Scientia Agricultura Sinica ›› 2026, Vol. 59 ›› Issue (13): 2919-2932.doi: 10.3864/j.issn.0578-1752.2026.13.013

• HORTICULTURE • Previous Articles     Next Articles

Analysis of Sugar and Organic Acid Components and Related Metabolic Gene Expression During Fruit Development in 14 Blueberry Cultivars

HUANG MengChi(), CHEN LiNa, GAO DengTao, LIU Li, HAN JingYi, DUAN ShuYun, LIU ZhaoNing, LI HaoXian, LU ZhenHua()   

  1. Zhengzhou Fruit Tree Research Institute, Chinese Academy of Agricultural Sciences, Zhengzhou 450009
  • Received:2026-02-10 Accepted:2026-05-09 Online:2026-07-01 Published:2026-07-01
  • Contact: LU ZhenHua

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

Fig. 1

Fruit phenotype of 14 varieties at differfent developmental stages for blueberry"

Table 1

Linear correlation of sugar and acid components"

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

Table 2

Primers information used for qRT-PCR"

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

Table 3

Total acid and total soluble solids content of 14 blueberry varieties at different maturity stages"

品种
Cultivars
总酸Total acid (Acid%) 总可溶性固形物Total soluble solids (Brix%)
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

Fig. 2

Distribution of organic acid types and contents in 14 blueberry varieties at different maturity stages"

Fig. 3

Distribution of soluble sugar types and contents in 14 blueberry varieties at different maturity stages"

Fig. 4

Cluster relationships of organic acids and soluble sugars in 14 blueberry varieties A: Cluster diagram of organic acids and soluble sugars for 14 blueberry varieties; B: Relative proportions of organic acids and soluble sugar content in Cluster I to Cluster IV"

Table 4

Correlation of sugar and organic acid components in blueberry fruits at the blue fruit stage"

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

Fig. 5

Analysis of citric acid, malic acid content, and PEPC expression in blueberry fruits at different maturation stages"

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