14个蓝莓品种不同成熟期果实糖酸组分测定及代谢基因表达分析

doi:10.3864/j.issn.0578-1752.2026.13.013

摘要/Abstract

摘要：

【目的】明确我国目前生产中不同熟期且口感差异明显的14个蓝莓主栽品种在3个关键发育阶段糖酸组分含量的动态变化，揭示PEPC与蓝莓有机酸代谢之间的关系，为蓝莓的生产和风味改良提供支持。【方法】以14个蓝莓品种的青果期、粉果期和蓝果期3个关键成熟期的果实为试材，采用HPLC测定可溶性糖与有机酸组分含量，并利用qRT-PCR分析PEPC表达模式。【结果】在果实成熟过程中，不同蓝莓品种TA与TSS的积累动态存在差异，可归纳为持续下降、平稳-下降和先升后降3种酸类型；以及持续上升、平稳-上升和先降后升3种糖类型。蓝莓果实可溶性糖的主要组分是葡萄糖与果糖，有机酸的主要组分是柠檬酸、奎宁酸和苹果酸，其中，奎宁酸变异系数较高。依据糖酸组分特征的聚类分析，将14个品种划分为4个类群，其中，4个品种有机酸以柠檬酸、奎宁酸和苹果酸为主；7个品种以柠檬酸和苹果酸为主；1个品种以柠檬酸和奎宁酸为主，上述12个品种的葡萄糖和果糖比例均衡，另有2个品种的奎宁酸占绝对优势，果糖含量显著高于葡萄糖。蓝莓果实总糖含量与葡萄糖和果糖含量显著正相关；总有机酸含量与柠檬酸、奎宁酸、草酸和乳酸含量极显著正相关。不同酸类型的3种蓝莓在青果期到粉果期PEPC表达量上升，而在粉果期到蓝果期PEPC表达量下调，此时，果实中苹果酸和柠檬酸含量同样下降。【结论】在蓝莓果实发育过程中，TA、TSS含量变化模式与品种密切相关。其可溶性糖的主要组分是葡萄糖和果糖，有机酸主要组分是柠檬酸、奎宁酸和苹果酸，可分为高柠檬酸型和高奎宁酸型两类。在粉果期到蓝果期，3种不同酸类型的蓝莓果实PEPC表达量下调，与苹果酸和柠檬酸含量下降趋势一致，参与了蓝莓有机酸的代谢。

关键词: 蓝莓, 有机酸, 可溶性糖, 成熟阶段, 代谢基因

Abstract:

【Objective】To characterize the dynamic changes in sugar and organic acid composition across three key developmental stages in 14 major blueberry cultivars currently cultivated in China, which differ markedly in ripening period and flavor profile, and to elucidate the relationship between PEPC and organic acid metabolism in blueberry fruit, thereby providing theoretical support for blueberry production and flavor quality improvement.【Method】Fruits of 14 blueberry cultivars at three key ripening stages (green, pink, and blue) were used as materials. Soluble sugar and organic acid components were determined by HPLC, and the expression patterns of the PEPC gene were analyzed by qRT-PCR.【Result】During fruit ripening, the accumulation dynamics of TA and TSS varied among different blueberry cultivars and could be classified into three acid-accumulation patterns (continuous decline, stable-decline, and increase-decline) and three sugar-accumulation patterns (continuous increase, stable- increase, and decline-increase). Glucose and fructose were the major soluble sugar components in blueberry fruits, whereas citric acid, quinic acid, and malic acid were the predominant organic acids. Among them, quinic acid exhibited a relatively high coefficient of variation. Cluster analysis based on sugar and organic acid composition divided the 14 cultivars into four groups. In four cultivars, the major organic acids were citric acid, quinic acid, and malic acid; seven cultivars were dominated by citric acid and malic acid; and one cultivar was characterized primarily by citric acid and quinic acid. In these 12 cultivars, the proportions of glucose and fructose were relatively balanced. In contrast, the remaining two cultivars showed a clear predominance of quinic acid, with fructose content significantly higher than that of glucose. Total sugar content in blueberry fruits was significantly positively correlated with glucose and fructose contents, while total organic acid content was extremely significantly positively correlated with the contents of citric acid, quinic acid, oxalic acid, and lactic acid. In the three blueberry cultivars representing different acid-accumulation patterns, PEPC expression increased from the green fruit stage to the pink fruit stage, but was downregulated from the pink fruit stage to the blue fruit stage. During this latter period, the contents of malic acid and citric acid in the fruits also decreased correspondingly.【Conclusion】During blueberry fruit development, the variation patterns of TA and TSS were closely associated with cultivar characteristics. The major soluble sugar components were glucose and fructose, while the predominant organic acids were citric acid, quinic acid, and malic acid. Based on organic acid composition, the cultivars could be categorized into two types: high-citric-acid and high-quinic-acid types. In three blueberry cultivars representing different acid-accumulation patterns, PEPC expression was downregulated from the pink fruit stage to the blue fruit stage, which was consistent with the declining trends of malic acid and citric acid contents, indicating that PEPC is involved in the metabolism of organic acids in blueberry fruits.

Key words: blueberry, oganic acid, soluble sugar, maturity stage, metabolism-related gene

黄梦池, 陈利娜, 高登涛, 刘丽, 韩静熠, 段书云, 刘钊宁, 李好先, 鲁振华. 14个蓝莓品种不同成熟期果实糖酸组分测定及代谢基因表达分析[J]. 中国农业科学, 2026, 59(13): 2919-2932.

HUANG MengChi, CHEN LiNa, GAO DengTao, LIU Li, HAN JingYi, DUAN ShuYun, LIU ZhaoNing, LI HaoXian, LU ZhenHua. Analysis of Sugar and Organic Acid Components and Related Metabolic Gene Expression During Fruit Development in 14 Blueberry Cultivars[J]. Scientia Agricultura Sinica, 2026, 59(13): 2919-2932.

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参考文献 42

[1]	余莹, 杨燕林, 和建平, 王宇萍, 吴永斌, 杨正松. 不同发育阶段喷施外源激素对蓝莓果实品质的影响[J/OL]. 果树学报, 2025:1-25. DOI: 10.13925/j.cnki.gsxb.20250590.
	Yu Y, Yang Y L, He J P, Wang Y P, Wu Y B, Yang Z S. Effects of exogenous plant growth hormones at different growth stage on fruit quality of blueberry[J/OL]. Journal of Fruit Science, 2025:1-25. DOI: 10.13925/j.cnki.gsxb.20250590. (in Chinese)
[2]	Jiao X Y, Wang Y H, Lin Y, Lang Y X, Li E H, Zhang X Y, Zhang Q, Feng Y, Meng X J, Li B. Blueberry polyphenols extract as a potential prebiotic with anti-obesity effects on C57BL/6 J mice by modulating the gut microbiota[J]. The Journal of Nutritional Biochemistry, 2019, 64: 88-100.
[3]	宋江涛, 谌丹丹, 公旭晨, 商祥明, 李春龙, 蔡永喜, 岳建平, 王帅玲, 张卜芬, 谢宗周, 刘继红. 人工疏果对'爱媛28'橘橙果实糖酸含量及代谢基因表达的影响[J]. 中国农业科学, 2022, 55(23): 4688-4701. DOI:10.3864/j.issn.0578-1752.2022.23.010.
	Song J T, Chen D D, Gong X C, Shang X M, Li C L, Cai Y X, Yue J P, Wang S L, Zhang B F, Xie Z Z, Liu J H. Effects of artificial fruit thinning on sugar and acid content and expression of metabolism- related genes in fruit of Beni-Madonna tangor[J]. Scientia Agricultura Sinica, 2022, 55(23): 4688-4701. DOI:10.3864/j.issn.0578-1752.2022.23.010. (in Chinese)
[4]	Doty T E. Fructose sweetness: A new dimension[J]. Cereal Foods World, 1976, 21(2): 62-63.
[5]	王鹏飞, 薛晓芳, 穆晓鹏, 张建成, 曹琴, 杜俊杰. 不同酸度欧李果实有机酸积累特性与相关代谢酶活性分析[J]. 中国农业科学, 2013, 46(19): 4101-4109. DOI: 10.3864/j.issn.0578-1752.2013.19.016.
	Wang P F, Xue X F, Mu X P, Zhang J C, Cao Q, Du J J. Analysis of organic acid accumulation characteristics and organic acid- metabolizing enzyme activities of chinese dwarf cherry (Cerasus humilis Bunge) fruit[J]. Scientia Agricultura Sinica, 2013, 46(19): 4101-4109. DOI: 10.3864/j.issn.0578-1752.2013.19.016. (in Chinese)
[6]	Pangborn R M. Relative taste intensities of selected sugars and organic acids[J]. Journal of Food Science, 1963, 28(6): 726-733.
[7]	Liu Y L, Zhang X J, Zhao Z Y. Effects of fruit bagging on anthocyanins, sugars, organic acids, and color properties of ‘Granny Smith’ and ‘Golden Delicious’ during fruit maturation[J]. European Food Research and Technology, 2013, 236(2): 329-339.
[8]	Cao M K, Zeng S C, Wang J X, Guo W C. Assessment of SSC and soluble sugar content of three pear cultivars during storage using dielectric method[J]. Postharvest Biology and Technology, 2024, 212: 112906.
[9]	Wu B, Quilot B, Kervella J, Génard M, Li S. Analysis of genotypic variation of sugar and acid contents in peaches and nectarines through the principle component analysis[J]. Euphytica, 2003, 132(3): 375-384.
[10]	刘硕, 刘有春, 刘宁, 张玉萍, 章秋平, 徐铭, 张玉君, 刘威生. 李属(Prunus)果树品种资源果实糖和酸的组分及其构成差异[J]. 中国农业科学, 2016, 49(16): 3188-3198. DOI: 10.3864/j.issn.0578-1752.2016.16.012.
	Liu S, Liu Y C, Liu N, Zhang Y P, Zhang Q P, Xu M, Zhang Y J, Liu W S. Sugar and organic acid components in fruits of plum cultivar resources of genus Prunus[J]. Scientia Agricultura Sinica, 2016, 49(16): 3188-3198. DOI: 10.3864/j.issn.0578-1752.2016.16.012. (in Chinese)
[11]	Gao Y, Kan C N, Wan C P, Chen C Y, Chen M, Chen J Y. Effects of hot air treatment and chitosan coating on citric acid metabolism in ponkan fruit during cold storage[J]. PLoS ONE, 2018, 13(11): e0206585.
[12]	Zhang Y Z, Li P M, Cheng L L. Developmental changes of carbohydrates, organic acids, amino acids, and phenolic compounds in ‘Honeycrisp’ apple flesh[J]. Food Chemistry, 2010, 123(4): 1013-1018.
[13]	Liu L, Chen C X, Zhu Y F, Xue L, Liu Q W, Qi K J, Zhang S L, Wu J. Maternal inheritance has impact on organic acid content in progeny of pear (Pyrus spp.) fruit [J]. Euphytica, 2016, 209(2): 305-321.
[14]	Zheng B B, Zhao L, Jiang X H, Cherono S, Liu J J, Ogutu C, Ntini C, Zhang X J, Han Y P. Assessment of organic acid accumulation and its related genes in peach[J]. Food Chemistry, 2021, 334: 127567.
[15]	Zhang J, Nie J Y, Li J, Zhang H, Li Y, Farooq S, Bacha S A S, Wang J. Evaluation of sugar and organic acid composition and their levels in highbush blueberries from two regions of China[J]. Journal of Integrative Agriculture, 2020, 19(9): 2352-2361.
[16]	刘晓荣, 刘有春, 刘成, 陆晓春, 陶承光. 越橘果实发育过程中糖酸积累及抗氧化性形成的动态变化[J]. 沈阳农业大学学报, 2020, 51(4): 462-469.
	Liu X R, Liu Y C, Liu C, Lu X C, Tao C G. Dynamic changes of sugar-acid accumulation and antioxidant formation in blueberry (Vaccinium corymbosum) during fruit development[J]. Journal of Shenyang Agricultural University, 2020, 51(4): 462-469. (in Chinese)
[17]	Smrke T, Veberic R, Hudina M, Zitko V, Ferlan M, Jakopic J. Fruit quality and yield of three highbush blueberry (Vaccinium corymbosum L.) cultivars grown in two planting systems under different protected environments [J]. Horticulturae, 2021, 7(12): 591.
[18]	Acharya T P, Malladi A, Nambeesan S U. Sustained carbon import supports sugar accumulation and anthocyanin biosynthesis during fruit development and ripening in blueberry (Vaccinium ashei)[J]. Scientific Reports, 2024, 14: 24964.
[19]	Ayaz F A, Kadioglu A, Bertoft E, Acar C, Turna I. Effect of fruit maturation on sugar and organic acid composition in two blueberries (Vaccinium arctostaphylos and V. myrtillus) native to Turkey [J]. New Zealand Journal of Crop and Horticultural Science, 2001, 29(2): 137-141.
[20]	Gupta V, Estrada A D, Blakley I, Reid R, Patel K, Meyer M D, Andersen S U, Brown A F, Lila M A, Loraine A E. RNA-Seq analysis and annotation of a draft blueberry genome assembly identifies candidate genes involved in fruit ripening, biosynthesis of bioactive compounds, and stage-specific alternative splicing[J]. GigaScience, 2015, 4(1): 5.
[21]	Das P R, Darwish A G, Ismail A, Haikal A M, Gajjar P, Balasubramani S P, Sheikh M B, Tsolova V, Soliman K F A, Sherif S M, El-Sharkawy I. Diversity in blueberry genotypes and developmental stages enables discrepancy in the bioactive compounds, metabolites, and cytotoxicity[J]. Food Chemistry, 2022, 374: 131632.
[22]	陈发兴, 刘星辉, 陈立松. 果实有机酸代谢研究进展[J]. 果树学报, 2005, 22(5): 526-531.
	Chen F X, Liu X H, Chen L S. Advances in research on organic acid metabolism in fruits[J]. Journal of Fruit Science, 2005, 22(5): 526-531. (in Chinese)
[23]	覃红梅, 杨国顺, 梁晓文, 罗飞雄, 何建军. 野酿2号毛葡萄果实发育进程中苹果酸代谢规律及相关基因表达分析[J]. 南方农业学报, 2024, 55(8): 2332-2341.
	Qin H M, Yang G S, Liang X W, Luo F X, He J J. Malic acid metabolism regulation and related genes expression during berry development in Vitis heyneana cv. Yeniang No.2[J]. Journal of Southern Agriculture, 2024, 55(8): 2332-2341. (in Chinese)
[24]	廖香娇, 张王妮, 赵凌吉, 刘勇, 杨莉. 桃溪蜜柚果实发育期有机酸含量变化及柠檬酸代谢相关基因表达分析[J]. 江苏农业科学, 2025, 53(12): 183-188.
	Liao X J, Zhang W N, Zhao L J, Liu Y, Yang L. Changes of organic acid content and expression of genes related to citric acid metabolism in Taoxi pomelo during fruit development period[J]. Jiangsu Agricultural Sciences, 2025, 53(12): 183-188. (in Chinese)
[25]	邵姁, 王月, 应炎标, 朱友银, 刘向蕾, 葛凡, 郭卫东. 蓝莓磷酸烯醇式丙酮酸羧化酶基因cDNA克隆及表达分析[J]. 基因组学与应用生物学, 2016, 35(1): 166-171.
	Shao X, Wang Y, Ying Y B, Zhu Y Y, Liu X L, Ge F, Guo W D. Cloning and expression analysing of phosphoenolpyruvate carboxylase cDNA in blueberry[J]. Genomics and Applied Biology, 2016, 35(1): 166-171. (in Chinese)
[26]	蒋际谋, 陈秀萍, 胡文舜, 姜帆, 邓朝军, 郑少泉. 枇杷种质资源果实糖组分及含量特征[J]. 园艺学报, 2015, 42(9): 1781-1788.
	Jiang J M, Chen X P, Hu W S, Jiang F, Deng C J, Zheng S Q. Characteristics of components and contents of soluble sugars in mature fruits of loquat germplasm[J]. Acta Horticulturae Sinica, 2015, 42(9): 1781-1788. (in Chinese)
[27]	董畅, 覃杨, 曾琪琪, 霍珏璇. 不同蓝莓品种果实发育过程中的品质指标变化趋势研究[J]. 现代农业科技, 2025(9): 43-47.
	Dong C, Qin Y, Zeng Q Q, Huo J X. Change trends in quality indicators of different blueberry varieties during fruit development[J]. XianDai NongYe KeJi, 2025(9): 43-47. (in Chinese)
[28]	陈招芳, 李恒刚, 许轲, 黄金秋, 汪志辉, 吴林, 王迅. 四川主栽蓝莓品种果实生长发育过程中的品质变化规律[J]. 食品与发酵工业, 2020, 46(12): 244-250.
	Chen Z F, Li H G, Xu K, Huang J Q, Wang Z H, Wu L, Wang X. Quality change patterns during fruit development in blueberries mainly planted in Sichuan[J]. Food and Fermentation Industries, 2020, 46(12): 244-250. (in Chinese)
[29]	陈凯莉. 蓝莓果实发育中果实主要品质变化及果胶分解关键酶活性及表达量分析[D]. 雅安: 四川农业大学, 2018.
	Chen K L. Changes of main fruit quality and activity of key enzymes in pectin decomposition and their expression in blueberry fruit development process[D]. Yaan: Sichuan Agricultural University, 2018. (in Chinese)
[30]	李明瑞. 蓝莓果实发育中色素及糖酸含量变化研究[D]. 沈阳: 沈阳农业大学, 2024.
	Li M R. Changes of pigment, sugar and acid contents during fruit development in blueberry[D]. Shenyang: Shenyang Agricultural University, 2024. (in Chinese)
[31]	徐玉涛, 李珂珂, 王贺新, 陈丽荣, 弓晓杰. 高效液相色谱法对蓝莓果实中8个有机酸含量的测定[J]. 食品科学, 2015, 36(18): 127-131.
	Xu Y T, Li K K, Wang H X, Chen L R, Gong X J. Determination of eight organic acids in blueberries by HPLC[J]. Food Science, 2015, 36(18): 127-131. (in Chinese)
[32]	吴林, 张强, 臧慧明, 徐振彪, 徐德冰. 云南丽江和吉林靖宇蓝莓糖酸组分差异化分析[J]. 中国果树, 2019(6): 54-58, 64.
	Wu L, Zhang Q, Zang H M, Xu Z B, Xu D B. Determination of sugar and acid components in blueberries from Lijiang, Yunnan and Jingyu, Jilin[J]. China Fruits, 2019(6): 54-58, 64. (in Chinese)
[33]	王瑜, 杨浩博, 仲帅, 刘鑫, 盛煜钧, 李彤, 宗成文. HPLC-ELSD法测定长白山笃斯越桔果实中可溶性糖含量[J]. 食品工业科技, 2020, 41(2): 238-243.
	Wang Y, Yang H B, Zhong S, Liu X, Sheng Y J, Li T, Zong C W. Determination of soluble sugar in Vaccinium uliginosum fruit in Changbai Mountain by HPLC-ELSD[J]. Science and Technology of Food Industry, 2020, 41(2): 238-243. (in Chinese)
[34]	张素敏, 杨巍, 魏鑫, 高文民, 刘成. 蓝莓果实外观与糖、酸、酚组成特征[J]. 食品研究与开发, 2023, 44(10): 53-61.
	Zhang S M, Yang W, Wei X, Gao W M, Liu C. Appearance and sugar, acid and phenol composition of blueberry fruit[J]. Food Research and Development, 2023, 44(10): 53-61. (in Chinese)
[35]	Chen L, Lou T J, Dong K X, Dong M, Sun H Y, Che J L, Yang S F, Li J, Li Y D. Inheritance analysis of sugar and organic acid components in blueberry hybrid progenies[J]. Scientia Horticulturae, 2025, 350: 114345.
[36]	霍宏亮, 任伟春, 许伟华, 叶芳, 林娅沁, 王金燕, 杜媛媛, 凌益春, 杨晓乐, 曹玉芬. 云和雪梨主要营养素分析[J]. 中国南方果树, 2024, 53(6): 187-193.
	Huo H L, Ren W C, Xu W H, Ye F, Lin Y Q, Wang J Y, Du Y Y, Ling Y C, Yang X L, Cao Y F. Analysis of the primary nutrients in Yunhexueli (Pyrus pyrifolia)[J]. South China Fruits, 2024, 53(6): 187-193. (in Chinese)
[37]	于淼, 阮成江, 李贺, 张少凤, 徐弘灏, 刘向前. HPLC法测定10个引自北美蓝莓品种果实中的糖酸组分及含量[J]. 食品工业科技, 2016, 37(10): 77-81.
	Yu M, Ruan C J, Li H, Zhang S F, Xu H H, Liu X Q. Components and contents of sugar and acid in 10 blueberry cultivars from the North American using HPLC[J]. Science and Technology of Food Industry, 2016, 37(10): 77-81. (in Chinese)
[38]	Yang Y J, Liu X, Wu H R, He X F, Bi Y R, Zhu Y, Liu Z L. Radical scavenging activity and cytotoxicity of active quinic acid derivatives from Scorzonera divaricata roots[J]. Food Chemistry, 2013, 138(2/3): 2057-2063.
[39]	Benali T, Bakrim S, Ghchime R, Benkhaira N, El Omari N, Balahbib A, Taha D, Zengin G, Hasan M M, Bibi S, Bouyahya A. Pharmacological insights into the multifaceted biological properties of quinic acid[J]. Biotechnology and Genetic Engineering Reviews, 2024, 40(4): 3408-3437.
[40]	Etienne A, Génard M, Lobit P, Mbeguié-A-Mbéguié D, Bugaud C. What controls fleshy fruit acidity? A review of malate and citrate accumulation in fruit cells[J]. Journal of Experimental Botany, 2013, 64(6): 1451-1469.
[41]	游鸿瑞, 武运, 马文瑞, 杨华峰, 白晓雪, 张利利, 唐江北, 黄文书. 微量元素对马瑟兰葡萄果实糖酸代谢及基因表达的影响[J]. 食品与发酵工业, 2026, 52(8): 170-180.
	You H R, Wu Y, Ma W R, Yang H F, Bai X X, Zhang L L, Tang J B, Huang W S. Effects of trace elements on sugar-acid metabolism and gene expression in Marselan grape fruits[J]. Food and Fermentation Industries, 2026, 52(8): 170-180. (in Chinese)
[42]	杨文渊, 谢红江, 陶炼, 宦云敏, 陈善波, 林立金, 廖明安. '金冠'苹果及其优系(SGP-1) 果实发育期间有机酸代谢特征比较[J]. 西北植物学报, 2022, 42(5): 829-836.
	Yang W Y, Xie H J, Tao L, Huan Y M, Chen S B, Lin L J, Liao M A. Comparison of organic acid metabolism characteristics between’Golden Delicious’Apple and its excellent variation Material’SGP-1'during fruit development[J]. Acta Botanica Boreali-Occidentalia Sinica, 2022, 42(5): 829-836. (in Chinese)

组分 Components	回归方程 Regression equation	相关系数 R²	线性范围 Linear range (g·kg^-1)
果糖Fructose	y=151709x-27135	0.996	0.010—20
葡萄糖Glucose	y=134296x-1781.1	0.999	0.010—20
蔗糖Sucrose	y=136951x-6747.4	0.999	0.005—20
柠檬酸Citarate	y=651989x-73438	1.000	0.060—60
奎宁酸Quinic	y=276651x-3371.1	1.000	0.030—3
苹果酸Malate	y=473139x-29753	0.999	0.020—20
草酸Oxalate	y=7×10⁶x-3036.4	0.999	0.004—2
莽草酸Shikimate	y=3×10⁷x-339060	0.999	0.002—1
乳酸Lactate	y=497673x-12516	1.000	0.020—20

基因Gene	正向引物Forward prime (5′-3′)	反向引物Reverse prime (5′-3′)
GAPDH	ACTACCATCCACTCTATCACCG	AACACCTTACCAACAGCCTTG
PEPC	CATTGAGAGTCGTTCCATTGTTTGA	AATCTGAGTACCCAATCATCACCTC

品种 Cultivars	总酸Total acid (Acid%)			总可溶性固形物Total soluble solids (Brix%)
品种 Cultivars	青果期Green	粉果期Pink	蓝果期Blue	青果期Green	粉果期Pink	蓝果期Blue
H5	5.00±0.17a	3.46±1.96b	0.90±0.07c	7.60±0.46b	7.60±0.20b	15.53±1.00a
L25 Eureka Sunrise	2.88±0.36a	2.87±0.29a	0.94±0.13b	6.63±0.21b	6.60±0.17b	9.33±0.06a
L	3.14±0.18b	4.57±0.33a	1.15±0.01c	8.80±0.10b	8.33±0.29b	15.87±0.42a
西班牙49 Blue Marina	2.43±0.33a	2.03±0.64ab	1.42±0.16b	8.43±0.15b	7.50±0.26c	17.57±0.23a
绿宝石Emerald	3.21±0.58a	2.44±0.78ab	0.77±0.44b	8.30±0.10b	8.47±0.15b	11.93±0.64a
F6 Magica	3.23±0.37a	3.10±0.06a	1.09±0.09b	7.37±0.55b	7.37±0.23b	13.17±0.40a
L11 Arana	4.45±0.15a	4.22±0.03a	1.84±0.67b	7.40±0.44c	7.77±0.23b	12.60±1.35a
西班牙45 Blue Manila	3.61±0.14a	2.50±0.72b	1.20±0.06c	6.70±0.10c	8.23±0.38b	10.73±1.46a
L26 Eureka Sunset	3.22±0.29a	2.99±0.28a	0.97±0.09b	6.43±0.45c	9.33±0.38b	13.97±1.46a
L10 Costa	5.57±0.95a	1.25±0.03b	0.91±0.09b	7.90±0.36c	10.77±0.90b	12.33±0.32a
西班牙02 Blue Madeira	4.30±0.64a	2.79±0.28b	0.88±0.02c	9.47±0.76c	14.77±1.03b	17.20±0.70a
西班牙42 Blue Maldiva	4.32±0.31a	2.28±0.03b	0.82±0.03c	7.43±0.15c	10.47±0.06b	12.47±0.32a
法新Fargo	5.37±0.13a	2.76±0.07b	1.08±0.03c	7.40±0.61c	10.97±0.40b	13.60±0.62a
泰坦Titan	5.33±0.19a	4.73±0.13a	2.53±0.38b	8.50±0.62b	10.80±1.13a	13.33±0.74a

组分 Components	总糖 Total soluble sugars	蔗糖 Sucrose	葡萄糖 Glucose	果糖 Fructose	总有机酸 Total organic acids	奎宁酸 Quinic	草酸 Oxalate	苹果酸 Malate	柠檬酸 Citrate	莽草酸 Shikimate	乳酸 Lactate
总糖 Total soluble sugars	1.00
蔗糖Sucrose	0.08	1.00
葡萄糖Glucose	0.68*	0.04	1.00
果糖Fructose	0.73*	0.10	0.87**	1.00
总有机酸 Total organic acids	-0.37	-0.14	-0.12	-0.30	1.00
奎宁酸Quinic	-0.05	-0.08	-0.01	0.02	0.90**	1.00
草酸Oxalate	-0.34	-0.06	-0.35	-0.42	0.91**	0.70**	1.00
苹果酸Malate	-0.26	-0.01	-0.18	-0.16	0.47	0.10	0.30	1.00
柠檬酸Citarate	0.02	-0.19	0.28	0.20	0.65*	0.45	0.35	-0.29	1.00
莽草酸Shikimate	-0.29	0.05	-0.23	-0.24	0.28	-0.10	0.10	0.78**	-0.63*	1.00
乳酸Lactate	-0.34	-0.06	-0.38	-0.43	0.92**	0.73**	0.99**	0.29	0.36	0.10	1.00