Scientia Agricultura Sinica ›› 2026, Vol. 59 ›› Issue (7): 1400-1419.doi: 10.3864/j.issn.0578-1752.2026.07.003

• CROP GENETICS & BREEDING·GERMPLASM RESOURCES·MOLECULAR GENETICS • Previous Articles     Next Articles

Effects of Different Types of Salt Stress on Seed Germination of Pennisetum alopecuroides and Study on Sodium-Regulated Transcriptome

LU XueLi1,2,3(), GILLANI SyedaWajeeha1(), MENG Chen1,2,3, LI XiaoBin2,4, SONG YiRu1,5, BAI Yu1,5, WANG JuYing2, FENG XiaoFei2, LIU ChenChen2, LI YiQiang1,2,3,*(), XU ZongChang1,2,*()   

  1. 1 Marine Agriculture Research Center, Institute of Tobacco Research, Chinese Academy of Agricultural Sciences/Qingdao Key Laboratory of Resources Mining and Biological Breeding in Coastal Saline-alkali Land, Qingdao 266100, Shandong
    2 National Center of Technology Innovation for Comprehensive Utilization of Saline-Alkali Land, Dongying 257345, Shandong
    3 Shandong Center of Technology Innovation for Flower Technology, Weifang 261000, Shandong
    4 Institute of Agricultural Resources and Regional Planning, Chinese Academy of Agricultural Sciences, Beijing 100081
    5 College of Agriculture, Qingdao Agricultural University, Qingdao 266109, Shandong
  • Received:2025-09-11 Accepted:2025-12-01 Online:2026-04-08 Published:2026-04-08
  • Contact: LI YiQiang, XU ZongChang

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Abstract:

【Objective】This study aimed to explore the effects of different types and concentrations of salt stress on the seed germination of Pennisetum alopecuroides, and clarify the molecular mechanism underlying the “low-promotion and high-inhibition” effect of Na+ concentration on seed germination. The fingdinga are expected to provide a theoretical basis for the cultivation of P. alopecuroides in saline-alkali land and the breeding of salt-tolerant varieties.【Method】Seeds of the ornamental-fodder dual-purpose P. alopecuroides line "Langyan No. 1" were used as experimental materials. Different concentration gradients were set for six types of salts, inciuding NaCl, MgSO4, NaHCO3, KCl, Na2SO4, and mixed salt (NaHCO3:NaCl:Na2SO4=1:15:84). Germination-related indicators (germination energy, germination rate, bud length, and root length) were determined, and the membership function method was employed to evaluate the impact of different salt treatments on seed germination. Additionally, stress physiological indicators were measured, and transcriptome sequencing was performed on P. alopecuroides seeds treated with CK (control), 10 mmol·L-1 NaCl, and 100 mmol·L-1 NaCl for 3 days. Metabolic pathways and candidate genes involved in low-concentration Na+-promoted germination were analyzed, and the reliability of sequencing data was verified by qRT-PCR.【Result】Low concentrations of NaCl (10-25 mmol·L-1) and Na2SO4 (10 mmol·L-1) could promote seed germination, with significantly higher germination energy and germination rate than the control group, and the relative salt injury rate was 0. In contrast, high-concentration salts (≥100 mmol·L-1) inhibited germination, among which NaHCO3 showed the strongest inhibitory effect (no seed germination was observed at 200 mmol·L-1). For stress physiological indicators, low-concentration NaCl stress significantly increased the activities of superoxide dismutase (SOD) and catalase (CAT), as well as the content of proline (Pro); while high-concentration NaCl stress significantly induced the accumulation of malondialdehyde (MDA). Transcriptome analysis identified 14 259 differentially expressed genes (DEGs) in total. DEGs associated with low-concentration NaCl-promoted germination were mainly enriched in the oxidative phosphorylation pathway. Genes such as PPA and ATPeV1D in this pathway were positively correlated with germination indicators, whereas genes like ATPeF0A and ND1 were positively correlated with relative salt injury rate and MDA content. The inhibitory effect of high Na⁺ concentration on seed germination was mainly achieved by downregulating the expression of genes related to germination-promoting hormone synthesis pathways and upregulating the expression of genes involved in germination-inhibiting hormone synthesis pathways. qRT-PCR verification confirmed the reliability of the transcriptome data.【Conclusion】Under salt stress, Na+ concentration exerts a “low-promotion and high-inhibition” effect on seed germination of Pennisetum alopecuroides. Low-concentration of neutral sodium salts (NaCl, Na2SO4) can promote germination, while high-concentration salts (especially the alkaline salt NaHCO3) exhibit inhibitory effects. The salt tolerance mechanism of Pennisetum alopecuroides is associated with the regulation of antioxidant enzyme activities, proline accumulation, and the expression of genes in pathways such as oxidative phosphorylation.

Key words: salt stress, Pennisetum alopecuroides, seed germination, physiological response, differentially expressed gene, oxidative phosphorylation

Table 1

qRT-PCR primer required for this study"

基因Gene 引物序列Primer sequence (5′-3′) 产物长度Product length (bp)
ACTIN2 F:AGCACCAGAAGAGCATCCAGTT 138
R:AGTGAAAGGACCGCTTGAATAG
Alcohol Dehydrogenase 1
（ADH1）		 F:ATCGAGGAGGTGGAGGTTG 102
R:CTTGGCCTCCCAGAAGTAGA
Plasma membrane ATPase 1
（PMA1）		 F:GATCTACCTTGTCACCTCCATG 123
R:TCAGAGTGAGCGTTGTCGTAA
ATP synthase F1 subunit alpha
（ATPeF1A）		 F:GCCCTTCCCATCATTGAGA 132
R:GTTAATGGCGGGACGGAT
Pyrophosphatase
（PPA）		 F:ATCCCGAGTACCGCCACTA 117
R:CGTCTACAGCCACTTCTTTG
Cytochrome c Oxidase Subunit 6A
（COX6A）		 F:TTGGATTGCCGACAACGA 107
R:AGGGAATGACAGCGAAGATC
ATP synthase F1 subunit delta
（ATPF1D）		 F:CCTCAAGACGCAGCTCAAC 118
R:CTCTGATGCGATCTCCTCAAT
ATP synthase F0 subunit D
（ATPeF0D）		 F:GACCGTTGACTTCTCCCACTAC 111
R:TGGCGGCTGACATCGTA
ATP synthase F1 subunit beta
（ATPeF1B）		 F:GTCTCAACCTTTCGCTGTCG 121
R:TTCGGGCAAAGCATCGT

Fig. 1

Statistical chart of daily germination of Langyan No.1 seeds under different types and concentrations of salt stress"

Table 2

Effects of different types and concentrations of salt stress treatments on the germination indices of Langyan No.1 seeds"

指标
Index		 处理
Treatments		 NaCl MgSO4 NaHCO3 KCl Na2SO4 混合盐
Salt mixtures
发芽势
Germination energy (%)		 CK 75.56±0.08b 75.56±0.08a 75.56±0.08a 75.56±0.08a 75.56±0.08a 75.56±0.08a
1 82.22±0.05a 71.11±0.13ab 66.67±0.09b 66.67±0.07ab 77.78±0.02a 71.11±0.10a
2 80.00±0.00a 65.56±0.02b 62.22±0.05b 65.56±0.07b 61.11±0.02b 68.89±0.05ab
3 71.11±0.15b 62.22±0.13b 28.89±0.11c 61.11±0.05b 51.11±0.12bc 67.78±0.04ab
4 44.44±0.02c 54.44±0.14c 7.78±0.02d 57.78±0.05bc 15.56±0.07c 41.11±0.13c
5 4.44±0.08d 38.89±0.15d 0.00±0.00d 23.33±0.10c 0.00±0.00d 13.33±0.06d
发芽率
Germination rate (%)		 CK 75.56±0.08b 75.56±0.08a 75.56±0.08a 75.56±0.08a 75.56±0.08b 75.56±0.08a
1 82.22±0.05a 72.22±0.12a 66.67±0.09b 70.00±0.03ab 82.22±0.04a 68.89±0.05b
2 80.00±0.00a 66.67±0.03ab 63.33±0.06b 67.78±0.10b 64.44±0.02bc 73.33±0.12a
3 71.11±0.15b 64.44±0.15b 31.11±0.10c 64.44±0.04b 54.44±0.10c 68.89±0.05b
4 47.78±0.04c 56.67±0.12bc 8.89±0.02d 58.89±0.04bc 36.67±0.09d 42.22±0.12c
5 5.56±0.10d 40.00±0.13c 0.00±0.00d 33.33±0.09c 2.22±0.04e 15.56±0.08d
相对盐害率
Relative salt injury rate		 1 0（1） 0.04（3） 0.12（8） 0.07（5） 0.00（1） 0.09（6）
2 0（1） 0.12（8） 0.16（10） 0.10（7） 0.15（9） 0.03（2）
3 0.06（4） 0.15（9） 0.59（19） 0.15（9） 0.28（13） 0.09（6）
4 0.37（14） 0.25（12） 0.88（21） 0.22（11） 0.51（17） 0.44（15）
5 0.93（22） 0.47（16） 1.00（24） 0.56（18） 0.97（23） 0.79（20）

Fig. 2

Effects of NaCl stress at different concentrations on physiological indices of Langyan No.1 seeds"

Fig. 3

Transcriptome data analysis of Pennisetum alopecuroides samples under different concentrations of NaCl stress for 3 days A: PCA analysis; B: Statistics of the number of up-regulated and down-regulated differentially expressed genes; C: Venn diagram analysis; D: Cluster analysis of differentially expressed genes"

Table 3

Analysis of the expression levels of differentially expressed genes related to stress-responsive physiological indicators and their correlation with stress-responsive physiological indicators"

基因编号
Gene ID		 基因
Gene		 基因表达量(FPKM) Gene expression level 逆境响应指标
Stress response indicators		 相关性
Correlation
CK 10 mmol·L-1 NaCl 100 mmol·L-1 NaCl
TRINITY_DN5743_c1_g1 Catalase isozyme 3 116.24 377.01 213.03 CAT 0.9871**
TRINITY_DN9351_c0_g1 Catalase 1 88.02 145.15 103.17 CAT 0.9992**
TRINITY_DN1109_c0_g1 Superoxide dismutase 1.1 89.32 136.52 130.71 SOD 0.9865**
TRINITY_DN45156_c0_g1 Superoxide dismutase 2 159.32 321.35 287.34 SOD 0.9686**
TRINITY_DN42450_c0_g1 Superoxide dismutase 7 7.55 17.68 20.45 SOD 0.9883**
TRINITY_DN30550_c0_g1 Superoxide dismutase 22 72.34 108.59 98.32 SOD 0.9461**
TRINITY_DN21193_c0_g1 Superoxide dismutase 35 18.34 37.62 30.34 SOD 0.9066**
TRINITY_DN10919_c0_g1 Glutathione S-transferase 2 808.61 813.16 391.12 MDA 0.6322*
TRINITY_DN190_c0_g2 Glutathione S-transferase GSTU1 105.13 105.84 31.42 MDA -0.9999**
TRINITY_DN5710_c0_g1 Glutathione S-transferase 4 124.79 65.09 17.51 MDA -0.8268**
TRINITY_DN4196_c0_g1 Glutathione S-transferase 3 34.67 41.25 117.54 MDA 0.9968**
TRINITY_DN11259_c0_g1 Glutathione S-transferase GSTF1 63.09 71.37 8.68 MDA -0.9936**
TRINITY_DN17343_c0_g1 Glutathione S-transferase GSTU6 21.36 31.66 70.26 MDA 0.9781**
TRINITY_DN7740_c0_g3 Glutathione synthetase isoform X2 55.69 51.87 5.87 MDA -0.9969**
TRINITY_DN7032_c0_g1 Glutathione S-transferase 10 3.94 5.03 2.12 MDA -0.9319**
TRINITY_DN9190_c0_g1 Proline dehydrogenase 2 71.01 54.10 123.09 Pro -0.7231*
TRINITY_DN2129_c0_g1 Proline transporter 1 90.42 120.04 67.14 Pro 0.9173**
TRINITY_DN17973_c0_g1 Proline transporter 7 10.09 14.65 7.66 Pro 0.9544**

Fig. 4

GO and KEGG pathway enrichment analysis of differentially expressed genes"

Fig. 5

The expression patterns analysis of differentially expressed genes in the oxidative phosphorylation pathway"

Fig. 6

Correlation analysis between candidate germination-promoting genes and agronomic physiological indicators"

Fig. 7

The expression patterns analysis of differentially expressed genes in plant hormone signal transduction pathways"

Fig. 8

Validation of gene expression patterns by qRT-PCR"

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