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红树莓果实发育过程中抗氧化物质含量变化及相关基因表达特性分析
张姗姗, 吴雅琼, 黄鑫, 吴文龙, 闾连飞, 李维林
南京林业大学学报（自然科学版） ›› 2025, Vol. 49 ›› Issue (6) : 125-134.
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红树莓果实发育过程中抗氧化物质含量变化及相关基因表达特性分析
Analysis of changes in antioxidant contents and related gene expression characteristics during red raspberry fruit development
【目的】红树莓(Rubus idaeus)富含多种营养成分,其中,总酚、类黄酮和维生素C(VC)等天然抗氧化活性物质备受关注。为更有效开发利用该资源,本研究拟对红树莓不同发育阶段的外观品质、抗氧化物含量变化及抗氧化酶相关基因表达规律进行系统研究。【方法】以优良红树莓栽培品种‘秋福'(‘Autumn Bliss')为试材,运用蒽酮比色法、氢氧化钠滴定法、羟胺法和qRT-PCR等方法对其不同发育阶段果实的外观、品质、抗氧化指标和抗氧化酶活性进行测定,对其抗氧化酶基因的表达特性进行分析,并对各指标进行相关性和主成分分析。【结果】红树莓果实的质量和横纵径随着果实的成熟而逐渐增加,果实亮度则逐渐降低,饱和度的变化趋势为先增加后减少。随着红树莓果实的逐渐成熟,糖酸比呈现逐渐升高的趋势,在35 DAF(开花后天数)时达到最高,为5.91。总酚、类黄酮和维生素C(VC)含量在红树莓果实发育过程中整体呈下降趋势,青果时期含量最高,与总抗氧化能力呈正相关;花色苷含量呈上升趋势,成熟期达到最高,质量分数为802.42 μg/g。此外,果实的抗氧化酶中POD活性(62.58 U/g)和GSH质量分数(1.01 mg/g)均在青果时最高,与总抗氧化能力呈极显著正相关;而SOD活性、MDA含量和 产生速率则在成熟时最高,与总抗氧化能力呈负相关。SOD活性相关基因RuSOD1和RuSOD2在果实发育中的表达量整体呈上升趋势,RuSODCP.2表达量前期也呈上升趋势,成熟阶段略有下降,RuFSD的表达量整体变化幅度不大,与SOD活性均呈正相关;POD活性的相关基因RuPOD表达量逐渐下降,RuPER5在7~14 DAF时期表达量下降,14~35DAF时期上升,RuPER25则是7~21 DAF时期表达量上升,21~35 DAF时期下降,而RuPROX6表达量逐渐上升,RuPER17表达量变化并不显著,POD活性与RuPOD、RuPER5、RuPER17和RuPER25表达量均呈正相关,而与RuPROX6呈负相关。【结论】红树莓青果时期抗氧化能力最强,其中总酚、类黄酮、POD和GSH是主要的抗氧化物成分,研究结果将为后续深入探讨红树莓果实抗氧化物质的变化规律及作用机制奠定基础。
【Objective】Rubus idaeus (red raspberry) fruits are renowned for their rich nutritional profile, particularly their high content of natural antioxidant compounds such as total phenols, flavonoids, and vitamin C (VC). To optimize the development and utilization of this valuable resource, this study aims to systematically investigate the appearance quality, dynamic changes in antioxidant content, and expression patterns of antioxidant enzyme-related genes of red raspberry at different developmental stages.【Method】The study employed the superior red raspberry cultivar ‘Autumn Bliss' as the experimental material. A comprehensive analysis was conducted to measure the fruit's appearance, quality, antioxidant indicators and antioxidant enzyme activity at different developmental stages. Methods such as anthrone colorimetry, sodium hydroxide titration, hydroxylamine method and quantitative real-time PCR (qRT-PCR) were utilized. The expression characteristics of antioxidant enzyme genes were analyzed, and the correlation and principal component analysis were performed to elucidate the relationships among the measured indicators.【Result】The fresh weight and transverse and longitudinal diameter of red raspberry fruit gradually increased with fruit maturity, while the fruit luminosity gradually decreased, and the trend of chroma change was initially increasing and then decreasing. As the red raspberry fruit gradually matures, the sugar acid ratio showed a gradually increasing trend, reaching its highest value of 5.91 at 35 days after flowering (DAF). The contents of total phenol, flavonoid and VC content showed an overall downward trend during the development of red raspberry fruits, with the highest content during the green fruit stage and a positive correlation with total antioxidant capacity. The content of anthocyanins showed an upward trend and reached its highest point during the mature stage, with a mass fraction of 802.42 μg/g. In terms of antioxidant enzyme activity, the POD activity (62.58 U/g) and glutathione (GSH) mass fraction (1.01 mg/g) were highest during the green fruit stage, respectively, showing an extremely significant positive correlation with the total antioxidant capacity. In contrast, SOD activity, MDA content, and production rate were highest at maturity and negatively correlated with total antioxidant capacity. The expression levels of SOD activity related genes RuSOD1 and RuSOD2 showed overall upward trends during fruit development, while the expression levels of RuSODCP.2 also showed an upward trend in the early stage and slightly decreased in the mature stage. The overall change in the expression level of RuFSD was not significant. And they were positively correlated with SOD activity. The expression level of the genes related to POD activity, RuPOD, gradually decreased. The expression level of RuPER5 decreased during the 7-14 DAF period and increased during the 14-35 DAF period, while the expression level of RuPER25 increased during the 7-21 DAF period and decreased during the 21-35 DAF period. The expression level of RuPROX6 gradually increased, while the expression level of RuPER17 did not change significantly. POD activity was positively correlated with the expression levels of RuPOD, RuPER5, RuPER17 and RuPER25, while it was negatively correlated with RuPROX6.【Conclusion】The findings indicate that red raspberry fruits exhibit the highest antioxidant capacity during the green fruit stage, with total phenols, flavonoids, POD and GSH identified as the primary antioxidant components. This research provides a foundation for further exploration into the dynamics changes and action mechanisms of antioxidant substances in red raspberry fruits, offering valuable insights for their potential applications in nutrition and health.
Rubus idaeus (red raspberry) / fruit development / quality / antioxidant enzyme / gene expression
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In order to explore the changes in the content of phenolic compounds and the activity of antioxidant enzymes during the growth and metabolism of red raspberry leaves, the leaves of ‘Heritage’ and ‘Autumn Bliss’ red raspberry were analyzed and compared for the contents of total flavonoids and seven phenolic substances and the activities of three antioxidant enzymes. The results showed that from May to December, the content of ellagic acid and peroxidase (POD) activity in the upper, middle and lower leaves of the two varieties showed a double peak curve during the process of growth and metabolism, the content of salicylic acid showed a downward trend, the contents of ferulic acid, catechin, epicatechin, rutin and total flavones showed a single peak curve, the superoxide dismutase (SOD) activity remained at a high level, and the catalase (CAT) activity decreased, then increased and finally decreased again. The content of caffeic acid in the upper leaves of the two varieties showed a downward trend, while those in the middle and lower leaves showed a single peak curve. The highest contents of salicylic acid, ferulic acid, ellagic acid and epicatechin were determined in the upper leaves of ‘Autumn Bliss’ in May, upper leaves in June, middle leaves in July and upper leaves in July, which were 59.33 mg/100 g, 158.18 mg/100 g, 250.84 mg/g and 350.04 mg/100 g, respectively. The highest contents of rutin, caffeic acid, catechin and total flavonoids were determined in the lower leaves of ‘Heritage’ in June, middle leaves in July, middle leaves in July and upper leaves in July, which were 8.00 mg/100 g, 100.48 mg/100 g, 18.71 mg/g and 34.18 mg/g, respectively. The highest activities of POD, SOD and CAT were determined in the upper leaves of ‘Autumn Bliss’ in July, the lower leaves of ‘Heritage’ in July and the lower leaves of ‘Heritage’ in July, which were 13.05 U/(g·min), 37.00 U/g and 41.12 U/(g·min), respectively. In conclusion, the contents of phenolic substances and antioxidant enzyme activities in the leaves of ‘Heritage’ and ‘Autumn Bliss’ red raspberry are highest from May to July. This study provides an important theoretical basis for the timely harvest and better development and utilization of red raspberry leaves.
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The composition and content of anthocyanins in mulberry extract and their <i>in vitro</i> antioxidant activities were determined. The composition of anthocyanins was identified by HPLC-MS, and the content of anthocyanins was determined by using cyanidin-3-<i>O</i>-glucoside as a standard, followed by investigating their antioxidant activity. The results showed that the content of total anthocyanins in mulberry extract was 314.30 μg/mg, which were mainly cyanidin-3-<i>O</i>-glucoside, cyanidin-3-<i>O</i>-rutinoside, and cyanidin. The <i>in vitro</i> antioxidant activity experiments showed that the IC<sub>50</sub> of the mulberry extract for scavenging DPPH·, ABTS<sup>+</sup>· and ·OH were 22.33 mg/mL, 12.90 mg/mL, and 1.74 mg/mL, respectively. Besides, 250 μg/mL mulberry extract had 0.744 reducing power. Therefore, mulberry extract has high anthocyanin content and good antioxidant activity, which provides a theoretical basis for developing and utilizing mulberry extract.
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The antioxidant properties of blueberries have been examined only in ripe fruit, although fruit of different maturities are used in processed food products. In this study, highbush blueberry cultivars Bergitta, Bluegold, and Nelson highbush blueberry fruit at different stages of ripeness were examined to characterize differences in oxygen radical absorbing capacity (ORAC) and the phenolic components responsible for ORAC. Underripe fruit at different stages of maturity were also stored at 20 °C for up to 8 days to assess changes in ORAC and phenolic content. Anthocyanin content was substantially higher in fruit of more advanced stages of ripeness. In contrast, the phenolic content and ORAC were lower in the riper fruit. Anthocyanins continued to form during storage, although rate of pigment formation declined after about 4 days. Less anthocyanin pigment was formed in the less ripe fruit. After 8 days of storage, the anthocyanin content of fruit harvested 5% to 50% or 50% to 95% blue exceeded that of ripe fruit. Up to 60% of the total phenolic content could be accounted for by anthocyanins. ORAC was positively correlated with total phenolic content (R2 = 0.78), but not with anthocyanin content.
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Anthocyanins are secondary metabolites derived from the specific branch of the flavonoid pathway, responsible for red, purple and blue coloration display in the flowers and fruits. The functions of anthocyanins are diverse, including acting as visual signals to pollinators, defense against biotic and abiotic stresses. Thus, anthocyanins have been the most intensely studied secondary metabolite pathway. From model plants to horticultural crops, numerous studies have resulted in the discovery of highly conserved MYB-bHLH-WDR (MBW) transcriptional complex for the regulation of anthocyanin biosynthesis in plants. Recent discoveries have revealed that the anthocyanin biosynthesis pathway is also controlled by MYB repressors. Here we focus on the research progress into the role of MYB repressors in anthocyanin biosynthesis. In particular, we will discuss their functions and relationship to the MBW complex in the control of anthocyanin accumulation. In addition, an integrated regulatory network of anthocyanin biosynthesis controlled by MYB repressors and MBW activation complex is built based on the significant progress.Copyright © 2019 Elsevier Masson SAS. All rights reserved.
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In this review we report the recent advances in anticancer activity of the family of natural occurring flavonoids, covering the time span of the last five years. The bibliographic data will be grouped, on the basis of biological information, in two great categories: reports in which the extract plants bioactivity is reported and the identification of each flavonoid is present or not, and reports in which the anticancer activity is attributable to purified and identified flavonoids from plants. Wherever possible, the targets and mechanisms of action as well as the structure-activity relationships of the molecules will be reported. Also, in the review it was thoroughly investigated the recent discovery on flavonoids containing the 2-phenyl-4H-chromen-4-one system even if some examples of unusual flavonoids, bearing a non-aromatic B-ring or other ring condensed to the base structure are reported.Copyright © 2017 Elsevier Masson SAS. All rights reserved.
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Blueberry is one of the richest phenolic sources, providing health benefits. To study blueberry phenolic biosynthesis, we investigated phenolics and proteomics at three typical fruit maturation phases. Multiple isoforms of enzymes and multiple members of transcription factors involved in phenolic biosynthesis were divergent and differently regulated. Regulation of some proteins resulted in change of phenolic content. During fruit maturation, down-regulation of VcOMT (CUFF.177.1) and VcLAR2 (CUFF.16780.1) was associated with decreases of ferulic acid and catechin, respectively; Up-regulation of VcFLS (CUFF.41155.1), and VcF3'5'H (CUFF.51711.1) and VcF3'5'H (gene.g10884.t1.1) likely drove increases of their products (quercetin and myricetin); Up-regulation of VcUFGALT (CUFF.20951.1) and VcUFGT73 (4333_g.1) and down-regulation of VcU5GT (CUFF.51258.1) were correlated to accumulation of anthocyanins with 3-glucoside/galactoside. Additionally, four TFs, VcAPRR2 (CUFF.24826.1), VcbHLH3 (CUFF.37765.1), VcWD (CUFF.28282.2) and VcWD (CUFF.28273.1) were probably related to regulation of anthocyanin biosynthesis. These proteins were potential targets for genetic improvement in a breeding program.Copyright © 2019. Published by Elsevier Ltd.
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Bitter gourd is one of the important cucurbits and highly liked among both farmers and consumers due to its high net return and nutritional value. However, being monoecious, it exhibits substantial variation in flower bearing pattern. Plant growth regulators (PGRs) are known to influence crop phenology while gibberellic acid (GA3) is one of the most prominent PGRs that influence cucurbits phenology. Therefore, a field trial was conducted at University of Agriculture Faisalabad to evaluate the impact of a commercial product of gibberellic acid (GA3) on growth, yield and quality attributes of two bitter gourd (Momordica charantiaL.) cultivars. We used five different concentrations (0.4 g, 0.6 g, 0.8 g, 1.0 g, and 1.2 g per litre) of commercial GA3 product (Gibberex, 10% Gibberellic acid). Results showed that a higher concentration of gibberex (1.0 and 1.20 g L−1 water) enhanced the petiole length, intermodal length, and yield of bitter gourd cultivars over control in Golu hybrid and Faisalabad Long. A significant decrease in the enzyme superoxidase dismutase, peroxidase, and catalase activities were observed with an increasing concentration of gibberex (1.0 and 1.20 gL−1 water) as compared to control. These results indicate that the exogenous application of gibberex at a higher concentration (1.2 g L−1) has a dual action in bitter gourd plant: i) it enhances the plant growth and yield, and ii) it also influenced the antioxidant enzyme activities in fruits. These findings may have a meaningful, practical use for farmers involved in agriculture and horticulture.
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Environmental stresses trigger a wide variety of plant responses, ranging from altered gene expression and cellular metabolism to changes in growth rates and crop yields. A plethora of plant reactions exist to circumvent the potentially harmful effects caused by a wide range of both abiotic and biotic stresses, including light, drought, salinity, high temperatures, and pathogen infections. Among the environmental stresses, drought stress is one of the most adverse factors of plant growth and productivity. Understanding the biochemical and molecular responses to drought is essential for a holistic perception of plant resistance mechanisms to water-limited conditions. Drought stress progressively decreases CO2 assimilation rates due to reduced stomatal conductance. Drought stress also induces reduction in the contents and activities of photosynthetic carbon reduction cycle enzymes, including the key enzyme, ribulose-1,5-bisphosphate carboxylase/oxygenase. The critical roles of proline and glycine-betaine, as well as the role of abscisic acid (ABA), under drought stress conditions have been actively researched to understand the tolerance of plants to dehydration. In addition, drought stress-induced generation of active oxygen species is well recognized at the cellular level and is tightly controlled at both the production and consumption levels in vivo, through increased antioxidative systems. Knowledge of sensing and signaling pathways, including ABA-mediated changes in response to drought stress, is essential to improve crop management. This review focuses on the ability and strategies of higher plants to respond and adapt to drought stress.
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In this study, nutritional composition (protein, lipids, carbohydrates, ash, and moisture), physicochemical properties (soluble solid content, titratable acidity, texture and instrumental colour on surface, and internal section), phytochemicals (total phenolic content and anthocyanin content), and antioxidant capacity (DPPH—2,2′-diphenyl-1-picrylhydrazyl radical scavenging capacity and ferric-reducing antioxidant power) of three strawberry (Fragaria × ananassa Duch.) cultivars (cv. “Primoris”, cv. “Endurance”, and cv. “Portola”) produced in the western region of Portugal (Caldas da Rainha) were evaluated. From the obtained, results no significant differences (P > 0.05) in nutritional composition were detected in all of the cultivars; with the exception of lower protein content observed in cv. “Portola” (0.57 g/100 g ± 0.04; P < 0.05). Regarding the a* value of whole strawberry fruits, no significant differences (P > 0.05) were found in any of the cultivars, which revealed a similar redness. The cv. “Endurance” revealed the highest bioactivity content compared to the other cultivars. Overall, these results provide important information about the high quality of strawberry produced in the western region of Portugal and may be used as a tool for adding value to a functional food in the Mediterranean diet due to the phytochemical composition and nutritional value of strawberry fruits
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王友升, 谷祖臣, 张帆. 不同品种和成熟度树莓和黑莓果实的氧化和抗氧化活性比较[J]. 食品科学, 2012, 33(9):81-86.
采用多变量分析方法探讨不同品种和成熟度对树莓和黑莓果实氧化和抗氧化活性的影响。单因素方差分析表明，“三冠王”黑莓果实的谷胱甘肽(GSH)、总酚和总黄酮含量均显著高于其他品种，且随果实成熟度的升高，总抗氧化能力下降。品种和成熟度对树莓和黑莓果实的GSH、总酚和丙二醛(MDA)含量，以及清除羟自由基( ·OH)能力的影响更为显著。主成分分析表明，相比于成熟度而言，品种对树莓的抗氧化能力影响更为显著。相关性分析和偏最小二乘回归分析表明，过氧化氢与MDA、GSH和清除超氧阴离子自由基(O2－ ·)均呈正相关性，而总酚与总抗氧化能力、清除 ·OH能力、清除DPPH自由基能力呈正相关性；通径分析表明，树莓和黑莓果实的总酚含量和总黄酮对H2O2含量的直接效应最强。
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王睿婷, 王鑫, 张富敏, 等. 树莓的抗氧化活性与主要化学成分的相关分析[J]. 沈阳药科大学学报, 2013, 30(11):901-905.
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