模拟涝害胁迫下大豆生理和转录组分析

doi:10.7668/hbnxb.20195959

摘要/Abstract

摘要：

涝害胁迫严重影响大豆生产,为探讨涝害胁迫对大豆生长发育的影响,解析其响应机制,以淮北地区种植的4个大豆种质为试验材料,通过模拟三叶期涝害处理,结合产量、生理指标和转录组学分析探究大豆耐涝机制。研究表明,涝害胁迫显著降低涝害敏感品种徐豆18的干物质积累和氮吸收效率,单株产量损失30.23%,而耐涝品种淮豆13单株产量只损失16.77%,通过稳定根系氮吸收和维持根系干物质积累表现出较强适应性。转录本测定结果表明,涝害胁迫后耐涝品种淮豆13差异表达基因显著富集于次生代谢产物的生物合成、光合系统和抗氧化相关通路;而涝害敏感型徐豆18差异表达基因主要富集于激素转导、光合作用、过氧化物酶体等相关通路等。共鉴定到148个在耐涝与敏感品种间存在显著表达差异的涝害胁迫响应基因(DEGs),主要富集在光合作用、次生代谢物和脂肪酸代谢途径,转录因子包含AP2/ERF-ERF、C3H、ARR-B、NAC、WRKY等,综上筛选到次生代谢途径中可能存在与耐涝特性相关的转录因子或基因。

关键词: 大豆, 涝害胁迫, 生理代谢, 转录组

Abstract:

Waterlogging stress severely impacts soybean production.To investigate the effects of waterlogging stress on soybean growth and development and elucidate its response mechanisms,this study employed four soybean germplasms cultivated in the Huaibei region as experimental materials.Through simulated waterlogging treatment at the V2 stages,combined with yield analysis,physiological indices,and transcriptomic profiling,the study explores the waterlogging tolerance mechanisms in soybean.The results demonstrated that waterlogging stress significantly reduced dry matter accumulation and nitrogen uptake efficiency in the waterlogging-sensitive cultivar Xudou 18,resulting in a yield loss of 30.23%.In contrast,the waterlogging-tolerant cultivar Huaidou 13 exhibited stronger adaptability with only a 16.77% yield loss,attributed to stabilized root nitrogen absorption and maintained root dry matter accumulation.Transcriptomic analysis revealed that differentially expressed genes(DEGs)in Huaidou 13 under waterlogging stress were significantly enriched in pathways that related to the biosynthesis of secondary metabolites,photosynthetic systems,and antioxidant activity.Conversely,DEGs in Xudou 18 were predominantly associated with hormone transduction,photosynthesis,and peroxisome-related pathways.A total of 148 genotype-specific DEGs were identified between cultivars with contrasting waterlogging tolerance,primarily enriched in photosynthesis,secondary metabolite biosynthesis,and fatty acid metabolism.Identified transcription factors included members of the AP2/ERF-ERF,C3H,ARR-B,NAC,and WRKY families.In summary,transcription factors or genes in secondary metabolic pathways that may be related to waterlogging tolerance were screened.

Key words: Soybean, Waterlogging stress, Physiological metabolism, Transcriptome

中图分类号:

赵志鑫, 刘宁宁, 徐海风, 王亚琪, 傅蒙蒙, 李曙光, 余希文. 模拟涝害胁迫下大豆生理和转录组分析[J]. 华北农学报, 2025, 40(S1): 9-19. doi: 10.7668/hbnxb.20195959.

ZHAO Zhixin, LIU Ningning, XU Haifeng, WANG Yaqi, FU Mengmeng, LI Shuguang, YU Xiwen. Physiological and Transcriptomic Analysis of Soybean under Waterlogging Stress[J]. Acta Agriculturae Boreali-Sinica, 2025, 40(S1): 9-19. doi: 10.7668/hbnxb.20195959.

http://www.hbnxb.net/CN/Y2025/V40/IS1/9

图/表 13

表1 试验材料的处理组合

Tab.1 Treatment combination of test materials

品种名称 Test materials	正常水分处理 (CK) Normal treatment of water(CK)	涝害处理(LH) Waterlogging treatment (LH)
淮豆13 Huaidou 13	CK-G01	LH-G01
淮豆4号Huaidou 4	CK-G02	LH-G02
Forrest	CK-G03	LH-G03
徐豆18 Xudou 18	CK-G04	LH-G04

表2 涝害胁迫对大豆产量及产量相关因素的影响

Tab.2 Effects of water logging stress on soybean yield and yield-related factors

品种 Test materials	对照处理CK			涝害处理LH
品种 Test materials	百粒质量/g 100-seed weight	单株粒数 Seed number per plant	单株产量/g Yield per plant	百粒质量/g 100-seed weight	单株粒数 Seed number per plant	单株产量/g Yield per plant
G01	18.36±2.56a	67.25±11.69a	12.22±3.15a	18.24±4.98a	57.00±9.24b	10.17±4.26ab
G02	20.42±3.54a	79.80±13.59a	14.62±3.24a	20.40±3.47a	57.75±10.66b	10.18±4.12b
G03	16.05±2.72a	71.00±11.32a	11.35±2.87a	16.57±3.68a	53.40±8.63b	8.53±3.47b
G04	19.47±3.16a	81.00±17.65a	14.75±3.87a	16.58±3.29b	69.92±14.23b	10.29±2.69b

图1 涝害胁迫对大豆干物质积累量的影响不同小写字母表示不同品种处理间差异显著(P<0.05)。图2-3同。

Fig.1 Effects of water logging stress on dry matter accumulation of soybean The different lowercase letters indicate significant differences among materials and treatments(P<0.05).The same as Fig.2-3.

图2 涝害胁迫对大豆氮素积累量的影响

Fig.2 Effects of water logging stress on nitrogen accumulation in soybean

图3 涝害胁迫对大豆丙二醛含量的影响

Fig.3 Effects of water logging stress on MDA content in soybean

表3 涝害胁迫对大豆抗氧化酶活性的影响

Tab.3 Effect of water logging stress on antioxidant enzyme activity of soybean

品种 Test materials	SOD/(U/g)		POD/(U/g)		CAT/(nmol/mg)
品种 Test materials	CK	LH	CK	LH	CK	LH
G01	129.88±20.00a	102.40±26.90b	8 054.67±886.39a	6 153.33±416.64b	688.36±211.60a	571.56±136.17b
G02	197.93±10.62b	220.50±40.63a	10 066.67±3 035.85a	10 625.30±1 945.02a	723.48±6.28a	807.72±198.62a
G03	147.87±10.66a	122.98±6.83a	10 256.00±1 385.93a	10 524.33±682.64a	1 844.13±97.17b	2 136.88±99.06a
G04	354.25±6.02b	371.48±15.72a	6 096.00±75.70b	7 128.00±1 509.57a	347.58±17.69b	528.46±136.55a

表4 测序数据质量评估统计

Tab.4 Statistical table of sequencing data quality evaluation

样品 Samples	过滤后数据 Clean reads	匹配到的数据 Total mapped reads	匹配率/% Total mapped reads	GC含量/% GC content	Q30碱基比例/% Q30 content
CK-G01-1	44 387 312	42 697 614	96.19	44.54	96.70
CK-G01-2	53 714 074	51 809 156	96.45	44.60	96.84
CK-G04-1	45 732 788	43 709 270	95.58	43.78	97.04
CK-G04-2	55 330 950	53 427 847	96.56	44.56	95.04
LH-G01-1	52 922 238	48 273 942	91.22	45.05	90.34
LH-G01-2	41 479 404	36 768 240	88.64	46.38	96.91
LH-G04-1	40 252 692	38 244 336	95.01	44.07	96.84
LH-G04-2	50 595 966	48 542 084	95.94	43.98	96.65

图4 淮豆13和徐豆18叶片转录组差异表达分析 A.样本热图;B.差异表达基因(DEGs)的数量柱状图;C.差异基因韦恩图。

Fig.4 Differentially expressed analysis of Huaidou 13 and Xudou 18 of leaf transcriptomic A.Heatmap of sample;B.Histogram of the number of differentially expressed genes;C.Venn.

图5 淮豆13(A)和徐豆18(B)的DEGs GO功能富集分析 A:1.细胞过程;2.代谢过程;3.单机体过程;4.生物调控;5.定位;6.生物过程调控;7.刺激响应;8.细胞组分组织或装配;9.多细胞生物过程;10.多机体互作过程;11.信号转导;12.生殖;13.发育过程;14.生殖过程;15.解毒作用;16.生物过程负调控;17.运动;18.生物过程正调控;19.免疫系统过程;20.行为;21.生物黏附;22.生长;23.细胞杀伤;24.节律过程;25.催化活性;26.结合;27.转运蛋白活性;28.核酸结合转录因子活性;29.电子载体活性;30.结构分子活性;31.抗氧化活性;32.分子功能调控因子活性;33.信号转导因子活性;34.分子转导因子活性;35.翻译调控因子活性;36.转录因子活性(蛋白质结合);37.金属伴侣蛋白活性;38.细胞部分;39.细胞;40.膜;41.膜部分;42.大分子复合物;43.细胞器;44.细胞器部分;45.胞外区域;46.膜包被腔室;47.病毒体;48.病毒体部分;49.突触;50.病毒包含体;51.超分子纤维;52胞外区域部分;53.突触部分;54.类核;55.胞外基质;56.共质体;57.细胞连接。B:1.细胞过程;2.代谢过程;3.单机体过程;4.定位;5.刺激响应;6.生物调控;7.生物过程调控;8.细胞组分组织或装配;9.多机体互作过程;10.信号转导;11.生殖;12.多细胞生物过程;13.发育过程;14.生殖过程;15.生物过程负调控;16.解毒作用;17.生物过程正调控;18.运动;19.免疫系统过程;20.生长;21.生物黏附;22.节律过程;23.行为;24.突触前过程(参与突触传递);25.细胞杀伤;26.催化活性;27.结合活性;28.转运蛋白活性;29.核酸结合转录因子活性;30.结构分子活性;31.抗氧化活性;32.分子功能调控因子活性;33.电子载体活性;34.信号转导因子活性;35.分子转导因子活性;36.转录因子活性(蛋白质结合);37.金属伴侣蛋白活性;38.蛋白质标签;39.翻译调控因子活性;40.细胞;41.细胞部分;42.膜;43.大分子复合物;44.细胞器;45.膜部分;46.细胞器部分;47.胞外区域;48.膜包被腔室;49.病毒体;50.病毒体部分;51.胞外区域部分;52.超分子纤维;53.胞外基质;54.病毒包含体;55.共质体;56.细胞连接;57.其他生物体;58.其他生物体部分;59.突触;60.突触部分。

Fig.5 GO enrichment analysis of DEGs in Huaidou 13(A)and Xudou 18(B) A:1.Cellular process;2.Metabolic process;3.Single-organism process;4.Biological regulation;5.Localization;6.Regulation of biological process;7.Rresponse to stimulus;8.Cellular component organization or biogenesis;9.Multicellular organismal process;10.Multi-organism process;11.Signaling;12.Reproduction;13.Developmental process;14.Reproductive process;15.Detoxification;16.Negative regulation of biological process;17.Locomotion;18.Positive regulation of biological process;19.Immune system process;20.Behavior;21.Biological adhesion;22.Growth;23.Cell killing;24.Rhythmic process;25.Catalytic activity;26.Binding;27.Transporter activity;28.Nucleic acid binding transcription factor activity;29.Electron carrier activity;30.Structural molecule activity;31.Antioxidant activity;32.Molecular function regulator;33.Signal transducer activity;34.Molecular transducer activity;35.Translation regulator activity;36.Transcription factor activity,protein binding;37.Metallochaperone activity;38.Cell part;39.Cell;40.Membrane;41.Membrane part;42.Macromolecular complex;43.Organelle;44.Organelle part;45.Extracellular region;46.Membrane-enclosed lumen;47.Virion;48.Virion part;49.Synapse;50.Viral occlusion body;51.Supramolecular fiber;52.Extracellular region part;53.Synapse part;54.Nucleoid;55.Extracellular matrix;56.Symplast;57.Cell junction.B:1.Ccellular process;2.Metabolic process;3.Single-organism process;4.Localization;5.Response to stimulus;6.Biological regulation;7.Regulation of biological process;8.Cellular component organization or biogenesis;9.Multi-organism process;10.Signaling;11.Reproduction;12.Multicellular organismal process;13.Developmental process;14.Reproductive process;15.Negative regulation of biological process;16.Detoxification;17.Positive regulation of biological process;18.Locomotion;19.Immune system process;20.Growth;21.Biological adhesion;22.Rhythmic process;23.Behavior;24.Presynaptic process involved in synaptic transmission;25.Cell killing;26.Catalytic activity;27.Binding;28.Transporter activity;29.Nucleic acid binding transcription factor activity;30.Structural molecule activity;31.Antioxidant activity;32.Molecular function regulator;33.Electron carrier activity;34.Signal transducer activity;35.Molecular transducer activity;36.Transcription factor activity,protein binding;37.Metallochaperone activity;38.Protein tag;39.Translation regulator activity;40.Cell;41.Cell part;42.Membrane;43.Mcromolecular complex;44.Organelle;45.Membrane part;46.Organelle part;47.Extracellular region;48.Membrane-enclosed lumen;49.Virion;50.Virion part;51.Extracellular region part;52.Supramolecular fiber;53.Extracellular matrix;54.Viral occlusion body;55.Symplast;56.Cell junction;57.Other organism;58.Other organism part;59.Synapse;60.Synapse part.

图6 淮豆13(A)和徐豆18(B)富集前20的GO分类 A:1.GO:0016020膜;2.GO:0009535叶绿体类囊体;3.GO:0055035质体类囊体膜;4.GO:0045156电子转运蛋白活性(光合循环电子传递途径);5.GO:0016168叶绿素结合活性;6.GO:0016491氧化还原酶活性;7.GO:0009534叶绿体类囊体;8.GO:0003700转录因子活性(序列特异性DNA结合);9.GO:0001071核酸结合转录因子活性;10.GO:0031976质体类囊体;11.GO:0015267通道活性;12.GO:0022803被动跨膜转运蛋白活性;13.GO:0009772光系统Ⅱ光合电子传递;14.GO:0009539光系统Ⅱ反应中心;15.GO:0016021膜内在组分;16.GO:0031224膜内在组分;17.GO:0005216离子通道活性;18.GO:0055114氧化还原过程;19.GO:0022838底物特异性通道活性;20.GO:000442羟甲基戊二酰-CoA合酶活性。B:1.GO:0006468蛋白质磷酸化;2.GO:0016310磷酸化;3.GO:0004672蛋白激酶活性;4.GO:0016301激酶活性;5.GO:0016773磷酸转移酶活性(以醇基为受体);6.GO:0003824催化活性;7.GO:0016740转移酶活性;8.GO:0003700转录因子活性(序列特异性DNA结合);9.GO:0001071核酸结合转录因子活性;10.GO:0016772转移酶活性(转移含磷基团);11.GO:0016798水解酶活性(作用于糖苷键);12.GO:0016667氧化还原酶活性(作用于供体硫基);13.GO:0006629脂质代谢过程;14.GO:0006464细胞蛋白质修饰过程;15.GO:0036211蛋白质修饰过程;16.GO:0004553水解酶活性(水解O-糖苷化合物);17.GO:0006793磷代谢过程;18.GO:0009654光系统Ⅱ放氧复合物;19.GO:0015035蛋白质二硫键氧化还原酶活性;20.GO:0042221化学物质响应。

Fig.6 The top 20 functional classification of GO enrichment analysis of Huaidou 13(A)and Xudou 18(B) A:1.GO:0016020 Membrane;2.GO:0009535 Chloroplast thylakoid membrane;3.GO:0055035 Plastid thylakoid membrane;4.GO:0045156 Electron transporter,transferring electrons within the cyclic electron transport pathway of photosynthesis activity;5.GO:0016168 Chlorophyll binding;6.GO:0016491 Oxidoreductase activity;7.GO:0009534 Chloroplast thylakoid;8.GO:0003700 Transcription factor activity,sequence-specific DNA binding;9.GO:0001071 Nucleic acid binding transcription factor activity;10.GO:0031976 Plastid thylakoid;11.GO:0015267 Channel activity;12.GO:0022803 Passive transmembrane transporter activity;13.GO:0009772 Photosynthetic electron transport in photosystem Ⅱ;14.GO:0009539 Photosystem Ⅱ reaction center;15.GO:0016021 Integral component of membrane;16.GO:0031224 Intrinsic component of membrane;17.GO:0005216 Ion channel activity;18.GO:0055114 Oxidation-reduction process;19.GO:0022838 Substrate-specific channel activity;20.GO:0004421 Hydroxymethylglutaryl-CoA synthase activity.B:1.GO:0006468 Protein phosphorylation;2.GO:0016310 Phosphorylation;3.GO:0004672 Protein kinase activity;4.GO:0016301 Kinase activity;5.GO:0016773 Phosphotransferase activity,alcohol group as acceptor;6.GO:0003824 Catalytic activity;7.GO:0016740 Transferase activity;8.GO:0003700 Transcription factor activity,sequence-specific DNA binding;9.GO:0001071 Nucleic acid binding transcription factor activity;10.GO:0016772 Transferase activity,transferring phosphorus-containing groups;11.GO:0016798 Hydrolase activity,acting on glycosyl bonds;12.GO:0016667 Oxidoreductase activity,acting on a sulfur group of donors;13.GO:0006629 Lipid metabolic process;14.GO:0006464 Cellular protein modification process;15.GO:0036211 Protein modification process;16.GO:0004553 Hydrolase activity,hydrolyzing O-glycosyl compounds;17.GO:0006793 Phosphorus metabolic process;18.GO:0009654 Photosystem Ⅱ oxygen evolving complex;19.GO:0015035 Protein disulfide oxidoreductase activity;20.GO:0042221 Response to chemical.

图7 淮豆13(A)和徐豆18(B)富集前20的KEGG分类 A:1.Ko04626植物-病原体互作通路;2.Ko00195光合作用通路;3.Ko00909倍半萜及三萜类生物合成通路;4.Ko04141内质网蛋白质加工通路;5.Ko01100代谢通路;6.Ko00380色氨酸代谢通路;7.Ko00561甘油酯代谢通路;8.Ko01110次级代谢物生物合成通路;9.Ko02010 ABC转运蛋白通路;10.Ko00901吲哚生物碱生物合成通路;11.Ko00073角质/木栓质/蜡质生物合成通路;12.Ko00591亚油酸代谢通路;13.Ko00062脂肪酸链延伸通路;14.Ko00903柠檬烯降解通路;15.Ko00941黄酮类生物合成通路;16.Ko00750维生素B6代谢通路;17.Ko00053抗坏血酸及糖醛酸代谢通路;18.Ko00250丙氨酸/天冬氨酸/谷氨酸代谢通路;19.Ko00071脂肪酸降解通路;20.Ko00330精氨酸及脯氨酸代谢通路。B:1.Ko04075植物激素信号转导通路;2.Ko00196光合作用-天线蛋白通路;3.Ko00860卟啉代谢通路;4.Ko04016 MAPK信号通路(植物);5.Ko00592α-亚麻酸代谢通路;6.Ko01110次级代谢物生物合成通路;7.Ko04626植物-病原体互作通路;8.Ko01212脂肪酸代谢通路;9.Ko00195光合作用通路;10.Ko00061脂肪酸生物合成通路;11.Ko00430牛磺酸及亚牛磺酸代谢通路;12.Ko00071脂肪酸降解通路;13.Ko00941黄酮类生物合成通路;14.Ko00410 β-丙氨酸代谢通路;15.Ko00945芪类、二芳基庚烷类及姜辣素生物合成通路;16.Ko04146过氧化物酶体通路;17.Ko04136自噬-其他类型通路;18.Ko00480谷胱甘肽代谢通路;19.Ko00280缬氨酸/亮氨酸/异亮氨酸降解通路;20.Ko01040不饱和脂肪酸生物合成通路。

Fig.7 The top 20 pathways of KEGG enrichment analysis of Huaidou 13(A)and Xudou 18(B) A:1.Ko04626 Plant-pathogen interaction;2.Ko00195 Photosynthesis;3.Ko00909 Sesquiterpenoid and triterpenoid biosynthesis;4.Ko04141 Protein processing in endoplasmic reticulum;5.Ko01100 Metabolic pathways;6.Ko00380 Tryptophan metabolism;7.Ko00561 Glycerolipid metabolism;8.Ko01110 Biosynthesis of secondary metabolites;9.Ko02010 ABC transporters;10.Ko00901 Indole alkaloid biosynthesis;11.Ko00073 Cutin,suberine and wax biosynthesis;12.Ko00591 Linoleic acid metabolism;13.Ko00062 Fatty acid elongation;14.Ko00903 Limonene degradation;15.Ko00941 Flavonoid biosynthesis;16.Ko00750 Vitamin B6 metabolism;17.Ko00053 Ascorbate and aldarate metabolism;18.Ko00250 Alanine,aspartate and glutamate metabolism;19.Ko00071 Fatty acid degradation;20.Ko00330 Arginine and proline metabolism.B:1.Ko04075 Plant hormone signal transduction;2.Ko00196 Photosynthesis-antenna proteins;3.Ko00860 Porphyrin metabolism;4.Ko04016 MAPK signaling pathway-plant;5.Ko00592 alpha-Linolenic acid metabolism;6.Ko01110 Biosynthesis of secondary metabolites;7.Ko04626 Plant-pathogen interaction;8.Ko01212 Fatty acid metabolism;9.Ko00195 Photosynthesis;10.Ko00061 Fatty acid biosynthesis;11.Ko00430 Taurine and hypotaurine metabolism;12.Ko00071 Fatty acid degradation;13.Ko00941 Flavonoid biosynthesis;14.Ko00410 beta-Alanine metabolism;15.Ko00945 Stilbenoid,diarylheptanoid and gingerol biosynthesis;16.Ko04146 Peroxisome;17.Ko04136 Autophagy-other;18.Ko00480 Glutathione metabolism;19.Ko00280 Valine,leucine and isoleucine degradation;20.Ko01040 Biosynthesis of unsaturated fatty acids.

图8 涝害胁迫响应基因转录因子家族分类

Fig.8 Family classification of transcription factors of differential expression genes

图9 148个涝害胁迫响应基因的GO和KEGG富集分析 A:1.GO:0015035蛋白质二硫键氧化还原酶活性;2.GO:0004364谷胱甘肽转移酶活性;3.GO:0004784超氧化物歧化酶活性;4.GO:0016721氧化还原酶活性(以超氧阴离子为受体);5.GO:0008686 3,4-二羟基-2-丁酮-4-磷酸合酶活性;6.GO:0015542糖外流转运跨膜转运蛋白活性;7.GO:0001871模式分子结合;8.GO:0030247多糖结合;9.GO:0030729乙酰乙酸-CoA连接酶活性;10.GO:0050218丙酸-CoA连接酶活性;11.GO:0008668 2,3-二羟基苯甲酰基腺苷酸合酶活性;12.GO:0008756邻琥珀酰苯甲酸-CoA连接酶活性;13.GO:0003987乙酸-CoA连接酶活性;14.GO:0047473 D-丙氨酸-聚(磷酸核糖醇)连接酶活性;15.GO:0005351糖:质子同向转运蛋白活性;16.GO:0030246碳水化合物结合;17.GO:0003700转录因子活性(序列特异性DNA结合);18.GO:0001071核酸结合转录因子活性;19.GO:0016405CoA连接酶活性;20.GO:0016878酸-巯基连接酶活性。B:1.Ko04141内质网蛋白质加工;2.Ko04626植物-病原体互作;3.Ko00591亚油酸代谢;4.Ko04146过氧化物酶体;5.Ko00061脂肪酸生物合成;6.Ko00592 α-亚麻酸代谢;7.Ko00250丙氨酸/天冬氨酸/谷氨酸代谢;8.Ko00195光合作用;9.Ko03420核苷酸切除修复;10.Ko01212脂肪酸代谢;11.Ko00909倍半萜及三萜类生物合成;12.Ko00480谷胱甘肽代谢;13.Ko00062脂肪酸链延伸;14.Ko00053抗坏血酸及糖醛酸代谢;15.Ko00280缬氨酸/亮氨酸/异亮氨酸降解;16.Ko00130泛醌及其他萜醌类生物合成;17.Ko00071脂肪酸降解;18.Ko00260甘氨酸/丝氨酸/苏氨酸代谢;19.Ko04070磷脂酰肌醇信号系统;20.Ko00562肌醇磷酸代谢。

Fig.9 GO and KEGG enrichment analysis of 148 response genes to water logging stress A:1.GO:0015035 Protein disulfide oxidoreductase activity;2.GO:0004364 Glutathione transferase activity;3.GO:0004784 Superoxide dismutase activity;4.GO:0016721 Oxidoreductase activity,acting on superoxide radicals as acceptor;5.GO:0008686 3,4-dihydroxy-2-butanone-4-phosphate synthase activity;6.GO:0015542 Sugar efflux transmembrane transporter activity;7.GO:0001871 Pattern binding;8.GO:0030247 Polysaccharide binding;9.GO:0030729 Acetoacetate-CoA ligase activity;10.GO:0050218 Propionate-CoA ligase activity;11.GO:0008668(2,3-dihydroxybenzoyl)adenylate synthase activity;12.GO:0008756 O-succinylbenzoate-CoA ligase activity;13.GO:0003987 Acetate-CoA ligase activity;14.GO:0047473 D-alanine-poly(phosphoribitol)ligase activity;15.GO:0005351 Sugar:proton symporter activity;16.GO:0030246 Carbohydrate binding;17.GO:0003700 Transcription factor activity,sequence-specific DNA binding;18.GO:0001071 Nucleic acid binding transcription factor activity;19.GO:0016405 CoA-ligase activity;20.GO:0016878 Acid-thiol ligase activity.B:1.Ko04141 Protein processing in endoplasmic reticulum;2.Ko04626 Plant-pathogen interaction;3.Ko00591 Linoleic acid metabolism;4.Ko04146 Peroxisome;5.Ko00061 Fatty acid biosynthesis;6.Ko00592 alpha-Linolenic acid metabolism;7.Ko00250 Alanine,aspartate and glutamate metabolism;8.Ko00195 Photosynthesis;9.Ko03420 Nucleotide excision repair;10.Ko01212 Fatty acid metabolism;11.Ko00909 Sesquiterpenoid and triterpenoid biosynthesis;12.Ko00480 Glutathione metabolism;13.Ko00062 Fatty acid elongation;14.Ko00053 Ascorbate and aldarate metabolism;15.Ko00280 Valine,leucine and isoleucine degradation;16.Ko00130 Ubiquinone and other terpenoid-quinone biosynthesis;17.Ko00071 Fatty acid degradation;18.Ko00260 Glycine,serine and threonine metabolism;19.Ko04070 Phosphatidylinositol signaling system;20.Ko00562 Inositol phosphate metabolism.

参考文献 29

[1]	王宝山. 逆境植物生物学[M]. 北京: 高等教育出版社, 2010.
	Wang B S. Plant stress biology[M]. Beijing: Higher Education Press, 2010.
[2]	公报编写组. 2023年中国水旱灾害及其防御成效综述[J]. 中国防汛抗旱, 2024, 34(12):35-40.doi:10.16867/j.issn.1673-9264.2024486.
	Compilation group of bulletin. Overview of water and drought disasters and defense effectiveness in China in 2023[J]. China Flood & Drought Management, 2024, 34(12):35-40.
[3]	Henshaw T L, Gilbert R A, Scholberg J M S, Sinclair T R. Soya bean(Glycine max L.merr.) genotype response to early-season flooding:I.root and nodule development[J]. Journal of Agronomy and Crop Science, 2007, 193(3):177-188.doi:10.1111/j.1439-037x.2007.00257.x.
[4]	Nguyen V L, Binh V T, Hoang D T, Mochizuki T, Nguyen V L. Genotypic variation in morphological and physiological response of soybean to waterlogging at flowering stage[J]. International Journal of Agricultural Science Research, 2015, 4(8):150-157.
[5]	朱建强, 张文英, 欧光华, 刘德福, 程伦国, 程玲. 夏大豆花荚期受渍胁迫对农艺性状、产量与品质的影响[J]. 大豆科学, 2001, 20(1):71-74.doi:10.3969/j.issn.1000-9841.2001.01.016.
	Zhu J Q, Zhang W Y, Ou G H, Liu D F, Cheng L G, Cheng L. Influence upon agronomic properties,yields and qualities of summer soybean in period of soybean with flowers and pods under subsurface waterloging of soybean field[J]. Soybean Science, 2001, 20(1):71-74.
[6]	Wu C J, Zeng A L, Chen P Y, Hummer W, Mokua J, Shannon J G, Nguyen H T. Evaluation and development of flood-tolerant soybean cultivars[J]. Plant Breeding, 2017, 136(6):913-923.doi:10.1111/pbr.12542. URL
[7]	Wu C J, Zeng A L, Chen P Y, Florez-Palacios L, Hummer W, Mokua J, Klepadlo M, Yan L, Ma Q B, Cheng Y B. An effective field screening method for flood tolerance in soybean[J]. Plant Breeding, 2017, 136(5):710-719.doi:10.1111/pbr.12487. URL
[8]	董艳. 渍水胁迫对大豆生理特性和产量的影响[D]. 南京: 南京农业大学, 2011.doi:10.7666/d.Y2361325.
	Dong Y. Effects of waterlogging stress on physiological characteristics and yield of soybean[D]. Nanjing: Nanjing Agricultural University,2011.
[9]	Githiri S M, Watanabe S, Harada K, Takahashi R. QTL analysis of flooding tolerance in soybean at an early vegetative growth stage[J]. Plant Breeding, 2006, 125(6):613-618.doi:10.1111/j.1439-0523.2006.01291.x.
[10]	Sullivan M, VanToai T, Fausey N, Beuerlein J, Parkinson R, Soboyejo A. Evaluating on-farm flooding impacts on soybean[J]. Crop Science, 2001, 41(1):93-100.doi:10.2135/cropsci2001.41193x. URL
[11]	宋晓慧, 张智杰, 李春光, 张代平, 韩英鹏, 李冬梅, 李文滨. 淹水时间对不同耐涝性大豆品种苗期根部形态和叶部生理指标的影响[J]. 大豆科学, 2014, 33(1):70-72,102.doi:10.11861/j.issn.1000-9841.2014.01.0070.
	Song X H, Zhang Z J, Li C G, Zhang D P, Han Y P, Li D M, Li W B. Effect of waterlogging time on root morphology and foliar physiological indexes of soybean varieties[J]. Soybean Science, 2014, 33(1):70-72,102.
[12]	王婷婷, 陈鸽, 包悦琳, 金东淳. 大豆根系响应早期缺铁胁迫的转录组分析[J]. 南京农业大学学报, 2022, 45(2):224-234.doi:10.7685/7685/jnau.202107006.
	Wang T T, Chen G, Bao Y L, Jin D C. Transcriptome analysis of soybean roots in response to early iron deficiency stress[J]. Journal of Nanjing Agricultural University, 2022, 45(2):224-234.
[13]	李明霞, 周际, 胡勇军, 韩德复, 郭继勋, 李淑英, 张涛, 石连旋. 低氮胁迫下栽培大豆和野大豆幼苗适应性转录组学比较研究[J]. 东北师大学报(自然科学版), 2023, 55(2):116-125.doi:10.16163/j.cnki.dslkxb202110200001.
	Li M X, Zhou J, Hu Y J, Han D F, Guo J X, Li S Y, Zhang T, Shi L X. Comparative study on the adaptable transcriptomics of cultivated and wild soybean seedling under low nitrogen stress[J]. Journal of Northeast Normal University(Natural Science Edition), 2023, 55(2):116-125.
[14]	曹颖, 竹龙鸣, 邓平, 陈紫芸, 袁凤杰. 模拟干旱胁迫下大豆耐旱突变体的生理和转录组分析[J]. 大豆科学, 2024, 43(4):421-430.doi:10.11861/j.issn.1000-9841.2024.04.0421.
	Cao Y, Zhu L M, Deng P, Chen Z Y, Yuan F J. Physiological and transcriptomic aanlysis of a soybean drought-tolerant mutant under simulate drought stress[J]. Soybean Science, 2024, 43(4):421-430.
[15]	Oosterhuis D M, Scott H D, Hampton R E, Wullschleger S D. Physiological responses of two soybean [Glycine max(L.) Merr]cultivars to short-term flooding[J]. Environmental and Experimental Botany, 1990, 30(1):85-92.doi:10.1016/0098-8472(90)90012-S. URL
[16]	VanToai T T, Cuc Hoa T T, Ngoc Hue N T, Nguyen H T, Shannon J G, Rahman M A. Flooding tolerance of soybean [Glycine max(L.) merr.]germplasm from Southeast Asia under field and screen-house environments[J]. The Open Agriculture Journal, 2010, 4(1):38-46.doi:10.2174/1874331501004010038. URL
[17]	Chandra S, Satpute G K, Nagar S, Singh M, Kumawat G, Rajesh V, Gupta S. Reproductive stage water-logging tolerance:a critical assessment of traits in soybean[J]. Soybean Research, 2020, 18(2),93-101.
[18]	Sakazono S, Nagata T, Matsuo R, Kajihara S, Watanabe M, Ishimoto M, Shimamura S, Harada K, Takahashi R, Mochizuki T. Variation in root development response to flooding among 92 soybean lines during early growth stages[J]. Plant Production Science, 2014, 17(3):228-236.doi:10.1626/pps.17.228. URL
[19]	张金霞, 董德坤, 胡兴旺, 陈芬, 朱丹华. 三个大豆品种萌发期和苗期的耐盐性比较[J]. 浙江农业学报, 2016, 28(7):1101-1107.doi:10.3969/j.issn.1004-1524.2016.07.02.
	Zhang J X, Dong D K, Hu X W, Chen F, Zhu D H. Research on salt tolerance of three soybean varieties at both germination and seedling stage[J]. Acta Agriculturae Zhejiangensis, 2016, 28(7):1101-1107.
[20]	Silva P A, Silva J C F, Caetano H D N, Machado J P B, Mendes G C, Reis P A B, Brustolini O J B, Dal-Bianco M, Fontes E P B. Comprehensive analysis of the endoplasmic reticulum stress response in the soybean genome:conserved and plant-specific features[J]. BMC Genomics, 2015, 16(1):783.doi:10.1186/s12864-015-1952-z. URL
[21]	Walter P, Ron D. The unfolded protein response:from stress pathway to homeostatic regulation[J]. Science, 2011, 334(6059):1081-1086.doi:10.1126/science.1209038. pmid: 22116877
[22]	Noctor G, Mhamdi A, Chaouch S, Han Y, Neukermans J, Marquez-Garcia B, Queval G, Foyer C H. Glutathione in plants:an integrated overview[J]. Plant,Cell & Environment, 2012, 35(2):454-484.doi:10.1111/j.1365-3040.2011.02400.x.
[23]	Hasanuzzaman M, Bhuyan M H M B, Anee T I, Parvin K, Nahar K, Al Mahmud J, Fujita M. Regulation of ascorbate-glutathione pathway in mitigating oxidative damage in plants under abiotic stress[J]. Antioxidants, 2019, 8(9):384.doi:10.3390/antiox8090384. URL
[24]	Sharma P, Lakra N, Goyal A, Ahlawat Y K, Zaid A, Siddique K H M. Drought and heat stress mediated activation of lipid signaling in plants:a critical review[J]. Frontiers in Plant Science, 2023, 14:1216835.doi:10.3389/fpls.2023.1216835. URL
[25]	Chen Y Y, Wang L F, Dai L J, Yang S G, Tian W M. Characterization of HbEREBP1,a wound-responsive transcription factor gene in laticifers of Hevea brasiliensis Muell.Arg[J]. Molecular Biology Reports, 2012, 39(4):3713-3719.doi:10.1007/s11033-011-1146-y. URL
[26]	张红梅, 刘晓庆, 陈华涛, 袁星星, 崔晓艳, 张智民, 黄中文, 陈新. 大豆转录因子GmWRKY58亚细胞定位及在非生物胁迫下的表达分析[J]. 华北农学报, 2018, 33(2):49-57.doi:10.7668/hbnxb.2018.02.008.
	Zhang H M, Liu X Q, Chen H T, Yuan X X, Cui X Y, Zhang Z M, Huang Z W, Chen X. Subcellular localization and expression analysis of a soybean transcription factor GmWRKY58 in response to abiotic stresses[J]. Acta Agriculturae Boreali-Sinica, 2018, 33(2):49-57. doi: 10.7668/hbnxb.2018.02.008
[27]	黄友举, 于泳波, 逄翠晶, 孙轼絮, 路晨, 于延冲. 大豆GmWRKY44生物信息学分析及功能验证[J]. 华北农学报, 2025, 40(1):29-35.doi:10.7668/hbnxb.20195201.
	Huang Y J, Yu Y B, Pang C J, Sun S X, Lu C, Yu Y C. Cloning and biology function analysis of soybean GmWRKY44[J]. Acta Agriculturae Boreali-Sinica, 2025, 40(1):29-35.
[28]	张斌. 大豆GmGRAS69基因过表达增强拟南芥的耐旱性[J]. 华北农学报, 2023, 38(3):18-25.doi:10.7668/hbnxb.20193597.
	Zhang B. Overexpression of soybean GmGRAS69 gene enhances drought tolerance of transgenic Arabidopsis[J]. Acta Agriculturae Boreali-Sinica, 2023, 38(3):18-25.
[29]	Kim Y H, Hwang S J, Waqas M, Khan A L, Lee J H, Lee J D, Nguyen H T, Lee I J. Comparative analysis of endogenous hormones level in two soybean(Glycine max L.) lines differing in waterlogging tolerance[J]. Frontiers in Plant Science, 2015, 6:714.doi:10.3389/fpls.2015.00714.

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