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Various effects of monochromatic light wavelengths on anthocyanin biosynthesis have also been reported in other species. For example, in turnip hypocotyls, far-red light had the most prominent effect on anthocyanin biosynthesis, comparable with the amount reached under sunlight [ 26 ]. In Gerbera , anthocyanin accumulation in flowers was particularly stimulated by blue light [ 21 ].

Blue light has been found to significantly increase the biosynthesis of anthocyanins also in fruit species, such as strawberries [ 25 ] and grape fruits [ 22 ],[ 23 ], while in cranberry fruits, red and far-red light increased the anthocyanin accumulation over white light [ 20 ]. A possible explanation of the present results can be found from the gene expression analyses of flavonoid pathway genes. Moreover, in earlier studies it has been shown that flavonoid pathway genes, for instance CHS , can have a diurnal rhythm [ 33 ],[ 34 ].

This is one factor that can have affected the variation in the gene expression results between the different time points. On the contrary, the expression of VmANS , which is the key gene in the biosynthesis of anthocyanins, shows a clear increasing trend under monochromatic light treatments, while white light and dark treatment does not have influence.

According to Jaakola et al. However, the early stages of berry development appeared to be reactive to the light treatments in the present study. Monochromatic light treatments affected the accumulation of anthocyanins by increasing the expression of VmANS already at this early stage of berry development. Considering that in northern latitudes, summer nights are characterized by long twilight with high ratios of blue and far-red light [ 35 ], the present study emphasizes that northern light environment promote the accumulation of anthocyanins in bilberry already at the early stages of fruit ripening, by inducing qualitative and quantitative changes in anthocyanin content of ripe fruits.

We showed that the treatment of bilberry plants under monochromatic light wavelengths of the visible light spectrum, for even short times during the ripening period of the fruits, is enough to induce a significant increase in the anthocyanin content in ripe fruits. Moreover, the quality of light affected particularly the biosynthesis of delphinidin glycosides. Our results indicate that the spectral composition of light regulates the accumulation of anthocyanins in fruits, showing an interaction between the flavonoid biosynthetic pathway and the composition of the light spectrum received by the plant.

After pollination, berries take usually six to seven weeks to ripe in natural stands of Finland. Bilberry fruit ripening stages were identified according to Jaakola et al. The plants irradiated under blue light received a photon fluence rate of 8. Plants exposed to white light were considered as a positive control. A set of plants kept in total darkness was considered as negative control.

Pools of bilberry plants from each location I-III , were used for the treatments.

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The berry developmental stage 2 was selected for the experiments based on preliminary analyses data not shown which indicated stage 2 to be the most reactive one, among all the bilberry fruit ripening stages, in the expression of flavonoid pathway genes in response to the light illumination. The light treatments did not affect the process of ripening of the berries. After filtering 0. Flavonols, flavanones, hydroxycinnamic acids, proanthocyanidins and stilbenes were analysed as described in Vrhovsek et al.

Chromatography, mass spectrometry conditions and multiple reaction monitoring MRM transitions can be found in the referred literature. Quantification was made by external calibration curves, injecting authentic standards of each of the detected compounds at different concentrations. For some of the compounds, there were no standards available, but they could be tentatively identified on the basis of their MRM transitions and the relative retention time, in respect to known compounds and considering previous results [ 37 ]. For example, standards of the galactoside derivatives of cyanidin and peonidin were available, and these compounds seem to elute before but closely to the respective glucoside derivatives peaks 1, 2 and 22, 23 in Additional file 3.

As such, the peak eluting 0. For quantification, external calibration curves were prepared by injecting authentic standards of each compound at different concentrations.

Monochromatic light increases anthocyanin content during fruit development in bilberry

In case the authentic standard was not available, the anthocyanins were quantified relative to malvidin O -glucose, using the malvidin O -glucose calibration curve Additional file 3. A supplementary study was conducted in order to study if the increased amount of anthocyanins was related to the gene expression of flavonoid pathway genes in bilberries Additional file 1.

The RNA was isolated according to the method of Jaakola et al. RNA extraction and further gene expression analyses was repeated twice for each set of plants. The primers used for the amplification are listed in Additional file 4. VmACT gene Additional file 4 was used as a reference gene for relative quantification. Steyn WJ: Prevalence and functions of anthocyanins in fruits. Anthocyanins: Biosynthesis, Functions, and Applications.


Dai J, Mumper RJ: Plant phenolics: extraction, analysis and their antioxidant and anticancer properties. Int J Mol Sci. J Agric Food Chem. Acta Agric Scand B. Plant Physiol. Funct Plant Biol. Koes R, Verweij W, Quattrocchio F: Flavonoids: a colorful model for the regulation and evolution of biochemical pathways. Trends Plant Sci.

Lillo C, Lea US, Ruoff P: Nutrient depletion as a key factor for manipulating gene expression and product formation in different branches of the flavonoid pathway. Plant Cell Env. Jaakola L, Hohtola A: Effect of latitude on flavonoid biosynthesis in plants. Martz F, Jaakola L, Julkunen-Tiitto R, Stark S: Phenolic composition and antioxidant capacity of bilberry Vaccinium myrtillus leaves in northern europe following foliar development and along environmental gradients. J Chem Ecol. Front Plant Sci. Nat Rev Gens. Plant Molec Biol Rep.

J Plant Physiol. Plant Growth Regul. Li Q, Kubota C: Effects of supplemental light quality on growth and phytochemicals of baby leaf lettuce. Env Exp Bot. J Plant Res. J Exp Bot. Food Chem. Du QJG, Winterhalter P: Isolation of two anthocyanin sambubiosides from bilberry Vaccinium myrtillus by high-speed counter-current chromatography. J Chromatogr A. Int J Food Prop. Progr Bot Belgian J Botany. Tian QG, Giusti MM, Stoner GD, Schwartz SJ: Screening for anthocyanins using high-performance liquid chromatography coupled to electrospray ionization tandem mass spectrometry with precursor-ion analysis, product-ion analysis, common-neutral-loss analysis, and selected reaction monitoring.


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Our special thanks to Matti Rauman, for his professionalism and for the great contribution to this project by setting up the light systems. Correspondence to Laura Jaakola. LZ performed most of the experimental work together with interpretation of data, was involved in the design of the work, and most of writing and editing; MS contributed in performing the experiment and gene expression analyses; EC gave support with the metabolic analyses and contribution to the interpretation of the data; KK, SM and LG gave contribution with the interpretation of the data; LJ and HH provided contribution to the conception and the design of the work.

All authors attended to the writing of the manuscript and read and approved the final manuscript. Reprints and Permissions. Search all BMC articles Search.

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Abstract Background Light is one of the most significant environmental factors affecting to the accumulation of flavonoids in fruits. Results All the three monochromatic light treatments had significant positive effect on the accumulation of total anthocyanins in ripe fruits compared to treatment with white light or plants kept in darkness. Great condition for a used book! Minimal wear. Skip to main content. Email to friends Share on Facebook - opens in a new window or tab Share on Twitter - opens in a new window or tab Share on Pinterest - opens in a new window or tab.

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