Effects of Abiotic Factors on Phenolic Compounds in the Grape Berry – A Review

E.H. Blancquaert, A. Oberholster, J.M. Ricardo-da-Silva, A.J. Deloire


Grape berry phenolic compounds are widely described in literature. Phenolics can be divided into two main groups: flavonoids and non-flavonoids, of which the flavonoids are the most important. The two bestknown groups of flavonoids are the anthocyanins and condensed tannins (also called proanthocyanidins).  Anthocyanins are responsible for the red colour in grapes. The condensed tannins (proanthocyanidins) are responsible for some major wine sensorial properties (astringency, browning, and turbidity) and are
involved in the wine ageing processes. This review summarises flavonoid synthesis in the grape berry and the impact of environmental factors on the accumulation rate during ripening of each of the flavonoids.  The impact of the accumulated flavonoids in grapes and the resulting impact on the sensorial aspects of the wine are also discussed.

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Adams, D., 2006. Phenolics and ripening in grape berries. Am. J. Enol. Vitic. 57, 249–256.

Ali, K., Maltese, F, Choi, Y.H. & Verpoorte, R. 2010. Metabolic constituents of grapevine and grape-derived products. Phytochem Rev. 9, 357–378.

Arnold, R.A., Noble, A.C. & Singleton, V.L., 1980. Bitterness and astringency of phenolic fractions in wine. J. Agric. Food Chem. 28, 686–680.

Barnes, P.W., Flint, S.D. & Caldwell, M.M., 1990. Morphological responses of crop and weed species of different growth forms to ultraviolet-B radiation. Am. J. Bot. 77, 1354–1360.

Bergqvist, J., Dokoozlian, N. & Ebisuda, N. 2001. Sunlight exposure and temperature effects on berry growth and composition of Cabernet Sauvignon and Grenache in the central San Joaquin Valley of California. Am. J. Enol. Vitic. 5, 1–7.

Berli, F.J., Moreno, D., Piccoli, P., Hespanhol-Viana, L., Silva, M.F., Bressan-Smith, R., Cavagnaro, J.B. & Bottini, R., 2010. Abscisic acid is involved in the response of grape (Vitis vinifera L.) cv. Malbec leaf tissues to ultraviolet-B radiation by enhancing ultraviolet-absorbing compounds, antioxidant enzymes and membrane sterols. Plant Cell Environ. 33, 1–10.

Bogs, J., Downey, M.O, Harvey, J.S., Ashton, A.R., Tanner, G.J. & Robinson, S.P., 2005. Proanthocyanidin synthesis and expression of genes encoding leucoanthocyanidin reductase and anthocyanidin reductase in developing grape berries and grapevine leaves. Plant Physiol. 139, 652–663.

Bogs, J., Ebadi, A., McDavid, D. & Robinson, S.P., 2006. Identification of the flavonoid hydroxylases from grapevine and their regulation during fruit development. Plant Physiol. 140, 279–291.

Bornman, J.F. & Vogelmann, T.C., 1991. Effect of UV-B radiation on leaf optical properties measured with fibre optics. J. Exp. Bot. 42, 547–554.

Bucchetti, B., Matthews M.A., Falginella, L., Peterlunger, E. & Castellarin, S.D., 2011. Effect of water deficit on Merlot grape tannins and anthocyanins across four seasons. Sci Hortic. 128, 297–305.

Buttrose, M.S., Hale, C.R. & Kliewer, W.M., 1971. Effect of temperature on the composition of ‘Cabernet Sauvignon’ berries. Am. J. Enol. Vitic. 22, 71–75.

Castellarin, S.D., Di Gaspero, G., Marconi, R., Nonis, A., Peterlunger, E., Paillard, S., Adam-Blondon, A.F. & Testolin, R., 2006. Colour variation in red grapevines (Vitis vinifera L.): Genomic organization, expression of flavonoid 3’-hydroxylase, flavonoid 3’, 5’-hydroxylase genes and related metabolite profiling of red cyanidin-/blue delphinidin-based anthocyanins in berry skin. BMC Genomics, 7, 12, 1–17.

Castellarin, S.D, Mathews, M.A., Di Gaspero, G. & Gambetta, G.A., 2007. Water deficits accelerate ripening and induce changes in gene expression regulating flavonoid biosynthesis in grape berries. Planta 227, 101–112.

Chalmers, D.J. & Faragher, J.D., 1977. Regulation of anthocyanin synthesis in apple skin. III: Involvement of phenylalanine ammonia-lyase. Austral. J. Plant Physiol. 4:133–141.

Cheynier, V. & Rigaud, J., 1986. HPLC separation and characterisation of flavonols in the skin of Vitis vinifera var. Cinsault. Am. J. Enol. Vitic. 37, 248–252.

Cheynier, V., Fulcrand, H., Sarni, P. & Moutounet, M. 1997. Reactivity of phenolic compounds in wine: diversity of mechanisms and resulting products. In: Vino Analytica Scientia. Bordeaux. pp.143–154.

Cheynier, V., Fulcrand, H., Brossaud, F., Asselin, C. & Mountounet, M., 1998. Phenolic composition as related to red wine flavour. In: Waterhouse, A. & Ebeler, S. (eds.). Wine flavour. ACS Symposium Series 174, pp. 124–141.

Cheynier, V., Dueñas-Paton, M., Salas, E., Maury, C, Jean-Marc Souquet, J., Sarni-Manchado, P. & Fulcrand, H., 2006. Structure and properties of wine pigments and tannins. Am. J. Enol. Vitic. 57, 298–305.

Chira, K., Schmauch, G., Saucier, C., Fabre, S. & Teissedre, P., 2008. Grape variety effect on proanthocyanidin composition and sensory perception of skin and seed tannin extracts from Bordeaux wine grapes (Cabernet Sauvignon and Merlot) for two consecutive vintages (2006 and 2007). J. Agric. Food Chem. 57, 545–553.

Chorti, E., Guidoni, S., Ferrandino, A. & Novello, V., 2010. Effect of different cluster sunlight exposure levels on ripening and anthocyanin accumulation in Nebbiolo grapes. Am. J. Enol. Vitic. 61, 23–29.

Cohen, S.D., Tarara, J.M. & Kennedy, J.A., 2008. Assessing the impact of temperature on grape phenolic metabolism. Anal. Chim. Acta 621, 57–67.

Cohen, S.D. & Kennedy, J.A. 2010. Plant metabolism and the environment: Implications for managing phenolics. Crit. Rev. Food Sci. Nutr. 50, 620–643.

Cohen, S.D., Tarara, J.M., Gambetta, G.A., Matthews, M.A. & Kennedy, J.A. 2012. Impact of diurnal temperature variation on grape berry development, proanthocyanidin accumulation, and the expression of flavonoid pathway genes. J. Exp. Bot. 1–11.

Conde, C., Silva, P., Fontes, N., Dias, A.C.P., Tavares, R.M., Sousa, M.J., Agasse, A., Delrot, S. & Gerós, H., 2007. Biochemical changes throughout grape berry development and fruit and wine quality. In: Global Science Books, Food 1, 1, pp.1–22.

Coombe, B.G., 1987. Distribution of solutes within the developing grape berry in relation to its morphology. Am. J. Enol. Vitic.38, 120-127.

Cortell, J.M., Halbleib, M., Gallagher, A.V., Righetti, T.L. & Kennedy, J.A., 2005. Influence of vine vigor on grape (Vitis vinifera L. cv Pinot noir) and wine proanthocyanidins. J. Agric. Food Chem. 53, 5798–5808.

Crippen, D.D.J. & Morrison, J.C., 1986. The effects of sun exposure on the phenolic content of Cabernet Sauvignon berries during development. Am. J. Enol. Vitic. 37, 243–247.

Czochanska, Z., Foo, L.Y. & Porter, L.J. 1979. Compositional changes in lower molecular weight flavans during grape maturation. Phytochemistry 18, 1819–1822.

Davies, K.M. & Schwinn, K.E., 2006. Molecular biology and biotechnology of flavonoid biosynthesis. In: Flavonoids: Chemistry, biochemistry and applications. CRC Press, Boca Raton FL, pp. 143–218.

De Freitas, V.A.P. & Glories, Y. 1999. Concentration and compositional changes of procyanidins in grape seeds and skins of white Vitis vinifera varieties. J. Sci. Food Agric. 79, 1601–1606.

Deloire, A., Carbonneau, A., Wang, Z. & Ojeda, H., 2004 Vine and water, a short review, J. Int. Sci. Vigne vin. 38, 1, 1–13.

Dewick, M. & Haslam, E., 1969. The biosynthesis of gallic acid in higher plants. Biochem J. 113, 537–542.

Dokoozlian, N.K. & Kliewer, W.M., 1996. Influence of light on grape berry growth and composition varies during fruit development. J. Am. Soc. Hort. Sci. 121, 869–874.

Dong, YH., Beuning L., Davies, K., Mitra, D., Morris, B. & Kootstra, A., 1998. Expression of pigmentation genes and photo-regulation of anthocyanin biosynthesis in developing Royal Gala apple flowers. Aust. J. Plant Physiol. 25, 245–252.

Downey, M.O., Harvey, J.S. & Robinson, S.P., 2003a. Analysis of tannins in seeds and skins of Shiraz grapes throughout berry development. Aust. J. Grape Wine Res. 9, 15–27.

Downey, M.O., Harvey, J.S. & Robinson, S.P., 2003b. Synthesis of flavonols and expression of flavonol synthase genes in the developing grape berries of Shiraz and Chardonnay (Vitis vinifera L.). Aust. J. Grape Wine Res. 9, 1101–21.

Downey, M.O., Harvey, J.S. & Robinson, S.P., 2004. The effect of bunch shading on berry development and flavonoid accumulation in Shiraz grapes. Aust. J. Grape Wine Res. 10, 55–73.

Downey, M.O., Dokoozlian, N.K. & Krstic, M.P., 2006. Cultural practice and environmental impacts on the flavonoid composition of grapes and wine: A review of recent research. Am. J. Enol. Vitic. 57, 257–268.

Ebadi, A., May, P., Sedgley, M. & Coombe, B.G., 1995. Fruit set on small Chardonnay and Shiraz vines grown under varying temperature regimes between budburst and flowering. Aust. J. Grape Wine Res. 1, 1–10.

Gawel, R., 1998. Red wine astringency: a review. Aust. J. Grape Wine Res. 4, 74–95.

Gehrke, C., Johansson, U., Gwynn-Jones, D., Björn, L.O., Callaghan, T.V. & Lee, J.A., 1996. Effects of enhanced ultraviolet-B on terrestrial subartic ecosystems and implications for interactions with increased CO2 . Ecol Bull. 45, 192–203.

Gladstones, J., 1992. Viticulture and Environment. Winetitles, Adelaide.

Gregan, S.M., Wargent, J.J., Liu, L., Shinkle, J., Hofmann, Winefield, C., Trought, M. & Jordan, B., 2012. Effects of solar ultraviolet radiation and canopy manipulation on the biochemical composition of Sauvignon Blanc grapes. Austr. J. Grape Wine Res. 18, 227–238.

Habertson, F. & Adams, J. A., 2002. Tannins in skins and seeds of Cabernet Sauvignon, Syrah and Pinot noir berries during ripening. Am. J. Enol. Vitic. 53, 54–59.

Haselgrove, L., Botting, D., van Heeswijck, R., Høj, P.B., Dry, P.R., Ford, C. & Iland, P.G., 2000. Canopy microclimate and berry composition: The effect of bunch exposure on the phenolic composition of Vitis vinifera L cv. Shiraz grape berries. Aust. J. Grape Wine Res. 6, 141–149.

Haslam, E., 1998. Practical polyphenolics: from structure to molecular recognition and physiological action, Cambridge, University Press, UK.

Hawker, J.S. 1982. Effect of temperature on lipid, starch and enzymes of starch metabolism in grape, tomato and broad bean leaves. Phytochemistry. 21, 33–36.

Heller, W. & Forkmann, G., 1988. Biosynthesis. In: The Flavonoids: Advances in Research since 1980. Harborne J.B. (ed.).Chapman and Hall, London. pp. 399–425.

Hollósy, F., 2002. Effects of ultraviolet radiation on plant cells. Micron 33, 179–197.

Hunter, J.J., De Villiers, O.T. & Watts, J.E., 1991. The effect of partial defoliation on quality characteristics of Vitis vinifera L. cv. Cabernet Sauvignon grapes. II. skin color, skin sugar, and wine quality

Am. J. Enol. Vitic. 42, 13–18.

Jackson, D.I. & Lombard, P.B., 1993. Environment and management practices affecting grape composition and wine quality – A review. Am. J. Enol. Vitic. 44, 409–430.

Jansen, M.A.K., Gaba, V. & Greenberg, B.M., 1998. Higher plants and UV-B radiation: balancing damage, repair and acclimation. Trends Plant Sci. 3, 131–135.

Jordan, B.R., 1996. The effects of ultraviolet-B radiation on plants: a molecular perspective. In: Advances in Botanical Research 2 vol. 22 (ed. J.A. Callow). Academic Press Ltd, pp. 97–162.

Jordão, A.M., Ricardo-da-Silva, J.M. & Laureano, O., 2001. Evolution of catechins and oligomeric procyanidins during grape maturation of Castelão Francês and Touriga Francesa. Am. J. Enol. Vitic. 52, 230–234.

Katsnelson, A., 2015. From the tongue to the brain. A body of research by Charles Zuker explains how we distinguish bitter from sweet, salty from sour—and why we should care. Accessed: 1 June 2015. (http://www.columbiamedicinemagazine.org/features/spring-2015/tongue-brain).

Kennedy, J.A., Matthews, M.M. & Waterhouse, A.L., 2000a. Changes in grape seed polyphenols during fruit ripening. Phytochemistry 55, 77–85.

Kennedy, J.A. Troup, G.J., Pilbrow, J.R., Hutton, D.R., Hewitt, D., Hunter, C.R., Ristic, R., Iland, P.G. & Jones, G.P., 2000b. Development of seed polyphenols in berries from Vitis vinifera L. cv. Shiraz. Aust. J. Grape Wine Res. 6, 3, 244–254

Kennedy, J.A., Hayasaka, Y., Vidal, S., Waters, E.J. & Jones, G.P. 2001. Composition of grape skin proanthocyanidins at different stages of berry development. J. Agric. Food Chem. 49, 5348–5355.

Kennedy, J.A., 2002a. Understanding grape berry development. Practical Winery. Available: http://www.practicalwinery.com?JulyAugust02/julaug02p14.htm. (Accessed: 21/11/2012).

Kennedy, J.A., Matthews, M.A. & Waterhouse, A.L., 2002b. Effect of maturity and vine water status on grape skin and wine flavonoids. Am. J. Enol. Vitic. 53, 268–274.

Kliewer, W.M. & Antcliff, A.J., 1970. Influence of defoliation, lea darkening, and cluster shading on the growth and composition of ‘Sultana’ grapes. Am. J. Enol. Vitic. 21, 26–36.

Kliewer, W.M. & R.E. Torres., 1972. Effect of controlled day and night temperatures on grape coloration. Am. J. Enol. Vitic. 23, 71–77.

Kliewer, W.M., 1977. Influence of temperature, solar radiation and nitrogen on coloration and composition of Emporer grapes. . Am. J. Enol. Vitic. 28, 96-103.

Kolb, C.A., Käser, M.A., Kopecký, J, Zotz, G, Riederer, M. & Pfündel, E.E., 2001. Effects of natural intensities of visible and ultraviolet radiation on epidermal ultraviolet screening and photosynthesis in grape leaves. Plant Physiol. 127, 863–875.

Kolb, C.A., Kopecký, J Riederer, M. & Pfündel, E.E., 2003. UV screening by phenolics in berries of grapevine (Vitis vinifera). Funct Plant Biol. 30, 1177–1186.

Koundouras, S., Marinos, V., Gkoulioti, A., Kotseridis, Y. & Van Leeuwen, C., 2006. Influence of vineyard location and vine water status on fruit maturation of non-irrigated cv. Agiorgitiko (Vitis vinifera L.). Effects on wine phenolic and aroma components. J. Agric. Food Chem. 54, 5077–5086.

Koyama, K., Ikeda, H., Poudel, P.R. & Goto-Yamamoto, N., 2012. Light quality affects flavonoid biosynthesis in young berries of Cabernet Sauvignon grape. Phytochemistry 78, 54–64.

Labarbe, B., Cheynier, V., Borssaud, F., Souquet, J.M. & Moutounet, M., 1999. Quantitative fractionation of grape proanthocyanidins according to their degree of polymerisation. J. Agric. Food Chem. 2719–2723.

Lancaster, J.E. 1992. Regulation of skin color in apples. CRC Crit. Rev. Plant Sci. 10, 487–502.

Lea, A.G.H. & Arnold, G.M., 1978. The phenolics of ciders: bitterness and astringency. J. Sci. Food Agric. 29, 478–483.

Lee, C.Y. & Jaworski, A.W., 1987. Phenolic compounds in white grapes grown in New York. Am. J. Enol. Vitic. 38, 2772–2781.

Mané, C., Souquet, J.M., Ollé, D., Verriès, C., Véran, F., Mazerolles, G., Cheynier, V. & Fulcrand, H., 2007. Optimization of simultaneous flavanol, phenolic acid and anthocyanin extraction from grapes using an experimental design: application to the characterization of Champagne grape varieties. J. Agric. Food Chem. 55, 7224–7233.

Martínez-Lüscher, J., Chen, C.C.L., Brillante, L & Kurtural, S.H., 2017. Partial Solar Radiation Exclusion with Color Shade Nets Reduces the Degradation of Organic Acids and Flavonoids of Grape Berry (Vitis vinifera L.). J. Agric. Food Chem. 65, 49, 10693–10702.

Mattivi, F., Guzzon, R., Vrhovsek, U., Steffanini, M. & Velasco, R., 2006. Metabolite profiling of grape: flavonols and anthocyanins. J. Agric. Food Chem. 54, 7692–7702.

McRae, J. & Kennedy, J.A., 2011. Wine and grape tannin interactions with salivary proteins and their impact on astringency: A review of current research. Molecules 16, 2348–2364.

Mori, K., Sugaya, S. & Gemma, H., 2005. Decreased anthocyanin biosynthesis in grape berries grown under elevated night temperature condition. Sci. Hortic. 105, 319–330.

Mori, K., Goto-Yamamoto, N., Kitayama, M. & Hashizume, K., 2007. Loss of anthocyanins in red-wine grape under high temperature. J. Exp. Bot. 58, 1935–1945.

Morrison, J. C. & Noble, A.C., 1990. The effects of leaf and cluster shading on the composition of Cabernet Sauvignon grapes and on fruit and wine sensory properties. Am. J. Enol. Vitic. 41, 193–200.

Moutounet, M., Rigaud, J., Souquet, J. & Cheynier, V. 1996. Caractérisation structurale des tanins de la baie de raisin. Quelques exemples de l’incidence du cépage, du terroir et du mode de conduite de la vigne. Bulletin OIV, pp. 433–443.

Oberholster, A., Francis, I.L., Iland, P.G. & Waters, E.J., 2009. Mouthfeel of white wines made with and without pomace contact and added anthocyanins. Aust. J. Grape Wine Res.15, 59–69.

Ojeda, H., Deloire, A. & Carbonneau, A., 2001. Influence of water deficits on grape berry growth. Vitis, 40, 141–145.

Ojeda, H., Andary, C., Kraeva, E., Carbonneau, A. & Deloire, A. 2002. Influence of pre- and postvéraison water deficit on synthesis and concentration of skin phenolic compounds during berry growth of Vitis vinifera cv. Syrah. Am. J. Enol. Vitic 53, 261–267.

OIV (Organisation Internationale de la Vigne et du Vin), 2013. Global Economic Survey, http://www.google.co.za/url?url=http://www.oiv.int/oiv/files/EN_point%2520de%2520conj.pdf (Accessed: 02 December 2014).

Ó-Marques, J., Reguinga, R., Laureano, O. & Ricardo-da-Silva, J.M., 2005. Alterações ao longo da maturação nos taninos condensados da grainha, película e polpa: influência da monda de cachos. Cienc Tec Vitivinic. 20, 35–52.

Petrie, P.R., Cooley N.M. & Clingeleffer, P.R., 2004. The effect of post-veraison water deficit on yield components and maturation of irrigated Shiraz (Vitis vinifera L.) in the current and following season. Aust. J. Grape Wine Res. 10, 203–215.

Peleg, H., Gacon, K., Schlich, P. & Noble, A.C., 1999. Bitterness and astringency of flavan-3-ol monomers, dimers and trimers. J. Sci. Food Agric. 79, 1123–1128.

Piretti, M.V., Ghedini, M., & Serrazanetti, G., 1976. Isolation and identification of the polyphenolic and terpenoid constituents of Vitis vinifera. v. Trebbiano variety. Ann Chim-Rome, 66, 429–437.

Price, S.F., Breen, P.J, Valladao, M. & Watson, B.T., 1995. Cluster sun exposure and quercetin in Pinot noir grapes and wine. Am. J. Enol. Vitic. 46, 187–194.

Prieur, C., Rigaud, J., Cheynier, V. & Moutounet, M., 1994. Oligomeric and polymeric procyanidins form grape seeds. Phytochemistry 36, 781–784.

Pontin, M.A., Piccoli, P.N., Francisco, R., Bottini, R, Martinez- Zapater, J.M., Lijavetzky, D., 2010. Transcriptome changes in grapevine (Vitis vinifera L.) cv. Malbec leaves induced by ultraviolet-B radiation. BMC Plant Biol. 2010, 10.

Ribereau-Gayon, P., Dubourdieu, D., Doneche, B. & Lonvaud, A., 2000. Handbook of Enology, 2nd Edition. Volume 1. John Wiley and Sons, Ltd. West Sussex, England.

Ricardo-da-Silva, J.M., Rigaud, J., Cheynier, V., Cheminat, A. & Moutounet, M. 1991a. Procyanidin dimers and trimers from grape seeds. Phytochemistry 30, 1259–1264.

Ricardo-da-Silva, J.M., Bourzeix, M., Cheynier, V. & Moutounet, M., 1991b. Procyanidin composition of Chardonnay, Mauzac and Grenanche blanc grapes. Vitis 30, 245–252.

Ricardo-da-Silva, J.M., Cheynier, V., Souquet, J.M. & Moutounet, M., 1991c. Interaction of grape seed procyanidins with various proteins in relation to wine fining. J. Sci. Food Agric. 57, 111–125.

Ricardo-da-Silva, J.M., Rosec, J.P., Mourgues, J. & Moutounet, M., 1992a. Dimer and trimer procyanidins in Carignan and Mourvedre grapes and wines. Vitis 31, 55–63.

Ricardo-da-Silva, J.M., Belchior, A.P. & Spranger, M.I., 1992b. Oligomeric procyanidins of three grapevine varieties and wines from Portugal. Sci Aliment. 12, 223–237.

Ristic, R., Downey, M.O., Iland, P.G., Bindon, K., Francis, I.L., Herderich, M. & Robinson, S.P. 2007. Exclusion of sunlight from Shiraz grapes alters wine colour, tannin and sensory properties. Austr. J. Grape Wine Res. 13, 53–65.

Robichaud, J.L. & Noble, A.C., 1990. Astringency and bitterness of selected phenolics in wine. J. Sci. Food Agric. 53, 343–353.

Roby, G., Harbertson, J.F., Adams, D.O. & Matthews, M.A., 2004a. Berry size and vine water deficits as factors in wine grape composition: Anthocyanins and tannins. Aust. J. Grape Wine Res. 10,100–107.

Roby, G. & Matthews, M.A., 2004b. Relative proportions of seed, skin and flesh in ripe berries from Cabernet Sauvignon grapevines grown in a vineyard either well irrigated or under water deficit. Aust. J. Grape Wine Res. 10, 74–82.

Rozema, J., vd Staaij, J., Bjorn, L.O. & Caldwell, M., 1997. UV-B as an environmental factor in plant life: stress and regulation. Trends Ecol Evol. 12, 22–28.

Ryel, R.J., Barnes, P.W., Beyschlag, W., Caldwell, M.M. & Flint, S.D., 1990. Plant competition for light analyzed with a multispecies canopy model. I. Model development and influence of enhanced UV-B conditions on photosynthesis in mixed wheat and wild oat canopies. Oecologia 82, 304–310.

Salon, J.L., Chirivella, C. & Castel, C.R. 2005. Response of cv. Bobal to timing of deficit irrigation in Requena, Spain: Water relations, yield, and wine quality. Am. J. Enol. Vitic. 56, 1–8.

Schwinn, K.E. & Davies, K.M., 2004. Flavonoids. In: Davies, K.M. (ed). Plant Pigments and their Manipulation. Blackwell Publishing Ltd, Oxford. pp. 92–149.

Siegelman, H.W. & Hendricks, S.B., 1958. Photocontrol of anthocyanin synthesis in apple skin. Plant Physiol. 33, 185–190.

Singleton, V.L. & Trousdale, E.K., 1992. Anthocyanin-tannin interactions explaining differences in polymeric phenols between white and red wines. Am. J. Enol. Vitic. 43, 63-70.

Smart, R.E., Robinson, J.B., Due, G.R. & Brien, C.J., 1985. Canopy microclimate modification for the cultivar Shiraz. 2. Effects on must and wine composition. Vitis 24, 119-128.

Smart, R.E. 1987. Influence of light on composition and quality of grapes. Acta Hort. 206, 37–47.

Smith, H., 1975. Phytochrome and Photomorphogenesis: An introduction to the photocontrol of plant development. McGraw – Hill, London.

Somers, T.C. & Vérette, E., 1988. Phenolic composition of natural wine types. In: Linskens, H.F. & Jackson, J.F. (eds.). Wine analysis. Springer-Verlag: Berlin. pp. 219–257.

Souquet, J.M., Cheynier, V., Brossaud, F. & Moutounet, M., 1996. Polymeric proanthocyanidins from grape skins. Phytochemistry, 43, 509–512.

Spayd, S.E., Tarara, J.M., Mee, D.L. & Ferguson, J.C., 2002. Separation of sunlight and temperature effects on the composition of Vitis vinifera cv. Merlot berries. Am. J. Enol. Vitic. 53, 171–182.

Stafford, H.A., 1990 (1st ed). Flavonoid metabolism. CRC Press, Boca Raton, FL.

Sun, B.S., Pinto, T., Leandro, M.C., Ricardo da Silva, J.M. and Spranger, M.I., 1999. Transfer of catechins and proanthocyanidins from solid parts of the grape cluster into wine. Am. J. Enol. Vitic. 50, 179–184.

Sun, B.S., Spranger, M.I., Roque-do-Vale, F., Leandro, C. & Belchior, P., 2001. Effect of different winemaking technologies on phenolic composition of Tinta Miura red wines. J. Agric. Food Chem. 49, 5809–5816.

Tanner, G.J., Francki, K.T., Abrahams, S., Watson, J.M., Larkin, P.J. and Ashton, A.R. 2003. Proanthocyanidin biosynthesis in plants. Purification of legume leucoanthocyanidin reductase and molecular cloning of its own cDNA. J. Biol. Chem. 278, 31647–31656.

Tanner, G., 2006. Condensed tannins, In: Davies, K. (ed.). Plant pigments and their manipulation, CRC-Press, Blackwell Publishing, UK. pp. 150–176.

Tarara, J.M., Lee, J., Spayd, S.E. & Scagel, C., 2008. Berry temperature and solar radiation alter acylation, proportion, and concentration of anthocyanin in Merlot grapes. Am. J. Enol. Vitic. 59, 235–247.

Teramura, A.H. & Sullivan, J.H., 1994. Effects of UV-B radiation on photosynthesis and growth of terrestrial plants. Photosynt. Res. 39, 463–473.

Terrier, N., Ollé, D., Verriès, C. & Cheynier, V. 2009. Biochemical & Molecular Aspects of flavan-3-ol synthesis during berry development. In: K.A. Roubelakis-Angelakis (ed.). Grapevine molecular physiology & biotechnology, 2nd, pp. 365–388.

Tevini, M. & Steinmuller, D., 1987. Influence of light, UV-B radiation and herbicides on wax biosynthesis of cucumber seedlings. J. Plant Phys. 131, 111–121.

Thorngate, J.H. & Noble, A.C., 1995. Sensory evaluation of bitterness and astringency of 3R( -)-epicatechin and 3S( +)-catechin. J. Sci. Food Agric. 67, 531–535.

Vass, I., 1997. Adverse effects of UV-B light on the structure and function of the photosynthetic apparatus. In: Pessarakli, M. (ed). Handbook of photosynthesis. Marcel Dekker. New York, pp. 931–949.

Vidal, S., Francis, L., Guyot, S., Marnet, N., Kwiatkowski, M., Gawel, R., Cheynier, V. & Waters, E.J., 2003. The mouth-feel properties of grape and apple proanthocyanidins in a wine-like medium. J. Sci. Food Agric. 83, 564–573.

Vidal, S., Francis, L., Noble, A., Kwiatkowski, M., Cheynier, V. & Waters, E., 2004a. Taste and mouth-feel properties of different types of tannin-like polyphenolic compounds and anthocyanins in wine. Anal Chim Acta 513, 57–65.

Vidal, S., Francis, L., Williams, P., Kwiatkowski, M., Gawel, R., Cheynier, W. & Waters, E., 2004b. The mouth-feel properties of polysaccharides and anthocyanins in a wine like medium. Food Chem. 85, 519–525.

Weber, F., Greve, K., Durner, D., Fischer, U. & Winterhalter, P., 2013. Sensory and chemical characterization of phenolic polymers from red wine obtained by gel permeation chromatography. Am. J. Enol. Vitic. 61, 1, 15–25.

Wicks, A.S. & Kliewer, W.M., 1983. Further investigations into the relationship between anthocyanins, phenolics and soluble carbo-hydrates in grape berry skins. Am. J. Enol. Vitic. 34, 114–116.

Wollman, N & Hofmann, T., 2013. Compositional and sensory characterization of red wine polymers. J. Agric. Food Chem. 61, 2045–2061.

Xie, D.Y., Sharma, S.B., Paiva, N.L., Ferreira, D. & Dixon, R.A., 2003. Role of anthocyanidin reductase, encoded by BANYULS in plant flavonoid biosynthesis. Science 299, 396–399.

Yamane, T., Jeong, S.T., Goto-Yamamoto, N., Koshita, Y. & Kobayashi, S., 2006. Effects of temperature on anthocyanin biosynthesis in grape berry skins. Am. J. Enol. Vitic. 57, 54–59.

Zucker, M., 1965. Induction of phenylalanine deaminase by light and its relation to chlorogenic acid synthesis in potato tuber tissue. Plant Physiol. 40, 779–784.

DOI: https://doi.org/10.21548/40-1-3060


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