苹果果实营养成分松脂醇的鉴定与生态影响因子分析

doi:10.3864/j.issn.0578-1752.2026.18.013

摘要/Abstract

摘要：

【目的】苹果果实品质的形成与生态因素密切相关，明确关键品质形成与生态因子的关系，为功能果品的特色栽培提供参考。【方法】以新疆伊犁、喀什，陕西洛川和山东烟台4个代表地区的主栽品种‘嘎啦’和‘富士’苹果果实为试材，采用广泛靶向代谢组学技术进行苹果果实的代谢物检测，取各样本中鉴定结果可信度最高的代谢物，结合多元统计分析筛选伊犁、喀什两地相较于洛川和烟台均显著上调的差异代谢物，再通过叠加分析获得共同高含量营养成分。进一步针对共同高含量成分建立液相色谱串联质谱(LC-MS/MS)检测方法，对代谢组检测结果进行定量验证。开展干旱、温差、紫外线等生态因子模拟处理苹果果实，分析目标成分的积累及其合成关键基因的响应情况，明确促进目标成分积累的主要生态因子。【结果】基于代谢组学分析，从4个产地的‘嘎啦’和‘富士’苹果果实中分别检测到1 653和2 052种代谢物。伊犁和喀什地区的‘嘎啦’‘富士’果实中分别共同含有18种和19种代谢物含量显著高于洛川和烟台地区2个品种的苹果样本。其中，木脂素类代谢物松脂醇（pinoresinol）与表松脂醇（epipinoresinol）在伊犁和喀什地区两种果实中的含量均显著高于其他地区，为共同高含量营养成分。进一步建立了基于LC-MS/MS技术的松脂醇测定方法，检测显示伊犁与喀什地区的‘嘎啦’苹果松脂醇含量分别达110.2和80.4 mg·kg^-1，‘富士’果实的松脂醇含量分别为73.4和64.5 mg·kg^-1，均为烟台、洛川果实中含量的2倍以上。此外，检测显示松脂醇含量随果实发育逐渐累积，于成熟期达到最高；干旱、高昼夜温差和紫外线可促进松脂醇的积累，而高盐胁迫无显著促进作用。在苹果基因组中，共筛选获得了35个松脂醇合成关键酶指导蛋白（DIR）基因，其中，MdDIR-a1与MdDIR-a2在苹果果实中表达，且均受干旱胁迫显著正调控。【结论】干旱、高昼夜温差和强紫外的生态环境有利于苹果果实营养成分松脂醇含量的积累，其中干旱胁迫是促进其积累的主要生态因子。

关键词: 苹果, 果实, 营养成分, 松脂醇, 代谢组, 生态因子

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

岳静雯, 郭鑫鑫, 周佳, 史天乐, 杨雨璋, 张军科, 李兴亮. 苹果果实营养成分松脂醇的鉴定与生态影响因子分析[J]. 中国农业科学, 2026, 59(18): 4121-4134.

YUE JingWen, GUO XinXin, ZHOU Jia, SHI TianLe, YANG YuZhang, ZHANG JunKe, LI XingLiang. Identification and Ecological Impact Factor Analysis of Nutritional Component Pinoresinol in Apple Fruits[J]. Scientia Agricultura Sinica, 2026, 59(18): 4121-4134.

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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

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	木脂素 Lignans	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	-

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	-