Genetic Diversity of Mung Bean Germplasms with Drought-tolerance at Seed Germination Stage
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摘要:
目的 鉴定56份绿豆种质种子萌发期耐旱性，明确其遗传特性。 方法 采用10%甘露溶液模拟干旱胁迫，以相对发芽率、相对发芽势和相对根长等10个指标作为耐旱性评价指标，通过主成分分析、隶属函数分析和多元分析方法鉴定种质耐旱性。同时基于SSR扩增对56份绿豆种质资源进行分子聚类，明确其遗传基础。 结果 干旱胁迫对萌发期10个耐旱性评价指标具有不同程度影响。主成分分析显示前三个综合指标贡献率分别为57.87%、16.42%、和11.42%，累积贡献率达到85.71 %，可以代表10个评价指标的变异信息。根据主成分分析计算得到的综合评价值D对56份种质耐旱性进行评价，鉴定出2份高耐、10份耐旱、30份中耐、13份敏感和1份高敏种质。基于SSR聚类将56份种质划分为3个主要类群，第1类群包含25份种质，表现为较耐旱。第2类群包含26份种质，遗传基础复杂，表现出不同程度的耐旱性。第三类群包含5份种质，表现为偏敏感。 结论 明确了56份绿豆种质萌发期的耐旱性及其遗传特性，为利用这些种质开展绿豆耐旱育种奠定了基础。 Abstract:Objective Drought-tolerance at seed germination stage and genetic diversity of 56 mung bean varieties were determined for resource evaluation. Method Selected mung bean seeds were treated with either water as control or 10% mannitol solution to simulate drought stress. Ten indicators on the draught-tolerance of the seeds at germination stage, such as relative germination rate and potential as well as root length of the sprouts, were monitored. The results were compared to categorize the germplasms using principal component, membership function, and multivariate analyses. Then, the amplified SSRs of the categorized germplasms provided the genetic basis for classifying the draught-resistant property of the seeds at germination stage. Result The top 3 indicators of draught-tolerance comprised 57.87%, 16.42%, and 11.42% separately, and 85.71% accumulatively, in contrition from a principal component analysis. And, at the calculated comprehensive evaluation value D, two of the 56 germplasms were graded highly tolerant, 10 tolerant, 30 moderately tolerant, 13 susceptible, and 1 highly susceptible to drought. From the cluster analysis on the SSR data, the germplasms were classified into 25 of relatively drought-resistant, 26 of varying degrees of drought-resistant due to complex genetic properties, and 5 of drought-susceptible. Conclusions The drought-tolerance of the 56 mung bean germplasms at seed germination stage and their genetic characteristics were determined and classified to facilitate future breeding and utilization of the resource. -
图 1 56份绿豆基于SSR遗传相似性的聚类图
注：HT：高耐；T：耐；MT：中耐；S：敏感；HS：高敏。
Figure 1. Dendrogram of 56 mung bean germplasms clustered by SSR genetic similarity
Note: HT: Highly tolerant; T: Tolerant; MT: Moderately tolerant; S: Susceptible; HS: Highly susceptible.
表 1 56份绿豆材料编号及基本信息
Table 1. Codes and basic information of 56 mung bean germplasms
编号
No.品种名称
Germplasm来源地
Origin01 中绿1号 Zhonglyu No.1 北京 Beijing 02 中绿2号 Zhonglyu No.2 北京 Beijing 03 中绿3号 Zhonglyu No.3 北京 Beijing 04 中绿4号 Zhonglyu No.4 北京 Beijing 05 中绿5号 Zhonglyu No.5 北京 Beijing 06 中绿6号 Zhonglyu No.6 北京 Beijing 07 中绿8号 Zhonglyu No.8 北京 Beijing 08 中绿9号 Zhonglyu No.9 北京 Beijing 09 中绿10号 Zhonglyu No.10 北京 Beijing 10 中绿11号 Zhonglyu No.11 北京 Beijing 11 中绿12号 Zhonglyu No.12 北京 Beijing 12 中绿13号 Zhonglyu No.13 北京 Beijing 13 中绿14号 Zhonglyu No.14 北京 Beijing 14 中绿15号 Zhonglyu No.15 北京 Beijing 15 中绿16号 Zhonglyu No.16 北京 Beijing 16 中绿17号 Zhonglyu No.17 北京 Beijing 17 鄂绿1号 Elyu No.1 湖北 Hubei 18 鄂绿2号 El2u No.2 湖北 Hubei 19 鄂绿3号 Elyu No.3 湖北 Hubei 20 鄂绿4号 Elyu No.4 湖北 Hubei 21 鄂绿5号 Elyu No.5 湖北 Hubei 22 潍绿1号 Weilyu No.1 山东 Shandong 23 潍绿5号 Weilyu No.5 山东 Shandong 24 潍绿7号 Weilyu No.7 山东 Shandong 25 潍绿8号 Weilyu No.8 山东 Shandong 26 潍绿9号 Weilyu No.9 山东 Shandong 27 潍绿05-08Weilyu 05-08 山东 Shandong 28 鹦哥3号 Budgerigar No.3 河北 Hebei 29 辽绿3号 Liaolyu No.3 辽宁 Liaoning 30 辽绿5号 Liaolyu No.5 辽宁 Liaoning 31 辽绿6号 Liaolyu No.6 辽宁 Liaoning 32 辽绿7号 Liaolyu No.7 辽宁 Liaoning 33 LPB02 河北 Hebei 34 LPB08 河北 Hebei 35 10L701 河北 Hebei 36 10L717 河北 Hebei 37 10L739 河北 Hebei 38 10L741 河北 Hebei 39 10L743 河北 Hebei 40 冀绿2号 Jilyu No.2 河北 Hebei 41 冀绿11号 Jilyu No.11 河北 Hebei 42 冀绿14号 Jilyu No.14 河北 Hebei 43 桂绿豆 L74 Guilyu L74 广西 Guangxi 44 晋7 Jin No.7 山西 Shanxi 45 晋8 Jin No.8 山西 Shanxi 46 保绿942 Baolyu No.942 河北 Hebei 47 横山大明绿豆
Yokoyama Daming Mung Bean陕西 Shaanxi 48 绿丰2号 Lyufeng No.2 吉林 Jilin 49 绿丰5号 Lyufeng No.5 吉林 Jilin 50 白绿9号 Bailyu No.9 吉林 Jilin 51 白绿10号 Bailyu No.10 吉林 Jilin 52 白绿11号 Bailyu No.11 吉林 Jilin 53 白绿12号 Bailyu No.12 吉林 Jilin 54 吉绿5号 Jilyu No.5 吉林 Jilin 55 吉绿7号 Jilyu No.7 吉林 Jilin 56 吉绿8号 Jilyu No.8 吉林 Jilin 
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表 2 用于遗传多样性分析的34对引物信息
Table 2. Thirty-four pairs of primers used for genetic diversity analysis
引物名称
Primer name上游引物（5′-3′）
Forward Primer1（5′-3′）下游引物（5′-3′）
Reverse Primer1（5′-3′）产物大小
Estimate product sizeVr1-37 ACGATTCGTTCACAGCCTCT TCATTTGTATTCTTTTTGTTGACTCC 250 Vr3-789 TCTTACCCACAAAACCTCCG GAGGGGCATATCACAGAGGA 200 Vr3-303 AACAACATCACCACCACCAC TTGTTCTAGTTGTTTTAGCCGTG 150 Vr1-1370 CTGCACAGCCAATCAACATA CAAGTTGGAGCTGCTTTTGA 220 Vr3-510 AACTCCAACATGCCGTTAGG TACCCATTCCCACATTGGTT 150 Vr3-598 GAGCCATGGGTATGTATAGATTTTT CCTCTCCCTCTTTTTCAGCC 160 Vr3-599 GAGGAGCCATGGGGTATGTA CCTCTCCCTCTTTTTCAGCC 170 Vr3-714 ACCTGCCATTTTTGTTCCAA AAAAACGCAAATTCTGGCTG 180 Vr2-629 TCTCCACTGCCTCCCTCTTA AAGGTGCTAATCCCACTCCA 120 Vr2-520 GGAATCACTTACAGAACCTCAACA AACTTATAGTGTGTGTGCTTGGG 110 Vr3-223 TGAGTTTGCTGTGGTTGAGG ATGACCGAGGTCCTCCTCTC 100 Vr3-310 AAGGGGGCTAGAAGTGGAAA CCACAGCAACCACAAATGAC 140 Vr2-64 AAGGTTGAGAATCGCAGGTG ACACGACGTGCTGCTAACTG 120 Vr3-135 CGTCCCATGCTTCTTCATCT TCATGGCATCCATCGATAAG 100 Vr3-605 GCGGTAATTGCTGGTTTTGT CAACCACAGTTGCAGTTGCT 150 Vr3-203 TAGCTTGGCCTACTTGCGAT TGGTCACAGGAGGTGTGGTA 110 Vr2-624 TCCTCTGATAGAAGCAGCTACG CATGCTCACTACTTTGTTGATCC 120 Vr3-627 TCAGGTCCCTGTTCATCAAA CCCATCTTTTCAATCAACAACA 160 Vr2-502 GCGGCCCTGGTAATACATTT CATGTGGTACATTGTGCAGTTG 120 Vr2-454 GATCACGCCCAGAGAACCTA CAATAGAGGGGCTGCTTCAG 110 Vr2-539 GGGGACGTTGAACTCTGTGT TCTGCATGTTTTGTACCGTTG 100 Vr2-627 TCGTGTAGTGAATAAAGCCAAA GAAAGAAGAGAAGAAGGTTTTAGGG 100 Vr2-208 CAAGTGCAAAGGGAAAAGGA TGTGATCACTGTCACCCACTT 120 Vr3-136 CTACGCGAGAGGAGCGATAG GGAAGAAACGCTCGAACAAG 110 Vr3-204 TAGGCCTCCTTGGCTGAGTA TCCATGAAGGTCATGACGAG 110 Vr1-1367 CTCCATTATTCCACTGCCCT AATGCTCAGTTCAGCCCAAT 220 Vr2-41 AAATTAGGGGATCCGCATGT TGCGTGTGTGTTTGTGTTTG 150 Vr2-298 CACTCACACACACGCAAACA AAATGGGGATTTTGGTTTGG 120 Vr2-462 GCAAGAGCTTTACCTAGGATGC CTCCAAAGACCCTCCATTCA 100 Vr2-126 ACGCACACGCTCAAATACAA AGAGAGAGTGTGTGTGGGGG 100 Vr2-518 GGAAGAAATTTGCAAGGGTTT GCCCTACGTGGTTACTTCCA 120 Vr2-436 GACCATTGGTTCAACGAGAAA CCTCAGTCTGGTACCGTAACTACTTC 100 Vr2-44 AAATTGGGTTAAAGGCCCAG TTTTCCTATCTTCGCTCCCTC 130 Vr2-534 GGCCTGCTTGGAGCAGAT ACACTTCACCCCTCCACTTG 100
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表 3 56份绿豆材料萌发期蒸馏水和10%甘露醇干旱胁迫下各指标的变化与比较
Table 3. Evaluation indicators (mean+SD) on seeds of 56 mung bean germplasms germinated under water or 10% mannitol treatment
处理
Treatment指标
Index发芽势
GV/%发芽率
GR/%根长
RL/cm胚轴长
PL/cm芽长
BL/cm芽鲜重
SFW/g芽干重
SDW/g发芽指数
GI活力指数
VI萌发指数
GICK 最大值 Max 100.00 100.00 17.20 15.80 40.20 0.57 0.33 26.04 33.55 2.50 最小值 Min. 80.00 80.00 0.00 0.00 0.00 0.00 0.00 7.63 10.70 0.81 平均值 Average 96.61±4.18 97.61±0.03 6.80±1.91 8.02±1.49 20.52±4.41 0.31±0.05 0.05±0.02 24.22±1.39 20.07±4.52 2.34±0.12 变异系数 CV（%） 4.33 3.53 0.28 0.19 0.21 0.15 0.43 0.06 0.23 0.05 10%甘露醇
10% Mannitol最大值 Max. 88.00 100.00 1.20 1.00 1.90 0.03 0.02 12.38 1.18 1.38 最小值 Min. 4.00 28.00 0.00 0.00 0.00 0.00 0.00 0.83 0.09 0.09 平均值 Average 44.32±20.21 64.74±21.56 0.47±0.08 0.63±1.21 0.95±0.17 0.01±0.003 0.002±0.001 6.53±2.63 0.64±0.27 0.69±0.29 变异系数 CV（%） 45.59 33.30 0.18 1.90 0.18 0.23 0.42 0.40 0.41 0.41 变化率
Rate of change平均值 Average −54.12 −33.67 −93.09 −92.14 −95.40 −99.00 −96.00 −73.04 −96.81 −70.51 变异系数 CV/% 952.89 84.33 35.71 900.00 −14.29 53.33 −2.33 566.67 78.26 720.00
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表 4 主成分分析
Table 4. Results of principal component analysis
测定指标
Indices因子 component 1 2 3 相对发芽率 RGR 0.910 −0.239 −0.218 相对发芽势 RGE 0.918 −0.213 −0.146 相对根长 RRL 0.324 0.858 0.052 相对胚轴长 RPL 0.658 −0.060 0.083 相对芽长 RBL 0.637 0.725 0.109 相对芽鲜重 RSFW 0.713 0.060 0.513 相对芽干重 RSDW 0.155 −0.299 0.867 活力指数 VI 0.941 0.214 −0.125 萌发胁迫指数 GSI 0.938 −0.253 −0.103 萌发耐旱指数 GDRI 0.934 −0.265 −0.101 特征值 Eigenvalue 5.787 1.642 1.142 百分率 Percentage/% 57.871 16.420 11.423 累计贡献率 Accumulative percentage/% 57.871 74.291 85.714
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表 5 56个绿豆品种的综合指标、指标权重、隶属函数和综合评价（D值）
Table 5. Comprehensive indices, index weights, subordinate function, and comprehensive evaluation D value on 56 mung bean germplasms
No. CI1 CI2 CI3 U（X）1 U（X）2 U（X）3 D 值
D value1 2.679 2.509 0.441 0.742 0.742 0.498 0.710 2 0.622 −1.224 −2.349 0.577 0.220 0.000 0.432 3 −6.557 −0.007 −1.525 0.000 0.390 0.147 0.095 4 −1.259 0.627 −0.340 0.426 0.479 0.359 0.427 5 −2.227 0.425 −0.792 0.348 0.451 0.278 0.358 6 −0.704 −0.463 1.202 0.470 0.327 0.634 0.465 7 −2.216 1.548 0.758 0.349 0.608 0.555 0.426 8 2.245 1.435 −0.508 0.707 0.592 0.329 0.635 9 0.386 1.419 −0.483 0.558 0.590 0.333 0.534 10 −2.188 −0.915 3.248 0.351 0.264 1.000 0.421 11 1.246 1.249 −0.090 0.627 0.566 0.404 0.586 12 −1.251 −1.150 −0.454 0.426 0.231 0.339 0.377 13 5.889 0.972 1.120 1.000 0.527 0.620 0.859 14 1.573 −0.724 −0.758 0.653 0.290 0.284 0.534 15 −0.472 0.726 0.962 0.489 0.493 0.591 0.503 16 4.183 2.932 2.365 0.863 0.801 0.842 0.848 17 2.506 −1.082 −0.726 0.728 0.240 0.290 0.576 18 2.679 −0.106 −0.887 0.742 0.377 0.261 0.608 19 2.601 −0.770 −2.319 0.736 0.284 0.005 0.552 20 2.928 0.335 0.800 0.762 0.438 0.563 0.673 21 −0.286 4.358 2.989 0.504 1.000 0.954 0.659 22 1.250 1.698 2.055 0.627 0.628 0.787 0.649 23 −0.333 −0.223 −1.455 0.500 0.360 0.160 0.428 24 −1.649 −0.433 −0.812 0.394 0.331 0.275 0.366 25 −3.638 −0.468 −0.786 0.235 0.326 0.279 0.258 26 −0.593 −0.540 −1.489 0.479 0.316 0.154 0.405 27 −1.221 −0.741 −0.228 0.429 0.288 0.379 0.395 28 1.087 0.072 1.360 0.614 0.401 0.663 0.580 29 −4.190 1.217 0.121 0.190 0.561 0.441 0.295 30 −3.001 0.870 −0.845 0.286 0.513 0.269 0.327 31 −1.273 −0.043 −1.040 0.425 0.385 0.234 0.392 32 −3.289 −0.265 −1.016 0.263 0.354 0.238 0.277 33 −2.924 −0.811 0.934 0.292 0.278 0.587 0.328 34 1.990 −0.885 −0.533 0.687 0.268 0.325 0.558 35 −1.364 0.726 −0.080 0.417 0.493 0.405 0.430 36 −1.882 0.913 −0.301 0.376 0.519 0.366 0.402 37 4.383 0.672 −0.124 0.879 0.485 0.397 0.739 38 −3.291 0.317 −0.409 0.262 0.436 0.347 0.307 39 0.938 −0.466 −1.082 0.602 0.326 0.226 0.499 40 2.895 −0.861 0.073 0.759 0.271 0.433 0.622 41 0.085 0.243 0.224 0.534 0.425 0.460 0.503 42 0.096 −0.116 0.764 0.535 0.375 0.556 0.507 43 1.952 −0.495 −0.844 0.684 0.322 0.269 0.559 44 −1.346 −0.476 1.277 0.419 0.325 0.648 0.431 45 −0.635 −1.171 0.065 0.476 0.228 0.431 0.422 46 0.888 −1.767 −1.170 0.598 0.145 0.211 0.460 47 −2.859 1.450 0.812 0.297 0.594 0.565 0.390 48 2.584 −2.461 1.735 0.734 0.048 0.730 0.602 49 0.440 −0.220 0.389 0.562 0.361 0.489 0.514 50 −1.012 −0.232 −0.400 0.446 0.359 0.348 0.416 51 0.254 −0.505 −0.223 0.547 0.321 0.380 0.481 52 3.272 −0.132 −0.364 0.790 0.373 0.355 0.652 53 −1.569 −0.874 −0.419 0.401 0.269 0.345 0.368 54 2.136 −2.802 −0.492 0.698 0.000 0.332 0.516 55 −1.066 −0.737 −0.126 0.441 0.288 0.397 0.406 56 0.506 −2.552 1.775 0.567 0.035 0.737 0.488
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表 6 34对SSR引物在绿豆中的多态性
Table 6. Polymorphisms of 34 pairs of SSR primers in mung beans
引物
Primer扩增总带数
Total number of amplifications基因型数量
Genotype No.多态性信息含量
PIC引物
Primer扩增总带数
Total number of amplifications基因型数量
enotype No.多态性信息含量
PICVr1-37 2 2 0.24 Vr3-627 6 2 0.24 Vr3-789 2 2 0.03 Vr2-502 2 2 0.28 Vr3-303 4 2 0.07 Vr2-454 2 2 0.29 Vr1-1370 4 2 0.10 Vr2-539 4 2 0.30 Vr3-510 6 2 0.10 Vr2-627 8 2 0.32 Vr3-599 6 2 0.12 Vr2-208 7 3 0.33 Vr3-598 4 2 0.15 Vr3-136 8 2 0.36 Vr3-714 4 2 0.17 Vr3-204 6 2 0.36 Vr2-629 5 3 0.18 Vr1-1367 4 2 0.36 Vr2-520 7 2 0.19 Vr2-41 6 2 0.37 Vr3-223 5 2 0.19 Vr2-298 11 2 0.37 Vr3-310 4 2 0.19 Vr2-462 3 2 0.37 Vr2-64 4 2 0.20 Vr2-126 8 2 0.40 Vr3-135 9 3 0.21 Vr2-518 5 3 0.40 Vr3-605 4 2 0.21 Vr2-436 10 2 0.52 Vr3-203 12 4 0.22 Vr2-44 7 3 0.58 Vr2-624 4 2 0.23 Vr2-534 8 4 0.67
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