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作物学报 ›› 2026, Vol. 52 ›› Issue (6): 1859-1875.doi: 10.3724/SP.J.1006.2026.51086

• 耕作栽培·生理生化 • 上一篇    下一篇

水培盐碱复合胁迫下‘济麦60’苗期根系有机酸分泌动态变化及其转录调控机制

陈雪燕1,**(), 何华川2,**(), 李政嘉2, 董新盼2, 李藕琪2, 刘小云2, 李丹萍1, 陈志伟1, 刘国霞1, 吕胜源1, 吴印莹1, 赵振东1, 曹新有1,*(), 万何平2,*()   

  1. 1 山东省农业科学院, 山东济南 250100
    2 江汉大学生命科学学院, 湖北武汉 430056
  • 收稿日期:2025-10-15 接受日期:2026-02-27 出版日期:2026-06-12 网络出版日期:2026-03-09
  • 通讯作者: * 曹新有, E-mail: cxytvs@163.com; 万何平, E-mail: wanheping@jhun.edu.cn
  • 作者简介:陈雪燕, E-mail: chxytvs@163.com;何华川, E-mail: huachuanhe99@163.com

    **同等贡献

  • 基金资助:
    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-03-06);山东省重点研发计划项目(2023LZGC009);山东省重点研发计划项目(2024LZGCQY001);山东省泰山学者工程项目(tstp20240843)

Dynamic changes in root organic acid secretion and its transcriptional regulatory mechanisms in ‘Jimai 60’ seedlings under combined salinity-alkalinity stress in hydroponics

Chen Xue-Yan1,**(), He Hua-Chuan2,**(), Li Zheng-Jia2, Dong Xin-Pan2, Li Ou-Qi2, Liu Xiao-Yun2, Li Dan-Ping1, Chen Zhi-Wei1, Liu Guo-Xia1, Lyu Sheng-Yuan1, Wu Yin-Ying1, Zhao Zhen-Dong1, Cao Xin-You1,*(), Wan He-Ping2,*()   

  1. 1 Shandong Academy of Agricultural Sciences, Jinan 250100, Shandong, China
    2 College of Life Sciences, Jianghan University, Wuhan 430056, Hubei, China
  • Received:2025-10-15 Accepted:2026-02-27 Published:2026-06-12 Published online:2026-03-09
  • Contact: * Cao Xin-You, E-mail: cxytvs@163.com; Wan He-Ping, E-mail: wanheping@jhun.edu.cn
  • About author:

    **Contributed equally to this work

  • Supported by:
    China Agriculture Research System of MOF and MARA(CARS-03-06);Key Research and Development Program of Shandong Province(2023LZGC009);Key Research and Development Program of Shandong Province(2024LZGCQY001);Taishan Scholar Program(tstp20240843)

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摘要：

土壤盐碱化是全球农业生产面临的主要限制因素之一, 显著制约小麦产量并威胁粮食安全。植物根系分泌有机酸是应对非生物胁迫的重要生理策略。然而, 在盐碱复合胁迫下, 小麦根系有机酸分泌的动态特征及其背后的转录调控网络尚缺乏系统研究。本研究以耐盐碱小麦品种‘济麦60’为材料, 系统分析了其在盐碱复合胁迫不同时间点根系有机酸的代谢组与转录组变化特征, 并结合加权基因共表达网络分析(WGCNA)挖掘关键调控模块。结果表明, 盐碱复合胁迫显著改变了根系有机酸的组成与含量, 在4个比较组中共鉴定到44种差异积累的有机酸, 其中12种持续上调, 4种持续下调。转录组分析显示, 随着胁迫时间的延长, 差异表达基因数量逐步增加, 与光合作用、有性生殖和卟啉代谢等相关的基因持续被激活, 而细胞壁组织、氧化磷酸化等过程则表现出时间阶段性响应。WGCNA分析进一步鉴定出MEblue、MEturquoise和MEyellow 3个与关键有机酸代谢显著相关的功能模块, 这些模块中的基因主要富集于能量代谢、氨基酸合成与氧化还原平衡等过程。本研究系统描绘了耐盐碱小麦品种‘济麦60’在盐碱复合胁迫下根系有机酸代谢与转录调控的动态图谱, 这不仅加深了对小麦耐盐碱生理机制的认识, 也为通过分子设计育种靶向改良根系功能与逆境适应性提供了重要的理论基础和基因资源。

关键词: 小麦, 盐碱复合胁迫, 根系, 有机酸, 转录组, 加权基因共表达网络分析

Abstract:

Soil salinization is a major constraint on global agricultural production, substantially reducing wheat yield and threatening food security. Root secretion of organic acids is an important physiological strategy by which plants cope with abiotic stress. However, under combined salt-alkali stress, the temporal dynamics of organic acid secretion in wheat roots and the underlying transcriptional regulatory networks remain poorly understood. Here, the salt-alkali-tolerant wheat cultivar ‘Jimai 60’ was used to systematically characterize time-resolved metabolomic and transcriptomic profiles of root organic acids under combined salt-alkali stress, and key regulatory modules were identified using WGCNA. Combined salt-alkali stress markedly reshaped the composition and abundance of root organic acids. Across four comparison groups, 44 differentially accumulated organic acids were identified, including 12 that were consistently upregulated and 4 that were consistently downregulated. Transcriptome analysis showed that the number of differentially expressed genes increased progressively with longer stress exposure. Genes related to photosynthesis, sexual reproduction, and porphyrin metabolism were persistently activated, whereas cell wall organization and oxidative phosphorylation exhibited stage-specific responses. WGCNA further identified the MEblue, MEturquoise, and MEyellow modules as significantly associated with major organic acid metabolic traits; genes in these modules were mainly enriched in pathways related to energy metabolism, amino acid biosynthesis, and redox homeostasis. Overall, this study delineates the dynamic landscape of root organic acid metabolism and transcriptional regulation in salt-alkali-tolerant wheat (‘Jimai 60’) under combined salt-alkali stress. These findings advance our understanding of the physiological basis of wheat salt-alkali tolerance and provide a foundation, as well as candidate gene resources, for molecular breeding aimed at improving root function and stress resilience.

Key words: wheat, combined saline-alkaline stress, roots, organic acids, transcriptome, WGCNA

图1

根系分泌有机酸收集流程 本图的制作借助了人工智能工具。"

附表1

本研究测定的有机酸种类"

序号
No.		 名称
Name		 序号
No.		 名称
Name
1 3-吲哚乙酸 3-indolelactic acid 34 丙酮酸 Pyruvic acid
2 氯氨酮 Kynurenic acid 35 反式-乌头酸 Trans-aconitic acid
3 苯乙酰甘氨酸 Phenaceturic acid 36 高香草酸 Homovanillic acid
4 肉桂酸 Cinnamic acid 37 马尿酸 Hippuric acid
5 山楂酸 Maslinic acid 38 莽草酸 Shikimic acid
6 吲哚-2-羧酸 2-indolecarboxylic acid 39 犬尿氨酸 Kynurenine
7 5-羟基吲哚-3-乙酸 5-hydroxyindoleacetic acid 40 邻羟基苯乙酸 2-hydroxyphenylacetic acid
8 4-氨基丁酸 4-aminobutyric acid 41 对羟基苯乙酸 4-hydroxyphenylacetic acid
9 邻氨基苯甲酸 Aminobenzoic acid 42 乳酸 Lactic acid
10 吲哚-3-乙酸 Indole-3-acetic acid 43 甲基丙二酸 Methylmalonic acid
11 3-羟基苯乙酸 3-hydroxyphenylacetic acid 44 琥珀酸 Succinic acid
12 3-羟基马尿酸 3-hydroxyhippuric acid 45 2-羟基-2-甲基丁酸 2-hydroxy-2-methylbutyric acid
13 4-羟基马尿酸 4-hydroxyhippuric acid 46 2-羟基-3-甲基丁酸 2-hydroxyisovaleric acid
14 齐墩果酸 Oleanolic acid 47 3-羟基异戊酸 3-hydroxyisovaleric acid
15 L-焦谷氨酸 Pyroglutamic acid 48 甲基丁二酸 2-methylsuccinic acid
16 3-甲基己二酸 3-methyladipic acid 49 戊二酸 Glutaric acid
17 癸二酸 Sebacic acid 50 乙基丙二酸 Ethylmalonic acid
18 氢化肉桂酸 Benzenepropanoic acid 51 己二酸 Adipic acid
19 3-羟基-3-甲基谷氨酸 3-hydroxymethylglutaric acid 52 α-酮戊二酸 Oxoglutaric acid
20 4-香豆酸 4-coumaric acid 53 新绿原酸 Neochlorogenic acid
21 泛酸 Pantothenic acid 54 隐绿原酸 Cryptochlorogenic acid
22 咖啡酸 Caffeic acid 55 富马酸 Fumaric acid
23 马来酸 Maleic acid 56 乙酰丙酸 Levulinic acid
24 没食子酸 Gallic acid 57 顺式-乌头酸 Cis-aconitic acid
25 壬二酸 Azelaic acid 58 柠康酸 Citraconic acid
26 鼠尾草酸 Carnosic acid 59 水杨酸 Salicylic acid
27 辛二酸 Suberic acid 60 对羟基苯甲酸 4-hydroxybenzoic acid
28 牛磺酸 Taurine 61 3,4-二羟基苯乙酸 3,4-dihydroxyphenylacetic acid
29 3-(3-羟基苯基)-3-羟基丙酸 3-hydroxyphenyl-hydracrylic acid 62 3-(4-羟基苯基)乳酸 3-(4-hydroxyphenyl) lactic acid
30 (R)-3-羟基丁酸 3-D-hydroxybutyric acid 63 5-羟甲基-2-呋喃甲酸 5-hydroxymethyl-2-furoic acid
31 DL-3-苯基乳酸 3-phenyllactic acid 64 L-苹果酸 L-malic acid
32 阿魏酸 Ferulic acid 65 酒石酸 Tartaric acid
33 苯甲酸 Benzoic acid

图2

盐碱复合胁迫对小麦根系有机酸分泌的影响 A-D: 盐碱复合胁迫6 h (A)、12 h (B)、24 h (C)和48 h (D)时, 差异分泌有机酸数量的火山图; E: 44种有机酸分泌的差异热图, *表示与CK相比, 差异显著的有机酸; Class 1-Class 5为K-means聚类分组结果; F: 44种差异有机酸K-means聚类。"

图3

差异分泌有机酸的KEGG富集分析 A-D: 盐碱复合胁迫6 h (A)、12 h (B)、24 h (C)和48 h (D)时, 差异分泌有机酸的KEGG富集分析。"

图4

差异有机酸的分布模式 A-B: 4个比较组中差异有机酸上调(A)和下调(B)数量的韦恩图; C-F: CK vs 6 h (C)、CK vs 12 h (D)、CK vs 24 h (E)和CK vs 48 h (F) 4个比较组中差异变化Top 20的有机酸; G, H: 4个比较组中差异变化Top 20的有机酸上调(G)和下调(H)数量的韦恩图。"

附表2

RNA-seq质量"

Sample Clean reads Q30 rate (%) GC (%) Total mapped (%)
0 h_1 57,429,278 96.14 52.82 94.16
0 h_2 49,625,856 95.85 53.02 94.26
0 h_3 43,491,670 96.78 52.48 93.45
6 h_1 42,536,020 96.34 53.87 94.88
6 h_2 52,132,006 95.77 53.48 93.95
6 h_3 44,629,420 95.27 53.94 93.73
12 h_1 58,740,522 95.13 54.78 88.73
12 h_2 52,958,382 95.76 53.49 89.27
12 h_3 50,815,118 96.17 53.25 92.87
24 h_1 48,190,720 96.00 53.69 93.28
24 h_2 55,706,772 95.71 53.43 89.49
24 h_3 51,966,976 96.44 52.97 89.52
48 h_1 52,006,420 96.09 52.12 92.92
48 h_2 48,201,206 95.67 53.18 93.72
48 h_3 36,711,764 95.07 52.48 91.33

图5

盐碱复合胁迫影响小麦根系转录组的变化 A-D: 盐碱复合胁迫6 h (A)、12 h (B)、24 h (C)和48 h (D)时, 差异表达基因数量的火山图; E, F: 4个比较组中上调差异表达基因(E)和下调差异表达基因(F)数量的韦恩图。"

图6

差异表达基因的GO富集分析比较气泡图 A-B: 4个比较组中上调差异表达基因(A)和下调差异表达基因(B)的GO富集分析Top 20的比较气泡图。"

图7

差异表达基因的KEGG富集分析比较气泡图 A-B: 4个比较组中上调差异表达基因(A)和下调差异表达基因(B)的KEGG富集分析比较气泡图。"

图8

有机酸和转录组的WGCNA分析 A: WGCNA分析将DEGs划分为14个模块; B-F: Class 1 (B)、Class 2 (C)、Class 3 (D)、Class 4 (E)和Class 5 (F)的矩阵热图。"

图9

3个模块基因的富集分析和重要候选基因的筛选 A: MEblue、MEturquoise和MEyellow模块基因的显著KEGG富集条目比较气泡图; B-D: 4个比较组中共有的上调差异表达基因和MEblue (B)、MEturquoise (C)和MEyellow (D) 3个模块差异表达基因数量的韦恩图; E-G: MEblue (E)、MEturquoise (F)和MEyellow (G) 3个模块中重要的候选基因热图。FPKM: 每百万比对片段中, 每千碱基长度上映射到的片段数。"

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