外源硫化氢调控盐碱胁迫下裸燕麦叶片糖和酚酸代谢反应

刘建新; 刘瑞瑞; 刘秀丽; 贾海燕; 卜婷; 李娜

doi:10.12357/cjea.20220649

外源硫化氢调控盐碱胁迫下裸燕麦叶片糖和酚酸代谢反应

doi: 10.12357/cjea.20220649

甘肃省陇东生物资源保护利用与生态修复重点实验室/陇东学院生命科学与技术学院　庆阳　745000

基金项目: 国家自然科学基金项目(31960375)和甘肃省自然科学基金项目(20JR5RA491)资助

详细信息

通讯作者:
刘建新, 主要从事植物逆境生理生态研究。E-mail: liujx1964@163.com

中图分类号: S512.6
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出版历程
- 收稿日期: 2022-08-22
- 录用日期: 2022-11-26
- 网络出版日期: 2023-02-10

Exogenous hydrogen sulfide modulates metabolic responses of sugar and phenolic acid in naked oat leaves under saline-alkali stress

Gansu Key Laboratory of Protection and Utilization for Biological Resources and Ecological Restoration in East Gansu / College of Life Sciences and Technology, Longdong University, Qingyang 745000, China

Funds: This work was supported by the National Natural Science Foundation of China (31960375) and the Natural Science Foundation of Gansu Province (20JR5RA491).

More Information

Corresponding author: LIU Jianxin, E-mail: liujx1964@163.com

摘要

摘要: 为明确硫化氢信号对盐碱胁迫下植物代谢组的调控作用, 揭示其增强植物耐盐碱性的机理, 以裸燕麦(Avena nuda L.)为材料进行盆栽土培试验, 设置不添加盐碱和添加3.00 g·kg⁻¹盐碱(摩尔比NaCl∶Na₂SO₄∶Na₂CO₃∶NaHCO₃=12∶8∶1∶9)与裸燕麦抽穗期叶面喷施蒸馏水和50 µmol·L⁻¹硫化氢供体硫氢化钠溶液交叉共4组处理, 利用超高效液相色谱-串联质谱技术结合正交偏最小二乘判别分析方法, 研究外源硫化氢对盐碱胁迫下裸燕麦叶片糖分解代谢产物水平、氧化还原平衡、酚酸含量和产量性状的影响。结果表明: 1)非盐碱条件下, 喷施硫氢化钠对裸燕麦叶片还原型谷胱甘肽/氧化型谷胱甘肽、还原型辅酶Ⅱ/氧化型辅酶Ⅱ、腺苷三磷酸含量和产量性状的影响微弱, 但显著上调柠檬酸、琥珀酸和6-磷酸葡萄糖酯含量, 显著下调葡萄糖-6-磷酸、丙酮酸、乳酸、α-酮戊二酸、谷氨酸、天冬酰胺、赤藓糖-4-磷酸、景天庚酮糖-7-磷酸含量; 盐碱胁迫导致裸燕麦叶片葡萄糖、葡萄糖-6-磷酸、果糖-6-磷酸、果糖-1,6-二磷酸、3-磷酸甘油醛、3-磷酸甘油酸、丙酮酸、乳酸、α-酮戊二酸、谷氨酸、谷氨酰胺、天冬酰胺、赤藓糖-4-磷酸、景天庚酮糖-7-磷酸、核糖-5-磷酸等糖酵解、三羧酸循环和戊糖磷酸途径中间代谢物及还原型谷胱甘肽、氧化型谷胱甘肽、还原型辅酶Ⅱ、氧化型辅酶Ⅱ含量显著降低, 而还原型谷胱甘肽/氧化型谷胱甘肽显著提高; 喷施硫氢化钠显著提高盐碱胁迫下裸燕麦叶片葡萄糖、果糖-6-磷酸、3-磷酸甘油酸、乳酸、α-酮戊二酸、延胡索酸、苹果酸、谷氨酰胺、6-磷酸葡萄糖酯、景天庚酮糖-7-磷酸含量, 显著降低天冬酰胺含量。2)喷施硫氢化钠显著下调非盐碱条件下裸燕麦叶片反式肉桂酸和丁香醛含量; 盐碱胁迫导致裸燕麦叶片反式肉桂酸含量显著降低, 而苯甲酸、对香豆酸和反式阿魏酸含量显著提高; 喷施硫氢化钠显著提高盐碱胁迫下裸燕麦叶片4-羟基苯甲酸和香草醛含量, 显著降低水杨酸、芥子酸含量。3)喷施硫氢化钠对盐碱胁迫造成的裸燕麦穗粒数量和籽粒产量下降具有显著缓解作用, 但对穗数量、穗铃数量、千粒重量和生物学产量的影响不显著。由此表明, 外源硫化氢参与裸燕麦糖分解代谢和酚酸水平调控, 能够增强裸燕麦耐受盐碱胁迫的能力, 它对糖分解途径有机酸水平的提升作用和酚酸独特的调节效应可能在其增强裸燕麦耐盐碱性中发挥着重要作用。
- 硫化氢 /
- 盐碱胁迫 /
- 裸燕麦 /
- 糖代谢 /
- 酚酸
Abstract: In order to clarify the regulatory effect of hydrogen sulfide signaling on the plant metabolome under saline-alkali stress and to reveal its mechanism of enhancing plant saline-alkali tolerance, a pot experiment was conducted with naked oat (Avena nude) as the material. Four treatments were applied to potted naked oat plants in a 2 × 2 factorial combination, including 0 or 3.00 g·kg⁻¹ saline-alkali (molar ratio of NaCl∶Na₂SO₄∶Na₂CO₃∶NaHCO₃at 12∶8∶1∶9) added to the potting soil and spraying with distilled water or 50 µmol·L⁻¹sodium hydrosulfide (a hydrogen sulfide donor) on leaves at the heading stage. The effects of exogenous hydrogen sulfide on glycolytic metabolite levels, redox balance, and phenolic acid content in leaves and on the yield traits of naked oats under the four treatments were investigated using ultra-performance liquid chromatography-tandem mass spectrometry technology combined with orthogonal partial least squares discriminant analysis. Under non-saline-alkali conditions, sodium hydrosulfide application did not have a significant effect on the ratios of (reduced glutathione)/(oxidized glutathione) and (reduced coenzyme Ⅱ)/(oxidized coenzyme Ⅱ), adenosine triphosphate content in leaves, and yield traits of naked oats; however, the levels of citrate, succinate, and 6-phosphogluconolactone were significantly upregulated and those of glucose-6-phosphate, pyruvate, lactate, α-ketoglutaric acid, glutamate, asparagine, erythrose-4-phosphate, and sedoheptulose-7-phosphate were significantly downregulated in the leaves. Saline-alkali stress significantly reduced the levels of glucose, glucose-6-phosphate, fructose-6-phosphate, fructose-1,6-diphosphate, 3-phosphate glyceraldehydes, 3-phosphoglyceric acid, pyruvate, lactate, α-ketoglutaric acid, glutamate, glutamine, asparagine, erythrose-4-phosphate, sedoheptulose-7-phosphate, ribose-5-phosphate, reduced glutathione, oxidized glutathione, reduced coenzyme Ⅱ, and oxidized coenzyme Ⅱ in the leaves of naked oats; whereas the ratio of reduced glutathione to oxidized glutathione was increased significantly. Spraying with sodium hydrosulfide significantly increased the levels of glucose, fructose-6-phosphate, 3-phosphoglyceric acid, lactate, α-ketoglutaric acid, fumarate, malate, glutamine, 6-phosphogluconolactone, and sedoheptulose-7-phosphate in the leaves of naked oats under saline-alkali stress, and significantly decreased the asparagine content. The levels of trans-cinnamic acid and syringaldehyde in the leaves of naked oats under non-saline-alkali conditions were significantly decreased by spraying with sodium hydrosulfide. Saline-alkali stress significantly reduced the content of trans-cinnamic acid in the leaves of naked oats, and markedly increased the levels of benzoic acid, p-hydroxycinnamic acid, and trans-ferulic acid. Spraying with sodium hydrosulfide significantly increased the levels of 4-hydroxybenzoic acid and vanillin in the leaves of naked oats under saline-alkali stress, and remarkably decreased the levels of salicylic acid and 4-hydroxy-3,5-dimethoxycinnamic acid.There was no significant increase in spike number, spike boll number, thousand-grain weight, and biological yield of naked oats under saline-alkali stress as a result of spraying with sodium hydrosulfide; however, sodium hydrosulfide significantly alleviated the decrease in spike grain number and grain yield induced by saline-alkali stress. These results indicate that exogenous hydrogen sulfide participates in the regulation of sugar catabolism and phenolic acid levels in naked oats, which can enhance the saline-alkali tolerance of naked oats. The increasing effect of exogenous hydrogen sulfide on organic acid levels in the sugar decomposition pathway and the unique regulatory effect on phenolic acids may play an important role in enhancing the saline-alkali tolerance of naked oats.
- Hydrogen sulfide /
- Saline-alkali stress /
- Naked oat (Avena nuda) /
- Carbohydrate catabolism /
- Phenolic acid

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图 1 不同外源硫化氢喷施和盐碱胁迫处理下裸燕麦叶片糖分解代谢物聚类热图分析

CK: 无盐碱胁迫下喷水; SA: 盐碱胁迫下喷水; SA+NaHS: 盐碱胁迫下喷NaHS; NaHS: 无盐碱胁迫下喷NaHS。Citrate: 柠檬酸; Asn: 天冬酰胺; Fructose: 果糖; Fumarate: 延胡索酸; Malate: 苹果酸; 6PG: 6-磷酸葡萄糖酯; Succinate: 琥珀酸; S7P: 景天庚酮糖-7-磷酸; Pyruvate: 丙酮酸; Asp: 天冬氨酸; G6P: 葡萄糖-6-磷酸; R5P: 核糖-5-磷酸; FBP: 果糖-1,6-二磷酸; Lactate: 乳酸; E4P: 赤藓糖-4-磷酸; Glu: 谷氨酸; F6P: 果糖-6-磷酸; Gln: 谷氨酰胺; α-KG: α-酮戊二酸; Glucose: 葡萄糖; GAP: 3-磷酸甘油醛; PEP: 磷酸烯醇式丙酮酸; DHAP: 磷酸二羟丙酮; 3PG: 3-磷酸甘油酸。

Figure 1. Hierarchical cluster analysis-heat map of sugar catabolites in leaves of naked oat under different treatments of saline-alkali stress and NaHS spraying

CK: no salt-alkali stress and no NaHS; SA: salt-alkali stress and no NaHS; SA+NaHS: NaHS spraying under salt-alkali stress; NaHS: NaHS spraying under no salt-alkali stress. Asn: asparagine; 6PG: 6-phosphogluconolactone; S7P: sedoheptulose-7-phosphate; Asp: asparate; G6P: glucose-6-phosphate; R5P: ribose-5-phosphate; FBP: fructose-1,6-diphosphate; E4P: erythrose-4-phosphate; Glu: glutamate; F6P: fructose-6-phosphate; Gln: glutamine; α-KG: α-ketoglutaric acid; GAP: 3-phosphate glyceraldehyde; PEP: enolphosphopyruvate; DHAP: phosphate dihydroxyacetone; 3PG: 3-phosphoglyceric acid.

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图 2 不同外源硫化氢喷施和盐碱胁迫处理下裸燕麦叶片糖分解代谢物PLS-DA得分图(a)及200次模型的置换检验(b)

CK: 无盐碱胁迫下喷水; SA: 盐碱胁迫下喷水; SA+NaHS: 盐碱胁迫下喷NaHS; NaHS: 无盐碱胁迫下喷NaHS。CK: no salt-alkali stress and no NaHS; SA: salt-alkali stress and no NaHS; SA+NaHS: NaHS spraying under salt-alkali stress; NaHS: NaHS spraying under no salt-alkali stress.

Figure 2. PLS-DA score plot (a) and 200 permutation test of the model (b) of sugar catabolites in naked oat leaves under different treatments of saline-alkali stress and NaHS spraying

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图 3 不同外源硫化氢喷施和盐碱胁迫处理下裸燕麦叶片糖分解代谢物OPLS-DA得分图(a: CK vs. SA; b: SA vs. SA+NaHS; c: CK vs. NaHS)、S形图(d: CK vs. SA; e: SA vs. SA+NaHS; f: CK vs. NaHS)和VIP图(g: CK vs. SA; h: SA vs. SA+NaHS; i: CK vs. NaHS)

CK: 无盐碱胁迫下喷水; SA: 盐碱胁迫下喷水; SA+NaHS: 盐碱胁迫下喷NaHS; NaHS: 无盐碱胁迫下喷NaHS。VIP: 变量投射重要度; 各变量名称解释说明见图1。CK: no salt-alkali stress and no NaHS; SA: salt-alkali stress and no NaHS; SA+NaHS: NaHS spraying under salt-alkali stress; NaHS: NaHS spraying under no salt-alkali stress. VIP: variable importance for the projection. The explainaiton of each variable is shown in the Figure 1.

Figure 3. OPLS-DA score plots (a: CK vs. SA; b: SA vs. SA+NaHS; c: CK vs. NaHS), S-plots (d: CK vs. SA; e: SA vs. SA+NaHS; f: CK vs. NaHS) and VIP plots (g: CK vs. SA; h: SA vs. SA+NaHS; i: CK vs. NaHS) of sugar catabolites in naked oat leaves under different treatments of saline-alkali stress and NaHS spraying

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图 4 不同外源硫化氢喷施和盐碱胁迫处理下裸燕麦叶片酚酸PLS-DA得分图(a)及200次模型的置换检验(b)

Figure 4. PLS-DA score plot (a) and 200 permutation test of the model (b) of phenolic acids in naked oat leaves under different treatments of saline-alkali stress and NaHS spraying

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图 5 不同外源硫化氢喷施和盐碱胁迫处理下裸燕麦叶片酚酸OPLS-DA得分图(a: CK vs. SA; b: SA vs. SA+NaHSl; c: CK vs. NaHS), S形图(d: CK vs. SA; e: SA vs. SA+NaHS; f: CK vs. NaHS)和VIP图(g: CK vs. SA; h: SA vs. SA+NaHS; i: CK vs. NaHS)

CK: 无盐碱胁迫下喷水; SA: 盐碱胁迫下喷水; SA+NaHS: 盐碱胁迫下喷NaHS; NaHS: 无盐碱胁迫下喷NaHS。VIP: 变量投射重要度; GA: 没食子酸; Phe: 苯丙氨酸; DBA:原儿茶酸; PCH: 原儿茶醛; CTC: 儿茶素; VA: 香草酸; CA: 咖啡酸; SGA:丁香酸; Epi: 表儿茶素; p-HCA: 对香豆酸; 4-HBZA: 4-羟基苯甲酸; VNL: 香草醛; SGH: 丁香醛; 4-HDCA: 芥子酸; tFA: 反式阿魏酸; SA: 水杨酸; BZA: 苯甲酸; HCA: 氢化肉桂酸; tCA: 反式肉桂酸。CK: no salt-alkali stress and no NaHS; SA: salt-alkali stress and no NaHS; SA+NaHS: NaHS spraying under salt-alkali stress; NaHS: NaHS spraying under no salt-alkali stress. VIP: variable importance for the projection; GA: Gallic acid; Phe: Phenylalanine; DBA: 3,4-Dihydroxybenzoic acid; PCH: Protocatechualdehyde; CTC: Catechin; VA: Vanillic acid; CA: Caffeic acid; SGA: Syringic acid; Epi: Epicatechin; p-HCA: p-Hydroxycinnamic acid; 4-HBZA: 4-Hydroxybenzoic acid; VNL: Vanillin; SGH: Syringaldehyde; 4-HDCA: 4-Hydroxy-3,5-dimethoxycinnamic acid; tFA: Trans-Ferulic acid; SA: Salicylic acid; BZA: Benzoic acid; HCA: Hydrocinnamic acid; tCA: Trans-Cinnamic acid.

Figure 5. PLS-DA score plots (a: CK vs. SA; b: SA vs. SA+NaHS; c: CK vs. NaHS), S-plots (d: CK vs. SA; e: SA vs. SA+NaHS; f: CK vs. NaHS) and VIP plots (g: CK vs. SA; h: SA vs. SA+NaHS; i: CK vs. NaHS) of phenolic acids in naked oat leaves under different treatments of saline-alkali stress and NaHS spraying

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图 6 OPLS-DA分析得出的外源硫化氢调控盐碱胁迫下裸燕麦叶片糖和酚酸代谢途径网络变化图

Figure 6. Proposed metabolic pathway network changes of sugars and phenolic acids in naked oat leaves under saline-alkali stress regulated by exogenous H₂S obtained from OPLS-DA analysis

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表 1 外源硫化氢对盐碱胁迫下裸燕麦叶片糖代谢物含量的影响

Table 1. Effects of exogenous H₂S on the contents of sugar metabolites in leaves of naked oat under saline-alkali stress

µmol∙g⁻¹
代谢路径 Metabolic pathway	代谢物 Metabolite	处理 Treatment
代谢路径 Metabolic pathway	代谢物 Metabolite	CK	SA	SA+NaHS	NaHS
糖酵解 Glycolysis	葡萄糖 Glucose	26 014.73±5023.43a	16 239.66±1357.08b	22 714.71±1929.09a	25 710.15±2980.95a
	果糖 Fructose	54 486.51±8760.76a	57 984.30±8750.24a	68 690.81±8393.35a	69 090.74±17 291.12a
	葡萄糖-6-磷酸 Glucose-6-phosphate	298.49±61.90a	145.73±15.64b	140.35±24.96b	162.36±38.80b
	果糖-6-磷酸 Fructose-6-phosphate	65.57±8.75a	22.55±3.00c	36.69±1.84b	70.18±9.96a
	果糖-1,6-二磷酸 Fructose-1,6-diphosphate	37.83±7.59a	25.02±4.23b	28.37±2.94b	30.82±6.73ab
	3-磷酸甘油醛 3-Phosphate glyceraldehyde	22.27±3.39a	16.50±2.33b	19.17±2.80ab	19.96±3.75ab
	磷酸二羟丙酮 Phosphate dihydroxyacetone	33.01±5.64a	25.81±3.32a	28.54±0.81a	30.38±8.13a
	3-磷酸甘油酸 3-Phosphoglyceric acid	49.56±5.38a	34.95±3.03c	41.99±1.01b	45.83±3.06ab
	磷酸烯醇式丙酮酸 Enolphosphopyruvate	32.86±6.05a	24.55±3.77b	28.26±3.62ab	28.98±5.20ab
	丙酮酸 Pyruvate	3186.10±291.05a	1467.18±255.22c	1518.68±221.93c	2417.37±375.22b
	乳酸 Lactate	71.27±7.54a	27.33±1.68d	37.62±8.58c	48.65±9.30b
戊糖磷酸途径 Pentose phosphate pathway (PPP)	6-磷酸葡萄糖酯 6-Phosphogluconolactone	45.67±7.56bc	41.20±6.81c	52.62±6.27ab	58.05±4.69a
	核糖-5-磷酸 Ribose-5-phosphate	60.80±14.18a	39.61±2.77b	42.90±7.03ab	49.72±9.33ab
	赤藓糖-4-磷酸 Erythrose-4-phosphate	304.45±19.96a	181.31±42.78b	210.11±20.75b	220.39±14.22b
	景天庚酮糖-7-磷酸 Sedoheptulose-7-phosphate	46.39±5.87a	34.93±4.28b	47.89±5.29a	37.21±2.07b
三羧酸循环及相关氨基酸 Tricarboxylic acid cycle and related amino acid (TCA)	柠檬酸 Citrate	1650.71±245.77b	2218.46±287.21a	2317.69±288.41a	2467.26±282.27a
	α-酮戊二酸 α-Ketoglutaric acid	63.48±2.78a	24.25±2.95d	54.14±2.01b	48.74±4.82c
	琥珀酸 Succinate	875.27±30.09a	823.95±56.74a	939.18±144.02a	1013.98±94.26a
	延胡索酸 Fumarate	445.85±104.85a	368.98±36.48a	505.78±121.97a	524.09±94.66a
	苹果酸 Malic acid	3544.86±924.13a	2933.51±351.98a	3924.18±942.21a	4039.05±620.36a
	谷氨酸 Glutamate	276.07±10.66a	130.63±11.95b	144.73±38.71b	174.53±24.16b
	谷氨酰胺 Glutamine	1400.46±244.50ab	592.36±129.80c	1086.11±64.60b	1850.98±531.38a
	天冬氨酸 Asparate	3802.91±319.93a	3419.40±319.93a	4124.16±319.93a	3874.08±319.93a
	天冬酰胺 Asparagine	31.63±1.21a	35.69±1.04a	35.25±1.43a	39.46±2.60a
CK: 非盐碱胁迫下喷水; SA: 盐碱胁迫下喷水; SA+NaHS: 盐碱胁迫下喷NaHS; NaHS: 非盐碱胁迫下喷NaHS。表中数据为平均值±标准差, 同行不同字母表示处理间在P<0.05水平差异显著。CK: no saline-alkali stress and no NaHS; SA: saline-alkali stress and no NaHS; SA+NaHS: NaHS spraying under saline-alkali stress; NaHS: NaHS spraying under no saline-alkali stress. Data are means±S.D. Different letters in the same row indicate significant differences among treatments at P<0.05 level.

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表 2 外源硫化氢对盐碱胁迫下裸燕麦叶片还原型谷胱甘肽(GSH)、氧化型谷胱甘肽(GSSG)、还原型辅酶Ⅱ(NADPH)、氧化型辅酶Ⅱ(NADP)含量及比值和腺苷三磷酸(ATP)含量的影响

Table 2. Effects of exogenous H₂S on the contents and ratios of reduced glutathione (GSH), oxidized glutathione (GSSG), reduced coenzyme Ⅱ (NADPH), oxidized coenzyme Ⅱ (NADP) and adenosine triphosphate (ATP) content in naked oat leaves under saline-alkali stress

处理 Treatment	GSH (µmol·g⁻¹)	GSSG (µmol·g⁻¹)	GSH/GSSG	NADPH (µmol·g⁻¹)	NADP (µmol·g⁻¹)	NADPH/NADP	ATP (µmol·g⁻¹)
CK	27.56±5.28a	103.80±15.36a	0.268±0.048b	26.27±4.64a	14.03±2.75a	1.878±0.039a	12.22±3.05a
SA	19.95±3.01b	53.81±9.86b	0.378±0.073a	19.47±2.81b	9.94±0.46b	1.961±0.281a	13.76±1.13a
SA+NaHS	22.92±3.54ab	58.97±5.55b	0.387±0.028a	21.96±2.51b	11.37±1.43ab	1.934±0.037a	14.89±1.34a
NaHS	23.50±3.86ab	68.27±6.27b	0.345±0.059ab	22.05±3.24b	12.08±1.97ab	1.843±0.258a	14.80±2.47a
CK: 无盐碱胁迫下喷水; SA: 盐碱胁迫下喷水; SA+NaHS: 盐碱胁迫下喷NaHS; NaHS: 无盐碱胁迫下喷NaHS。表中数据为平均值±标准差, 同列不同字母表示处理间在P<0.05水平差异显著。CK: no salt-alkali stress and no NaHS; SA: salt-alkali stress and no NaHS; SA+NaHS: NaHS spraying under salt-alkali stress; NaHS: NaHS spraying under no salt-alkali stress. Data are means±S.D. Different letters in the same column indicate significant differences among treatments at P<0.05 level.

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表 3 外源硫化氢对盐碱胁迫下裸燕麦叶片酚酸含量的影响

Table 3. Effects of exogenous H₂S on the contents of phenolic acids in leaves of naked oat under saline-alkali stress µg·g⁻¹

酚酸 Phenolic acid	处理 Treatment
酚酸 Phenolic acid	CK	SA	SA+NaHS	NaHS
没食子酸 Gallic acid	0.031±0.009b	0.043±0.015a	0.043±0.005a	0.033±0.006ab
苯丙氨酸 Phenylalanine	0.013±0.003a	0.013±0.003a	0.010±0.001a	0.013±0.006a
原儿茶酸 3,4-Dihydroxybenzoic acid	0.238±0.053b	0.278±0.060b	0.354±0.083a	0.280±0.041b
原儿茶醛 Protocatechualdehyde	1.767±0.192c	2.018±0.131b	2.282±0.427a	1.654±0.141c
儿茶素 Catechin	0.001±0.000a	0.001±0.000a	0.001±0.000a	0.001±0.000a
香草酸 Vanillic acid	10.327±0.890a	10.714±0.633a	10.883±0.901a	10.168±0.582a
咖啡酸 Caffeic acid	1.860±0.629a	1.795±0.523a	1.784±0.566a	1.882±0.502a
丁香酸 Syringic acid	4.475±0.286b	4.801±0.266a	4.643±0.235ab	4.453±0.325b
表儿茶素 Epicatechin	0.001±0.001a	0.001±0.000a	0.001±0.001a	0.001±0.001a
对香豆酸 p-Hydroxycinnamic acid	80.049±8.481b	85.333±3.229ab	86.691±7.315a	80.550±4.960b
4-羟基苯甲酸 4-Hydroxybenzoic acid	17.332±1.125ab	17.719±1.052a	18.230±0.727a	16.709±0.866b
香草醛 Vanillin	19.411±1.214b	19.943±1.382ab	20.686±1.004a	18.757±1.020b
丁香醛 Syringaldehyde	3.557±0.743a	2.947±0.510ab	2.700±1.176b	3.072±0.516ab
芥子酸 4-Hydroxy-3,5-dimethoxycinnamic acid	12.034±1.507a	11.444±1.071a	10.066±1.977a	11.759±2.360a
反式阿魏酸 Trans-Ferulic acid	162.599±11.015b	173.123±5.836a	177.174±14.843a	163.635±9.689b
水杨酸 Salicylic acid	0.579±0.115a	0.615±0.090a	0.520±0.075a	0.534±0.099a
苯甲酸 Benzoic acid	12.399±1.117bc	14.236±1.479a	13.401±1.321ab	11.900±1.361c
氢化肉桂酸 Hydrocinnamic acid	0.008±0.005a	0.005±0.002ab	0.005±0.002ab	0.004±0.001b
反式肉桂酸 Trans-Cinnamic acid	0.201±0.065a	0.145±0.018b	0.168±0.022ab	0.135±0.021b
酚酸总量 Total phenolic acid content	326.882±20.816b	345.170±9.309a	349.951±20.613a	325.538±13.128b
CK: 无盐碱胁迫下喷水; SA: 盐碱胁迫下喷水; SA+NaHS: 盐碱胁迫下喷施NaHS; NaHS: 无盐碱胁迫下喷施NaHS。表中数据为平均值±标准差, 同行不同字母表示处理间在P<0.05水平差异显著。CK: no salt-alkali stress and no NaHS; SA: salt-alkali stress and no NaHS; SA+NaHS: NaHS spraying under salt-alkali stress; NaHS: NaHS spraying under no salt-alkali stress. Data are means±S.D. Different letters in the same row indicate significant differences among treatments at P<0.05 level.

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表 4 外源硫化氢对盐碱胁迫下裸燕麦产量性状的影响(平均值±标准差)

Table 4. Effects of exogenous H₂S on the yield traits of naked oat under saline-alkaline stress

处理 Treatment	穗数量 Number of spike per plant	穗铃数量 Number of bolls per spike	穗粒数量 Number of grains per spike	千粒重量 1000-grain weigh (g)	单株籽粒产量 Grain yield per plant (g)	单株生物学产量 Biological yield per plant (g)
CK	6.33±0.49a	14.11±4.65a	17.73±4.37a	15.62±0.76a	1.79±0.40a	23.41±3.96a
SA	5.86±1.41a	13.01±0.88a	11.05±3.03b	15.22±2.16a	1.01±0.19b	21.08±2.60a
SA+NaHS	5.98±1.08a	13.96±1.74a	17.07±3.84a	14.32±0.52a	1.54±0.38a	23.10±6.25a
NaHS	6.44±1.70a	16.84±2.48a	22.58±3.63a	13.76±1.15a	1.91±0.18a	24.14±2.86a
CK: 无盐碱胁迫下喷水; SA: 盐碱胁迫下喷水; SA+NaHS: 盐碱胁迫下喷NaHS; NaHS: 无盐碱胁迫下喷NaHS。表中数据为平均值±标准差, 同列不同字母表示处理间在P<0.05水平差异显著。CK: no salt-alkali stress and no NaHS; SA: salt-alkali stress and no NaHS; SA+NaHS: NaHS spraying under salt-alkali stress; NaHS: NaHS spraying under no salt-alkali stress. Data are means±S.D. Different letters in the same column indicate significant differences among treatments at P<0.05 level.

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