5‑AzaC缓解紫花苜蓿盐碱胁迫的生理效应及其对DNA甲基化酶基因表达的影响

doi:10.3864/j.issn.0578-1752.2025.21.017

Abstract

Abstract:

【Background】Soil salinization is one of the major ecological challenges threatening global agricultural production, severely restricting crop growth, yield formation, and quality improvement. Alfalfa (Medicago sativa), as an important perennial legume forage, is particularly constrained by salt-alkali stress, while the epigenetic regulatory mechanisms underlying its response remain largely unknown. DNA methylation, as a key epigenetic modification, plays an essential role in plant adaptation to abiotic stresses. 【Objective】This study aimed to systematically identify DNA methylation-related gene families in alfalfa, characterize their expression patterns under salt-alkali stress, and further explore the role of DNA methylation in salt-alkali tolerance by applying the DNA methylation inhibitor 5-azacytidine (5-AzaC), thereby providing the theoretical insights for the genetic improvement of salt-alkali-tolerant alfalfa. 【Method】Based on the reference genome of alfalfa, DNA methyltransferase and demethylase genes were identified genome-wide, and their functions were inferred through phylogenetic analysis and conserved domain annotation. RT-qPCR was employed to analyze the expression patterns of these genes under salt-alkali stress. Using the cultivar Gannong No. 3 as plant material, a hydroponic salt-alkali stress system was established. Different concentrations of 5-AzaC were applied as pretreatments, and the optimal concentration was selected for subsequent assays. Plant growth, physiological, and photosynthetic parameters were then measured to evaluate the regulatory role of 5-AzaC in alfalfa salt-alkali tolerance. 【Result】A total of 13 DNA methyltransferase genes and 4 DNA demethylase genes were identified in alfalfa, all of which were localized to the nucleus with complete conserved domains. Expression analysis revealed that MsCMT4, MsCMT6, MsCMT8, and MsDML2 were significantly upregulated under salt-alkali stress, indicating the involvement of both methylation and demethylation processes in stress responses. Physiological analyses showed that 100 μmol·L^-1 5-AzaC significantly alleviated growth inhibition caused by salt-alkali stress, so plant height, fresh weight, and dry weight increased by 12.62%, 23.50%, and 18.67%, respectively, compared with the control. Regarding chlorophyll metabolism, 5-AzaC suppressed the expression of chlorophyll degradation-related genes (PAO, CAO, NYC), thereby delaying pigment degradation. Photosynthetic analysis indicated that 5-AzaC treatment markedly increased net photosynthetic rate (Pn), stomatal conductance (Gs), and transpiration rate (Tr), as well as the quantum efficiency of photosystem II (YII) and photochemical quenching (qP), suggesting enhanced stability and efficiency of the photosynthetic system. In terms of osmotic adjustment, 5-AzaC promoted soluble sugar accumulation (+37.21%) but had no significant effect on soluble protein. Reactive oxygen species (ROS) measurements showed that 5-AzaC reduced H₂O₂ and O₂^-·levels by 22.8% and 35.8%, respectively, while superoxide dismutase (SOD) and catalase (CAT) activities increased by 13.58% and 21.82%, respectively, indicating that 5-AzaC enhanced antioxidant capacity and alleviated oxidative damage. 【Conclusion】This study systematically characterized DNA methylation-related gene families in alfalfa and their responses to salt-alkali stress, revealing the pivotal role of DNA methylation in shaping salt-alkali tolerance. Exogenous application of 5-AzaC improved alfalfa tolerance by maintaining photosystem stability, enhancing photosynthetic efficiency, promoting osmolyte accumulation, and strengthening ROS scavenging capacity. These findings provided the new experimental evidence for understanding the epigenetic mechanisms of forage adaptation to abiotic stress and offer theoretical guidance for the improvement and utilization of salt-alkali-tolerant alfalfa germplasm.

Key words: salt-alkali stress, Medicago sativa, epigenetics, photosynthetic pigments

GAO Rong, LI HengYu, CHEN LiJuan, MA HuiLing. Physiological Effects of 5-AzaC on Alleviating Salt‑Alkali Stress in Alfalfa and Its Impact on the Expression of DNA Methylation Enzyme Genes[J].Scientia Agricultura Sinica, 2025, 58(21): 4482-4496.

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References 51

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基因名称 Gene name	正向引物 Forward primer（5′-3′）	反向引物 Reverse primer（5′-3′）
MsActin	GACAATGGAACTGGAATGG	CAATACCGTGCTCAATGG
MsCMT1	CGCCTGAGTTTGAGCGCA	TGGCATTTCGTCCCGAGT
MsCMT2	CCCCAGCGTTGTACCTGG	TCAGTATGAGGAGTGGTCCCA
MsCMT3	GCTGACTCTCAAAGTGCACC	GCACTCCCTTGAGGTCCC
MsCMT4	GACATACTGCCCAGCTGCA	GTCAAAGGAGGTCCGCCA
MsCMT5	AGGTTCTTCTCCACCCGGA	ACGGGAACAGCAACTGCA
MsCMT6	AGCTACGGGGCTTCTCCT	GCATTCCACTTGGTGCGC
MsCMT7	TGGCCTCAACGTTACGCA	TCCCTTCCTCGCCCTTGA
MsCMT8	GGACCTGGGCCCTTTGAA	GTCCACTGGGGTGAGTGC
MsCMT9	GCTGTTCCTGTGGCCCTT	CGGGCTAGACAACTTGGGT
MsDNMT1	TTCACCGAAGCTTGGCGT	CACTCGATCAACGGCGGA
MsMET1	GCTGCCGTCCGTAGTACC	CCATTGCCTACAGCTGGGA
MsMET2	CGTCCGCAGTACCGCAAA	CCATTGCCTACAGCTGGGA
MsMET3	CAGTCCGCAGTACCGCAA	CCATTGCCTACAGCTGGGA
MsDML1	TCTGGCAGAGATTTGTCCTGT	GCTGTTTGGCATTACTGTGGC
MsDML2	AACGAACGGTGCATGGGT	TGGTCTGCTTGCCAGTCTG
MsDML3	CAGGACTGGAGCAGAAGAACA	TCAGGGAGAGACAAGGGCA
MsDML4	TGGGATTGTACCTGCGGC	CCGAAGCCGCCTCTGTC
MsCAO	CCGGCATCTGGTCTGCAA	CCGGGCTTCGAAATCCCA
MsCLH	AGCTGCAATAACAAGTTGGCT	TTTCCGCCGCGACTATGG
MsNYC	GTTCCACGAATGCGAGTCG	GACCTCGCCGTACCCAAG
MsPAO	CCGAGAATGGCGGTGACA	CGTGTCGCCTCAGCCAAT

基因名称 Gene name	基因ID Gene ID	染色体 Chromosome	氨基酸数 Number of amino acids	分子量Molecular weigh	等电点 Point isoelectric	不稳定系数Instability index	亲水性 GRAVY value	亚细胞定位 Subcellular localization
MsMET1	MS.gene071054.t1	chr4.1	1653	185714.22	5.53	44.39	-0.498	Nuclear
MsMET2	MS.gene043208.t1	chr4.2	1506	169133.86	5.98	44.2	-0.492	Nuclear
MsMET3	MS.gene010135.t1	chr5.2	1311	146538.82	5.95	43.9	-0.381	Nuclear
MsCMT1	MS.gene056526.t1	chr8.3	1124	126873.89	5.78	42.68	-0.576	Nuclear
MsCMT2	MS.gene010674.t1	chr4.4	850	96315.79	5.22	43.52	-0.563	Nuclear
MsCMT3	MS.gene64852.t1	chr5.1	835	94196.68	5.36	41.38	-0.549	Nuclear
MsCMT4	MS.gene72088.t1	chr6.3	931	103979.18	5.67	44.82	-0.548	Nuclear
MsCMT5	MS.gene51465.t1	chr3.4	894	100820.99	6.28	36.94	-0.427	Nuclear
MsCMT6	MS.gene92341.t1	chr5.1	859	96724.01	7.93	38.79	-0.526	Nuclear
MsCMT7	MS.gene50806.t1	chr4.1	550	62072.73	4.67	37.23	-0.488	Nuclear
MsCMT8	MS.gene009335.t1	chr3.1	481	53450.93	5.99	36.33	-0.28	Nuclear
MsCMT9	MS.gene89792.t1	chr4.4	652	74165.94	5.85	42.92	-0.54	Nuclear
MsDNMT1	MS.gene002215.t1	chr2.3	427	48313.21	4.89	40.8	-0.397	Nuclear
MsDML1	MS.gene050207.t1	chr1.1	1497	168803.93	6.07	44.77	-0.732	Nuclear
MsDML2	MS.gene60556.t1	chr1.2	1795	202529.87	6.28	43.97	-0.776	Nuclear
MsDML3	MS.gene020552.t1	chr7.2	1798	201809.19	6.06	47.28	-0.747	Nuclear
MsDML4	MS.gene47350.t1	chr1.1	2232	247919.17	5.88	52.08	-0.776	Nuclear

Physiological Effects of 5-AzaC on Alleviating Salt‑Alkali Stress in Alfalfa and Its Impact on the Expression of DNA Methylation Enzyme Genes

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