTTGTTCTTTTTCCCAACTCACC
OS900181_K01 11 11901489 CATTAGGGGTAATTCCATTGTAGGG GAAGGTGACCAAGTTCATGCTGACTACCTATTATTGTTTTCAGTCAATTCT GAAGGTCGGAGTCAACGGATTCCTATTATTGTTTTCAGTCAATTCC T C CC TT TT CC CTCTCTGGGTCCATCCATAAGCATGCATACTGGCGCTCTCTTATTGGCTATACACACACACACACACACATATATATACATATATATATATATATATGTGTGTGTGTGTGTGTGTATAGTACTTTTAGATTTGATTAAATTACTTCCTATACATTCATAATGCTACGAGGTGACTACCTATTATTGTTTTCAGTCAATTC[C/T]GTAATTTGTGCTGATTAATTTAGTTTCTAGATAGAGTTATGTTTTTATCCCTACAATGGAATTACCCCTAATGTTATGACAAAGTTACTTTCGTTTTGTCTGAAGTTACTTCTATAATATATGTAAATTACATTTAAACTTTATTAAATTTACTTTTGTATGTCTAAGAAGTAATTTAGTGATATCCAAAAGTAATTTAG
OS900196_K01 1 38383221 TAGTAGTTGCACCGCATGATTGAGC GAAGGTGACCAAGTTCATGCTGGCGTGGAGCGAGGCTACTAT GAAGGTCGGAGTCAACGGATTGGCGTGGAGCGAGGCTACTAC T C C T GATGAAGGAGCTGTCGCTGACGATCATGGAACTCCTGGAGCTGAGCCTGGGCGTGGAGCGAGGCTACTA[C/T]AGGGAGTTCTTCGCGGACAGCAGCTCAATCATGCGGTGCAACTACTACCCGCCATGCCCGGAGCCGGAGCGGACGCTCGGCAC

附表2

候选基因引物信息"

基因名称Gene name 起始Start(bp) 终止End(bp) 正向引物Forward primer(5'-3') 反向引物Reverse primer (5'-3')
β-Actin AGTGTCTGGATTGGAGGAT TCTTGGCTTAGCATTCTTG
GAPDH AAGCCAGCATCCTATGATCAGATT CGTAACCCAGAATACCCTTGAGTTT
UBC1 CCGTTTGTAGAGCCATAATTGCA AGGTTGCCTGAGTCACAGTTAAGTG
LOC_Os03g49400 28116620 28124217 TCTTTCCCGTACCCTTGGAC TGCCGATACACATCGACAGA
LOC_Os03g49500 28169807 28173984 AGCCCTCAGGATCTACCTCA AGATGTGCCCTCCCATCAAA
LOC_Os03g49620 28254419 28259880 CCAACGTCCTGCTTGATGAG GCGCAATGTGACCCATAGTT
LOC_Os03g53150 30480660 30483331 GGTGAATGGCTCTGATGCTG CCTCAAGCGTTTGCATGACT
LOC_Os03g56050 31921933 31925513 TTCCACGCACATCAACTTCC CCTGCCTGCTCATGTTCTTC
LOC_Os03g57640 32851425 32852782 CAAACTGAGCTGCCTGAGTC TAGTCCTCGAAGCGGATCAC
LOC_Os03g52450 30094292 30099092 AAAGGCCAGTTTGCTGGAAG CCACAGGCATTACAGAGGGA
LOC_Os03g56060 31930541 31934624 CGTCGAGTGCCAATGTGAAT GAGCTTCCGGAGAAGGAAGA
LOC_Os03g53880 30890100 30899911 TACAGACTTTGGCCTCAGCA GTTCCAGCTCCCTGAGAAGT
LOC_Os03g55290 31465625 31464840 CCTCCTCTTCCCTCCAAGTT TACTTGAGGCAGTCGTCGTG
LOC_Os03g58260 33193735 33198078 CAGGTGCACGCTCAAATGTA CCGATGATCACACCATCTGC
LOC_Os03g58290 33207531 33210400 AAAGAAGCTGGTGTGCAAGG TCATCCTCTCTCCCGGTGTA
LOC_Os03g58300 33217418 33214533 GGCATGCTCTTTCAGGAGTG CCTTCATCCTGTCTGGTGGT
LOC_Os03g58320 33226460 33228543 GCTGGTGCTGCTTACTACAC AAAGCCAACACACACAGCTT

图1

‘IR 145’与‘Changai’的中胚轴长度比较 红色标识处为对应品种中胚轴部位。左侧为‘IR 145’, 中胚轴长度为0.18 cm左右。右侧为‘Changai’,中胚轴长度为4.75 cm左右。"

图2

IR 145×Changai F2群体中胚轴长度分布频率 X轴表示F2群体中胚轴长度, 分为0.18~1.21, 1.21~2.23, 2.23~3.26, 3.26~4.29 和4.29~5.32 cm 5个水平。Y轴表示各个水平的个体数。"

附表3

测序结果统计"

样品编号 Changai L-Pool S-Pool IR 145
Clean reads 160733360 189587718 223654650 145947156
Clean bases(bp) 24110004000 28438157700 33548197500 21892073400
Mapped reads 157946429 186412438 220135923 142034376
Mapped bases(bp) 23691964350 27762740240 32098485368 21305156400
Mapping rate 98.27% 98.33% 98.43% 97.32%
Duplicate reads 26216469 35168893 38980737 18236886
Duplicate rate 16.60% 18.87% 17.71% 12.84%
Mismatch bases(bp) 309119777 369422281 396469736 310450630
Mismatch rate 1.30% 1.33% 1.24% 1.46%
Average sequencing depth 51.32 57.83 66.53 47.49
Coverage 91.95% 95.90% 98.82% 92.53%
Coverage at least 4X 88.98% 93.47% 95.62% 89.25%
Coverage at least 10X 85.93% 90.72% 91.91% 86.24%
Coverage at least 20X 82.13% 86.27% 87.90% 82.48%
Coverage at least 50X 36.81% 46.49% 61.40% 15.04%

图3

水稻中胚轴极端池间的Δ(SNP-index)与G-value的基因组分布 A: Δ(SNP-index)在水稻12条染色体上的分布, 其中, 红色和绿色代表的置信区间为95%和99%; B: G-value在12条染色体上的分布, 阈值为FDR < 0.01."

图4

KASP标记在遗传图谱上的位置 左侧为3号染色体KASP标记遗传图谱, 右侧为对应LOD曲线, LOD阈值设为2.5。"

图5

候选基因在亲本间的表达量差异 图A~N依次表示14个候选基因在‘IR 145’与‘Changai’中胚轴组织中的相对表达量。内参基因为GAPDH。LOC_Os03g56050、LOC_Os03g57640、LOC_Os03g52450、LOC_Os03g56060、LOC_Os03g58290、LOC_Os03g58300和LOC_Os03g58320在短中胚轴品种‘IR 145’和长中胚轴品种‘Changai’中的表达量呈现显著差异(P < 0.05)。"

附图1

以管家基因β-Actin、GAPDH、UBC1作为参照的14个候选基因在‘IR 145’与‘Changai’中胚轴组织中的相对表达量 X轴表示不同的管家基因,分别为β-Actin、GAPDH、UBC1。Y轴表示各基因在‘IR 145’与‘Changai’中胚轴组织中的相对表达量。"

表1

水稻中胚轴伸长相关候选基因"

候选基因
Candidate gene		 基因名称
Gene name		 注释信息
Annotation
LOC_Os03g56050 OsAP23 AP2-like ethylene-responsive transcription factor Aintegumenta
LOC_Os03g57640 OsGID1L2 Gibberellin receptor GID1L2
LOC_Os03g52450 OsGATA19a GATA transcription factor 25
LOC_Os03g56060 OsCSLC9 CSLC9 - cellulose synthase-like family C
LOC_Os03g58300 OsIGL1 indole-3-glycerol phosphate lyase, chloroplast precursor
LOC_Os03g58320 OsIGL2 indole-3-glycerol phosphate lyase, chloroplast precursor
LOC_Os03g58290 OsIGL3 indole-3-glycerol phosphate lyase, chloroplast precursor

附表4

水稻3号染色体上已报道的中胚轴伸长相关QTL/基因"

位点/基因 标记区间 位置（bp） LOD值 解释变异 群体 参考文献
QTL/Gene Marker interval Position LOD value R2（%） Population Reference
- RZ403-RZ448 - 4.73 17.5 - Redona et al., 1996
- RZ403-RZ630 - 3.33 10.3 - Redona et al., 1996
qml3 RZ403-RG179 - 3.67 11.32 DH Cao et al., 2002
- G232-R1927 - 3.11 -0.59 RILs Cai et al., 2002
- R1927-BCD450 - 5.51 0.67 RILs Cai et al., 2002
- BCD450-RZ393 - 3.04 -0.56 RILs Cai et al., 2002
qml3-1 RZ517-RZ399 - 4.67 7.1 RILs OUYANG et al., 2005
qml3-2 RZ328-RZ575 - 3.45 6 RILs OUYANG et al., 2005
qml3-3 RZ696-RZ22 - 4.43 8.1 RILs OUYANG et al., 2005
qLOE-3 RM3919-RM520 - 2.548 10.7 DH Zhang et al., 2006
- RM426-RM448 - 8.93 24.97 RILs Huang et al., 2010
- RM426-RM514 - 15.52 33.36 RILs Huang et al., 2010
- R3226/R1927/R1925 - 4.2 11.5 BILs Lee et al., 2012b
- C595/R1927/R1925 - 7.5 20.8 BILs Lee et al., 2012b
- RM8208-RM8277 - 2.8 6.5 - Lee et al., 2012a
- R1618 - 12.09 37.56 BILs Lee et al., 2017
- R1618 - 7.18 26.97 BILs Lee et al., 2017
LOC_Os03g51340 - 29376077-29378021 - - - Zhao et al., 2018
LOC_Os03g53320 - 30603235-30602885 - - - Zhao et al., 2018
LOC_Os03g53340 - 30607159-30604067 - - - Zhao et al., 2018
LOC_Os03g06070 - 3039736-3045376 - - - Wu et al., 2015
LOC_Os03g06139 - 3085989-3075919 - - - Wu et al., 2015
LOC_Os03g39790 - 22138216-22135801 - - - Wu et al., 2015
LOC_Os03g40040 - 22244649-22243187 - - - Wu et al., 2015
LOC_Os03g40084 - 22268809-22270021 - - - Wu et al., 2015
LOC_Os03g40110 - 22292562-22285683 - - - Wu et al., 2015
LOC_Os03g40194 - 22351524-22347660 - - - Wu et al., 2015
LOC_Os03g40390 - 22440834-2248628 - - - Wu et al., 2015

附表5

已克隆的调控中胚轴伸长的基因"

染色体 基因名称 基因登录号 位置（bp） 参考文献
Chromosome Gene name Gene accession Position Reference
1 OsGY1 LOC_Os01g67430 39177169-39178676 Xiong et al., 2017
5 OsGSK2 LOC_Os05g11730 6661493-6657481 Sun et al., 2018
8 OsSMAX1 LOC_Os08g15230 9228110-9224161 Zheng et al., 2020
[1] Zhao G, Fu J, Wang G, Ma P, Wu L, Wang J. Gibberellin-induced mesocotyl elongation in deep-sowing tolerant maize inbred line 3681-3684. Plant Breed, 2010, 129:87-91.
doi: 10.1111/pbr.2010.129.issue-1
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