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作物学报

作物学报 ›› 2026, Vol. 52 ›› Issue (2): 376-388.doi: 10.3724/SP.J.1006.2026.54061

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

绿豆种质资源苗期耐盐性鉴定及相关基因发掘

李诗晴1,2(), 王茜2, 王素华1, 张耀文2, 王丽侠1,*()   

  1. 1作物基因资源与育种全国重点实验室 / 中国农业科学院作物科学研究所, 北京 100081
    2山西农业大学农学院, 山西太谷030801
  • 收稿日期:2025-05-16 接受日期:2025-10-30 出版日期:2026-02-12 网络出版日期:2025-11-12
  • 通讯作者: *王丽侠 E-mail:wanglixia03@caas.cn
  • 作者简介:E-mail: leesqaq@163.com
  • 基金资助:
    国家重点研发计划项目(2023YFD1200705);国家重点研发计划项目(2019YFD1001303);国家自然科学基金项目(32241042);财政部和农业农村部国家现代农业产业技术体系建设专项(CARS-08)

Evaluation of salt tolerance at the seedling stage and related gene mining in mung bean germplasm resources

Li Shi-Qing1,2(), Wang Qian2, Wang Su-Hua1, Zhang Yao-Wen2, Wang Li-Xia1,*()   

  1. 1National Key Laboratory of Crop Gene Resources and Breeding / Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, Beijing 100081, China
    2College of Agronomy, Shanxi Agricultural University, Taigu 030801, Shanxi, China
  • Received:2025-05-16 Accepted:2025-10-30 Published:2026-02-12 Published online:2025-11-12
  • Contact: *Wang Li-Xia E-mail:wanglixia03@caas.cn
  • Supported by:
    National Key Research & Development Program of China(2023YFD1200705);National Key Research & Development Program of China(2019YFD1001303);National Natural Science Foundation of China(32241042);China Agriculture Research System of MOF and MARA(CARS-08)

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摘要：

土壤盐渍化对农业生产构成严峻威胁, 发掘绿豆耐盐种质对充分利用我国盐渍土地和发展绿豆产业均具有重要意义。本研究基于株高、地上部鲜重和根鲜重等13个性状, 对200份绿豆种质进行了苗期耐盐性(150 mmol L-1 NaCl)鉴定。结果表明, 单位体积根长的平均耐盐系数最大(0.806), 根分枝数最小(0.591); 盐胁迫下不同种质对根干重、单位体积根长和根鲜重的响应存在明显差异。根据盐害症状划分的盐害级别, 高耐盐种质16份, 盐高敏感种质12份, 华北区的耐盐种质占比最高; 结合重测序数据的全基因组关联分析(GWAS)共检测到67个与绿豆耐盐性显著关联的SNP位点, 共涉及348个候选基因, 包括99个具有功能注释的基因; 经单倍型分析验证, 基因Vradi04g09980、Vradi01g06290和Vradi08g02890可能与苗期耐盐性相关。上述结果为进一步开展绿豆耐盐种质创制及解析耐盐调控机制奠定了基础。

关键词: 绿豆, 苗期, 耐盐性, 全基因组关联分析, 种质资源, 单倍型分析

Abstract:

Soil salinization poses a serious threat to agricultural production. Exploring salt-tolerant germplasm in mung bean is crucial for the effective utilization of saline-alkali soils in China and for promoting the sustainable development of the mung bean industry. In this study, 200 mung bean germplasm accessions were evaluated for salt tolerance (150 mmol L-1 NaCl) at the seedling stage based on 13 phenotypic traits, including plant height, above-ground fresh weight, and root fresh weight. The results showed that the salt tolerance coefficient for root length per unit volume was the highest (0.806), while that for root branching was the lowest (0.591). Significant differences were observed among germplasms in their responses to salt stress, particularly in root dry weight, root length per unit volume, and root fresh weight. Based on salt damage symptoms, 16 highly salt-tolerant and 12 highly salt-sensitive germplasms were identified, with the highest proportion of salt-tolerant accessions found in the North China region. A genome-wide association study (GWAS) based on resequencing data identified 67 SNP loci significantly associated with salt tolerance, corresponding to 348 candidate genes, of which 99 had annotated functions. Among these, Vradi04g09980, Vradi01g06290, and Vradi08g02890 were validated through haplotype analysis as potentially associated with salt tolerance at the seedling stage. These findings provide a valuable foundation for the genetic improvement of salt tolerance in mung bean and for further elucidating the underlying regulatory mechanisms.

Key words: mung bean, seedling stage, salt tolerance, genome-wide association study (GWAS), germplasm resources, haplotype analysis

表1

200份绿豆种质的来源地信息"

地区
Region		 来源地
Location		 种质数
Number of germplasm		 地区
Region		 来源地
Location		 种质数
Number of germplasm
东北区
Northeast region		 中国黑龙江 Heilongjiang, China 3 华东区
East China region		 中国安徽 Anhui, China 10
中国吉林 Jilin, China 13 中国江苏 Jiangsu, China 22
中国辽宁 Liaoning, China 5 中国江西 Jiangxi, China 1
中国内蒙古 Inner Mongolia, China 25 中国山东 Shandong, China 8
华北区
North China
region		 中国北京 Beijing, China 2 中国台湾 Taiwan, China 1
中国河北 Hebei, China 11 华南区
South China region		 中国广西 Guangxi, China 2
中国山西 Shanxi, China 14 西北区
Northwest region		 中国宁夏 Ningxia, China 1
华中区
Central China region		 中国河南 Henan, China 30 中国陕西 Shaanxi, China 2
中国湖北 Hubei, China 15 中国新疆 Xinjiang, China 1
中国湖南 Hunan, China 8 南亚区
South Asia region		 世界蔬菜中心(原亚洲蔬菜研究发展中心)
World Vegetable Center (the former Asian Vegetable Research and Development Center)		 17
西南区
Southwest region		 中国重庆 Chongqing, China 1 东亚区
East Asia region		 日本 Japan 8

图1

盐胁迫下不同绿豆种质苗期盐害症状差异 1: 高耐盐; 3: 耐盐; 5: 中耐盐; 7: 盐敏感; 9: 盐高敏感。"

表2

盐胁迫下基于绿豆各性状的耐盐系数描述性统计"

性状
Trait		 最小值
Min.		 最大值
Max.		 平均值
Mean		 变异系数
CV (%)
株高 PH 0.386 1.172 0.778 24.01
地上部鲜重 AFW 0.207 1.125 0.712 29.01
根鲜重 RFW 0.157 1.193 0.693 33.23
地上部干重 ADW 0.196 1.160 0.636 30.12
根干重 RDW 0.134 1.194 0.709 35.40
根长 RL 0.310 1.097 0.750 26.73
根表面积 RSA 0.410 1.192 0.682 28.21
根平均直径 RAD 0.165 1.189 0.660 29.62
单位体积根长 RLPV 0.470 1.392 0.806 31.10
根体积 RV 0.229 1.113 0.711 29.07
根尖数 RT 0.442 1.198 0.675 30.90
根分枝数 RF 0.403 1.061 0.591 26.43
根交叉数 RC 0.500 1.083 0.656 16.06

表3

盐胁迫下绿豆表型性状及盐害级别间的相关性分析"

性状
Trait		 PH AFW RFW ADW RDW RL RSA RAD RLPV RV RT RF RC
AFW 0.76**
RFW 0.23** 0.46**
ADW 0.37** 0.72** 0.63**
RDW -0.05 0.24** 0.71** 0.55**
RL 0.19** 0.43** 0.72** 0.57** 0.58**
RSA 0.29** 0.50** 0.77** 0.60** 0.60** 0.84**
RAD 0.03 -0.06 -0.18* -0.21** -0.21** -0.62** -0.46**
RLPV 0.26** 0.46** 0.71** 0.58** 0.58** 0.84** 0.97** -0.59**
RV 0.30** 0.54** 0.83** 0.61** 0.61** 0.80** 0.93** -0.21** 0.85**
RT 0.12 0.31** 0.49** 0.46** 0.45** 0.68** 0.75** -0.61** 0.81** 0.61**
RF 0.19** 0.36** 0.63** 0.51** 0.55** 0.74** 0.86** -0.56** 0.88** 0.72** 0.71**
RC 0.25** 0.37** 0.53** 0.45** 0.46** 0.67** 0.75** -0.52** 0.78** 0.62** 0.61** 0.93**
SDL -0.23** -0.38** -0.76** -0.41** -0.50** -0.74** -0.84** 0.24** -0.78** -0.87** -0.62** -0.59** -0.45**

图2

绿豆苗期各耐盐系数全基因组关联分析的曼哈顿图和Q-Q图 a: 株高; b: 地上部鲜重; c: 根鲜重; d: 地上部干重; e: 根干重; f: 根长; g: 根表面积; h: 根平均直径; i: 单位体积根长; j: 根体积; k: 根尖数; l: 根分枝数; m: 根交叉数。"

附表1

绿豆种质耐盐鉴定候选基因"

关联性状
Associated traits		 基因ID
Gene ID		 功能注释
Functional annotation
RAD Vradi01g04810 非特异性脂质转移蛋白2样蛋白 Non-specific lipid-transfer protein 2-like
RAD Vradi01g04850 未知功能蛋白LOC106773804 Uncharacterized protein LOC106773804
RAD Vradi01g04920 未知功能蛋白LOC106769195 Uncharacterized protein LOC106769195
RLPV Vradi01g06280 未知功能蛋白LOC106769919 Uncharacterized protein LOC106769919
RLPV Vradi01g06290 ABC转运蛋白G家族成员11 ABC transporter G family member 11
RLPV Vradi01g06370 未知功能蛋白LOC106766299 Uncharacterized protein LOC106766299
RL Vradi01g08580 未知功能蛋白LOC106765526 Uncharacterized protein LOC106765526
RL Vradi01g08590 60S核糖体蛋白L36 60S ribosomal protein L36
RL Vradi01g08600 未知功能蛋白LOC106764536 Uncharacterized protein LOC106764536
RL Vradi01g08640 核仁复合体蛋白2同源物 Nucleolar complex protein 2 homolog
RL Vradi01g14660 CEN样蛋白1 CEN-like protein 1
RAD Vradi04g04000 木葡聚糖内切转葡糖基酶/水解酶 Xyloglucan endotransglucosylase/hydrolase
RAD Vradi04g04010 ω-羟基棕榈酸O-阿魏酰转移酶样蛋白
Omega-hydroxypalmitate O-feruloyl transferase-like
RAD Vradi04g04030 GPI转氨酶组分PIG-S GPI transamidase component PIG-S
RAD Vradi04g04040 未知功能蛋白LOC106758832 Uncharacterized protein LOC106758832
RV Vradi04g09650 转录因子BIM2 Transcription factor BIM2
RFW/RV Vradi04g09660 O-岩藻糖基转移酶27 O-fucosyltransferase 27
RFW/RV Vradi04g09670 磷脂酰肌醇4-激酶γ 7样蛋白 Phosphatidylinositol 4-kinase gamma 7-like
RFW/RV Vradi04g09700 F-box/kelch重复蛋白At5g15710 F-box/kelch-repeat protein At5g15710
RFW/RV Vradi04g09740 含PPR (五肽重复)结构域的蛋白At2g31400, 叶绿体型
Pentatricopeptide repeat-containing protein At2g31400, chloroplastic
RFW/RV Vradi04g09790 含PPR (五肽重复)结构域的蛋白At3g02330, 线粒体型
Pentatricopeptide repeat-containing protein At3g02330, mitochondrial
RFW/RV Vradi04g09800 碱性亮氨酸拉链蛋白43 Basic leucine zipper 43
RFW/RV Vradi04g09810 6-磷酸葡萄糖酸脱氢酶, 脱羧型
6-phosphogluconate dehydrogenase, decarboxylating
RFW/RV Vradi04g09880 含PPR (五肽重复)结构域的蛋白At3g02490, 线粒体样
Pentatricopeptide repeat-containing protein At3g02490, mitochondrial-like
RFW/RV Vradi04g09890 短链脱氢酶/还原酶ATA1 Short-chain dehydrogenase reductase ATA1
RFW/RV Vradi04g09920 YIF1B样蛋白 Protein YIF1B-like
RFW/RV Vradi04g09940 可能的2,3-二磷酸甘油酸非依赖性磷酸甘油酸变位酶
Probable 2,3-bisphosphoglycerate-independent phosphoglycerate mutase
RFW/RV Vradi04g09970 FLX样蛋白2 Protein FLX-like 2
RFW/RV Vradi04g09980 毛状体双折射样蛋白19 Protein trichome birefringence-like 19
RC/RF Vradi05g05350 绒毡层决定子蛋白1异构体X2
Protein tapetum determinant 1 isoform X2
AFW Vradi05g14520 WUSCHEL相关同源异形盒8样蛋白 WUSCHEL-related homeobox 8-like
AFW Vradi05g14540 RING型E3泛素连接酶 RING-type E3 ubiquitin transferase
AFW Vradi05g14570 可能的蛋白磷酸酶2C 27 Probable protein phosphatase 2C 27
AFW Vradi05g14580 3-磷酸甘油醛脱氢酶 Glyceraldehyde-3-phosphate dehydrogenase
AFW Vradi05g14630 多叶绿体分裂位点蛋白1
Protein multiple chloroplast division site 1
AFW Vradi05g14660 AT-hook基序核定位蛋白10 AT-hook motif nuclear-localized protein 10
AFW Vradi05g14670 含psbP结构域的蛋白3, 叶绿体型
psbP domain-containing protein 3, chloroplastic
AFW Vradi05g14680 SWI/SNF复合物亚基SWI3B SWI/SNF complex subunit SWI3B
AFW Vradi05g14700 未知功能蛋白LOC106759854 Uncharacterized protein LOC106759854
AFW Vradi05g14750 BURP结构域蛋白USPL1 BURP domain protein USPL1
RC/RF Vradi06g08010 PRA1家族蛋白 PRA1 family protein
RC Vradi07g04190 含CBS结构域的蛋白CBSX2, 叶绿体型
CBS domain-containing protein CBSX2, chloroplastic
RC Vradi07g04200 多聚腺苷酸结合蛋白 Polyadenylate-binding protein
RC Vradi07g04260 At1g66480样未知功能蛋白 Uncharacterized protein At1g66480-like
RC Vradi07g04280 推定的1-磷脂酰肌醇-3-磷酸-5-激酶FAB1C
Putative 1-phosphatidylinositol-3-phosphate 5-kinase FAB1C
RC Vradi07g04300 几丁质结合凝集素1 Chitin-binding lectin 1
RC Vradi07g04430 SEC12样蛋白2异构体X1 SEC12-like protein 2 isoform X1
RC Vradi07g04470 10 kD伴侣蛋白样蛋白 10 kD chaperonin-like
RC Vradi07g04490 植烷酰辅酶A双加氧酶 Phytanoyl-CoA dioxygenase
RAD Vradi08g02890 铜转运蛋白CCH Copper transport protein CCH
RAD Vradi08g02900 胞吐复合体组分 Exocyst complex component
RAD Vradi08g02910 Polycomb (多梳蛋白)增强子样蛋白 Enhancer of polycomb-like protein
RAD Vradi08g02950 开花位点D样蛋白 Protein flowering locus D-like protein
RAD Vradi08g02960 未知功能蛋白LOC106772352 Uncharacterized protein LOC106772352
RAD Vradi08g03010 香豆酰辅酶A: 花青素3-O-葡萄糖苷-6''-O-香豆酰转移酶1样异构体X2
Coumaroyl-CoA: anthocyanidin 3-O-glucoside-6''-O-coumaroyltransferase 1-like isoform X2
RC/RT Vradi08g10930 未知功能蛋白LOC106771460异构体X1
Uncharacterized protein LOC106771460 isoform X1
RC/RT Vradi08g10970 MYB108样转录因子 Transcription factor MYB108-like
RC/RT Vradi08g10990 类似于ADP核糖基化因子1 Similar to ADP-ribosylation factor 1
RT Vradi08g11000 NIM1互作蛋白1样蛋白 Protein NIM1-INTERACTING 1-like
RT Vradi08g11010 未知功能蛋白 LOC106771693 Uncharacterized protein LOC106771693
PH Vradi09g02010 CPN60样分子伴侣蛋白2, 线粒体型
Chaperonin CPN60-like 2, mitochondrial
PH Vradi09g02040 未知功能蛋白 LOC106773708 Uncharacterized protein LOC106773708
PH Vradi09g02060 推定的F-box/FBD/LRR重复蛋白At1g78840异构体X1
Putative F-box/FBD/LRR-repeat protein At1g78840 isoform X1
PH Vradi09g02080 丝氨酸/苏氨酸蛋白磷酸酶7长型同源物
Serine/threonine-protein phosphatase 7 long form homolog
PH Vradi09g02120 含IQ结构域的IQM1样蛋白 IQ domain-containing protein IQM1-like
PH Vradi09g02170 酪蛋白激酶I样蛋白 Casein kinase I-like
PH Vradi09g02190 甘氨酰肽N-十四烷酰基转移酶 Glycylpeptide N-tetradecanoyltransferase
RAD Vradi09g05880 含PPR (五肽重复)结构域的蛋白At1g02150
Pentatricopeptide repeat-containing protein At1g02150
RAD Vradi09g05900 含PPR (五肽重复)结构域的蛋白At1g71460, 叶绿体样
Pentatricopeptide repeat-containing protein At1g71460, chloroplastic-like
RAD Vradi09g05920 延长子复合物蛋白4异构体X1 Elongator complex protein 4 isoform X1
RAD Vradi09g05930 含PPR (五肽重复)结构域的蛋白 At1g15510, 叶绿体型
Pentatricopeptide repeat-containing protein At1g15510, chloroplastic
RAD Vradi09g05940 色氨酸-tRNA连接酶, 细胞质型 Tryptophan-tRNA ligase, cytoplasmic
RAD Vradi09g05950 赖氨酸-tRNA连接酶, 叶绿体/线粒体型
Lysine-tRNA ligase, chloroplastic/mitochondrial
RAD Vradi09g05970 含ELMO结构域的A样蛋白异构体X1
ELMO domain-containing protein A-like isoform X1
RAD Vradi09g05980 细胞周期蛋白 Cyclin
RAD Vradi09g06060 S.2样受体蛋白激酶 Receptor like protein kinase S.2-like
RL Vradi09g06870 重金属P1B型ATP酶, 铜转运ATP酶, 控制水稻籽粒中铜的积累
Heavy metal P1B-type ATPase, Cu-transporting ATPase, control of Cu accumulation in rice grai
RL Vradi09g06880 可能的葡聚糖内切-1,3-β-葡萄糖苷酶A6
Probable glucan endo-1,3-beta-glucosidase A6
RL Vradi09g06900 蛋白激酶超家族蛋白 Protein kinase superfamily protein
RC Vradi11g00960 未知功能蛋白 LOC106776934 Uncharacterized protein LOC106776934
RC Vradi11g00980 基质加工肽酶, 叶绿体型 Stromal processing peptidase, chloroplastic
RC Vradi11g00990 组蛋白乙酰转移酶MCC1异构体X1
Histone acetyltransferase MCC1 isoform X1
RC Vradi11g01000 未知功能蛋白 LOC106776944 Uncharacterized protein LOC106776944
RC Vradi11g01010 热应激转录因子A-1b样蛋白 Heat stress transcription factor A-1b-like
RC Vradi11g01030 TOM1样蛋白2 TOM1-like protein 2
RC Vradi11g01040 纤维素合酶样蛋白D2 Cellulose synthase-like protein D2
RC Vradi11g01060 DPB样转录因子 Transcription factor-like protein DPB
RC Vradi11g01200 未知功能蛋白LOC106777204 Uncharacterized protein LOC106777204
RC Vradi11g01210 细胞分裂周期蛋白20.1, APC复合体辅助因子样蛋白
Cell division cycle 20.1, cofactor of APC complex-like
RC Vradi11g01220 HVA22样蛋白 HVA22-like protein
RC Vradi11g01230 U4/U6小核核糖核蛋白Prp3样蛋白
U4/U6 small nuclear ribonucleoprotein Prp3-like
RC Vradi11g01240 氧还铁蛋白家族蛋白 Rubredoxin family protein
RC Vradi11g01270 未知功能蛋白 LOC106777046 Uncharacterized protein LOC106777046
RC Vradi11g01280 植物镉抗性蛋白10 Protein plant cadmium resistance 10
RC Vradi11g01290 含PPR (五肽重复)结构域的蛋白At1g71420
Pentatricopeptide repeat-containing protein At1g71420
RC Vradi11g01300 含PPR (五肽重复)结构域的蛋白 At3g18110, 叶绿体型
Pentatricopeptide repeat-containing protein At3g18110, chloroplastic
RC Vradi11g01310 琥珀酸脱氢酶组装因子1, 线粒体型
Succinate dehydrogenase assembly factor 1, mitochondrial
RC Vradi11g01340 细胞色素c Cytochrome c
RC Vradi11g01350 乙烯应答转录因子ERF022 Ethylene-responsive transcription factor ERF022

图3

Vradi04g09980编码区的核苷酸多态性(a)、单倍型间根鲜重差异(b)及根体积差异(c) RFW: 根鲜重; RV: 根体积。*和**分别表示在0.05和0.01水平差异显著。"

图4

Vradi01g06290编码区的核苷酸多态性(a)与单倍型间单位体积根长差异(b) RLPV: 单位体积根长。**表示在0.01水平差异显著。"

图5

Vradi08g02890编码区的核苷酸多态性(a)与单倍型间根平均直径差异(b) RAD: 根平均直径。*表示在0.05水平差异显著。"

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