非靶向代谢组学揭示盐胁迫下两种菌根真菌介导的核桃根系激素响应研究

doi:10.16420/j.issn.0513-353x.2025-1332

摘要/Abstract

摘要：

为了解菌根真菌对盐胁迫下核桃幼苗根系生理机制的响应，选取摩西斗管囊霉（Funneliformis mosseae）和印度梨形孢（Piriformospora indica）为菌种，‘盐源早'核桃（Juglans regia ‘Yanyuanzao'）温室盆栽苗为材料，设置不接种、单一接种和复合接种，利用非靶向代谢组学手段，分析150 mmol · L^-1NaCl溶液在短期胁迫下两种菌根真菌及不同接种方式对核桃幼苗根系植物内源激素（茉莉酸、水杨酸、生长素和脱落酸）含量的影响。结果表明，两种菌根真菌均可定殖于核桃幼苗根部，且接菌侵染率无显著差异；盐胁迫下，菌根真菌可显著影响核桃幼苗根系代谢物数量，其中接种印度梨形孢（单独接种和复合接种）引起的差异代谢物数量最多，分别为505种和527种；代谢通路富集分析结果显示，盐胁迫下摩西斗管囊霉影响氮代谢与谷胱甘肽代谢，印度梨形孢激活磷酸戊糖途径与泛酸合成通路，复合接种处理调控2-氧代羧酸代谢、糖酵解等多条通路；盐胁迫下核桃幼苗根系内源激素水平发生变化，其中JA、SA和IAA含量下降，ABA含量上升；预接种菌根真菌使盐胁迫下核桃幼苗根系的JA、SA和IAA水平提升，其中印度梨形孢对JA和IAA的提升能力更好，复合接种使ABA含量显著下降。综上，摩西斗管囊霉和印度梨形孢的单一和复合接种可在核桃幼苗根系定殖和形成共生体，促进核桃幼苗激素合成和激素信号传导等不同途径，增强核桃幼苗的耐盐性，其中复合接种处理的协同效应更明显。

关键词: 核桃, 盐胁迫, 菌根真菌, 非靶向代谢组学, 植物激素

Abstract:

To investigate the response of mycorrhizal fungi to the physiological mechanisms of walnut seedling roots under salt stress，Funneliformis mosseae and Piriformospora indica were selected as the fungal strains，and potted greenhouse seedlings of Juglans regia‘Yanyuanzao'were used as the plant material. Treatments included non-inoculation，single inoculation，and dual inoculation. Non-targeted metabolomics was employed to analyze the effects of the two mycorrhizal fungi and different inoculation methods on the content of plant endogenous hormones（jasmonic acid，salicylic acid，auxin，and abscisic acid）in the roots of walnut seedlings under short-term stress induced by 150 mmol · L^-1 NaCl solution. The results showed that both mycorrhizal fungi could colonize the roots of walnut seedlings，with no significant difference in the colonization rate between the inoculation treatments. Under salt stress，mycorrhizal fungi significantly affected the number of metabolites in the roots of walnut seedlings. Among these，inoculation with P. indica（single or dual inoculation）resulted in the highest number of differential metabolites，with 505 and 527 species，respectively. Enrichment analysis of metabolic pathways revealed that under salt stress，F. mosseae influenced nitrogen metabolism and glutathione metabolism，P. indica activated the pentose phosphate pathway and pantothenate biosynthesis pathway，and the dual inoculation treatment regulated multiple pathways，including 2-oxocarboxylic acid metabolism and glycolysis. Under salt stress，the levels of endogenous hormones in walnut seedling roots changed，with a decrease in JA，SA，and IAA content and an increase in ABA content. Pre-inoculation with mycorrhizal fungi increased the levels of JA，SA，and IAA in the roots of walnut seedlings under salt stress. Among these，P. indica showed a greater ability to enhance JA and IAA levels，while dual inoculation significantly reduced ABA content. In summary，single and dual inoculations with F. mosseae and P. indica can colonize and form symbionts in the roots of walnut seedlings，promoting hormone synthesis and hormone signaling pathways，thereby enhancing the salt tolerance of walnut seedlings. The synergistic effect was more pronounced in the dual inoculation treatment.

Key words: walnut, salt stress, mycorrhizal fungi, untargeted metabolomics, plant hormones

吴承勖, 聂瑞宁, 姬新颖, 唐佳佳, 李敖, 郑旭, 张俊佩. 非靶向代谢组学揭示盐胁迫下两种菌根真菌介导的核桃根系激素响应研究[J]. 园艺学报, 2026, 53(4): 1057-1072.

WU Chengxu, NIE Ruining, JI Xinying, TANG Jiajia, LI Ao, ZHENG Xu, ZHANG Junpei. Non-Targeted Metabolomics Reveals Hormonal Responses in Walnut Roots Mediated by Two Mycorrhizal Fungi Under Salt Stress[J]. Acta Horticulturae Sinica, 2026, 53(4): 1057-1072.

图/表 11

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代谢物Metabolites	数量Number	代谢物Metabolites	数量Number
有机酸 Organic acids	722	木脂素和香豆素 Lignans and Coumarins	40
苯及其衍生物 Benzene and substituted derivatives	310	甘油磷脂 Glycerophospholipids	29
氨基酸及其衍生物 Amino acids and derivatives	272	鞣质 Tannins	21
黄酮 Flavonoids	151	醌类 Quinones	12
核苷酸及其衍生物 Nucleotides and derivatives	117	甘油脂 Glyceride	11
酚酸类 Phenolic acids	90	脂肪酰类 Fatty acyls	8
醇、胺类 Alcohol and amines	67	甾体 Steroids	7
生物碱 Alkaloids	64	鞘脂类 Sphingolipids	4
杂环化合物 Heterocyclic compounds	56	色胺、胆碱、色素 Tryptamines，Cholines，Pigments	2
脂质 Lipids	50	其他类 Others	339
萜类 Terpenoids	41

代谢物Metabolites	数量Number	代谢物Metabolites	数量Number
有机酸 Organic acids	722	木脂素和香豆素 Lignans and Coumarins	40
苯及其衍生物 Benzene and substituted derivatives	310	甘油磷脂 Glycerophospholipids	29
氨基酸及其衍生物 Amino acids and derivatives	272	鞣质 Tannins	21
黄酮 Flavonoids	151	醌类 Quinones	12
核苷酸及其衍生物 Nucleotides and derivatives	117	甘油脂 Glyceride	11
酚酸类 Phenolic acids	90	脂肪酰类 Fatty acyls	8
醇、胺类 Alcohol and amines	67	甾体 Steroids	7
生物碱 Alkaloids	64	鞘脂类 Sphingolipids	4
杂环化合物 Heterocyclic compounds	56	色胺、胆碱、色素 Tryptamines，Cholines，Pigments	2
脂质 Lipids	50	其他类 Others	339
萜类 Terpenoids	41

处理 Treatment	激素相关化合物 Hormone-related compounds	P	VIP	倍数变化 Fold change	类型 Type
C-150 vs. F-150	冠状酸 Coriolic acid	0.001	2.08	2.78	上调Up
	磷脂酸 Phosphatidic acid	0.001	2.31	10.44	上调Up
	塞达明 Sedamine	0.001	2.06	8.15	上调Up
	2-羟基肉桂酸 2-Hydroxycinnamic acid	0.005	2.14	0.18	下调Down
	1,2-二乙酰氧基-4,7,8-三羟基-3-（4-羟基苯基）二苯并呋喃 1,2-Diacetoxy-4,7,8-trihydroxy-3-（4-hydroxyphenyl）dibenzofuran	0.003	2.19	0.08	下调Down
C-150 vs.P-150	双没食子酸 Digallic acid	0.011	1.99	6.49	上调Up
	（1E）-1-（4-羟基-3-甲氧基苯基）dec-1-烯-3-酮（E）-1-（4-hydroxy-3-methoxyphenyl）dec-1-en-3-one	0.004	2.01	12.55	上调Up
	橡精 Datiscetin	0.013	1.98	6.54	上调Up
	4-氧代戊酸 4-Oxopentanoate	0.001	2.03	3.10	上调Up
	3-巯基丙酮酸 3-Mercaptopyruvic acid	0.001	2.06	4.88	上调Up
	1-（2,4-二羟苯基）-3-{3-[（3,3-二甲基环氧乙烷-2-基）甲基]- 4-羟基苯基}丙烷-1-酮 1-（2,4-dihydroxyphenyl）-3-{3-[（3，3-dimethyloxiran-2-yl）methyl]-4-hydroxyphenyl}propan-1-one	0.001	2.07	0.46	下调Down
	3-{[3,4-二羟基-5-（羟甲基）氧杂环戊烷-2-基]氧}-5,7-二羟基-2-（3-羟基苯基）-4H-苯并吡喃-4-酮 3-{[3，4-dihydroxy-5-（hydroxymethyl）oxolan-2-yl]oxy}-5,7-dihydroxy-2-（3-hydroxyphenyl）-4H-chromen-4-one	0.001	2.05	0.14	下调Down
	3-（11,12-二羟基-15,19-三十二碳二烯基）-5-甲基-2（5H）-呋喃酮 3-（11，12-Dihydroxy-15，19-dotriacontadienyl）-5-methyl-2（5H）-furanone	0.003	1.97	0.22	下调Down
	3-（咪唑-5-基）乳酸 3-（Imidazol-5-yl）lactate	0.001	2.06	0.39	下调Down
	表没食子儿茶素没食子酸酯 Epigallocatechin gallate	0.003	2.02	0.16	下调Down
	脱氧鸟苷5'-1磷酸（dGMP）Deoxyguanosine 5'-monophosphate（dGMP）	0.001	2.02	0.18	下调Down
C-150 vs. FP-150	磷脂酸 Phosphatidic acid	0.001	1.89	6.62	上调Up
	（1E）-1-（4-羟基-3-甲氧基苯基）dec-1-烯-3-酮（E）-1-（4-hydroxy-3-methoxyphenyl）dec-1-en-3-one	0.001	1.94	29.82	上调Up
	4-氧代戊酸 4-Oxopentanoate	0.001	1.89	2.86	上调Up
	γ-谷氨酰谷氨酸 γ-Glutamylglutamate	0.001	1.86	7.23	上调Up
	双-γ-谷氨酰半胱氨酸 Bis-γ-L-glutamyl-L-cystine	0.001	1.89	2.44	上调Up
	芥子酸酯 Sinapoyltartronic acid	0.001	1.88	0.25	下调Down
	1-（2,4-二羟苯基）-3-{3-[（3,3-二甲基环氧乙烷-2-基）甲基]-4-羟基苯基}丙烷-1-酮 1-（2,4-dihydroxyphenyl）-3-{3-[（3,3- dimethyloxiran-2-yl）methyl]-4-hydroxyphenyl}propan-1-one	0.001	1.89	0.38	下调Down
	3-{[3,4-二羟基-5-（羟甲基）氧杂环戊烷-2-基]氧}-5,7-二羟基-2-（3-羟基苯基）-4H-苯并吡喃-4-酮 3-{[3,4-dihydroxy-5-（hydroxymethyl）oxolan-2-yl]oxy}-5,7-dihydroxy-2-（3-hydroxyphenyl）-4H-chromen-4-one	0.001	1.89	0.38	下调Down
	2'',6''-二没食子酰基异里夫洛苯酮3-C-葡萄糖苷 2'',6''-Digalloyliriflophenone 3-C-glucoside	0.001	1.88	0.38	下调Down
	水飞蓟亭 Silicristin	0.001	1.89	0.31	下调Down

处理 Treatment	激素相关化合物 Hormone-related compounds	P	VIP	倍数变化 Fold change	类型 Type
C-150 vs. F-150	冠状酸 Coriolic acid	0.001	2.08	2.78	上调Up
	磷脂酸 Phosphatidic acid	0.001	2.31	10.44	上调Up
	塞达明 Sedamine	0.001	2.06	8.15	上调Up
	2-羟基肉桂酸 2-Hydroxycinnamic acid	0.005	2.14	0.18	下调Down
	1,2-二乙酰氧基-4,7,8-三羟基-3-（4-羟基苯基）二苯并呋喃 1,2-Diacetoxy-4,7,8-trihydroxy-3-（4-hydroxyphenyl）dibenzofuran	0.003	2.19	0.08	下调Down
C-150 vs.P-150	双没食子酸 Digallic acid	0.011	1.99	6.49	上调Up
	（1E）-1-（4-羟基-3-甲氧基苯基）dec-1-烯-3-酮（E）-1-（4-hydroxy-3-methoxyphenyl）dec-1-en-3-one	0.004	2.01	12.55	上调Up
	橡精 Datiscetin	0.013	1.98	6.54	上调Up
	4-氧代戊酸 4-Oxopentanoate	0.001	2.03	3.10	上调Up
	3-巯基丙酮酸 3-Mercaptopyruvic acid	0.001	2.06	4.88	上调Up
	1-（2,4-二羟苯基）-3-{3-[（3,3-二甲基环氧乙烷-2-基）甲基]- 4-羟基苯基}丙烷-1-酮 1-（2,4-dihydroxyphenyl）-3-{3-[（3，3-dimethyloxiran-2-yl）methyl]-4-hydroxyphenyl}propan-1-one	0.001	2.07	0.46	下调Down
	3-{[3,4-二羟基-5-（羟甲基）氧杂环戊烷-2-基]氧}-5,7-二羟基-2-（3-羟基苯基）-4H-苯并吡喃-4-酮 3-{[3，4-dihydroxy-5-（hydroxymethyl）oxolan-2-yl]oxy}-5,7-dihydroxy-2-（3-hydroxyphenyl）-4H-chromen-4-one	0.001	2.05	0.14	下调Down
	3-（11,12-二羟基-15,19-三十二碳二烯基）-5-甲基-2（5H）-呋喃酮 3-（11，12-Dihydroxy-15，19-dotriacontadienyl）-5-methyl-2（5H）-furanone	0.003	1.97	0.22	下调Down
	3-（咪唑-5-基）乳酸 3-（Imidazol-5-yl）lactate	0.001	2.06	0.39	下调Down
	表没食子儿茶素没食子酸酯 Epigallocatechin gallate	0.003	2.02	0.16	下调Down
	脱氧鸟苷5'-1磷酸（dGMP）Deoxyguanosine 5'-monophosphate（dGMP）	0.001	2.02	0.18	下调Down
C-150 vs. FP-150	磷脂酸 Phosphatidic acid	0.001	1.89	6.62	上调Up
	（1E）-1-（4-羟基-3-甲氧基苯基）dec-1-烯-3-酮（E）-1-（4-hydroxy-3-methoxyphenyl）dec-1-en-3-one	0.001	1.94	29.82	上调Up
	4-氧代戊酸 4-Oxopentanoate	0.001	1.89	2.86	上调Up
	γ-谷氨酰谷氨酸 γ-Glutamylglutamate	0.001	1.86	7.23	上调Up
	双-γ-谷氨酰半胱氨酸 Bis-γ-L-glutamyl-L-cystine	0.001	1.89	2.44	上调Up
	芥子酸酯 Sinapoyltartronic acid	0.001	1.88	0.25	下调Down
	1-（2,4-二羟苯基）-3-{3-[（3,3-二甲基环氧乙烷-2-基）甲基]-4-羟基苯基}丙烷-1-酮 1-（2,4-dihydroxyphenyl）-3-{3-[（3,3- dimethyloxiran-2-yl）methyl]-4-hydroxyphenyl}propan-1-one	0.001	1.89	0.38	下调Down
	3-{[3,4-二羟基-5-（羟甲基）氧杂环戊烷-2-基]氧}-5,7-二羟基-2-（3-羟基苯基）-4H-苯并吡喃-4-酮 3-{[3,4-dihydroxy-5-（hydroxymethyl）oxolan-2-yl]oxy}-5,7-dihydroxy-2-（3-hydroxyphenyl）-4H-chromen-4-one	0.001	1.89	0.38	下调Down
	2'',6''-二没食子酰基异里夫洛苯酮3-C-葡萄糖苷 2'',6''-Digalloyliriflophenone 3-C-glucoside	0.001	1.88	0.38	下调Down
	水飞蓟亭 Silicristin	0.001	1.89	0.31	下调Down

处理 Treatment	激素相关化合物 Hormone-related compounds	关键代谢通路 Key metabolic pathways	调控的内源激素 Regulated endogenous hormones
C-150 vs. CK	赤藓糖 Erythrose ↑ L-赖氨酸-L-色氨酸 Lys-Trp ↓	苯丙氨酸代谢 Phenylalanine metabolism 亚油酸代谢 Linoleic acid metabolism 泛醌和其他萜类醌生物合成 Biosynthesis of ubiquinone and other terpenoid quinones	IAA ↓ ABA ↑
F-150 vs. C-150	磷脂酸Phosphatidic acid ↑ 莽草酸 Shikimic acid ↓	氮代谢 Nitrogen metabolism 谷胱甘肽代谢 Glutathione metabolism C5-支链二元酸代谢 C5-Branched dicarboxylic acid metabolism	SA ↑ IAA ↑ ABA ↓
P-150 vs. C-150	3-巯基丙酮酸 3-mercaptopyruvic acid ↑ 4-氧代戊酸 4-Oxopentanoate ↑ 双没食子酸 Digallic acid ↑ 3-（咪唑-5-基）乳酸 3-（Imidazol-5-yl）lactate ↓	磷酸戊糖途径 Pentose phosphate pathway 泛酸和辅酶A生物合成 Pantothenic acid and coenzyme A biosynthesis 酪氨酸代谢 Tyrosine metabolism	JA ↑ SA ↑ IAA ↑
FP-150 vs. C-150	麦芽三糖 Maltotriose ↑ 双-γ-L-谷氨酰-L-半胱氨酸 Bis（γ-L-glutamyl）-L-cysteine ↑ 氧化的γ-L-谷氨酰-L-半胱氨酸 γ-L-Glutamyl-L-cysteine disulfide ↑ 4-氧代戊酸 4-Oxopentanoate ↑ 苯丙氨酸-萘酚羟基 Hydroxynaphthol phenylalanine ↓	泛酸和辅酶A生物合成 Antothenate and Coenzyme A biosynthesis 2-氧代羧酸代谢 2-Oxocarboxylic acid metabolism 糖酵解/糖异生 Glycolysis/Gluconeogenesis 硫代葡萄糖苷生物合成 Biosynthesis of glucosinolates 萜类化合物骨架生物合成 Biosynthesis of terpenoid backbone	JA ↑ SA ↑ IAA ↑ ABA ↓