Scientia Agricultura Sinica ›› 2026, Vol. 59 ›› Issue (18): 4121-4134.doi: 10.3864/j.issn.0578-1752.2026.18.013

• HORTICULTURE • Previous Articles     Next Articles

Identification and Ecological Impact Factor Analysis of Nutritional Component Pinoresinol in Apple Fruits

YUE JingWen(), GUO XinXin, ZHOU Jia, SHI TianLe, YANG YuZhang, ZHANG JunKe, LI XingLiang()   

  1. Institute of Forestry and Pomology, Beijing Academy of Agriculture and Forestry Sciences/Key Laboratory of Biology and Genetic Improvement of Horticultural Crops (North China), Ministry of Agriculture and Rural Affairs, Beijing 100093
  • Received:2026-01-27 Accepted:2026-06-08 Online:2026-09-16 Published:2026-09-20
  • Contact: LI XingLiang

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Abstract:

【Objective】The formation of apple fruit quality is closely associated with ecological factors. This study aimed to clarify the relationship between key quality attributes and ecological factors, so as to provide a reference for the specialty cultivation of functional fruits.【Method】Apple fruits of the main cultivars Gala and Fuji were collected from four representative regions, namely Ili and Kashgar in Xinjiang, Luochuan in Shaanxi, and Yantai in Shandong. Widely targeted metabolomics was employed to detect metabolites in apple fruits. Metabolites with the highest identification confidence from each sample were selected, and multivariate statistical analysis was used to screen for differential metabolites that were significantly up-regulated in both Ili and Kashgar compared with Luochuan and Yantai. Overlay analysis was then performed to obtain the common high-content nutritional components. A liquid chromatography-tandem mass spectrometry (LC-MS/MS) method was further established for the identified common high-content components to quantitatively validate the metabolomics results. Simulation treatments of ecological factors, including drought, diurnal temperature difference, and UV-B, were conducted on apple fruits to analyze the accumulation of target components and the expression responses of key genes involved in their biosynthesis, thereby identifying the predominant ecological factor promoting the accumulation of target components.【Result】Based on metabolomics analysis, 1 653 and 2 052 metabolites were detected in Gala and Fuji apple fruits from four regions, respectively. A total of 18 and 19 metabolites were significantly more abundant in Gala and Fuji fruits from the Ili and Kashgar regions than in samples from the Luochuan and Yantai regions, respectively. Among these, the lignan metabolites pinoresinol and epipinoresinol were significantly more abundant in both apple cultivars from the Ili and Kashgar regions than that from the other two regions, which were identified as the common high-content nutritional components. An LC-MS/MS method for pinoresinol determination was further established. Quantitative analysis showed that pinoresinol content in Gala apples from Ili and Kashgar reached 110.2 and 80.4 mg·kg-1, respectively, and those in Fuji apples reached 73.4 and 64.5 mg·kg-1, respectively, all of which were more than twice the content in apples from Yantai and Luochuan. Furthermore, pinoresinol content gradually increased during fruit development and reached the highest level at the mature stage. Drought, large diurnal temperature difference, and UV-B promoted pinoresinol accumulation, whereas high salt stress had no significant promoting effect. A total of 35 dirigent protein (DIR) genes involved in pinoresinol biosynthesis were identified in the apple genome. Among them, MdDIR-a1 and MdDIR-a2 were expressed in apple fruits, and both were significantly up-regulated by drought stress.【Conclusion】Drought, large diurnal temperature difference, and strong UV-B constituted an ecological environment favorable for the accumulation of the nutritional component pinoresinol in apple fruits, among which drought stress was the predominant ecological factor promoting its accumulation.

Key words: apple, fruit, nutritional components, pinoresinol, metabolomics, ecological factors

Table 1

Primers used for Real-time qPCR"

序号No. 基因Gene 引物序列Primer sequence (5′-3′) 基因号Accession No.
1 MdDIR-a1 F: ATCACCACCATGACCGCCACC
R: CCCATCTAAGCTGGATGTGACAGA		 MD02G1118100
2 MdDIR-a2 F: CCATGACCGTCGCCACCACCATAAC
R: TCCATCTAAGCTGGATGAGACGGT		 MD02G1118400
3 MdDIR-a3 F: GACTGATGAGGACGGTCACGTCG
R: TCATCCCAGAACAGTCTGAGACTGA		 MD03G1292000
4 MdDIR-a4 F: CTAATGCAACATCTGCTGCAAC
R: CTGAGTAGGTGTCTTTCCTGTCG		 MD06G1096600
5 MdDIR-a5 F: ATACATTCAACAACCACCCAACAAC
R: TGTCCGGTTGCGGTTGTTTTCCG		 MD11G1306400
6 β-Actin F: GAATTAAGTGTTGTTCGATATCGCTC
R: GTAGTTGCTTTTGATCCCCTGTTCTA		 XM008356922

Fig. 1

Overlaid total ion chromatogram (TIC) from mass spectrometry and correlation analysis among samples A: Positive ion mode (P); B: Negative ion mode (N); C: Correlation analysis among samples; D: Principal component analysis. YT: Yantai; YL:Yili; KS:Kashi; LC: Luochuan; G: Gala apple. The same as below"

Fig. 2

Analysis of common up-regulated metabolites in Gala apple fruits A: Classification of 1 653 metabolites, red columns represent the primary metabolites, and blue columns represent the secondary metabolites; B: Venn diagram of up-regulated metabolites in YL-G and KS-G samples"

Table 2

High-content differential metabolites in Gala apple"

序号No. 编号ID 物质Compound 类别Category CAS
1 mws0097 松脂醇 (+)-Pinoresinol 木脂素 Lignans 487-36-5
2 MWSHC20189 表松脂醇 Epipinoresinol 24404-50-0
3 mws1587 L-正亮氨酸 L-Norleucine 氨基酸及其衍生物
Amino acids and derivatives		 327-57-1
4 MWS04555 别异亮氨酸 D-Allo-Isoleucine 1509-34-8
5 mws0227 亮氨酸 L-Leucine 328-39-2
6 mws0258 L-异亮氨酸 L-Isoleucine 73-32-5
7 pme0274 6-氨基己酸 6-Aminohexanote 有机酸 Organic acids 60-32-2
8 Walbn08556 二甲基磷脂-OA DMAP-OA 脂质 Lipids -
9 WaYn011395 二甲基磷脂-甘油-肉蔻酸酯 DMAEP-GMo -
10 Walbn08698 二甲磷-OA (异构体) DMAEP-OA -
11 Lcfn066903 3-羟基辛二酸3-Hydroxyoctanedioicacid 73141-47-6
12 Zken00392 油酰溶血磷脂酰胆碱 18:1-9z LPC -
13 pmp001281 溶血磷脂酰胆碱 LysoPC 18:1 -
14 Safp021800 溶血卵磷脂 (18:1) LPC (18:1/0:0) 3542-29-8
15 Lmhp10190 溶血磷脂酰胆碱 LysoPC 18:1 2n isomer -
16 Walbn011704 1位甘油-二甲磷-肉蔻酸酯 sn-1-DMAEP-Mo -
17 pmp001198 6-脱氧荞麦碱 6-Deoxyfagomine 生物碱 Alkaloids 197449-09-5
18 Wabn006607 双羟甲氧色满二聚体 3, 3′-Bi-HMC 其他类 Others -

Fig. 3

Screening of common up-content metabolites in fruits of two apple varieties A: Classification of Fuji apple fruit metabolites; B: Venn diagram of up-regulated metabolites in YL-F and KS-F samples; C: Screening and content comparison of up-regulated metabolites common to both varieties; D: The biosynthetic pathway of lignan. The “F” in figure A-C stand for Fuji apple"

Table 3

High-content differential metabolites in Fuji apple"

序号No. 编号ID 物质Compound 类别Category CAS
1 mws0097 松脂醇 (+)-Pinoresinol 木脂素
Lignans		 487-36-5
2 MWSHC20189 表松脂醇 Epipinoresinol 24404-50-0
3 MWStz451 Clemaphenol A 362606-60-8
4 Yxxp005044 蛇菰宁; 蛇菰脂醛素 Balanophonin 118916-57-7
5 pme1086 谷胱甘肽还原型 Glutathione 氨基酸及其衍生物
Amino acids and derivatives		 70-18-8
6 Sasn003250 Valinopine -
7 Zbqn003748 N-丙二酰亮氨酸 N-malonylleucine -
8 WaXYp00973 谷胱甘肽 Glutathione -
9 pmb2826 2-甲基苹果酸(S)-2-Methylmalate 有机酸
Organic acids		 -
10 Zmyn000247 2-羟基戊二酸 2-Hydroxyglutaric acid 13095-48-2
11 Lmbn000216 3-甲基苹果酸 D-erythro-3-methylmalate 152204-30-3
12 mws0091 槲皮素-3-O-葡萄糖苷(异槲皮苷) Isoquercitrin 黄酮
Flavonoids		 482-35-9
13 PD0485458 6-羟基木犀草素-6-葡萄糖苷 6-Hydroxyluteolin 6-glucoside -
14 Lcfn086739 野黄岑素-7-O-葡萄糖苷 Plantaginin 26046-94-6
15 PDN020315 红景天金丝桃苷 Rhodiolatuntoside 萜类
Terpenoids		 -
16 Hmcn001884 6-羟基木犀草素-5-葡萄糖苷 6-Hydroxyluteolin 5-glucoside -
17 mws0856 槲皮素-4'-O-葡萄糖苷 (绣线菊苷) Quercetin-4'-O-glucoside (Spiraeoside) 20229-56-5
18 Safn005826 还阳参酸苷 Napiferoside -
19 Zmyn000230 2-脱氢-3-脱氧-L-阿拉伯糖2-Dehydro-3-deoxy-L-arabinonate 其他类Others -

Fig. 4

Detection of pinoresinol content and ecological factor analysis A: Establishment of the LC-MS/MS method for pinoresinol quantification; B: Determination of pinoresinol content in apple fruits from different regions; C: Changes in pinoresinol content during the development of Fuji apple fruits; D: Effects of different ecological factors on pinoresinol accumulation. Different lowercase letters indicate significant difference (P<0.05). The same as below"

Fig. 5

Effect of different ecological factors on the expression of MdDIR-a genes A: Evolutionary and conserved domain analysis of DIR proteins; B: Functional domain analysis of DIR-a proteins; C: Expression detection of MdDIR-a genes in response to different ecological factors"

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