{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T17:32:25Z","timestamp":1792085545545,"version":"4.4.0"},"reference-count":50,"publisher":"MDPI AG","issue":"1","license":[{"start":{"date-parts":[[2021,12,28]],"date-time":"2021-12-28T00:00:00Z","timestamp":1640649600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001053","name":"Wine Australia","doi-asserted-by":"publisher","award":["CSU1801"],"award-info":[{"award-number":["CSU1801"]}],"id":[{"id":"10.13039\/501100001053","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Biomolecules"],"abstract":"<jats:p>Sunburn is a physiological disorder that reduces grape quality and vineyard yield. It is the result of excessive sunlight and high temperatures. As climate change continues to increase air temperatures, reports of sunburn damage in vineyards worldwide are becoming more frequent. Grapes produce secondary metabolites (carotenoids, polyphenols and aroma compounds) to counter photooxidative stress and acclimate to higher radiation environments. This study evaluated changes in these compounds in during ripening when grapes were exposed post-flowering (ED) and at v\u00e9raison (LD), and compared them to a nondefoliated control (ND). ND contained more \u03b1-terpineol and violaxanthin, and the defoliated treatments contained more zeaxanthin, \u03b2-carotene, C6 compounds and flavonoids. ED berries adapted better to higher-light environments, displayed larger changes in secondary metabolite concentrations and lower levels of sunburn damage than LD berries did. The composition of berries with increasing sunburn damage was evaluated for the first time. Berries with no damage had the lowest concentrations of flavonoids and oxidized glutathione, and the highest concentrations of chlorophyll and \u03b1-terpineol. As damage increased, destruction of photosynthetic pigments, increase in polyphenols and loss of aroma compounds were evidenced. A significant effect of temperature and developmental stage on grape composition was also observed. This study provides a holistic overview of changes in secondary metabolites experienced by grape berries when exposed to excessive light, how these vary along development and how they affect sunburn incidence.<\/jats:p>","DOI":"10.3390\/biom12010042","type":"journal-article","created":{"date-parts":[[2021,12,28]],"date-time":"2021-12-28T06:55:03Z","timestamp":1640674503000},"page":"42","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":29,"title":["Secondary Metabolites Coordinately Protect Grapes from Excessive Light and Sunburn Damage during Development"],"prefix":"10.3390","volume":"12","author":[{"given":"Joanna M.","family":"Gambetta","sequence":"first","affiliation":[{"name":"School of Agricultural, Environmental and Veterinary Sciences, National Wine and Grape Industry Centre, Charles Sturt University, Locked Bag 588, Wagga Wagga, NSW 2678, Australia"},{"name":"Waite Research Institute, School of Agriculture, Food and Wine, University of Adelaide, PMB1, Glen Osmond, SA 5064, Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Valentina","family":"Romat","sequence":"additional","affiliation":[{"name":"School of Agricultural, Environmental and Veterinary Sciences, National Wine and Grape Industry Centre, Charles Sturt University, Locked Bag 588, Wagga Wagga, NSW 2678, Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9765-5510","authenticated-orcid":false,"given":"Leigh M.","family":"Schmidtke","sequence":"additional","affiliation":[{"name":"School of Agricultural, Environmental and Veterinary Sciences, National Wine and Grape Industry Centre, Charles Sturt University, Locked Bag 588, Wagga Wagga, NSW 2678, Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bruno P.","family":"Holzapfel","sequence":"additional","affiliation":[{"name":"Wagga Wagga Agriculture Institute, New South Wales Department of Primary Industries, Wagga Wagga, NSW 2650, Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2021,12,28]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"630","DOI":"10.3389\/fpls.2017.00630","article-title":"Transcriptional Responses to Pre-Flowering Leaf Defoliation in Grapevine Berry from Different Growing Sites, Years, and Genotypes","volume":"8","author":"Zenoni","year":"2017","journal-title":"Front. Plant Sci."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"604691","DOI":"10.3389\/fpls.2020.604691","article-title":"Sunburn in Grapes: A Review","volume":"11","author":"Gambetta","year":"2021","journal-title":"Front. Plant Sci."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1016\/j.scienta.2015.01.029","article-title":"Chlorophyll Role in Berry Sunburn Symptoms Studied in Different Grape (Vitis vinifera L.) Cultivars","volume":"185","author":"Rustioni","year":"2015","journal-title":"Sci. Hortic."},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Solovchenko, A.E. (2010). Photoprotection in Plants\u2014Optical Screening Based Mechanisms, Springer.","DOI":"10.1007\/978-3-642-13887-4"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"1261","DOI":"10.3389\/fpls.2017.01261","article-title":"The Transcriptional Responses and Metabolic Consequences of Acclimation to Elevated Light Exposure in Grapevine Berries","volume":"8","author":"Young","year":"2017","journal-title":"Front. Plant Sci."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"140","DOI":"10.1080\/07352689.2019.1613320","article-title":"Physiological Mechanisms Underlying Fruit Sunburn","volume":"38","author":"Vincent","year":"2019","journal-title":"Crit. Rev. Plant Sci."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"27","DOI":"10.21273\/JASHS.133.1.27","article-title":"Changes in Pigment Concentrations Associated with the Degree of Sunburn Browning of \u2018Fuji\u2019 Apple","volume":"133","author":"Felicetti","year":"2008","journal-title":"J. Am. Soc. Hortic. Sci."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"786","DOI":"10.3389\/fpls.2016.00786","article-title":"Field-Grown Grapevine Berries Use Carotenoids and the Associated Xanthophyll Cycles to Acclimate to UV Exposure Differentially in High and Low Light (Shade) Conditions","volume":"7","author":"Joubert","year":"2016","journal-title":"Front. Plant Sci."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1071\/FP03076","article-title":"UV Screening by Phenolics in Berries of Grapevine (Vitis vinifera)","volume":"30","author":"Kolb","year":"2003","journal-title":"Funct. Plant Biol."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"He, L., Xu, X.-Q., Wang, Y., Chen, W.-K., Sun, R.-Z., Cheng, G., Liu, B., Chen, W., Duan, C.-Q., and Wang, J. (2020). Modulation of Volatile Compound Metabolome and Transcriptome in Grape Berries Exposed to Sunlight under Dry-Hot Climate. BMC Plant Biol., 20.","DOI":"10.1186\/s12870-020-2268-y"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"1337","DOI":"10.3389\/fpls.2016.01337","article-title":"Transcriptomic and Metabolomic Networks in the Grape Berry Illustrate That It Takes More than Flavonoids to Fight against Ultraviolet Radiation","volume":"7","author":"Matus","year":"2016","journal-title":"Front. Plant Sci."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"1235","DOI":"10.1104\/pp.15.01775","article-title":"Grapevine Plasticity in Response to an Altered Microclimate: Sauvignon Blanc Modulates Specific Metabolites in Response to Increased Berry Exposure","volume":"170","author":"Young","year":"2016","journal-title":"Plant Physiol."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Carbonell-Bejerano, P., Diago, M.P., Mart\u00ednez-Abaigar, J., Mart\u00ednez-Zapater, J.M., Tard\u00e1guila, J., and N\u00fa\u00f1ez-Olivera, E. (2014). Solar Ultraviolet Radiation Is Necessary to Enhance Grapevine Fruit Ripening Transcriptional and Phenolic Responses. BMC Plant Biol., 14.","DOI":"10.1186\/1471-2229-14-183"},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Pastore, C., Zenoni, S., Fasoli, M., Pezzotti, M., Tornielli, G.B., and Filippetti, I. (2013). Selective Defoliation Affects Plant Growth, Fruit Transcriptional Ripening Program and Flavonoid Metabolism in Grapevine. BMC Plant Biol., 13.","DOI":"10.1186\/1471-2229-13-30"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1004","DOI":"10.1104\/pp.17.01127","article-title":"Photo-Oxidative Stress during Leaf, Flower and Fruit Development","volume":"176","year":"2018","journal-title":"Plant Physiol."},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"M\u00fcller, P., Li, X.P., and Niyogi, K.K. (2001). Non-Photochemical Quenching. A Response to Excess Light Energy. Plant Physiol., 125.","DOI":"10.1104\/pp.125.4.1558"},{"key":"ref_17","first-page":"1","article-title":"The Interactive Effects of Temperature and Light Intensity on Vitis vinifera cv. \u201cSemillon\u201d Grapevines. II. Berry Ripening and Susceptibility to Sunburn at Harvest","volume":"79","author":"Hulands","year":"2014","journal-title":"Eur. J. Hortic. Sci."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"3043","DOI":"10.1021\/jf405772f","article-title":"Characterization of Grape (Vitis vinifera L.) Berry Sunburn Symptoms by Reflectance","volume":"62","author":"Rustioni","year":"2014","journal-title":"J. Agric. Food Chem."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"93","DOI":"10.20870\/oeno-one.2018.52.2.2158","article-title":"Intensity and Timing of Defoliation on White Cultivar Chasselas under the Temperate Climate of Switzerland","volume":"52","author":"Verdenal","year":"2018","journal-title":"Oeno One"},{"key":"ref_20","unstructured":"Northcote, K.H. (1979). A Factual Key for the Recognition of Australian Soils, Rellim Technical Publications. [4th ed.]."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"104","DOI":"10.1111\/j.1755-0238.1995.tb00086.x","article-title":"Growth Stages of the Grapevine: Adoption of a System for Identifying Grapevine Growth Stages","volume":"1","author":"Coombe","year":"1995","journal-title":"Aust. J. Grape Wine Res."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"171","DOI":"10.20870\/oeno-one.2019.53.2.2392","article-title":"Single and Cumulative Effects of Whole-Vine Heat Events on Shiraz Berry Composition","volume":"53","author":"Gouot","year":"2019","journal-title":"Oeno One"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"2842","DOI":"10.1007\/s12161-016-0467-9","article-title":"Towards the Creation of a Wine Quality Prediction Index: Correlation of Chardonnay Juice and Wine Compositions from Different Regions and Quality Levels","volume":"9","author":"Gambetta","year":"2016","journal-title":"Food Anal. Methods"},{"key":"ref_24","unstructured":"Iland, P., Bruer, N., Edwards, G., Weeks, S., and Wilkes, E. (2004). Chemical Analysis of Grapes and Wine: Techniques and Concepts, Patrick Iland Wine Promotions."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"5075","DOI":"10.1007\/s00216-012-6555-9","article-title":"High-Throughput Carotenoid Profiling Using Multivariate Curve Resolution","volume":"405","author":"Wehrens","year":"2013","journal-title":"Anal. Bioanal. Chem."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"349","DOI":"10.1111\/j.1755-0238.2010.00097.x","article-title":"The Development of a Method for the Extraction of Carotenoids and Chlorophylls from Grapevine Leaves and Berries for HPLC Profiling","volume":"16","author":"Lashbrooke","year":"2010","journal-title":"Aust. J. Grape Wine Res."},{"key":"ref_27","first-page":"301","article-title":"ASCA: The implementation of design of experiments into multivariate modelling in chemometrics","volume":"Volume 82","author":"Jaumot","year":"2018","journal-title":"Comprehensive Analytical Chemistry"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"144","DOI":"10.1111\/j.1755-0238.1996.tb00102.x","article-title":"The Plastids of the Grape Pericarp and Their Significance in Isoprenoid Synthesis","volume":"2","author":"Hardie","year":"1996","journal-title":"Aust. J. Grape Wine Res."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"287","DOI":"10.20870\/oeno-one.2019.53.2.2424","article-title":"The Influence of Temperature and Solar Radiation on Phenols in Berry Skin and Maturity Parameters of Vitis vinifera L. cv. Riesling","volume":"53","author":"Brandt","year":"2019","journal-title":"Oeno One"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"171","DOI":"10.5344\/ajev.2002.53.3.171","article-title":"Separation of Sunlight and Temperature Effects on the Composition of Vitis Vinifera Cv. Merlot Berries","volume":"53","author":"Spayd","year":"2002","journal-title":"Am. J. Enol. Vitic."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"10","DOI":"10.3389\/fpls.2019.00010","article-title":"Flavonol Profile Is a Reliable Indicator to Assess Canopy Architecture and the Exposure of Red Wine Grapes to Solar Radiation","volume":"10","author":"Brillante","year":"2019","journal-title":"Front. Plant Sci."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"54","DOI":"10.1016\/j.phytochem.2012.02.026","article-title":"Light Quality Affects Flavonoid Biosynthesis in Young Berries of Cabernet Sauvignon Grape","volume":"78","author":"Koyama","year":"2012","journal-title":"Phytochemistry"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"70","DOI":"10.3389\/fpls.2017.00070","article-title":"Sunlight Modulates Fruit Metabolic Profile and Shapes the Spatial Pattern of Compound Accumulation within the Grape Cluster","volume":"8","author":"Reshef","year":"2017","journal-title":"Front. Plant Sci."},{"key":"ref_34","unstructured":"Hixson, J., Grebneva, Y., Glameyer, N., Vollmer, K., Black, C., Krstic, M., and Herderich, M. (2017, January 18\u201322). Shedding Light on the Modulation of Key Riesling Wine Aroma Compounds in a Changing Climate. Proceedings of the XV Weurman Flavour Research Symposium, Graz, Austria."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"6841","DOI":"10.1021\/jf904555p","article-title":"Timing of Cluster Light Environment Manipulation during Grape Development Affects C13 Norisoprenoid and Carotenoid Concentrations in Riesling","volume":"58","author":"Kwasniewski","year":"2010","journal-title":"J. Agric. Food Chem."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"6512","DOI":"10.1021\/jf501945s","article-title":"Factors Influencing the Aroma Composition of Chardonnay Wines","volume":"62","author":"Gambetta","year":"2014","journal-title":"J. Agric. Food Chem."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"16301","DOI":"10.1038\/s41598-017-16529-5","article-title":"Ozone Improves the Aromatic Fingerprint of White Grapes","volume":"7","author":"Vilanova","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1204","DOI":"10.1111\/pce.12643","article-title":"Effects of Heat and Drought Stress on Post-Illumination Bursts of Volatile Organic Compounds in Isoprene-Emitting and Non-Emitting Poplar","volume":"39","author":"Jud","year":"2016","journal-title":"Plant Cell Environ."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"5975","DOI":"10.1093\/jxb\/eru343","article-title":"Metabolic Effects of Elevated Temperature on Organic Acid Degradation in Ripening Vitis vinifera Fruit","volume":"65","author":"Sweetman","year":"2014","journal-title":"J. Exp. Bot."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"1935","DOI":"10.1093\/jxb\/erm055","article-title":"Loss of Anthocyanins in Red-Wine Grape under High Temperature","volume":"58","author":"Mori","year":"2007","journal-title":"J. Exp. Bot."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1038\/nchembio.158","article-title":"A Unified Mechanism of Action for Volatile Isoprenoids in Plant Abiotic Stress","volume":"5","author":"Vickers","year":"2009","journal-title":"Nat. Chem. Biol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"664","DOI":"10.1016\/j.jplph.2011.12.019","article-title":"Emissions of Green Leaf Volatiles and Terpenoids from Solanum lycopersicum Are Quantitatively Related to the Severity of Cold and Heat Shock Treatments","volume":"169","author":"Copolovici","year":"2012","journal-title":"J. Plant Physiol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.envexpbot.2016.08.003","article-title":"Mono- and Sesquiterpene Release from Tomato (Solanum lycopersicum) Leaves upon Mild and Severe Heat Stress and through Recovery: From Gene Expression to Emission Responses","volume":"132","author":"Pazouki","year":"2016","journal-title":"Environ. Exp. Bot."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"5017","DOI":"10.1093\/jxb\/erz255","article-title":"Lethal Heat Stress-Dependent Volatile Emissions from Tobacco Leaves: What Happens beyond the Thermal Edge?","volume":"70","author":"Turan","year":"2019","journal-title":"J. Exp. Bot"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"270","DOI":"10.1016\/j.plaphy.2020.03.003","article-title":"Oxidations in White Grape (Vitis vinifera L.) Skins: Comparison between Ripening Process and Photooxidative Sunburn Symptoms","volume":"150","author":"Rustioni","year":"2020","journal-title":"Plant Physiol. Biochem."},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Deluc, L.G., Grimplet, J., Wheatley, M.D., Tillett, R.L., Quilici, D.R., Osborne, C., Schooley, D.A., Schlauch, K.A., Cushman, J.C., and Cramer, G.R. (2007). Transcriptomic and Metabolite Analyses of Cabernet Sauvignon Grape Berry Development. BMC Genom., 8.","DOI":"10.1186\/1471-2164-8-429"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"367","DOI":"10.5344\/ajev.2012.11116","article-title":"Impact of Prebloom and Fruit-Set Basal Leaf Removal on the Flavonol and Anthocyanin Composition of Tempranillo Grapes","volume":"63","author":"Diago","year":"2012","journal-title":"Am. J. Enol. Vitic."},{"key":"ref_48","unstructured":"Bureau of Meteorology (2019, February 08). Summer 2019 Sets New Benchmarks for Australian Temperatures, Available online: http:\/\/media.bom.gov.au\/social\/blog\/2020\/summer-2019-sets-new-benchmarks-for-australian-temperatures\/?utm_source=edm&utm_medium=org&utm_campaign=sm-003-0187&utm_content=Nbr%C2%A0."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"2","DOI":"10.1111\/j.1467-7652.2008.00377.x","article-title":"Molecular Engineering of Resveratrol in Plants","volume":"7","author":"Delaunois","year":"2009","journal-title":"Plant Biotechnol. J."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"95","DOI":"10.1016\/j.jplph.2009.07.011","article-title":"Changes of Resveratrol and Antioxidant Enzymes during UV-Induced Plant Defense Response in Peanut Seedlings","volume":"167","author":"Tang","year":"2010","journal-title":"J. Plant Physiol."}],"container-title":["Biomolecules"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2218-273X\/12\/1\/42\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T07:54:49Z","timestamp":1760169289000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2218-273X\/12\/1\/42"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,12,28]]},"references-count":50,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2022,1]]}},"alternative-id":["biom12010042"],"URL":"https:\/\/doi.org\/10.3390\/biom12010042","relation":{},"ISSN":["2218-273X"],"issn-type":[{"value":"2218-273X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,12,28]]}}}