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Journal of Integrative Agriculture
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Spermidine treatment promotes soybean germination under bicarbonate stress

Tingting Wang*, Haikuo Song*, Kexin Zhang, Yongjing Sun, Yongbin Zhuang, Xiaoming Li, Jinfei Zhang, Yiran Meng, Chenfei Shan, Yue Xin, Baoyin Chen#, Dajian Zhang#

National Key Laboratory of Wheat Improvement, College of Agronomy, Shandong Agricultural University, Tai’an, China

 Highlights 

l Spermidine treatment alleviates bicarbonate stress in germinating soybean seeds.

l Spermidine upregulates the phenylpropanoid and flavonoid pathways to enhance stress tolerance.

GmPAL1.1-Hap1 and GmCCR1-Hap3 are linked to adaptation to alkaline stress.

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摘要  

由碳酸氢盐胁迫驱动的土壤碱化严重抑制大豆(Glycine max L.)种子萌发。尽管已知亚精胺(Spd)能够增强植物对非生物胁迫的耐受性,但其是否以及如何缓解碳酸氢盐胁迫尚不明确。本研究表明,Spd处理可显著缓解大豆萌发过程中所遭受的碳酸氢盐胁迫,表现为更高的萌发率和更大的根系生物量。生理指标表明,Spd可降低活性氧和丙二醛的积累,同时促进脯氨酸的积累。转录组及加权共表达网络分析(WGCNA)表明,Spd在萌发种子中持续上调苯丙烷和类黄酮生物合成途径的相关基因。这些途径中的两个关键基因GmPAL1.1Phenylalanine Ammonia-Lyase 1.1)和GmCCR1Cinnamoyl-CoA Reductase 1与种子萌发相关。此外,GmPAL1.1GmCCR1的自然变异与对盐碱地的适应性相关:优良单倍型GmPAL1.1-Hap1GmCCR1-Hap3与有益的农艺性状及对盐碱地的适应性相关。这些发现揭示了Spd通过协调抗氧化防御机制并激活苯丙烷和类黄酮生物合成途径来增强碳酸氢盐耐受性。此外,本研究结果还表明GmPAL1.1GmCCR1可作为培育耐碱大豆品种的潜在优良基因。



Abstract  

Soil alkalization, driven by bicarbonate stress, severely inhibits soybean (Glycine max L.) seed germination. While spermidine (Spd) is known to enhance plant tolerance to abiotic stress, whether and how it alleviates bicarbonate stress remains unclear. Here, we demonstrate that Spd treatment significantly alleviates bicarbonate stress experienced by in germinating soybeans, as evidenced by higher germination rates and greater root biomass. Physiological assays revealed that Spd diminishes accumulation of reactive oxygen species accumulation and malondialdehyde while promoting proline accumulation. Transcriptome and weighted gene co-expression network analysis (WGCNA) indicated that Spd continuously upregulates genes in the phenylpropanoid and flavonoid biosynthesis pathways in germinating seeds. Two key genes in these pathways, GmPAL1.1 (Phenylalanine Ammonia-Lyase 1.1) and GmCCR1 (Cinnamoyl-CoA Reductase 1), are associated with seed germination. Furthermore, natural variation in GmPAL1.1 and GmCCR1 is associated with adaptation to saline-alkaline soils: the elite haplotypes GmPAL1.1-Hap1 and GmCCR1-Hap3 are associated with favorable agronomic traits and improved adaptation to these soils. These findings reveal that Spd enhances bicarbonate tolerance by coordinating antioxidant defense mechanisms and activating the phenylpropanoid and flavonoid biosynthesis pathways. In addition, our results identify GmPAL1.1 and GmCCR1 as potential elite genes for breeding alkaline-tolerant soybean cultivars.

Keywords:  soybean       seed germination              spermidine              bicarbonate stress              alkaline tolerance  
Online: 09 May 2026  
Fund: 

This work was supported by the Agricultural Variety Improvement Project of Shandong Province, China (2024LZGC030 and 2023LZGC006), the National Natural Science Foundation of China (32572365, 32322062, and 32441057), the National Key R&D program of China (2024YFF1000502), the Scientific Research Innovation Capability Support Project for Young Faculty (SRICSPYF-BS2025109), the Natural Science Foundation of Shandong Province, China (ZR2023JQ009), and the Shandong Agriculture Research System, China (SDARS-28-01 and SDARS-28-06).

About author:  #Correspondence Baoyin Chen, E-mail: bychen@sdau.edu.cn; Dajian Zhang, E-mail: dajianzhang@sdau.edu.cn *These authors contributed equally to this work.

Cite this article: 

Tingting Wang, Haikuo Song, Kexin Zhang, Yongjing Sun, Yongbin Zhuang, Xiaoming Li, Jinfei Zhang, Yiran Meng, Chenfei Shan, Yue Xin, Baoyin Chen, Dajian Zhang. 2026. Spermidine treatment promotes soybean germination under bicarbonate stress. Journal of Integrative Agriculture, Doi:10.1016/j.jia.2026.05.008

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