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作物学报

作物学报 ›› 2020, Vol. 46 ›› Issue (3): 330-340.doi: 10.3724/SP.J.1006.2020.94077

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

利用EST-SSR标记评价羽扇豆属(Lupinus L.)遗传多样性

张红岩1,5,*,杨涛1,*,刘荣1,*,晋芳2,张力科2,于海天3,胡锦国4,杨峰3,王栋1,何玉华3,*(),宗绪晓1,*()   

  1. 1. 中国农业科学院作物科学研究所 / 国家农作物基因资源与基因改良重大科学工程, 北京 100081
    2. 全国农业技术推广服务中心, 北京 100125
    3. 云南省农业科学院粮食作物研究所, 云南昆明 650205
    5. 青海大学, 青海西宁 810016;
  • 收稿日期:2019-05-21 接受日期:2019-09-26 出版日期:2020-03-12 网络出版日期:2019-11-11
  • 通讯作者: 张红岩,杨涛,刘荣,何玉华,宗绪晓
  • 作者简介:张红岩, E-mail: 1006248823@qq.com|杨涛, E-mail: yangtao02@caas.cn|刘荣, E-mail: liurong@caas.cn
  • 基金资助:
    本研究由国家重点研发计划项目(2017YFE0105100);农作物种质资源保护与利用专项(2019NWB036-07);中国农业科学院科技创新工程项目和中国农业科学院作物科学研究所中央级公益性科研院所基本科研业务费专项资助(S2018XC04)

Assessment of genetic diversity by using EST-SSR markers in Lupinus

Hong-Yan ZHANG1,5,*,Tao YANG1,*,Rong LIU1,*,Fang JIN2,Li-Ke ZHANG2,Hai-Tian YU3,Jin-Guo HU4,Feng YANG3,Dong WANG1,Yu-Hua HE3,*(),Xu-Xiao ZONG1,*()   

  1. 1. Institute of Crop Sciences, Chinese Academy of Agricultural Sciences / The National Key Facility for Crop Gene Resources and Genetic Improvement, Beijing 100081, China
    2. National Agro-Tech Extension & Service Centre, Beijing 100125, China;
    3. Institute of Food Crops, Yunnan Academy of Agricultural Sciences, Kunming 650205, Yunnan, China
    4. USDA-ARS Western Regional Plant Introduction Station (WRPIS), Pullman, WA 99164, USA
  • Received:2019-05-21 Accepted:2019-09-26 Published:2020-03-12 Published online:2019-11-11
  • Contact: Hong-Yan ZHANG,Tao YANG,Rong LIU,Yu-Hua HE,Xu-Xiao ZONG
  • Supported by:
    This study was supported by the National Key Research and Development Program of China (2017YFE0105100);the Crop Germplasm Resources Protection(2019NWB036-07);the Agricultural Science and Technology Innovation Program (ASTIP) in CAAS, and the Fundamental Research Funds for Central Non-Profit of Institute of Crop Sciences, CAAS.(S2018XC04)

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摘要：

利用基于窄叶羽扇豆转录组开发并筛选出的95对多态性EST-SSR标记, 对羽扇豆属的22个种133份资源进行全基因组扫描, 初步探究羽扇豆属下种间的进化关系, 分析来源于”旧世界”羽扇豆种间的遗传多样性, 为羽扇豆优异资源的挖掘和创新利用提供理论依据。结果表明, 用95对SSR标记共检测出1318个等位变异, 每对标记平均检测出3~37个等位变异, 平均为13.87个; 多态性信息量(PIC)变化范围0.39~0.91, 平均为0.75; 基因多样性变化范围0.41~0.92, 平均为0.78。基于邻接法(NJ)的系统发育初步探究了羽扇豆种间的进化关系, 22个羽扇豆种分别来源于”旧世界”和”新世界”, 与之前研究结果相对一致。聚类分析、群体结构分析和主成分分析结果均表明, 来源于”旧世界”的7个羽扇豆种被划分为4个组群, 各组群所包含的参试资源没有种间交叉重叠。

关键词: EST-SSR, 羽扇豆属, 转录组, 遗传多样性

Abstract:

In order to explore the evolutionary relationship of Lupinus preliminarily as well as to excavate and utilize lupin resources from the “Old World” effectively, the genetic diversity among the species under Lupinus genus was analyzed. Ninety-five polymorphic pairs of EST-SSR markers developed based on the transcriptome of narrow-leaved lupin (Lupinus angustifolius L.) were used to scan 133 lupin accessions from 22 species. A total of 1318 alleles were detected with 13.87 alleles per locus on average, ranging from 3 to 37 alleles; the polymorphism information content (PIC) ranged from 0.39 to 0.91 with the mean value of 0.63; the genetic diversity ranged from 0.41 to 0.92 with the mean value of 0.78. This study showed evolutionary relations among the 22 species under Lupinus genus from the “Old World” and the “New World” based on Neighbor-Joining (NJ) method, which is consistent with previous studies. Moreover, seventy-seven lupin accessions of seven Lupinus species from the “Old World” were divided into 4 groups; there was no overlap of accession from different species contained in each identified group, detected by all the three analysis methods like population structure, cluster analysis based on UPGMA and principal component analysis (PCA).

Key words: EST-SSR, Lupinus genus, transcriptome, genetic diversity

图1

基于不同地理来源的羽扇豆资源分布图"

图2

95对EST-SSR标记在窄叶羽扇豆染色体上分布情况"

表1

95多态性标记遗传参数"

标记
Marker		 等位基因
No. of allele		 基因多样性
Gene diversity		 PIC 标记
Marker		 等位基因
No. of allele		 基因多样性
Gene diversity		 PIC
lup-est4467 22.00 0.88 0.87 lup-est6567 31.00 0.91 0.91
lup-est4113 17.00 0.80 0.78 lup-est2194 14.00 0.80 0.77
lup-est3140 14.00 0.73 0.68 lup-est4304 25.00 0.89 0.88
lup-est4349 10.00 0.65 0.62 lup-est3549 10.00 0.68 0.63
lup-est6531 7.00 0.42 0.39 lup-est2949 15.00 0.89 0.88
lup-est5958 11.00 0.73 0.69 lup-est2185 13.00 0.77 0.74
lup-est2267 17.00 0.88 0.86 lup-est3185 14.00 0.85 0.84
lup-est2076 16.00 0.82 0.80 lup-est2385 14.00 0.79 0.76
lup-est6513 16.00 0.85 0.84 lup-est3894 14.00 0.83 0.82
lup-est6558 19.00 0.91 0.90 lup-est1449 14.00 0.80 0.78
lup-est6258 16.00 0.86 0.85 lup-est2140 13.00 0.87 0.85
lup-est6104 17.00 0.71 0.69 lup-est3458 10.00 0.68 0.64
lup-est6049 11.00 0.73 0.69 lup-est6158 13.00 0.79 0.77
lup-est1985 12.00 0.76 0.73 lup-est2822 12.00 0.80 0.77
lup-est2531 30.00 0.89 0.88 lup-est5340 9.00 0.77 0.74
lup-est976 18.00 0.82 0.80 lup-est5440 12.00 0.73 0.69
lup-est2013 14.00 0.76 0.74 lup-est4549 18.00 0.87 0.86
lup-est840 7.00 0.73 0.68 lup-est5513 10.00 0.77 0.73
lup-est1640 12.00 0.76 0.73 lup-est1149 8.00 0.82 0.79
lup-est931 18.00 0.88 0.87 lup-est4576 7.00 0.70 0.65
lup-est1749 21.00 0.89 0.88 lup-est2694 18.00 0.87 0.86
lup-est2558 16.00 0.85 0.84 lup-est3494 12.00 0.85 0.83
lup-est458 9.00 0.72 0.67 lup-est158 7.00 0.79 0.75
lup-est4640 16.00 0.88 0.87 lup-est5049 10.00 0.80 0.77
lup-est5985 11.00 0.81 0.79 lup-est1058 8.00 0.68 0.64
lup-est2849 22.00 0.84 0.82 lup-est3994 21.00 0.91 0.90
lup-est3649 37.00 0.91 0.90 lup-est1594 12.00 0.80 0.78
lup-est2340 23.00 0.92 0.91 lup-est4322 16.00 0.85 0.84
lup-est4522 9.00 0.66 0.63 lup-est4185 12.00 0.83 0.81
lup-est5240 13.00 0.83 0.81 lup-est5749 7.00 0.70 0.64
lup-est2567 7.00 0.41 0.39 lup-est5667 9.00 0.64 0.60
lup-est58 10.00 0.73 0.69 lup-est3067 21.00 0.88 0.87
lup-est4458 12.00 0.86 0.84 lup-est1649 12.00 0.66 0.63
lup-est4131 9.00 0.72 0.67 lup-est5622 9.00 0.75 0.71
lup-est5740 13.00 0.63 0.58 lup-est3576 5.00 0.71 0.66
lup-est164 8.00 0.63 0.59 lup-est6213 9.00 0.78 0.75
lup-est6031 14.00 0.81 0.79 lup-est2358 11.00 0.88 0.87
lup-est922 13.00 0.82 0.80 lup-est2540 11.00 0.79 0.76
lup-est2313 20.00 0.84 0.83 lup-est4249 10.00 0.53 0.49
lup-est249 17.00 0.77 0.75 lup-est4949 13.00 0.67 0.61
lup-est2704 13.00 0.44 0.43 lup-est4158 14.00 0.80 0.77
lup-est1085 13.00 0.80 0.78 lup-est822 14.00 0.86 0.84
lup-est1622 19.00 0.81 0.79 lup-est3340 10.00 0.81 0.79
lup-est6504 9.00 0.67 0.61 lup-est1313 17.00 0.81 0.80
lup-est613 12.00 0.77 0.75 lup-est2404 12.00 0.80 0.78
lup-est449 12.00 0.69 0.66 lup-est713 25.00 0.80 0.77
lup-est4022 7.00 0.72 0.67 lup-est5967 13.00 0.80 0.78
lup-est5831 15.00 0.84 0.82 平均Mean 13.87 0.78 0.75

图3

22个羽扇豆种的系统发育分析"

图4

基于7个羽扇豆不同种间(“旧世界”)的 UPGMA 聚类图"

表2

7个羽扇豆种间(“旧世界”)的Nei’s (1978)遗传距离和遗传一致度"

物种名称
Species ID		 L. atlanticus L. cosentinii
(沙质平原羽扇豆)		 L. albus
(白羽扇豆)		 L. angustifolius
(窄叶羽扇豆)		 L. hispanicus L. luteus
(黄羽扇豆)		 L. pilosus
L. atlanticus 0.663 0.235 0.167 0.187 0.197 0.561
L. cosentinii
(沙质平原羽扇豆)		 0.411 0.199 0.158 0.187 0.167 0.556
L. albus
(白羽扇豆)		 1.45 1.613 0.244 0.205 0.263 0.254
L. angustifolius
(窄叶羽扇豆)		 1.789 1.845 1.412 0.196 0.254 0.223
L. hispanicus 1.676 1.679 1.586 1.628 0.487 0.213
L. luteus
(黄羽扇豆)		 1.625 1.792 1.336 1.372 0.719 0.218
L. pilosus 0.578 0.586 1.369 1.502 1.547 1.524

图5

77份羽扇豆资源(“旧世界”)遗传多样性分析 (a)基于ln P(D)平均值, 绘制散点图; (b)基于ln (ΔK)值, 绘制散点图; (c)当k = 4时, 77份羽扇豆资源遗传结构; (d)基于Nei’s遗传距离, 77份羽扇豆资源NJ树; (e) 77份羽扇豆资源主成分分析。"

表3

7个羽扇豆种之间的分子方差分析"

变异来源
Source		 自由度
df		 方差
SS		 均方差
MS		 Est. Var. % P-value
组群内Within pops 6 6584.338 1097.39 108.0748 80.00 0.01
组群间Among pops 70 1852.636 26.466 26.466 20.00 0.01

表4

7个羽扇豆种间遗传分化分析"

物种名称
Species ID		 L. atlanticus L. cosentinii
(沙质平原羽扇豆)		 L. albus
(白羽扇豆)		 L. angustifolius
(窄叶羽扇豆)		 L. hispanicus L. luteus
(黄羽扇豆)		 L. pilosus
L. atlanticus 0
L. cosentinii
(沙质平原羽扇豆)		 0.475 0
L. albus
(白羽扇豆)		 0.617 0.661 0
L. angustifolius
(窄叶羽扇豆)		 0.614 0.639 0.546 0
L. hispanicus 0.688 0.714 0.636 0.602 0
L. luteus
(黄羽扇豆)		 0.707 0.743 0.635 0.597 0.573 0
L. pilosus 0.320 0.341 0.470 0.476 0.535 0.531 0

附表 1

133份羽扇豆资源来源地信息"

Accession Species Continents Longitude (E) Latitude (N)
PI 284712a,c L. arbustus North America -121.45 38.57
PI 284714a L. arbustus North America -121.45 38.57
PI 232569a,c L. argenteus North America -77.03 38.88
PI 504372a L. argenteus North America -111.87 40.77
PI 504374a L. argenteus North America -123.03 44.93
PI 577284a L. argenteus North America -116.22 43.62
PI 477930a L. argenteus North America -123.03 44.93
PI 477931a L. argenteus North America -123.03 44.93
PI 477936a L. argenteus North America -123.03 44.93
PI 504330a L. argenteus North America -77.03 38.88
PI 384612a,b,c L. atlanticus Africa -6.85 34.03
PI 384613a,b L. atlanticus Africa -6.85 34.03
W6 17965a,b L. atlanticus Africa -6.85 34.03
PI 284715a,c L. concinnus North America -121.45 38.57
PI 284716a L. concinnus North America -121.45 38.57
PI 660715a,b,c L. cosentinii Oceania 149.13 -35.30
W6 17967a,b L. cosentinii Europe -9.08 38.70
PI 185099a,c L. elegans North America -99.15 19.47
W6 15613a L. elegans North America -121.45 38.57
PI 344647a,c L. garfieldensis North America -77.03 38.88
PI 344648a L. garfieldensis North America -77.03 38.88
PI 477932a,c L. lepidus North America -123.03 44.93
PI 477934a L. lepidus North America -123.03 44.93
PI 504325a L. lepidus North America -123.03 44.93
PI 504391a,c L. leucophyllus North America -77.03 38.88
PI 504397a L. leucophyllus North America -123.03 44.93
PI 504398a L. leucophyllus North America -123.03 44.93
PI 504399a L. leucophyllus North America -123.03 44.93
PI 504400a L. leucophyllus North America -123.03 44.93
PI 602371a,c L. mexicanus Europe 19.25 47.43
PI 615395a L. mexicanus North America -121.45 38.57
PI 615398a L. mexicanus North America -121.45 38.57
PI 615399a L. mexicanus North America -121.45 38.57
W6 14749a L. mexicanus Europe 19.25 47.43
PI 602373a,c L. microcarpus North America -121.45 38.57
PI 602376a L. microcarpus North America -121.45 38.57
PI 344664a,c L. polyphyllus North America -116.22 43.62
PI 442515a L. polyphyllus Europe 4.35 50.85
PI 477122a L. polyphyllus Europe 13.03 52.52
PI 504411a L. polyphyllus North America -116.22 43.62
PI 504412a L. polyphyllus North America -77.03 38.88
PI 504413a L. polyphyllus North America -77.03 38.88
PI 504414a L. polyphyllus North America -123.03 44.93
PI 452129a,c L. rivularis North America -123.03 44.93
PI 660700a L. rivularis North America -123.03 44.93
PI 660701a L. rivularis North America -123.03 44.93
PI 660720a L. rivularis North America -123.03 44.93
PI 284726a L. stiversii North America -121.45 38.57
PI 602374a,c L. stiversii North America -121.45 38.57
PI 284728a,c L. succulentus North America -121.45 38.57
PI 577290a L. succulentus North America -77.03 38.88
PI 615405a,b,c L. albus North America -92.32 34.73
PI 615406a,b L. albus North America -92.32 34.73
PI 615409b L. albus North America -84.42 33.77
PI 660685b L. albus Europe -6.97 38.88
PI 660711b L. albus Europe -16.23 28.45
PI 660712b L. albus Africa 36.85 7.38
PI 660713b L. albus Asia 36.32 33.05
PI 660714b L. albus Africa 31.28 30.00
PI 660721a,b L. albus Europe 2.03 48.85
PI 660722b L. albus Europe 2.03 48.85
PI 660723b L. albus North America -86.03 32.37
PI 660724b L. albus Asia 35.93 31.95
PI 660726a,b L. albus Oceania 149.13 -35.30
PI 666380b L. albus Europe -4.03 39.86
PI 666381b L. albus Europe -6.94 37.29
PI 666382b L. albus Europe -4.03 39.86
PI 666385b L. albus Oceania 149.13 -35.30
PI 666386b L. albus Africa 3.22 36.70
W6 15551b L. albus Europe 2.03 48.85
W6 19548a,b L. albus South America -70.07 -33.43
W6 39795a,b L. albus Europe 13.03 52.52
W6 39796a,b L. albus Europe 13.03 52.52
W6 39797b L. albus Europe 13.03 52.52
W6 39798b L. albus Europe 13.03 52.52
W6 39800a,b L. albus Europe 13.03 52.52
W6 39801a,b L. albus Europe -9.08 38.70
W6 39802a,b L. albus Asia 35.22 31.78
PI 168527b,c L. angustifolius Europe -9.08 38.70
PI 180708b L. angustifolius Europe 13.03 52.52
PI 208682b L. angustifolius Africa 3.22 36.70
PI 255472b L. angustifolius Europe 20.47 44.82
PI 274821a,b L. angustifolius Europe 21.00 52.25
PI 274811a,b L. angustifolius Europe 21.00 52.25
PI 274812b L. angustifolius Europe 21.00 52.25
PI 274813b L. angustifolius Europe 21.00 52.25
PI 274814a,b L. angustifolius Europe 21.00 52.25
PI 274815a,b L. angustifolius Europe 21.00 52.25
PI 274817b L. angustifolius Europe 21.00 52.25
PI 274819a,b L. angustifolius Europe 21.00 52.25
PI 274820b L. angustifolius Europe 21.00 52.25
w6 39768b L. angustifolius Europe 13.03 52.52
w6 39769a,b L. angustifolius Europe 13.03 52.52
w6 39771b L. angustifolius Europe 13.03 52.52
w6 39772a,b L. angustifolius Europe -3.70 40.43
w6 39775a,b L. angustifolius Europe 13.03 52.52
w6 39776a,b L. angustifolius Europe 13.03 52.52
w6 39777a,b L. angustifolius Europe 13.03 52.52
PI 384554b,c L. hispanicus Europe -9.08 38.70
PI 384555b L. hispanicus Europe -9.08 38.70
PI 384556a,b L. hispanicus Europe -9.08 38.70
PI 384558a,b L. hispanicus Europe -9.08 38.70
PI 384559b L. hispanicus Europe -9.08 38.70
PI 384560a,b L. hispanicus Europe -9.08 38.70
PI 384561a,b L. hispanicus Europe -9.08 38.70
PI 384562b L. hispanicus Europe -9.08 38.70
PI 168539a,b,c L. luteus Europe -9.08 38.70
PI 168544b L. luteus Europe -9.08 38.70
PI 168546a,b L. luteus Europe -9.08 38.70
PI 189159b L. luteus Europe 4.87 52.35
PI 224493a,b L. luteus Europe 4.87 52.35
PI 289168a,b L. luteus Europe 19.25 47.43
PI 255475a,b L. luteus Europe 20.47 44.82
PI 266298b L. luteus Europe 12.99 55.61
PI 206508a,c L. mutabilis South America -76.92 -12.10
PI 432332a L. mutabilis South America -76.92 -2.10
PI 457966a L. mutabilis South America -71.98 -13.52
PI 457967a L. mutabilis Africa -6.85 34.03
PI 457972a L. mutabilis South America -76.92 -12.10
PI 249758a,b,c L. pilosus Europe 24.02 35.51
PI 338647b L. pilosus Africa -6.85 34.03
PI 491183b L. pilosus Asia 32.90 40.03
PI 602369b L. pilosus Asia 37.81 28.49
PI 602375a,b L. pilosus Asia 35.22 31.78
PI 664108a,b L. pilosus Europe 19.25 47.43
PI 664109b L. pilosus Europe 12.57 55.72
PI 664110b L. pilosus Asia 35.22 31.78
PI 664111b L. pilosus Europe -9.08 38.70
PI 344687a,c L. sericeus North America -116.22 43.62
PI 344694a L. sericeus North America -116.22 43.62
PI 452488a L. sericeus North America -112.03 46.58
PI 477950a L. sericeus North America -77.03 38.88
PI 477952a L. sericeus North America -77.03 38.88

附表 2

95对多态性EST-SSR信息"

引物
Primer		 重复单元
Repeat motif		 正向引物
Forward sequence (5°-3°)		 正向引物
Reverse sequence (5°-3°)		 退火温度
Tm (℃)
lup-est6104 (TTG)4 CTCAAGTTTTGGCTTCATGC CTCAAGTGCATTCATGGAAGT 54
lup-est4304 (TC)6 AGGGGTTTTGAAGAGAAAGG AGACAGTGATAGCTCAGTGGG 54
lup-est2704 (GA)5 TCAAGCCCTTCACTCAAAAC CCCCCTTTCTAAAAATCCAA 54
lup-est2313 (TC)4 GTGACAATGGCTCTTCAACC CCTTGTGCAAAAGTCCAAAT 54
lup-est713 (TAA)4 CAGCCATTAAAGCTTTTGGA AGAAAACACCAAAAGGGACC 54
lup-est613 (TTG)5 TTCCACCCATCATTGAAATC GGAGCAACAACTACACATCCA 54
lup-est1313 (TTA)5 GAAAACCACCACCACTTTTG GGGGGTAGTGCTAAGAAAGG 54
lup-est4458 (CAA)3 CAGAATCACACCCCAATCTC ATCCCAGGCCATTGACTAAT 54
lup-est5958 (AAC)4 CAACGGGGTCATTGAATTAG GTTTGTGACGGGTTAAGGTG 54
lup-est3458 (CAC)4 GACGACTCCAATGACACTCC GATGAGGGTTTCGAAGAGGT 54
lup-est1058 (GGT)4 TTTATCGGGAAGCATGTGTT ATAGGTGAGCCTGGGACTTC 54
lup-est2558 (CT)9 AGACCCACCAAACAATGAAA ATCCCGAGGCCAAATCTA 54
lup-est158 (CTT)3 TGGTTGTTGGGTCTAATAGCA TGAGGAGAACCATCATTCAAG 54
lup-est6558 (ATG)6 TGTTGATGCCAGAAACATTG GCCTTCCCTTCTCTACAACC 54
lup-est4949 (TGA)5 CTCAATCCCCTAGTGACCCT GGTAATTCCAAACTTGCCCT 54
lup-est1649 (TC)6 CAATCTTCTTCACTCTACTCCTCA CAAGTCTGAAATCCTTGCGT 54
lup-est1449 (TTG)6 GGTTTCCATCATAAGCATCC TCAGCACACTAGCAAGTACCC 54
lup-est5740 (AAT)5 ACATGTGCATGTTGAATTGC ATTTCGAGGACCAACAATGA 54
lup-est2540 (CAC)4 GAATCATTGAACCGTGATGG GAAAGGGTTCTCTGCTTCGT 54
lup-est4549 (TTG)4 AACCCCATTTTAAGCAAGGT CCACTCTTCCTCAGAAAAATCA 54
lup-est2949 (CAA)5 ATCAATGGCACAAGCAATG AACCAGTAGCAACAAGACCAA 54
lup-est3140 (CTT)4 CATTCAACCAAAACCCATGT ACACGCAGTTTGAAGAGAGC 54
lup-est2849 (TTC)5 CGATGGTCCATTGATATAGAAG AGACCTTCCGAAGTGAGACA 54
lup-est1149 (TGT)4 GCAATGTTGAGCTTCCAAAG CTTTCTTGGTGTTGGAGGAA 54
lup-est4640 (TTG)7 CCTTACCATTGCATCAGTCC GGTATAACCCCTTTTCAGCC 54
lup-est6031 (AAC)4 AAATCTAAGCCCATGGAACC CCCTGCTCTTCTCAAAGTCA 54
lup-est840 (ATC)5 CAACTTTTCAACGGGTTCAC GCAGAAGGGTAAAATACGCA 54
lup-est931 (GAA)4 CAATTTCACCCTCTTCGCTA CTTTAGCAACATCGAAGGGA 54
lup-est6531 (CCT)4 CAAGGTTTGCTTCGTAGGAG AGGGAGGAAATTGGATGTGT 54
lup-est1985 (TGC)4 AAGTGATGTCCTGAGCTTGC GCTCCACAGATGACACAACA 54
lup-est4185 (GTG)5 GCAACAGTCACCTTGCATTA CCCTTAAACTCCACTTCAACC 54
lup-est3185 (CAA)5 CTCAACCCAACAAAACAACA GCACTGCAATCAGAAGTGAA 54
lup-est6567 (CTA)5 CCAACTCCAGTCACTGCTCT GCTCTCAAATGGGGTTGTAG 54
lup-est2567 (TAG)6 AGTGGCCAGGATAGTGTTGA CAAACTCCACCTCATTGTCC 54
lup-est5667 (GGA)4 CAAGGGTAAGTACCCGATCC GAGAGATGAGTGAAGCACAGAA 54
lup-est2076 (GGT)3 ATAGTTGCTGTAACTCGGCG CTGTCCTCCTCCTCCTTCTC 54
lup-est4467 (ACA)4 TGCATTCATGGCTTCACTTA TTGTGACAACTTGGAGGGAT 54
lup-est976 (GAA)4 GCTTAAGGGAGGAGCAAGAT GCTTCTTGTGCTGCTCTTTC 54
lup-est1622 (CAC)4 GTACCCCTGATGATGCTGAC TGGTGGATTGTCTGATGATG 54
lup-est4022 (GAA)4 TCTACTTCCTTCACAAAACGC CAAAAGGTAACGAAGCTCCA 54
lup-est822 (ATA)4 CATATTCAAGGCTTTGGTGG TTTGGGGCAAGTAGAAATTG 54
lup-est4522 (ATG)5 AGGTAACGTTGATTCGTGGA CACCAACAACACCACTACCA 54
lup-est2822 (CCT)4 AGACCAAGAGAGTGGAGGCT CACTCTCTCAGTAGGAGGCG 54
lup-est6213 (AAG)5 GGCGTGGGTTAAGCTTATGT ACCCCACCTTCTTCAATCAT 54
lup-est2694 (ATC)6 GGGGCTGCTTCATCTTATTT CGCCTCAATTCAAGGATATG 54
lup-est1594 (ATC)6 GAATGTTCCAGTTGGTGGAG ATCATAACCCATTGACCCCT 54
lup-est3894 (TC)6 GAAACCCAACCAATTGTTCA AACCAACATTCTCCACTTGC 54
lup-est2013 (ATG)4 CTATGGAAGATGCGCAGAAT CCCACTGAATCTGACTTGCT 54
lup-est6504 (ACC)4 TGCCTTCTATTGTCACATGC AGATTCTGACTAGCCCCACC 54
lup-est2404 (TGG)4 AAGCACCACCTTCTGTGCTA TCACTTTGGTTCCACTAGCC 54
lup-est6158 (AAC)4 ACCAAAGAAAACAAAACCCC GGACGTCCTTGTGCTGTTAC 54
lup-est458 (CGC)4 AGAGAAGAAACCGGAGGAAA GCATGAACACTTTCAGCACA 54
lup-est4158 (GTT)4 AGAGAGACCAGTGGGAGGAT AAAACCTGTTCGGCTTCTTT 54
lup-est6049 (TGG)5 CTGGCAGCATGGTTTTTAAG AAATGGACTTGTCCCAAACC 54
lup-est58 (GA)7 TCCAGCAGATCCGTTAATCT TCAGATCGCACTCTCTGTGA 54
lup-est2358 (TTG)4 GGTTGATGCAGCAGAAAAGT ACAACTTGCAAATCCACCAT 54
lup-est5049 (AAC)4 TCAGTTAAATCCAACGCCAC GGAATTCTGCGAAGGTATCA 54
lup-est5749 (TGC)4 TGTCTCGATGGTTCAGCATA GCCTTAGATGCCCATCTTCT 54
lup-est6258 (AAT)5 GCTGAAATTGTAGCAGCCAT TGGACCTTGTTTTTCAATGC 54
lup-est3340 (ATG)7 AAGGTTCAGGACTTGTTGAAGA ATCAACCTGTGACCCAGAAA 54
lup-est449 (CAG)6 AGCTCCTCCCAATGCTTATT ATGTTCCAGAGAAATGCCAG 54
lup-est3649 (ACA)4 ACTCAAGTCCAACAATCCCA CACCTTAAACGTGAGAAAGCA 54
lup-est2340 (CTA)5 AAACCCAGCAAATTCACAAG TCTGGAACTCCATTCTTTGC 54
lup-est5440 (TAT)4 GGCTTTTCTTCCCATTGTTT TTTCAGAACAACGTGTCCAA 54
lup-est4349 (AAC)5 AAGAAATGCCCTAGCTGGTT TCTTGTTGCTCTTCCTTTGG 54
lup-est3549 (CT)6 CATGTGCACTCCGTTTATCA GACCTTGTTGTGGGTATTGC 54
lup-est249 (ATG)4 ACAGGTCCATGGAAAACAGA GGTGATGAATTGGGATCTGA 54
lup-est5340 (CCT)5 CGAGAAGCGATCGACATAAT CGACACAGCAGAAGTTGTGA 54
lup-est4249 (CTT)6 AGCTTTGATTTTTGACTGCG TCGTAGAAGTGGATACCCCA 54
lup-est2140 (CAA)4 AGGAACAATCTCAAGCCACA GAGATTATGCTGCCACTGCT 54
lup-est1749 (TGC)3 GCGTAGCTGCTCTCATTTTC AGCTCCACATTCATCCTCAG 54
lup-est5240 (AAC)3 GCAAACTCATTCCCTTCAAA CCTCATCAACGACACCTTCT 54
lup-est1640 (TTC)4 TTAGGGCATTGAACCTTTTG CCACCGTCGTAAGTAACACC 54
lup-est5831 (CTT)5 CACAGACTCAACCTTGGCTT TACTGGCTCCAGATCAAACC 54
lup-est4131 (GTT)6 CAAGCCCCTTAACAAGTCCT CTGGCCAGAGCTGGTAAATA 54
lup-est2531 (TC)6 CACGTTGGGTTTCTTTCATT GAGAGCTTTGCACGTTGATT 54
lup-est2185 (CCA)4 CACTCTTCCAAACGCAAGTT AGCTTCTTTAGGGTTCGGAG 54
lup-est5985 (GAA)5 CCCCAATCTCAGAAATCAAA CAAGTCAGCTTGGGAAAGAA 54
lup-est4576 (ATT)6 CCATATAGCTGCTTCCTCCA AATGCACTTCTCCACCACTC 54
lup-est1085 (CCA)4 AACCTATCTGCTGCTGATGC AGTTCTGCATGCAATCTGGT 54
lup-est2385 (ATG)4 CATGAAGAAGAAGCCAAGGA TTCACCACTTTCATCTTCAGC 54
lup-est3067 (TCA)4 CAAAGATGCTCATCAAGAAGC TAGGGGTTGTTCTTTGTCCA 54
lup-est2267 (GTT)5 AGCCAAGTAGGTGCTGTTTG TCACCACCTCTCTCAAAACC 54
lup-est3576 (CAC)4 CTGGGAATTGAGGTTTGTTG ACTGAAGACGGTGCTGATTC 54
lup-est5967 (AAC)4 AAATCACAATCACCACAGCA TGCATTAGGGAGAAAGTTGG 54
lup-est922 (TTC)4 AGTAGGGTGGAGATGGAAGG GCTTCTCCATGCTCTCTTCA 54
lup-est5622 (ACC)5 TGCTCCACTTTGTTCTTTCC TTGGCTATGGCAGTTTGAAT 54
lup-est6513 (CAA)5 ACCCTTCACAGACATCTCCA TAAGAGACAACATTGGGGGA 54
lup-est4113 (TGG)5 GGTGTTGTTGGAGAGAAGGA TGGTTTTGTTGGCTGAGATT 54
lup-est5513 (ACT)4 AATTCACACGTGGAGGATTC GAGGATCCAAGTTTTGTGGA 54
lup-est4322 (TCT)4 CTGTGCTGGACTTGGTATCC CTTTGCATATTGGTTTGCCT 54
lup-est3494 (CTT)6 TAGCACTGGTTTTCATTGCC CCTGGTGAACCTGATTGAAG 54
lup-est3994 (ACT)4 GGAAACTCCACAAAACCCTT TGTGATACACTGGAATCCCC 54
lup-est2194 (AAT)6 GTGATCAAGATGTGGCAGTG GTTTGTTGTTGGTGCAATGA 54
lup-est164 (CAA)3 AACAAGATTTGGGAGGTTCC CCATCCGGAAGACATATTCA 54
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