Acta Horticulturae Sinica ›› 2026, Vol. 53 ›› Issue (4): 1057-1072.doi: 10.16420/j.issn.0513-353x.2025-1332
• Research on Arbuscular Mycorrhizal Fungi in Horticultural Plants • Previous Articles Next Articles
WU Chengxu1, NIE Ruining1, JI Xinying1, TANG Jiajia1,2, LI Ao1, ZHENG Xu1, ZHANG Junpei1,*(
)
Received:2025-12-30
Revised:2026-03-16
Online:2026-04-25
Published:2026-04-20
Contact:
ZHANG Junpei
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.
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URL: https://www.ahs.ac.cn/EN/10.16420/j.issn.0513-353x.2025-1332
Fig. 1 Mycorrhizal colonization status of walnut seedlings under different inoculation treatments(A,B)and phenotypic changes of walnut seedlings after 10 days of salt stress(C) NE:Non-inoculated treatment;F.m:Funneliformis mosseae treatment;P.i:Piriformospora indica treatment;FP:F. mosseae + P. indica treatment. Different letters indicate significant differences at the P < 0.05 level. The same below
Fig. 3 PCA score of samples CK:Control group;C-150:Non-inoculated stress group;F-150:Stress group inoculated with Funneliformis mosseae;P-150:Stress group inoculated with Piriformospora indica;FP:Stress group with combined inoculation of F. mosseae and P. indica. The same below
| 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 |
Table 1 Metabolites in walnut roots mediated by endophytic fungi under salt stres
| 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 |
Table 2 Effects of mycorrhizal fungi on the relative concentrations and fold changes of hormone-related substances in walnut seedling roots under salt stress
| 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 ↓ |
Table 3 Correlation analysis of different metabolites,key pathways,and endogenous hormones under regulated by endophytic fungi
| 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 ↓ |
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