大豆耐盐与耐镉胁迫共性基因的挖掘

doi:10.3969/j.issn.1004-1524.20250086

浙江农业学报 ›› 2026, Vol. 38 ›› Issue (1): 1-16.DOI: 10.3969/j.issn.1004-1524.20250086

大豆耐盐与耐镉胁迫共性基因的挖掘

徐燕¹^,²(), 李素娟³, 陈光², 徐盛春¹^,²^,⁴, 王剑²^,^*()

1.浙江农林大学现代农学院,浙江杭州 311300
2.浙江省农业科学院数字农业研究所,浙江杭州 310021
3.浙江省农业科学院农产品质量安全全国重点实验室,浙江杭州 310021
4.湘湖实验室,浙江杭州 311258

收稿日期:2025-02-07 出版日期:2026-01-25 发布日期:2026-02-11
作者简介:*王剑,E-mail: wangj@zaas.ac.cn
徐燕,主要从事大豆抗性基因筛选与功能分析。E-mail:2932737880@qq.com
通讯作者: 王剑
基金资助:
中央引导地方科技发展资金(2023ZY1016)

Identification of common genes for salt and cadmium tolerance in soybean

XU Yan¹^,²(), LI Sujuan³, CHEN Guang², XU Shengchun¹^,²^,⁴, WANG Jian²^,^*()

1. College of Advanced Agricultural Science, Zhejiang A&F University, Hangzhou 311300, China
2. Institute of Digital Agriculture, Zhejiang Academy of Agricultural Sciences, Hangzhou 310021, China
3. State Key Laboratory for Quality and Safety of Agro-Products, Zhejiang Academy of Agricultural Sciences, Hangzhou 310021, China
4. Xianghu Laboratory, Hangzhou 311258, China

Received:2025-02-07 Online:2026-01-25 Published:2026-02-11
Contact: WANG Jian

摘要/Abstract

摘要：

盐碱化土地偶尔会伴有重金属污染。研究植物对盐、镉复合胁迫的生长与生理响应,并挖掘其共有耐性基因和调控通路,对作物抗逆遗传改良具有重要意义。本研究以50份大豆栽培种和野生种为材料,分别设置盐胁迫(200 mmol·L^-1 NaCl)、镉胁迫(0.3 mmol·L^-1 CdCl₂)与正常条件进行培养,测定萌发率、株高、根长、根长与株高比、地上部与地下部鲜重共6个指标,通过主成分分析筛选关键耐性指标;利用耐性最强的野生大豆种质W-3-12-90构建全长cDNA酵母表达文库,结合全长cDNA过表达(FOX)基因搜寻系统与二代测序,鉴定盐、镉共耐受相关基因。主成分分析结果显示,株高、根长、地上部鲜重和萌发率是评价大豆耐盐、耐镉能力的4项关键指标;野生大豆种质W-3-12-90在盐、镉胁迫下耐受性最强。基于该材料共鉴定出109个盐、镉共同响应基因。亚细胞定位预测显示,39个基因编码胞外蛋白,此类蛋白响应快、占比高;51个基因编码的蛋白质分布于细胞核、细胞质与细胞膜,主要参与蛋白质代谢、细胞信号转导、防御与应激反应,以及氧化还原酶活性等通路。基因表达分析表明,6个候选共耐性基因在盐、镉胁迫下均显著上调。综上,植物响应盐、镉胁迫的基因主要通过编码胞外蛋白,并借助其与质膜、细胞核及细胞质的相互作用协同调控植物的耐盐、耐镉能力。本研究为解析大豆耐盐、耐镉的分子机制提供了新思路,并为培育耐盐、耐镉新种质提供了基因资源与理论依据。

关键词: 大豆, 盐胁迫, 镉胁迫, 种质筛选, 基因挖掘

Abstract:

Saline-alkali land is occasionally accompanied by heavy metal pollution. Studying plant growth and physiological responses to combined salt and cadmium stress and identifying their common tolerance genes and regulatory pathways are of great significance for the genetic improvement of crop stress resistance. In this study, 50 soybean cultivars and wild soybean accessions were used as materials. They were cultivated under salt stress (200 mmol·L^-1 NaCl), cadmium stress (0.3 mmol·L^-1 CdCl₂), and normal conditions, respectively. Six indexes, including germination rate, plant height, root length, root-shoot ratio, and fresh weights of the aboveground and underground parts, were measured. Key tolerance indicators were screened through principal component analysis. The wild soybean accession W-3-12-90, which exhibited the strongest tolerance, was used to construct a full-length cDNA yeast expression library. Combined with the Full-length cDNA Over-expressor (FOX) gene hunting system and second-generation sequencing, genes related to co-tolerance to salt and cadmium were identified. Principal component analysis revealed that plant height, root length, aboveground fresh weight, and germination rate were the four key indexes for evaluating soybean tolerance to salt and cadmium. The wild soybean accession W-3-12-90 demonstrated the strongest tolerance under salt and cadmium stress. Using this material, a total of 109 common responsive genes to salt and cadmium were identified. Subcellular localization prediction showed that 39 of these genes encoded extracellular proteins, which respond rapidly and account for a high proportion. Proteins encoded by another 51 genes were distributed in the nucleus, cytoplasm, and cell membrane, and were primarily involved in pathways related to protein metabolism, cellular signal transduction, defense and stress responses, and oxidoreductase activity. Gene expression analysis indicated that six candidate co-tolerance genes were significantly upregulated under both salt and cadmium stress. In summary, genes responding to salt and cadmium stress in plants primarily function by encoding extracellular proteins and coordinately regulate plant tolerance through interactions of these proteins with the plasma membrane, nucleus, and cytoplasm. This study provides new insights for elucidating the molecular mechanism of soybean tolerance to salt and cadmium, and provides gene resources and theoretical basis for breeding new salt- and cadmium-tolerant germplasms.

Key words: soybean, salt stress, cadmium stress, germplasm screening, gene mining

中图分类号:

S565.1

徐燕, 李素娟, 陈光, 徐盛春, 王剑. 大豆耐盐与耐镉胁迫共性基因的挖掘[J]. 浙江农业学报, 2026, 38(1): 1-16.

XU Yan, LI Sujuan, CHEN Guang, XU Shengchun, WANG Jian. Identification of common genes for salt and cadmium tolerance in soybean[J]. Acta Agriculturae Zhejiangensis, 2026, 38(1): 1-16.

图/表 12

表1 耐盐、耐镉候选基因qRT-PCR分析所用的引物

Table 1 Primers for qRT-PCR analysis of candidate genes associated with salt- and cadmium-tolerance

基因Gene	正向引物序列(5'→3') Forward primer sequence(5'→3')	反向引物序列(5'→3') Reverse primer sequence(5'→3')
GmEF1b	CCACTGCTGAAGAAGATGATGATG	AAGGACAGAAGACTTGCCACTC
GmDehydrin	AGGAAGGAACATCGTCAGCA	TGACAAGACACTGTACGTACG
GmSSP	CCACCTCAGGAGTCTCAGAA	CCCGCAAAAGTTTCGTGACT
GmGF14	ACGTTGGGAGAGGAATCATACA	GCATTCAACACCTTCTCCCT
GmPAP85	AGCAGAAAGAGGAGGGGAAC	AGCAGACAGTTGAAGTACACA
GmHUP54	ATGCCTAGGATTGACAGCGA	AGCAGAGTCAGCACCATCAT
GmMET2	TCGAGAGTGCTGAAATGGGT	ACACACCCATCACAAGTCCA

图1 不同大豆种质幼苗在盐或镉胁迫下的生理表现盐、镉胁迫8 d后耐盐和耐镉品种,以及盐、镉敏感品种大豆幼苗的生理表现。NaCl,盐胁迫条件;CdCl2,镉胁迫条件;Control,正常生长条件。

Fig.1 Physiological performance of different soybean seedlings under salt or cadmium stress Physiological responses of soybean seedlings of salt/Cd-tolerant and salt/Cd-sensitive varieties under salt/Cd stress for 8 days. NaCl, Salt stress; Cd, Cadmium stress; Control, Normal growth.

表2 50份大豆种质苗期在正常条件和盐/镉胁迫下的生理表现

Table 2 Physiological performance of 50 soybean germplasms during seedling stage under normal conditions and salt/cadmium stress

处理 Treatment	PL/cm		RL/cm		RL/PL		SW/g		RW/g		GR
处理 Treatment	均值 Mean	CV/%	均值 Mean	CV/%	均值 Mean	CV/%	均值 Mean	CV/%	均值 Mean	CV/%	均值 Mean	CV/%
正常条件 CK	10.26± 0.31 a	24.91	7.52± 0.16 a	18.27	0.81± 0.02 b	23.83	0.76± 0.02 a	24.14	0.24± 0.01 a	31.58	0.97± 0.01 a	6.15
盐胁迫 Salt stress	1.03± 0.02 b	16.75	1.86± 0.05 b	23.06	1.92± 0.05 a	24.63	0.41± 0.01 b	31.97	0.05± 0.00 b	46.62	0.92± 0.01 b	10.56
镉胁迫 Cd stress	0.88± 0.03 b	37.75	0.30± 0.04 c	133.61	0.30± 0.04 c	134.28	0.41± 0.01 b	30.95	0.03± 0.01 b	353.39	0.75± 0.02 c	31.25

表3 50份大豆种质响应盐胁迫各变量对主成分的贡献度

Table 3 Contribution of variables to principal components under salt stress across 50 soybean germplasms

性状 Trait	主成分1 PC1	主成分2 PC2	主成分3 PC3
PL	0.86	-0.12	0.08
RL	0.35	0.82	-0.12
GR	-0.49	0.84	-0.09
RL/PL	0.42	0.67	0.24
RW	0.84	-0.04	0.20
SW	-0.37	0.04	0.90
特征值Eigenvalue	2.09	1.81	0.93
贡献率/%Contribution rate/%	34.85	30.12	15.52
累计贡献率/%	34.85	64.97	80.49
Cumulative contribution rate/%

表4 50份大豆种质响应镉胁迫各变量对主成分的贡献度

Table 4 Contribution of variables to principal components under cadmium stress across 50 soybean germplasms

性状 Trait	主成分1 PC1	主成分2 PC2	主成分3 PC3
PL	0.76	-0.19	0.52
RL	0.94	0.25	-0.16
RL/PL	0.55	-0.30	-0.14
GR	0.65	0.53	-0.51
RW	0.62	-0.48	0.20
SW	0.05	0.70	0.66
特征值Eigenvalue	2.56	1.20	1.05
贡献率/%Contribution rate/%	42.68	20.01	17.51
累计贡献率/%	42.68	62.69	80.20
Cumulative contribution rate/%

图2 转化大豆文库的酵母菌株在不同生长条件下的生长表现 A,大豆全长cDNA文库转化酵母G19菌株在含500 mmol·L-1 NaCl的SD-His-Ura培养基上的存活情况;B,大豆全长cDNA文库转化酵母Δycf1菌株在含50 μmol·L-1 CdCl2的SD-Ura培养基上的存活情况。照片拍摄于点板后第3天。

Fig.2 Growth performance of yeast strains transformed with soybean library under different growth conditions A, The survival performance of yeast strain G19 transformed with full-length soybean cDNA library on SD-His-Ura medium supplemented with 500 mmol·L-1 NaCl; B, The survival performance of yeast strain Δycf1 transformed with full-length soybean cDNA library on SD-Ura medium supplemented with 50 μmol·L-1 CdCl2. The photos were taken three days after plating.

图3 大豆耐盐候选基因的GO、KEGG分析 A,GO功能聚类分析;B,KEGG通路富集分析。A图注释名称自上而下依次为:翻译起始;翻译起始因子活性;RNA结合;胁迫响应;蛋白质结构域特异性结合;光合作用,光捕获;氧化还原酶活性,作用于单一供体并结合分子氧,结合两个氧原子;营养储存活性;单层膜包被的脂质储存体;糖酵解过程。B图通路名称自上而下依次为:剪接体;RNA转运;核糖体;光合作用-天线蛋白;代谢途径;亚油酸代谢;碳代谢;光合生物中的碳固定;次生代谢物的生物合成;氨基酸的生物合成。

Fig.3 GO and KEGG analysis of candidate genes for salt tolerance in soybean A, GO function clustering analysis; B, KEGG pathway enrichment analysis. The annotation names of Fig. A from top to bottom are as follows: Translational initiation; Translation initiation factor activity; RNA binding; Response to stress; Protein domain specific binding; Photosynthesis, light harvesting; Oxidoreductase activity, acting on single donors with incorporation of molecular oxygen, incorporation of two atoms of oxygen; Nutrient reservoir activity; Monolayer-surrounded lipid storage body; Glycolytic process. The annotation names of Fig. B from top to bottom are as follows: Spliceosome; RNA transport; Ribosome; Photosynthesis-antenna proteins; Metabolic pathways; Linoleic acid metabolism; Carbon metabolism; Carbon fixation in photosynthetic organisms; Biosynthesis of secondary metabolites; Biosynthesis of amino acids.

图4 大豆耐镉候选基因的GO、KEGG富集分析 A,GO功能的聚类分析;B,KEGG通路的富集分析。A图注释自上而下依次为:丝氨酸型内肽酶抑制活性;S-腺苷甲硫氨酸生物合成过程;RNA结合;胁迫响应;蛋白质结构域特异性结合;光合作用,光捕获;氧化还原酶活性(作用于供体的醛基或酮基,以NAD或NADP为受体);营养储存活性;甲硫氨酸腺苷转移酶活性;热休克蛋白结合。B图通路自上而下依次为:核糖体;光合作用-天线蛋白;代谢途径;乙醛酸和二羧酸代谢;糖酵解/糖异生;半胱氨酸和甲硫氨酸代谢;碳代谢;光合生物中的碳固定;次生代谢物生物合成;氨基酸生物合成。

Fig.4 GO and KEGG enrichment analysis of candidate genes for cadmium tolerance in soybean A, GO function clustering analysis; B, KEGG pathway enrichment analysis. The annotation names of Fig. A from top to bottom are as follows: Serine-type endopeptidase inhibitor activity; S-adenosylmethionine biosynthetic process; RNA binding; Response to stress; Protein domain specific binding; Photosynthesis, light harvesting; Oxidoreductase activity, acting on the aldehyde or oxo group of donors, NAD or NADP as acceptor; Nutrient reservoir activity; Methionine adenosyltransferase activity; Heat shock protein binding. The annotation names of Fig. B from top to bottom are as follows: Ribosome; Photosynthesis-antenna proteins; Metabolic pathways; Glyoxylate and dicarboxylate metabolism; Glycolysis/Gluconeogenesis; Cysteine and methionine metabolism; Carbon metabolism; Carbon fixation in photosynthetic organisms; Biosynthesis of secondary metabolites; Biosynthesis of amino acids.

图5 大豆耐盐和耐镉候选基因编码蛋白的亚细胞定位 A,已知基因编码蛋白的亚细胞定位;B,未知基因编码蛋白的亚细胞定位。

Fig.5 Subcellular localization of proteins encoded by soybean salt and cadmium tolerance candidate genes A, Subcellular localization of proteins encoded by known genes; B, Subcellular localization of proteins encoded by unknown genes.

图6 大豆盐、镉共同耐性候选基因的GO、KEGG分析 A,GO功能的聚类分析;B,KEGG通路的富集分析。A图注释名称自上而下依次为:丝氨酸型内肽酶抑制活性;胁迫响应;蛋白质结构域特异性结合;光合作用,光捕获;氧化还原酶活性(作用于供体的醛基或酮基,以NAD或NADP 为受体);营养储存活性;单层膜包被的脂质储存体;细胞外区域;内肽酶抑制活性;半胱氨酸型肽酶活性。B图通路名称自上而下依次为:核糖体;内质网中的蛋白质加工;蛋白酶体;光合作用-天线蛋白;单环β-内酰胺生物合成;代谢途径;亚油酸代谢;糖酵解/糖异生;碳代谢;光合生物中的碳固定。Gene ratio表示注释到特定条目的差异基因数与差异基因总数的比值;Richfactor为某通路中富集到的差异基因与该通路所有基因的比值。下同。

Fig.6 GO and KEGG analysis of candidate genes for salt-cadmium co-tolerance in soybean A, GO function clustering analysis; B, KEGG pathway enrichment analysis. The annotation names of Fig. A from top to bottom are as follows: Serine-type endopeptidase inhibitor activity; Response to stress; Protein domain specific binding; Photosynthesis, light harvesting; Oxidoreductase activity, acting on the aldehyde or oxo group of donors, NAD or NADP as acceptor; Nutrient reservoir activity; Monolayer-surrounded lipid storage body; Extracellular region; Endopeptidase inhibitor activity; Cysteine-type peptidase activity. The annotation names of Fig. B from top to bottom are as follows: Ribosome; Protein processing in endoplasmic reticulum; Proteasome; Photosynthesis-antenna proteins; Monobactam biosynthesis; Metabolic pathways; Linoleic acid metabolism; Glycolysis/Gluconeogenesis; Carbon metabolism; Carbon fixation in photosynthetic organisms. Gene ratio represents the ratio of differentially expressed genes annotated to a specific term to the total number of differentially expressed genes; Rich factor is the ratio of enriched differentially expressed genes in a pathway to all genes in that pathway. The same as below.

表5 大豆关键耐盐和镉基因信息

Table 5 Key salt and cadmium tolerant gene information of soybean

基因ID Gene ID	基因名 Gene name	cDNA大小/bp cDNA size/bp	预测功能 Function prediction
Glyma.09G185500	GmDehydrin	681	Dehydrin-like protein
Glyma.03G163533	GmSSP	1 488	Rmlc-like cupins superfamily protein
Glyma.02G208700	GmGF14	789	14-3-3-like protein
Glyma.10G246300	GmPAP85	1 866	Cupin family protein
Glyma.15G072400	GmHUP54	756	Aluminium induced protein with YGL and LRDR motifs
Glyma.07G132000	GmMET2	240	Metallothionein 2A

图7 盐、镉、盐+镉复合胁迫下大豆根、茎和叶中候选基因的相对表达水平无相同小写字母表示同一个时间点在不同处理间差异显著(p<0.05)。

Fig.7 Relative expression level of candidate genes in soybean roots, stems, and leaves under salt, Cd and salt-Cd stress Bars marked with the same lowercase letters in the figure indicate significant differences (p<0.05) among treatments at the same time.

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大豆耐盐与耐镉胁迫共性基因的挖掘

Identification of common genes for salt and cadmium tolerance in soybean

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