Skip to main content
NCBI home page
ACS AuthorChoice logoLink to ACS AuthorChoice
. 2026 Sep 25;74(40):31321–31328. doi: 10.1021/acs.jafc.6c08282

Regulation of the Biosynthesis of Phenolic Compounds in Vaccinium Berries

Laura Jaakola †,‡,*
PMCID: PMC13674239  PMID: 42916028

Abstract

Vaccinium berries are among the food ingredients with the highest levels and most diverse compositions of phenolic compounds, including anthocyanins, proanthocyanidins, and flavonols along with hydroxycinnamic acids. The biosynthesis of these compounds through the phenylpropanoid pathway is regulated via coordinated transcriptional control of the structural enzymes in the pathway by the MBW complex that involves interactions with DNA-binding R2R3MYB transcription factors, MYC-like basic helix loop helix (bHLH), and WD40-repeat proteins. Some upstream regulators of the pathway were also identified. Both the developmental process and environmental factors influence the metabolism of phenolic compounds during berry ripening. Recent studies have deepened our understanding of the complex regulatory networks controlling the berry ripening-related biosynthesis of phenolic compounds. This knowledge enables the development of new tools for the selection of cultivars with desirable traits, such as berries with enhanced health benefits or cultivars with greater stress resistance.

Keywords: anthocyanins, berries, flavonoids, phenolic compounds, Vaccinium

graphic file with name jf6c08282_0004.webp

1. Introduction

Vaccinium species encompass many economically important cultivated and wild berries, such as blueberries (Vaccinium corymbosum, V. virgatum, V. angustifolium), cranberries (V. macrocarpon), bilberries (V. myrtillus), and lingonberries (V. vitis-idaea). These species grow in acidic soils and mostly in cooler northern hemisphere climates. Multiple studies have shown evidence on the health beneficial effects of Vaccinium berries, for example, in reducing risk of metabolic syndrome and various degenerative and microbial diseases. , Health benefits are mostly attributed to high level of phenolic compounds present in these berries, even if they are also good sources of other health beneficial compounds, such as vitamins, carotenoids, and fibers. Recently, a substantial amount of new genetic and functional data has been produced on the regulation of phenolic compounds in Vaccinium species. This review considers the update on the biosynthesis of the phenolic compounds present in wild- and cultivated Vaccinium berries. The special focus is on the recent progress in understanding the regulation of the biosynthesis of anthocyanins and other key compounds of the flavonoid pathway during berry development and ripening and as response to environmental cues.

2. Vaccinium Phenolics

In Vaccinium species, anthocyanins, proanthocyanidins, and flavonols are flavonoid classes contributing to the main phenolics in addition to hydroxycinnamic acids. ,, Flavonoids in general are the most common phenolic compounds in fruits, all of which are built upon the same C15 (C6–C3–C6) basic skeleton in which the three-carbon bridge is cyclized with oxygen. Flavonoid classes differ in the oxidation level of the central pyran ring except chalcones, which have an open central pyran (Figure ).

1.

1

Open in a new tab

Chemical structures of the most common flavonoids inVaccinium berries.

Vaccinium species contain high levels of hydroxycinnamic acids (HCAs), which can be found in all organs and represent the main phenolics in the leaves. HCAs are found in plant tissues in combined forms, often linked through an ester bond between their carboxylic group and an alcohol group of an organic compound or through a bond with one of their phenolic groups. HCAs can also be associated with flavonoids, lignin, suberin, and cutin. In most Vaccinium berries, caffeic, ferulic, and p-coumaric acids have all been identified. ,,

Flavonols (3-hydroxyflavones) are distinguished by a double bond between the C-2 and C-3 positions along with a hydroxyl group at the C-3 position. These compounds are primarily found as glycosides, with the sugar moiety typically attached to the hydroxyl group at the 3-position. The possible sugar residues include d-glucose, d-galactose, l-rhamnose, l-arabinose, d-xylose, and d-glucuronic acid. Quercetin glycosides are the most common flavonols present in all Vaccinium berries and also in early developmental stages. Myricetin glycosides are found more abundantly in ripe, blue-colored berries, while kaempferol glycosides are present in smaller amounts (Figure ). , In leaves, quercetin and kaempferol glycosides are the dominant flavonols.

Anthocyanins are the primary pigments in Vaccinium species, responsible for the bluish and reddish hues of the berries. The accumulation of anthocyanins is rapid during ripening stage attracting seed dispersals and enhance the quality and value of ripe berries by influencing their appearance and nutritional properties. Structurally, anthocyanins consist of an anthocyanidin aglycone attached to one or more sugar molecules. To date, over 700 distinct anthocyanins have been identified, with the most significant being glucosides of six common aglycones: cyanidins, delphinidins, pelargonidins, petunidins, peonidins, and malvidins. Among these, cyanidins, delphinidins, petunidins, peonidins, and malvidins are typically found in blue-colored Vaccinium berries, while red-colored cranberries and lingonberries primarily contain cyanidin-derived anthocyanins, specifically cyanidin and peonidin glucosides (Figure ). , The levels of anthocyanins in Vaccinium berries vary largely between species but also within species affected by the ripening stage, environmental factors, and genetic variation. Bilberries generally contain 4–5 fold higher levels of anthocyanins compared to blueberries, due to the high anthocyanin content in flesh, whereas in most blueberry species anthocyanins are only found in skin. Moreover, further decorations of anthocyanins, such as acylation, have also been reported especially in blue-colored Vaccinium berries, such as blueberries, bilberries, and bog bilberries. − Numerous studies associate anthocyanins with protective effects against human diseases, including cardiovascular conditions, age-related degenerative diseases, obesity, metabolic syndrome, and cancer. , It has also been shown that a key mechanism behind these health benefits is the significant influence of an anthocyanin-rich diet on gut microflora, which further processes these compounds into functional end products. −

Proanthocyanidins (PAs), also termed condensed tannins, are the most structurally complex members of the flavonoid family, ranging from dimers to high-molecular-weight polymers. These polymers are composed of flavan-3-ols as their structural units, which also exist as monomers in the plant tissues. In Vaccinium species, the most commonly identified flavan-3-ols include (+)-catechin (2,3-trans-), (−)-epicatechin (2,3-cis-), (+)-gallocatechin, and (+)-epigallocatechin (Figure ). The structural diversity of proanthocyanidins arises from variations in the stereochemistry and hydroxylation patterns of the flavan-3-ol starter and extension units. The most prevalent flavanol–flavanol linkages are carbon–carbon bonds between the 4-position of the upper unit and the 6- or 8-position of the lower unit, known as B-type linkages. Less common A-type linkages involve a C4 → C8 bond along with an ether bond O7 → C2. Although A-type linkages are rare in fruits, both A- and B-type linkages have been identified in Vaccinium species. During berry development, the level of PAs is highest during the earlier developmental stages and decreases during the ripening. As astringent compounds, PAs provide protection for the developing seeds inside berry against early feeding of the berries. PAs are the most abundant flavonoids in cranberry fruits and are associated with their potential health benefits, such as preventing urinary tract infections and treating oral infections.

3. Biosynthesis of Phenolic Compounds

The shikimate pathway serves as the starting point for the biosynthesis of the phenolic compounds. The general phenylpropanoid pathway begins with phenylalanine, which is converted into cinnamate (trans-cinnamic acid) by the enzyme phenylalanine ammonia lyase (PAL) (Figure ). Cinnamic acid acts as a key intermediate in both the phenylpropanoid and shikimate pathways and serves as a precursor for its hydroxyl derivatives, including p-coumaric, caffeic, ferulic, sinapic, and chlorogenic acids. The biosynthesis of cinnamic acids is facilitated by cinnamate 4-hydroxylase (C4H), which catalyzes the conversion of cinnamate to p-hydroxycinnamate. Subsequently, 4-coumarate:CoA ligase (4CL) converts p-coumarates into their corresponding CoA esters. ,

2.

2

Open in a new tab

A schematic presentation of the phenylpropanoid and flavonoid pathway with emphasis on phenolic compounds present in Vaccinium berries. Red-colored Vaccinium berries contain primarily cyanidin branch anthocyanins whereas both cyanidin and delphinidin branch anthocyanins are present in blue-colored berries. Enzyme abbreviations: PAL, phenylalanine ammonia-lyase; C4H, cinnamate 4-hydroxylase; 4CL, 4-coumaroyl:CoA ligase; CAD, cinnamoyl-alcohol dehydrogenase; CCR, cinnamoyl- CoA reductase; STS, stilbene synthase; CHS, chalcone synthase; CHI, chalcone isomerase; F3H, flavanone 3-hydroxylase; F3′H, flavonoid 3′ hydroxylase; F3′5′H, flavonoid 3′5′ hydroxylase; FLS, flavonol synthase; DFR, dihydroflavonol 4-reductase; LAR, leucoanthocyanidin reductase; ANR, anthocyanidin reductase; ANS, anthocyanidin synthase; UFGT, UDP glucose-flavonoid 3-O-glucosyl transferase; MT, methyltransferase; AT, acyltransferase. Created in BioRender. Jaakola, L. (2026) https://BioRender.com/ap7fdk8.

The specific flavonoid pathway begins with the condensation of one molecule of 4-coumaroyl-CoA and three molecules of malonyl-CoA, resulting in the formation of naringenin chalcone (Figure ). This reaction is catalyzed by chalcone synthase (CHS). Chalcone is then isomerized into (2S)-flavanone by chalcone flavanone isomerase (CHI). From these central intermediates, the pathway branches into various side routes, producing diverse classes of flavonoids. Flavanone 3-hydroxylase (F3H) catalyzes the stereospecific 3β-hydroxylation of (2S)-flavanones to dihydrokaempferol, which serves as a precursor for flavonols, anthocyanins, flavan-3-ols, and proanthocyanidins. The hydroxylation of the B-ring of dihydrokaempferol is carried out by flavonoid 3′-hydroxylase (F3′H) and flavonoid 3′,5′-hydroxylase (F3′5′H), producing dihydroquercetin or dihydromyricetin, which direct the biosynthesis toward the cyanidin or delphinidin branches, respectively. Flavonol synthase (FLS) catalyzes the dehydrogenation of dihydroflavonols into their corresponding flavonols. Competing with FLS, dihydroflavonol 4-reductase (DFR) facilitates the conversion of dihydroflavonols into leucoanthocyanins, which are subsequently transformed into anthocyanidins by anthocyanidin synthase (ANS). Glycosylation of anthocyanidins by UDP-glucose:flavonoid-O-glucosyltransferases (UFGT) results in the formation of anthocyanins, while methyltransferases (MT) are responsible for producing O-methylated anthocyanins such as peonidin, petunidin, and malvidin. , Acylation of anthocyanins is carried out by acyltransferases (ATs), members of which have recently been characterized in blueberry. ,

The monomeric units of proanthocyanidins, flavan-3-ols, are synthesized through two pathways: leucocyanidins are converted into (+)-catechins by leucoanthocyanidin reductase (LAR), while anthocyanidins are converted into (−)-epicatechins by anthocyanidin reductase (ANR) (Figure ). Once synthesized, flavonoids are transported to vacuoles via vesicle trafficking, membrane transporters, and glutathione-S-transferase. The polymerization or oligomerization of proanthocyanidins occurs within the vacuoles. It has been unclear whether this process is enzymatic or nonenzymatic, however, recent studies support a nonenzymatic mechanism in which flavan-3-ol monomers serve as starter units for sequential trapping of extension units via nucleophilic attack. In this current PA polymerization model, LAR is a key enzyme in regulating the ratio of starter and extension units. Activity of both LAR and ANR has been reported in Vaccinium species corresponding to the chemical profiling of flavan-3-ols in these berries. , In bilberry, one ANR gene and three homologues for LAR were identified from the sequence databases, all of which showed different expression profiles during berry ripening and in leaf, stem, and rhizome tissues, indicating diverse roles in PA biosynthesis.

4. Transcriptional Regulation

In transcriptional level, biosynthesis of flavonoids is regulated via coordinated control of the structural enzymes in the pathway by the MYB-bHLH-WD40 (MBW) complex that includes interactions with DNA-binding R2R3MYB transcription factors, MYC-like basic helix loop helix (bHLH), and WD40-repeat proteins (Figure ). , The R2R3MYB is considered as the main component in the MBW complex specifying the target gene activation, even though bHLH partner can also have an role in the activation. The understanding of the transcriptional regulation of flavonoid biosynthesis has primarily been based on the studies performed in model species such as Arabidopsis thaliana, Petunia hybrida, Antirrhinum majus and red fruits of the Rosaceae family, which exhibit simpler anthocyanin profile compared with blue-colored berries. Studies on red grapes (Vitis spp.) and blue-colored Vaccinium berries have, however, advanced our understanding of the transcriptional regulation of the more complicated anthocyanin profiles including the delphinidin branch of the flavonoid pathway, with a particular focus on the role of MYB family members. −

3.

3

Open in a new tab

A schematic presentation on the regulation of the biosynthesis of phenolic compounds in Vaccinium berries focusing on flavonoid group phenolics. Created in BioRender. Jaakola, L. (2026) https://BioRender.com/jb6vs9e.

Research focused on blue-colored Vaccinium berries has demonstrated that MYBA, MYBPA1, MYBPA2 subgroups, and MYB suppressors collaboratively regulate anthocyanin biosynthesis. ,− In bilberry, 18 R2R3MYBs associated with flavonoid biosynthesis regulation have been identified. Among them, the MYBPA1-type transcription factor VmMYBPA1.1 was found to specifically regulate the delphinidin pathway by influencing the expression of the branching point gene VmF3′5′H and other genes involved in delphinidin biosynthesis, resulting in bluish anthocyanin pigments. Further studies revealed that in the early stage of fruit development, MYBPA2.3 activates MYBPA1.1 to promote PA biosynthesis, and at the onset of ripening, MYBA1 activates the MYBPA1.1 promoter to drive anthocyanin biosynthesis in both blueberry (V. corymbosum) and bilberry. Earlier, MYBPA1 transcription factors were generally associated with PA biosynthesis, even if increase in expression at the onset of ripening and anthocyanin production has been detected in blue-colored berries, such as Chinese bayberry (Myrica rubra) and grapevine (Vitis spp.) in addition to Vaccinium berries. ,,− The absence of these transcription factors in blueberry flesh explains its white coloration, in contrast to bilberry, which accumulates anthocyanins in both skin and flesh. In bilberry, altogether 8 PA type MYB transcription factor sequences were identified, representing MYBPA1, MYBPA2, and MYB3 subgroups. Besides the active role of MYBPA1.1 and MYBPA2.2 connected to anthocyanin biosynthesis in ripening berry, the precise function of all others has not yet been fully characterized. Expression of these regulators is higher in the early development of berries and in leaf, stem, and root tissues. It seems that VmMYBPA1.2 has the conventional role of MYBPA1 transcription factors in regulation of proanthocyanidin biosynthesis as overexpression of this gene in Nicotiana benthamiana leaves strongly upregulated NbANR gene.

Concerning other MYB regulators, also MYB5 members contributing to precursors of the flavonoid pathway have been identified in Vaccinium berries in addition to MYBC group repressors. , It has been shown earlier that MYBC-type repressors can provide feedback inhibition in the MBW complexes associated with anthocyanin biosynthesis for fine-tuning the pigment production. In Vaccinium species, precise function of the MYBC group repressors has not yet been studied in detail.

Additionally, the recent characterization of bilberry bHLH transcription factors provides new insights into the interaction between R2R3-MYBs and bHLH partners within the MBW complex. The results suggest that TT8-type VmbHLH2 is the main family member involved in anthocyanin biosynthesis during the bilberry ripening.

Concerning upstream regulators, a number of transcription factors from MADS-box, NAC, SPL, and WRKY families have been shown to regulate fruit development and ripening. Of these, SQUAMOSA MADS-box gene TDR4/FUL1 was shown to have a role in accumulation of anthocyanin during bilberry ripening. Moreover, several ripening regulator candidates have been identified from the recent Vaccinium transcriptomes and genomes. − Recently, transient injection of a blueberry ethylene response factor (ERF) transcription factor, a member of APETALA2/ETHYLENE RESPONSIVE FACTOR (AP2/ERF) family, on blueberry leaves and fruits increased anthocyanin biosynthesis. Similar response was found in transgenic adventitious shoots of the same gene.

Epigenetic regulation including DNA methylation and histone modifications has been shown to have a role in anthocyanin accumulation in fruits. For instance, in apple (Malus domestica) and pear (Pyrus spp.), DNA methylation in the promoter region of the R2R3-MYB transcription factor of the MBW complex was found to negatively regulate anthocyanin biosynthesis. In Vaccinium berries, DNA methylation of the promoter regions of structural genes VcCHS and VcANS was shown to inhibit anthocyanin biosynthesis in blueberry (V. corymbosum).

5. Hormonal Regulation

Plant hormones regulate fruit development and ripening, coordinating physiological and biochemical transitions between different stages. Auxins and gibberellins are involved in the regulation of fruit set and early development, whereas ethylene and abscisic acid (ABA) are the key-regulations of the ripening process. Based on the respiration burst rate and increase in ethylene production or the ABA peak at the onset of ripening, fruits are classified into climacteric or nonclimacteric fruits. Vaccinium berries are generally classified as nonclimacteric fruits, however, evidence on both ethylene and ABA accumulation has been detected at the onset of ripening, which indicates that both of these plant hormones have a role in the ripening process. Watanabe et al. pointed out the controversy over the classification of blueberries (Vaccinium spp.) as climacteric or nonclimacteric fruit because some of the earlier studies have detected increased respiration rate and ethylene production during the maturation period while some other studies did not observe ethylene evolution but only increase in ABA concentration in ripening blueberries. ,, Especially ABA has been found to be connected and to induce ripening-related anthocyanin accumulation in bilberries and blueberries. ,− It has also been shown that red light can stimulate anthocyanin biosynthesis regulated by MBW complex via induction of the ABA biosynthesis. , It seems that the burst of ethylene at the onset of berry ripening increases ABA biosynthesis, leading to increase in anthocyanin biosynthesis. Indeed, in rabbiteye blueberry (V. ashei) fruit, a treatment with ethephon, an ethylene-releasing compound, induced increase in the ABA levels and related anthocyanin biosynthesis.

Sugar-hormone cross talk has been shown to have a role in regulation of anthocyanin biosynthesis, and sugar accumulation has been found to stimulate anthocyanin biosynthesis in different models, such as in Arabidopsis leaves and in detached grape (Vitis spp.) berries or in vitro cultures. In strawberry (Fragaria x ananassa), exogenous sucrose treatments in postharvest fruit increased pelargonidin-derivative anthocyanins and related gene expression. In Vaccinium berries, accumulation of sugars fructose, glucose, and sucrose occurs simultaneously with anthocyanin accumulation during berry ripening. ,− However, Karppinen et al. tested exogenous sucrose and glucose treatments of green bilberry fruits in the same study with exogenous treatment of berries with two different ABA concentrations. ABA effectively induced anthocyanin accumulation in 4 days, whereas no statistical differences were detected in sucrose- or glucose-treated berries compared to controls.

6. Environmental Regulation

Environmental factors have a considerable role in the regulation of secondary metabolism in general and specifically of phenolic compounds. Light and temperature conditions in addition to nutritional status, water balance, diseases, and pest attacks have all been shown to affect the biosynthesis of phenolic compounds. However, interaction of different stressors affects differentially different species and cultivars because of which controlled studies can bring valuable information. Especially, the effect of light and temperature on the metabolism of phenolic compounds has been widely studied in both wild and cultivated Vaccinium species.

Light intensity and quality, in addition to photoperiod have all been shown to affect profile of phenolic compounds in Vaccinium species (Figure ). In many studies, higher light intensity has been shown to increase the content of phenolic compounds. − However, strong interaction has been detected between temperature, and light intensity as bilberries adapted to high altitudes produced lower levels of anthocyanins in field conditions at low altitude in northern Italy, compared to the plants under blue or black selective nets, which offered slightly lower temperature. In separate studies light quality has been shown to markedly affect anthocyanin biosynthesis. ,, Samkumar et al. showed that both red and blue light significantly increase anthocyanin biosynthesis in ripening bilberries, however, the response was different if berries were attached or detached on the mother plant during the light treatments. , In the same study, red light also increased levels of most flavonols in addition to catechin and gallocatechin levels but not as much as anthocyanins. Transcriptome analyses of the samples revealed that the red light signaling induced ABA-regulated anthocyanin biosynthesis. Longer day length has also been shown to increase accumulation of anthocyanins in ripening bilberry. Based on these studies, it seems that anthocyanins, especially delphinidin branch anthocyanins, react more in higher light intensity or increase in red and blue light, compared to other flavonoids or phenolic compounds in Vaccinium berries.

Vaccinium species and cultivars have adapted to different temperature regimes at different growth regions. Southern highbush blueberries can be grown in warmer temperatures, such as in Mediterranean climate region, whereas half-high and lowbush blueberries, as well as cranberries, lingonberries, and bilberries prefer cooler growth temperatures. Both bilberries and lingonberries have been shown to produce higher levels of phenolic compounds in lower temperatures under controlled conditions. , Moreover, many studies have indicated higher anthocyanin levels in bilberries, bog bilberries, and lingonberries growing in higher latitudes characterized with cooler temperature and longer day length, compared to the southern counterparts. ,− Same phenomenon has been detected toward higher altitudes. , Considering the ongoing global warming, it seems that in the future, the warmer climate during the growth season may change the composition and decrease the content of the key phenolic compounds especially of the wild Vaccinium berries adapted to northern climates. Moreover, in the southern areas, the wild Vaccinium species may be restricted even in higher altitudes, reducing the present distribution areas.

In conclusion, a great deal of information has become available on factors affecting the accumulation of phenolic compounds in Vaccinium species. Especially understanding the regulation of the biosynthesis of phenolic compounds in Vaccinium berries has advanced markedly in recent years. Candidates of the MBW complex have been identified and characterized in different species, and functional studies have increased understanding of the transcriptional regulation, especially of blue-colored Vaccinium berries. Many recent Vaccinium genomes and transcriptomes are available for blueberries, cranberries, bilberries, and lingonberries, which are detailedly described by Edger et al. These databases offer further possibilities to deepen understanding of the complex regulation and signaling networks regarding hormonal and environmental regulation. More detailed studies are needed to clarify the interaction of ethylene and ABA during berry ripening and the effect of the interaction on berry quality in different types of Vaccinium berries. Along with a deeper understanding of the environmental and hormonal regulation, it is possible to develop new tools for the selection of suitable cultivars for different climate regions. Improving the health properties and quality of Vaccinium berries can be targeted via the breeding programs but also by paying attention to growth conditions, for instance, by choosing an optimal growth location, optimizing light conditions in tunnel systems, or reflecting shields or shading nets.

Glossary

Abbreviations

AT

acetyltransferase

ABA

abscisic acid

ANR

anthocyanidin reductase

ANS

anthocyanidin synthase

bHLH

MYC-like basic helix loop helix

CHI

chalcone flavanone isomerase

CHS

chalcone synthase

4CL

4-coumarate:CoA ligase

DFR

dihydroflavonol 4-reductase

F3H

flavanone 3-hydroxylase

F3′H

flavonoid 3′-hydroxylase

F3′5′H

flavonoid 3′,5′-hydroxylase

FLS

flavonol synthase

FUL1

FRUITFULL 1

HCA

hydroxycinnamic acid

LAR

leucoanthocyanidin reductase

MYB

DNA-binding R2R3MYB transcription factor

MBW

MYB-bHLH-WD40

MT

methyltransferase

PA

proanthocyanidin

SPL

SQUAMOSA promoter-binding protein like

STS

stilbene synthase

TDR4

tomato FRUITFULL homologue 1

UFGT

UDP-glucose:flavonoid-O-glucosyltransferase

WD40

WD40-repeat proteins

The author declares no competing financial interest.

Published as part of Journal of Agricultural and Food Chemistry special issue “XXXII International Conference on Polyphenols (ICP2025)”.

References

  1. Nestby R., Percival D., Martinussen I., Opstad N., Rohloff J.. The European blueberry (Vaccinium myrtillus L.) and the potential for cultivation. Eur. J. Plant Sci. Biotechnol. 2011;5:5–16. [Google Scholar]
  2. Tundis R., Tenuta M. C., Loizzo M. R., Bonesi M., Finetti F., Trabalzini L., Deguin B.. Vaccinium Species (Ericaceae): From chemical composition to bio-functional activities. Appl. Sci. 2021;11:5655. doi: 10.3390/app11125655. [DOI] [Google Scholar]
  3. Ghosh A., Debnath S. C., Igamberdiev A. U.. Effects of Vaccinium-derived antioxidants on human health: the past, present and future. Front. in Mol. Biosci. 2024;11:1520661. doi: 10.3389/fmolb.2024.1520661. [DOI] [PMC free article] [PubMed] [Google Scholar]
  4. Kalt W., MacKinnon S., McDonald J., Vinqvist M., Craft C., Howell A.. Phenolics of Vaccinium berries and other fruit crops. J. Sci. Food Agric. 2008;88:68–76. doi: 10.1002/jsfa.2991. [DOI] [Google Scholar]
  5. Riihinen K., Jaakola L., Kärenlampi S., Hohtola A.. Organ-specific distribution of phenolic compounds in bilberry (Vaccinium myrtillus) and ’Northblue’ blueberry (V. corymbosum x V. angustifolium) Food Chem. 2008;110:156–160. doi: 10.1016/j.foodchem.2008.01.057. [DOI] [PubMed] [Google Scholar]
  6. Amundsen M., Jaakola L., Aaby K., Martinussen I., Kelanne N., Tuominen S., Laaksonen O., Yang B., Hykkerud A. L.. Effect of ripening temperature on the chemical composition of lingonberries (Vaccinium vitis-idaea L.) of northern and southern origin. Food Res. Int. 2023;167:112738. doi: 10.1016/j.foodres.2023.112738. [DOI] [PubMed] [Google Scholar]
  7. Harborne J. B., Williams C.. Advances in flavonoid research since 1992. Phytochemistry. 2000;55:481–504. doi: 10.1016/S0031-9422(00)00235-1. [DOI] [PubMed] [Google Scholar]
  8. Määttä-Riihinen K., Kamal-Eldin A., Mattila P. H., González-Paramás A., Törrönen R.. Distribution of contents of phenolic compounds in eighteen Scandinavian berry species. J. Agric. Food Chem. 2004;52:4477–4486. doi: 10.1021/jf049595y. [DOI] [PubMed] [Google Scholar]
  9. Wallace T. C., Giusti M. M.. Anthocyanins. Adv. Nutr. 2015;6(5):620–622. doi: 10.3945/an.115.009233. [DOI] [PMC free article] [PubMed] [Google Scholar]
  10. Karppinen K., Zoratti L., Nguyenquynh N., Häggman H., Jaakola L.. (2016). On the developmental and environmental regulation of secondary metabolism in Vaccinium spp. berries. Front. Plant Sci. 2016;7:655. doi: 10.3389/fpls.2016.00655. [DOI] [PMC free article] [PubMed] [Google Scholar]
  11. Wang Y., Fong S. K., Singh A. P., Vorsa N., Johnson-Cicalese J.. (2019). Variation of anthocyanins, proanthocyanidins, flavonols, and organic acids in cultivated and wild diploid blueberry species. HortSci. 2019;54(3):576–585. doi: 10.21273/HORTSCI13491-18. [DOI] [Google Scholar]
  12. Li R., Wang P., Guo Q., Wang Z.. Anthocyanin composition and content of the Vaccinium uliginosum berry. Food Chem. 2011;125(1):116–120. doi: 10.1016/j.foodchem.2010.08.046. [DOI] [Google Scholar]
  13. Zoratti L., Sarala M., Carvalho E., Karppinen K., Martens S., Giongo L., Häggman H., Jaakola L.. Monochromatic light increases anthocyanin content during fruit development in bilberry. BMC Plant Biol. 2014;14:377. doi: 10.1186/s12870-014-0377-1. [DOI] [PMC free article] [PubMed] [Google Scholar]
  14. Jucá M. M., Cysne Filho F. M. S., de Almeida J. C., Mesquita D. d. S., Barriga J. R. d. M., Dias K. C. F., Barbosa T. M., Vasconcelos L. C., Leal L. K. A. M., Ribeiro J. E., Vasconcelos S. M. M.. Flavonoids: biological activities and therapeutic potential. Nat. Prod. Res. 2020;34:692–705. doi: 10.1080/14786419.2018.1493588. [DOI] [PubMed] [Google Scholar]
  15. Ntemiri A., Ghosh T. S., Gheller M. E., Tran T. T. T., Blum J. E., Pellanda P., Vlckova K., Neto M. C., Howell A., Thalacker-Mercer A., O’Toole P. W.. Whole blueberry and isolated polyphenol-rich fractions modulate specific gut microbes in an in vitro colon model and in a pilot study in human consumers. Nutrients. 2020;12(9):2800. doi: 10.3390/nu12092800. [DOI] [PMC free article] [PubMed] [Google Scholar]
  16. Ou J. L. S., Yang D., Liu M. H.. (2020). Effects of Anthocyanins in Composite Meals on Cardiometabolic Outcomes-A Systematic review of randomized controlled feeding trials. Nutrients. 2020;12(12):3781. doi: 10.3390/nu12123781. [DOI] [PMC free article] [PubMed] [Google Scholar]
  17. Suvanto J., Karppinen K., Riihinen K., Jaakola L., Salminen J.-P.. Changes in the proanthocyanidin composition and related gene expression in bilberry (Vaccinium myrtillus L.) tissues. J. Agric. Food Chem. 2020;68:7378–7386. doi: 10.1021/acs.jafc.0c02158. [DOI] [PMC free article] [PubMed] [Google Scholar]
  18. Xie D. Y., Dixon R. A.. Proanthocyanidin biosynthesis – still more questions than answers? Phytochemistry. 2005;66:2127–2144. doi: 10.1016/j.phytochem.2005.01.008. [DOI] [PubMed] [Google Scholar]
  19. Feghali K., Feldman M., La V. D., Santos J., Grenier D.. Cranberry proanthocyanidins: Natural weapons against periodontal diseases. J. Agric. Food Chem. 2012;60(23):5728–5735. doi: 10.1021/jf203304v. [DOI] [PubMed] [Google Scholar]
  20. Weisshaar B., Jenkins G. I.. Phenylpropanoid biosynthesis and its regulation. Current Opin. Plant Biol. 1998;1:251–257. doi: 10.1016/S1369-5266(98)80113-1. [DOI] [PubMed] [Google Scholar]
  21. Vogt T.. Phenylpropanoid biosynthesis. Mol. Plant. 2010;3:2–20. doi: 10.1093/mp/ssp106. [DOI] [PubMed] [Google Scholar]
  22. Jaakola L.. New insights into the regulation of anthocyanin biosynthesis in fruits. Trends Plant Sci. 2013;18:477–483. doi: 10.1016/j.tplants.2013.06.003. [DOI] [PubMed] [Google Scholar]
  23. Dare A. P., Wu C., Carvajal J. I., Nguyen H. M., Günther C. S., Hamiaux C., Bailey S., Deng C., Mengist M. F., Iorizzo M., Foster T. M., Chagné D., Montanari S., Espley R. V.. Haplotyped genome mapping and functional characterization of a blueberry anthocyanin acetyltransferase (AAT) controlling the accumulation of acylated anthocyanins. J. Exp. Bot. 2025;76(6):1607–1624. doi: 10.1093/jxb/erae489. [DOI] [PMC free article] [PubMed] [Google Scholar]
  24. Mengist M. F., Abid M. A., Grace M. H., Seth R., Bassil N., Kay C. D., Dare A. P., Chagné D., Espley R. V., Neilson A., Lila M. A., Ferruzzi M., Iorizzo M.. Identification and functional characterization of BAHD acyltransferases associated with anthocyanin acylation in blueberry. Hortic. Res. 2025;12(5):uhaf041. doi: 10.1093/hr/uhaf041. [DOI] [PMC free article] [PubMed] [Google Scholar]
  25. Zhao J.. Flavonoid transport mechanisms: how to go and with whom. Trends Plant Sci. 2015;20(9):576–585. doi: 10.1016/j.tplants.2015.06.007. [DOI] [PubMed] [Google Scholar]
  26. Yu K., Song Y., Lin J., Dixon R. A.. The complexities of proanthocyanidin biosynthesis and its regulation in plants. Plant Commun. 2023;4(2):100498. doi: 10.1016/j.xplc.2022.100498. [DOI] [PMC free article] [PubMed] [Google Scholar]
  27. Lafferty D. J., Espley R. V., Deng C., Günther C. S., Plunkett B., Turner J., Jaakola L., Karppinen K., Allan A. C., Albert N. W.. Hierarchical regulation of MYBPA1 by anthocyanin and proanthocyanidin related MYB proteins is conserved in Vaccinium species. J. Exp. Bot. 2022;73:1344–1356. doi: 10.1093/jxb/erab460. [DOI] [PubMed] [Google Scholar]
  28. Allan A. C., Espley R. V.. MYBs drive novel consumer traits in fruits and vegetables. Trends Plant Sci. 2018;23:693–705. doi: 10.1016/j.tplants.2018.06.001. [DOI] [PubMed] [Google Scholar]
  29. Matus J. T., Cavallini E., Loyola R., Höll J., Finezzo L., Dal Santo S., Vialet S., Commisso M., Roman F., Schubert A., Alcalde J. A., Bogs J., Ageorges A., Tornielli G. B., Arce-Johnson P.. A group of grapevine MYBA transcription factors located in chromosome 14 control anthocyanin synthesis in vegetative organs with different specificities compared with the berry color locus. Plant J. 2017;91:220–236. doi: 10.1111/tpj.13558. [DOI] [PubMed] [Google Scholar]
  30. Jaakola L., Poole M., Jones M., Kämäräinen-Karppinen T., Koskimäki J. J., Hohtola A., Häggman H., Fraser P., Manning K., King G. J., Thomson H., Seymour G. B.. A SQUAMOSA MADS-box gene involved in the regulation of anthocyanin accumulation in bilberry fruits. Plant Physiol. 2010;153:1619–1629. doi: 10.1104/pp.110.158279. [DOI] [PMC free article] [PubMed] [Google Scholar]
  31. Primetta A. K., Karppinen K., Riihinen K., Jaakola L.. Metabolic and molecular analysis of white mutant Vaccinium berries show down-regulation of MYBPA1-type R2R3MYB regulatory factor. Planta. 2015;242:631–643. doi: 10.1007/s00425-015-2363-8. [DOI] [PubMed] [Google Scholar]
  32. Plunkett B. J., Espley R. V., Dare A. P., Warren B. A. W., Grierson E. R. P., Cordiner S., Turner J. L., Allan A. C., Albert N. W., Davies K. M., Schwinn K. E.. MYBA from blueberry (Vaccinium section Cyanococcus) is a subgroup 6 type R2R3MYB transcription factor that activates anthocyanin production. Front. Plant Sci. 2018;9:1300. doi: 10.3389/fpls.2018.01300. [DOI] [PMC free article] [PubMed] [Google Scholar]
  33. Günther C. S., Dare A. P., McGhie T. K., Deng C., Lafferty D. J., Plunkett B. J., Grierson E. R. P., Turner J. L., Jaakola L., Albert N. W., Espley R. V.. Spatiotemporal modulation of flavonoid metabolism in blueberries. Front. Plant Sci. 2020;11:545. doi: 10.3389/fpls.2020.00545. [DOI] [PMC free article] [PubMed] [Google Scholar]
  34. Karppinen K., Lafferty D. J., Albert N., Mikkola N., McGhie T., Allan A. C., Afzal B. M., Häggman H., Espley R. V., Jaakola L.. MYBA and MYBPA transcription factors co-regulate anthocyanin biosynthesis in blue-coloured berries. New Phytol. 2021;232:1350–1367. doi: 10.1111/nph.17669. [DOI] [PubMed] [Google Scholar]
  35. Lafferty D. J., Espley R. V., Deng C. H., Dare A. P., Günther C. S., Jaakola L., Karppinen K., Boase M. R., Wang L., Luo H., Allan A. C., Albert N. W.. The coordinated action of MYB activators and repressors controls proanthocyanidin and anthocyanin biosynthesis in Vaccinium . Front. Plant Sci. 2022;13:910155. doi: 10.3389/fpls.2022.910155. [DOI] [PMC free article] [PubMed] [Google Scholar]
  36. Shi L., Chen X., Chen W., Zheng Y., Yang Z.. 2018. Comparative transcriptomic analysis of white and red Chinese bayberry (Myrica rubra) fruits reveals flavonoid biosynthesis regulation. Sci. Hortic. 2018;235:9–20. doi: 10.1016/j.scienta.2018.02.076. [DOI] [Google Scholar]
  37. Bogs J., Jaffé F. W., Takos A. M., Walker A. R., Robinson S. P.. 2007. The grapevine transcription factor VvMYBPA1 regulates proanthocyanidin synthesis during fruit development. Plant Physiol. 2007;143:1347–1361. doi: 10.1104/pp.106.093203. [DOI] [PMC free article] [PubMed] [Google Scholar]
  38. Czemmel S., Heppel S. C., Bogs J.. R2R3MYB transcription factors: key regulators of the flavonoid biosynthetic pathway in grapevine. Protoplasma. 2012;249:S109–S118. doi: 10.1007/s00709-012-0380-z. [DOI] [PubMed] [Google Scholar]
  39. Zifkin M., Jin A., Ozga J. A., Zaharia I., Schernthaner J. P., Gesell A., Abrams S. R., Kennedy J. A., Constabel C. P.. (2012). Gene expression and metabolite profiling of developing highbush blueberry fruit indicates transcriptional regulation of flavonoid metabolism and activation of abscisic acid metabolism. Plant Physiol. 2012;158:200–224. doi: 10.1104/pp.111.180950. [DOI] [PMC free article] [PubMed] [Google Scholar]
  40. Albert N. W., Davies K. M., Lewis D. H., Zhang H., Montefiori M., Brendolise C., Boase M. R., Ngo H., Jameson P. E., Schwinn K. E.. A conserved network of transcriptional activators and repressors regulates anthocyanin pigmentation in eudicots. Plant Cell. 2014;26:962–980. doi: 10.1105/tpc.113.122069. [DOI] [PMC free article] [PubMed] [Google Scholar]
  41. Karppinen K., Raami L., Edema H., Ashraf M. F., Salo H., Espley R. V., Jaakola L.. Two bHLH transcription factors regulate anthocyanin biosynthesis in bilberries as a part of the MBW complex. Sci. Rep. 2025;15:29816. doi: 10.1038/s41598-025-15557-w. [DOI] [PMC free article] [PubMed] [Google Scholar]
  42. Bian Y., Ballington J., Raja A., Brouwer C., Reid R., Burke M., Wang X., Rowland L. J., Bassil N., Brown A.. Patterns of simple sequence repeats in cultivated blueberries (Vaccinium section Cyanococcus spp.) and their use in revealing genetic diversity and population structure. Mol. Breed. 2014;34(2):675–689. doi: 10.1007/s11032-014-0066-7. [DOI] [Google Scholar]
  43. 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. GigaScience. 2015;4(1):5. doi: 10.1186/s13742-015-0046-9. [DOI] [PMC free article] [PubMed] [Google Scholar]
  44. Polashock J., Zelzion E., Fajardo D., Zalapa J., Georgi L., Bhattacharya D., Vorsa N.. The American cranberry: first insights into the whole genome of a species adapted to bog habitat. BMC Plant Biol. 2014;14(1):165. doi: 10.1186/1471-2229-14-165. [DOI] [PMC free article] [PubMed] [Google Scholar]
  45. Colle M., Leisner C. P., Wai C. M., Ou S., Bird K. A., Wang J., Wisecaver J. H., Yocca A. E., Alger E. I., Tang H., Xiong Z., Callow P., Ben-Zvi G., Brodt A., Baruch K., Swale T., Shiue L., Song G.-Q., Childs K. L., Schilmiller A., Vorsa N., Buell C. R., VanBuren R., Jiang N., Edger P. P.. Haplotype-phased genome and evolution of phytonutrient pathways of tetraploid blueberry. GigaScience. 2019;8:giz012. doi: 10.1093/gigascience/giz012. [DOI] [PMC free article] [PubMed] [Google Scholar]
  46. Nguyen N., Suokas M., Karppinen K., Vuosku J., Jaakola L., Häggman H.. De novo transcriptomes of wild bilberry – a source of candidate transcription factors in fruit ripening. Sci. Rep. 2018;8:9943. doi: 10.1038/s41598-018-28158-7. [DOI] [PMC free article] [PubMed] [Google Scholar]
  47. Wu C., Deng C., Hilario E., Albert N. W., Lafferty D., Grierson E. R. P., Plunkett B. J., Elborough C., Saei A., Günther C. S., Ireland H., Yocca A., Edger P. P., Jaakola L., Karppinen K., Grande A., Kylli R., Lehtola V. P., Allan A. C., Espley R. V., Chagné D.. A chromosome-scale assembly of the bilberry genome identifies a complex locus controlling berry anthocyanin composition. Mol. Ecol. Resour. 2022;22:345–360. doi: 10.1111/1755-0998.13467. [DOI] [PubMed] [Google Scholar]
  48. Hirabayashi K., Debnath S. C., Owens G. L.. Unveiling the evolutionary history of lingonberry (Vaccinium vitis-idaea L.) through genome sequencing and assembly of European and North American subspecies. G3: Genes Genomes Genet. 2023;14(3):jkad294. doi: 10.1093/g3journal/jkad294. [DOI] [PMC free article] [PubMed] [Google Scholar]
  49. Ma S., Tang Q., Wang X., Chi F., Song Y.. Molecular mechanism of ethylene signaling response gene VcERF061 to promote anthocyanin synthesis in blueberry (Vaccinium corymbosum) BMC Plant Biol. 2025;26:46. doi: 10.1186/s12870-025-07712-2. [DOI] [PMC free article] [PubMed] [Google Scholar]
  50. Fenn M. A., Giovannoni J. J.. Phytohormones in fruit development and maturation. Plant J. 2021;105:446–458. doi: 10.1111/tpj.15112. [DOI] [PubMed] [Google Scholar]
  51. Jaakola L., Karppinen K., Nguyen N., Jokipii-Lukkari S.. Recent progress in understanding the regulation of fruit ripening in bilberry (Vaccinium myrtillus) Acta Hortic. 2025;1440:15–22. doi: 10.17660/ActaHortic.2025.1440.2. [DOI] [Google Scholar]
  52. Watanabe M., Goto R., Murakami M., Komori S., Suzuki A.. Interaction between ethylene and abscisic acid and maturation in highbush blueberry. Hortic. J. 2021;90:14–22. doi: 10.2503/hortj.UTD-210. [DOI] [Google Scholar]
  53. Chung S. W., Yu D. J., Oh H. D., Ahn J. H., Huh J. H., Lee H. J.. Transcriptional regulation of abscisic acid biosynthesis and signal transduction, and anthocyanin biosynthesis in ‘Bluecrop’ highbush blueberry fruit during ripening. PLoS One. 2019;14(7):0220015. doi: 10.1371/journal.pone.0220015. [DOI] [PMC free article] [PubMed] [Google Scholar]
  54. Karppinen K., Hirvelä E., Nevala T., Sipari N., Suokas M., Jaakola L.. Changes in the abscisic acid levels and related gene expression during fruit development and ripening in bilberry (Vaccinium myrtillus L.) Phytochemistry. 2013;95:127–134. doi: 10.1016/j.phytochem.2013.06.023. [DOI] [PubMed] [Google Scholar]
  55. Karppinen K., Tegelberg P., Häggman H., Jaakola L.. Abscisic acid regulates anthocyanin biosynthesis and gene expression associated with cell wall modification in ripening bilberry (Vaccinium myrtillus L.) fruits. Front. Plant Sci. 2018;9:1259. doi: 10.3389/fpls.2018.01259. [DOI] [PMC free article] [PubMed] [Google Scholar]
  56. Nguyen N., Bergmann U., Jaakola L., Häggman H., Jokipii-Lukkari S., Toth K.. (2024) Bilberry metabolomic and proteomic profiling during fruit ripening reveals key-dynamics affecting fruit quality. Physiol. Plant. 2024;176(5):e14534. doi: 10.1111/ppl.14534. [DOI] [PubMed] [Google Scholar]
  57. Samkumar A., Jones D., Karppinen K., Dare A. P., Sipari N., Espley R. V., Martinussen I., Jaakola L.. Red and blue light treatments of ripening bilberry fruits reveal differences in signaling through ABA regulated anthocyanin biosynthesis. Plant, Cell Environ. 2021;44:3227–3245. doi: 10.1111/pce.14158. [DOI] [PubMed] [Google Scholar]
  58. Wang Y. W., Nambeesan S. U.. Ethylene promotes fruit ripening initiation by downregulating photosynthesis, enhancing abscisic acid and suppressing jasmonic acid in blueberry (Vaccinium ashei) BMC Plant Biol. 2024;24:418. doi: 10.1186/s12870-024-05106-4. [DOI] [PMC free article] [PubMed] [Google Scholar]
  59. Espley R., Jaakola L.. The role of environmental stress in fruit pigment production. Plant, Cell Environ. 2023;46:3663–3679. doi: 10.1111/pce.14684. [DOI] [PubMed] [Google Scholar]
  60. Jovančević M., Balijagić J., Menković N., Šavikin K., Zdunić G., Janković T., Dekić-Ivanković M.. Analysis of phenolic compounds in wild populations of bilberry (Vaccinium myrtillus L.) from Montenegro. J. Med. Plants Res. 2011;5:910–914. doi: 10.5897/JMPR.9000080. [DOI] [Google Scholar]
  61. Mikulic-Petkovsek M., Schmitzer V., Slatnar A., Stampar F., Veberic R.. A comparison of fruit quality parameters of wild bilberry (Vaccinium myrtillus L.) growing at different locations. J. Sci. Food Agric. 2015;95:776–785. doi: 10.1002/jsfa.6897. [DOI] [PubMed] [Google Scholar]
  62. Guo X. L., Hu J. B., Wang D. L.. Effect of light intensity on blueberry fruit coloration, anthocyanin synthesis pathway enzyme activity, and gene expression. Russ. J. Plant Physiol. 2023;70:136. doi: 10.1134/s1021443723601064. [DOI] [Google Scholar]
  63. Zoratti L., Jaakola L., Häggman H., Giongo L.. Modification of sunlight radiation through colored photo-selective nets affects anthocyanin profile in Vaccinium spp. berries. PLoS One. 2015;10(8):e0135935. doi: 10.1371/journal.pone.0135935. [DOI] [PMC free article] [PubMed] [Google Scholar]
  64. Zoratti L., Karppinen K., Luengo Escobar A., Häggman H., Jaakola L.. Light-controlled flavonoid biosynthesis in fruits. Front. Plant Sci. 2014;5:00534. doi: 10.3389/fpls.2014.00534. [DOI] [PMC free article] [PubMed] [Google Scholar]
  65. Wei Z., Yang H., Shi J., Duan Y., Wu W., Lyu L., Li W.. Effects of Different Light Wavelengths on fruit quality and gene expression of anthocyanin biosynthesis in Blueberry (Vaccinium corymbosum) Cells. 2023;12:1225. doi: 10.3390/cells12091225. [DOI] [PMC free article] [PubMed] [Google Scholar]
  66. Samkumar A., Karppinen K., McGhie T. K., Espley R. V., Martinussen I., Jaakola L.. Flavonoid biosynthesis is differentially altered in detached and attached ripening bilberries in response to spectral light quality. Front. Plant Sci. 2022;13:969934. doi: 10.3389/fpls.2022.969934. [DOI] [PMC free article] [PubMed] [Google Scholar]
  67. Uleberg E., Rohloff J., Jaakola L., Trost K., Junttila O., Häggman H., Martinussen I.. Effects of temperature and photoperiod on yield and chemical composition of northern and southern clones of bilberry (Vaccinium myrtillus L) J. Agric. Food Chem. 2012;60:10406–10414. doi: 10.1021/jf302924m. [DOI] [PubMed] [Google Scholar]
  68. Lätti A. K., Riihinen K. R., Kainulainen P. S.. Analysis of anthocyanin variation in wild populations of bilberry (Vaccinium myrtillus L.) in Finland. J. Agric. Food Chem. 2008;56:190–196. doi: 10.1021/jf072857m. [DOI] [PubMed] [Google Scholar]
  69. Åkerstöm A., Jaakola L., Bång U., Jäderlund A.. (2010) Effects of latitude-related factors and geographical origin on anthocyanidin concentrations in fruits of Vaccinium myrtillus L. (bilberries) J. Sci. Food Agric. 2010;58:11939–11945. doi: 10.1021/jf102407n. [DOI] [PubMed] [Google Scholar]
  70. Lätti A. K., Jaakola L., Riihinen K. R., Kainulainen P. S.. (2010) Anthocyanin and flavonol variation in bog bilberries (Vaccinium uliginosum L.) in Finland. J. Agric. Food Chem. 2010;58:427–433. doi: 10.1021/jf903033m. [DOI] [PubMed] [Google Scholar]
  71. Zoratti L., Jaakola L., Häggman H., Giongo L.. Anthocyanin profile in berries of wild and cultivated Vaccinium spp. along altitudinal gradients in Alps. J. Agric. Food Chem. 2015;63:8641–8650. doi: 10.1021/acs.jafc.5b02833. [DOI] [PubMed] [Google Scholar]
  72. Zeng Q., Dong G., Tian L., Wu H., Ren Y., Tamir G., Huang W., Yu H.. High Altitude is beneficial for antioxidant components and sweetness accumulation of Rabbiteye Blueberry. Front. Plant Sci. 2020;11:573531. doi: 10.3389/fpls.2020.573531. [DOI] [PMC free article] [PubMed] [Google Scholar]
  73. Cui Q., Hu S., Chu R., Chen R., Taheri A., Yang F., Li X., He X., Zheng L., Zhou M., Liang Z., Yang D.. How DNA Methylation Regulates Plant Natural Product Biosynthesis: From Epigenetics to Secondary Metabolism. Plant, Cell Environ. 2026;49:1295–1308. doi: 10.1111/pce.70292. [DOI] [PubMed] [Google Scholar]
  74. Yu L., Zhou Y., Zhang Y., Liu W., Li Y., Lu M., Fan C., Shao N., Yang L., Guo W.. DNA methylation balance is involved in anthocyanin accumulation during Vaccinium corymbosum fruit ripening. J. Berry Res. 2020;10:651–663. doi: 10.3233/JBR-200553. [DOI] [Google Scholar]
  75. Zhao T., Li Q., Yan T., Yu B., Wang Q., Wang D.. Sugar and anthocyanins: A scientific exploration of sweet signals and natural pigments. Plant Sci. 2025;353:112409. doi: 10.1016/j.plantsci.2025.112409. [DOI] [PubMed] [Google Scholar]
  76. Li D., Zhang X., Xu Y., Li L., Aghdam M. S., Luo Z.. Effect of exogenous sucrose on anthocyanin synthesis in postharvest strawberry fruit. Food Chem. 2019;289:112–120. doi: 10.1016/j.foodchem.2019.03.042. [DOI] [PubMed] [Google Scholar]
  77. Samkumar A., Karppinen K., Dhakal B., Martinussen I., Jaakola L.. Insight into sugar metabolism during bilberry (Vaccinium myrtillus L.) fruit development. Physiol. Plant. 2022;174:e13657. doi: 10.1111/ppl.13657. [DOI] [PMC free article] [PubMed] [Google Scholar]
  78. Dare A. P., Günther C. S., Grey A. C., Guo G., Demarais N. J., Cordiner S., McGhie T. K., Boldingh H., Hunt M., Deng C., Karppinen K., Jaakola L., Espley R. V.. Resolving the developmental distribution patterns of polyphenols and related primary metabolites in bilberry (Vaccinium myrtillus) fruit. Food Chem. 2022;374:131703. doi: 10.1016/j.foodchem.2021.131703. [DOI] [PubMed] [Google Scholar]
  79. 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) Sci. Rep. 2024;14:24964. doi: 10.1038/s41598-024-74929-w. [DOI] [PMC free article] [PubMed] [Google Scholar]
  80. Edger P. P., Iorizzo M., Bassil N. V., Benevenuto J., Ferrão L. F. V., Giongo L., Hummer K., Lawas L. M. F., Leisner C. P., Li C.. et al. There and back again; historical perspective and future directions for Vaccinium breeding and research studies. Hortic. Res. 2022;9:uhac083. doi: 10.1093/hr/uhac083. [DOI] [PMC free article] [PubMed] [Google Scholar]

Articles from Journal of Agricultural and Food Chemistry are provided here courtesy of American Chemical Society

RESOURCES