Scientia Agricultura Sinica ›› 2026, Vol. 59 ›› Issue (13): 2815-2827.doi: 10.3864/j.issn.0578-1752.2026.13.005

• TILLAGE & CULTIVATION·PHYSIOLOGY & BIOCHEMISTRY·AGRICULTURE INFORMATION TECHNOLOGY • Previous Articles     Next Articles

Analysis of Physiological and Metabolic Response Mechanisms in Maize Varieties with Different Salt Tolerances Under Salt Stress

MA XiaoYing1(), QU XiaoYu1, LI Tao1, LIU WeiFan1, LIU Hao1, MA FengLan1, WAN MengHu1, WU Na1(), LIU JiLi2   

  1. 1 College of Agriculture, Ningxia University, Yinchuan 750021
    2 College of Ecology and Environment, Ningxia University, Yinchuan 750021
  • Received:2025-11-25 Accepted:2026-05-28 Online:2026-07-01 Published:2026-07-01
  • Contact: WU Na

Abstract:

【Objective】As a major food crop in China, maize (Zea mays L.) is relatively sensitive to salt stress, which significantly inhibits its photosynthesis and causes metabolic disorders. Therefore, this study utilized maize varieties with different levels of salt tolerance to systematically investigate their photosynthetic physiology and metabolic responses under salt stress, with the aim of elucidating the underlying causes of inter-varietal differences in salt tolerance at the physiological and metabolic levels. 【Method】A pot experiment was conducted using salt-tolerant maize varieties Yinyu 238 (YY238) and Heyu 157 (HY157), and the salt-sensitive variety Xianyu 335 (XY335). Three salt stress concentration gradients were established: 0 mmol NaCl·L-1 (CK), 120 mmol NaCl·L-1 (S1), and 240 mmol NaCl·L-1 (S2). Stress treatment was applied at the six-leaf stage of the maize seedlings. Samples were taken at 24 hours after stress treatment to measure the net photosynthetic rate (Pn), photosynthetic performance index (PI), ion content (K+, and Na+), and the content of malondialdehyde (MDA), proline (Pro), abscisic acid (ABA), and jasmonic acid (JA). Non-targeted metabolomics technology was used to analyze leaf metabolites. Differential metabolites were screened based on a variable importance in projection (VIP) > 1 and P<0.05, and pathway enrichment analysis was performed using the KEGG database. 【Result】Salt stress significantly reduced both Pn and PI. Compared with CK, the salt-sensitive variety XY335 exhibited a maximum Pn reduction of 39.8% under S2, while the salt-tolerant variety YY238 showed a relatively smaller reduction of 31.9%. In terms of ion homeostasis, salt stress significantly decreased the shoot K+/Na+ ratio, with XY335 experiencing a reduction of 76.4%, significantly higher than the 74.6% reduction in YY238. Salt-tolerant varieties demonstrated stronger osmotic adjustment capacity, with YY238 showing a 199.2% increase in Pro accumulation. In contrast, the salt-sensitive variety XY335 suffered more severe oxidative damage, with a 172.1% increase in MDA content. Hormonal responses indicated that XY335 had a higher increase in ABA (59.3%), while YY238 showed the largest increase in JA under S2, reaching 56.3%. Metabolomic analysis revealed that salt stress induced variety-specific metabolic reprogramming. The metabolic disorder in the salt-sensitive variety XY335 worsened with increasing stress intensity, with the number of differential metabolites surging to 127. In contrast, salt-tolerant varieties exhibited a stronger ability to maintain metabolic homeostasis. Salt-tolerant varieties specifically accumulated stress resistance-related metabolites: YY238 significantly upregulated amino acid derivatives and organic acids, including Nα-methylhistidine and trans-aconitate; HY157 specifically accumulated amino acids and their derivatives, such as γ-aminobutyric acid and L-glutamine, as well as phenolic acids like salicylic acid. In comparison, the salt-sensitive variety XY335 primarily accumulated nucleic acid metabolites related to energy metabolism, such as β-guanidinopropionic acid, and organic acids like isonicotinic acid. KEGG pathway enrichment analysis revealed that salt-tolerant varieties were commonly enriched in pathways, such as phenylpropanoid biosynthesis and ABC transporters. Additionally, YY238 specifically activated the biosynthesis of flavonoids and flavonols, while HY157 was significantly enriched in alanine, aspartate, and glutamate metabolism pathways. 【Conclusion】In summary, salt stress enhanced the adaptability of salt-tolerant maize varieties by maintaining photosynthetic performance and ion homeostasis, mitigating oxidative damage, and activating specific metabolic pathways. Salt-tolerant varieties maintained higher photosynthetic efficiency through precise ion compartmentalization and alleviate oxidative damage by rapidly accumulating osmotic adjustment substances and effectively scavenging reactive oxygen species. At the metabolic level, salt-tolerant varieties specifically activated pathways, such as phenylpropanoid biosynthesis and amino acid metabolism, providing the key material and signaling foundations for their salt tolerance.

Key words: maize, salt stress, photosynthetic characteristics, physiological response, metabolomics, metabolic pathways

Fig. 1

Seedling growth phenotypes of maize varieties under salt stress A: XY335 (Xianyu 335, salt-sensitive variety); B: YY238 (Yinyu 238, salt-tolerant variety); C: HY157 (Heyu 157, salt-tolerant variety). CK, S1, and S2 represent salt concentration treatments with 0, 120, and 240 mmol NaCl·L-1, respectively. The same as below"

Fig. 2

Responses of ion homeostasis and photosynthetic characteristics in maize varieties under salt stress *:P<0.05,**:P<0.01,***:P<0.001;ns表示无显著差异(P≥0.05) ns indicates no significant difference (P≥0.05)。下同 The same as below"

Fig. 3

Salt stress-induced oxidative damage, osmoregulation and hormonal signaling changes"

Fig. 4

Metabolomics data quality assessment of different salt-tolerant maize varieties A: RSD distribution of features in QC samples; B: OPLS-DA score plot of metabolites among groups. The RSD distribution shows that approximately 65% of features in QC samples had RSD<30%, indicating acceptable data quality. The OPLS-DA plot demonstrates the metabolic separation among salt-sensitive maize (XY335) and salt-tolerant maize (YY238, and HY157) under different salt concentration treatments (CK, S1, and S2)"

Fig. 5

Classification and statistics of differential metabolites and intergroup variation"

Fig. 6

Volcano plot of differentially altered metabolites The x-axis represents the log2-transformed fold change of metabolite quantification values between two samples; the y-axis represents the -log10 transformed P-value. Each point in the plot represents a metabolite. Red, blue, and gray points indicate significantly up-regulated, significantly down-regulated, and non-significant metabolites, respectively. Point size corresponds to the VIP value. The numbers in the top right corner indicate the counts of metabolites annotated by each color. The top 5 metabolites with the smallest P-values are labeled. Metabolites with -log10 (P value)≥1.3 were considered significantly expressed"

Fig. 7

Differential metabolites in maize cultivars in response to salt stress and KEGG enrichment analysis A and B are Venn diagrams of differential metabolites within and between maize varieties, respectively, while C and D are KEGG enrichment distribution diagrams of differential metabolites within and between maize varieties, respectively"

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