{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,4]],"date-time":"2026-05-04T12:35:44Z","timestamp":1777898144665,"version":"3.51.4"},"reference-count":39,"publisher":"MDPI AG","issue":"2","license":[{"start":{"date-parts":[[2013,1,25]],"date-time":"2013-01-25T00:00:00Z","timestamp":1359072000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/3.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Molecules"],"abstract":"<jats:p>In the present study, evolution of apple color (L* and a\/b), the accumulation of anthocyanins and the activity of the related enzymes, phenylalanine ammonia-lyase (PAL), chalcone isomerase (CHI), dihydroflavonol4-reductase (DFR) and UDP-Glucose: flavonoid-3-O-galactosyl transferase (UFGT), were investigated in bagged non-red apple cultivars (\u2018Granny Smith\u2019 and \u2018Golden Delicious\u2019) and red apple cultivars (\u2018Starkrimon\u2019 and \u2018Pink Lady\u2019). Young fruits were bagged 40\u201345 days after flowering (DAF), and fruits of \u2018Golden Delicious\u2019 and \u2018Starkrimon\u2019 were uncovered and exposed to light 120 DAF, while those of \u2018Granny Smith\u2019 and \u2018Pink Lady\u2019 were exposed for 160 DAF. Results showed that cyanidin 3-galactoside (cy3-gal) was the most abundant anthocyanin in both non-red and red cultivars. Level of anthocyanins was higher in \u2018Granny Smith\u2019 than in \u2018Golden Delicious\u2019, indicating that red color was easier to develop in green cultivar \u2018Granny Smith\u2019 than in yellow cultivar \u2018Golden Delicious\u2019 after bag removal. The cy3-gal accumulation of non-red cultivars tested was not significantly correlated with PAL, CHI and DFR activity, but was significantly correlated with UFGT activity. During the reddening of non-red apples, UFGT may be the more important factor in the anthocyanin biosynthesis.<\/jats:p>","DOI":"10.3390\/molecules18021549","type":"journal-article","created":{"date-parts":[[2013,1,25]],"date-time":"2013-01-25T11:06:55Z","timestamp":1359112015000},"page":"1549-1563","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":84,"title":["Fruit Coloration and Anthocyanin Biosynthesis after Bag Removal in Non-Red and Red Apples (Malus \u00d7 domestica Borkh.)"],"prefix":"10.3390","volume":"18","author":[{"given":"Yulian","family":"Liu","sequence":"first","affiliation":[{"name":"Department of Pomology, College of Horticulture, Northwest A&F University, Yangling 712100, Shaanxi, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Fei","family":"Che","sequence":"additional","affiliation":[{"name":"Department of Pomology, College of Horticulture, Northwest A&F University, Yangling 712100, Shaanxi, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Lixin","family":"Wang","sequence":"additional","affiliation":[{"name":"Department of Pomology, College of Horticulture, Northwest A&F University, Yangling 712100, Shaanxi, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Rui","family":"Meng","sequence":"additional","affiliation":[{"name":"Department of Pomology, College of Horticulture, Northwest A&F University, Yangling 712100, Shaanxi, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Xiaojun","family":"Zhang","sequence":"additional","affiliation":[{"name":"Department of Pomology, College of Horticulture, Northwest A&F University, Yangling 712100, Shaanxi, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Zhengyang","family":"Zhao","sequence":"additional","affiliation":[{"name":"Department of Pomology, College of Horticulture, Northwest A&F University, Yangling 712100, Shaanxi, China"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2013,1,25]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"236","DOI":"10.1016\/j.tplants.2005.03.002","article-title":"Flavonoids: A colorful model for the regulation and evolution of biochemical pathways","volume":"10","author":"Koes","year":"2005","journal-title":"Trends Plant Sci."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"955","DOI":"10.1016\/S0981-9428(02)01454-7","article-title":"Anthocyanin biosynthetic genes are coordinately expressed during red coloration in apple skin","volume":"40","author":"Honda","year":"2002","journal-title":"Plant Physiol. Biochem."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"487","DOI":"10.1080\/07352689209382324","article-title":"Regulation of skin color in apples","volume":"10","author":"Lancaster","year":"1992","journal-title":"Crit. Rev. Plant Sci."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"1381","DOI":"10.1111\/j.1365-3040.2007.01716.x","article-title":"Transcriptional regulation of anthocyanin biosynthesis in ripening fruits of grapevine under seasonal water deficit","volume":"30","author":"Castellarin","year":"2007","journal-title":"Plant Cell Environ."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"1059","DOI":"10.1104\/pp.111.4.1059","article-title":"Analysis of the expression of anthocyanin pathway genes in developing Vitis vinifera L. cv Shiraz grape berries and the implications for pathway regulation","volume":"111","author":"Boss","year":"1996","journal-title":"Plant Physiol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"1216","DOI":"10.1104\/pp.106.088104","article-title":"Light-induced expression of a MYB gene regulates anthocyanin biosynthesis in red apples","volume":"142","author":"Takos","year":"2006","journal-title":"Plant Physiol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"414","DOI":"10.1111\/j.1365-313X.2006.02964.x","article-title":"Red coloration in apple fruit is due to the activity of the MYB transcription factor, MdMYB10","volume":"49","author":"Espley","year":"2007","journal-title":"Plant J."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"285","DOI":"10.1007\/s00299-010-0821-0","article-title":"A WD40-repeat gene from Malus \u00d7 domestica is a functional homologue of Arabidopsis thaliana TRANSPARENT TESTA GLABRA1","volume":"29","author":"Brueggemann","year":"2010","journal-title":"Plant Cell Rep."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"181","DOI":"10.1016\/0304-4238(90)90082-P","article-title":"External control of anthocyanin formation in apple","volume":"42","author":"Saure","year":"1990","journal-title":"Sci. Hortic."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1023\/A:1013378702940","article-title":"Biosynthesis of phenolic compounds and its regulation in apple","volume":"34","author":"Treutter","year":"2001","journal-title":"Plant Growth Regul."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"487","DOI":"10.1016\/j.plantsci.2005.10.001","article-title":"Condensed tannin biosynthesis genes are regulated separately from other flavonoid biosynthesis genes in apple fruit skin","volume":"170","author":"Takos","year":"2006","journal-title":"Plant Sci."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"499","DOI":"10.1104\/pp.105.067231","article-title":"Characterization of a grapevine R2R3-MYB transcription factor that regulates the phenylpropanoid pathway","volume":"140","author":"Deluc","year":"2006","journal-title":"Plant Physiol."},{"key":"ref_13","first-page":"485","article-title":"Flavonoid Biosynthesis. A Colorful Model for Genetics, Biochemitry, Cell Biology, and Biotechnology","volume":"126","year":"2001","journal-title":"Postharvest Biol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1016\/S0925-5214(00)00089-2","article-title":"Influence of maturity and bagging on the relationship between anthocyanin accumulation and phenylalanine ammonia-lyase (PAL) activity in \u2018Jonathan\u2019 apples","volume":"19","author":"Wang","year":"2000","journal-title":"Postharvest Biol. Technol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"313","DOI":"10.1002\/(SICI)1097-0010(199607)71:3<313::AID-JSFA586>3.0.CO;2-N","article-title":"Developmental changes in enzymes of flavonoid biosynthesis in the skins of red and green apple cultivars","volume":"71","author":"Lister","year":"1996","journal-title":"J. Sci. Food Agric."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1016\/S0304-4238(97)00040-X","article-title":"Dihydroflavonol reductase activity and anthocyanin accumulation in \u2018Delicious\u2019, \u2018Golden Delicious\u2019 and \u2018Indo\u2019 apples","volume":"70","author":"Ju","year":"1997","journal-title":"Sci. Hortic."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"871","DOI":"10.1007\/s00425-009-0993-4","article-title":"UDP-sugar biosynthetic pathway: Contribution to cyanidin 3-galactoside biosynthesis in apple skin","volume":"230","author":"Ban","year":"2009","journal-title":"Planta"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1016\/S0304-4238(01)00308-9","article-title":"Orchard cooling with overtree microsprinkler irrigation to improve fruit color and quality of \u2018Topred Delicious\u2019 apples","volume":"93","author":"Iglesias","year":"2002","journal-title":"Sci. Hortic."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1016\/j.plantsci.2010.01.008","article-title":"Light-induced decrease of reflectance provides an insight in the photoprotective mechanisms of ripening apple fruit","volume":"178","author":"Solovchenko","year":"2010","journal-title":"Plant Sci."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"571","DOI":"10.1016\/j.plantsci.2005.10.009","article-title":"Expression analyis of anthocyanin biosynthetic genes in apple skin: Effect of UV-B and temperature","volume":"170","author":"Ubi","year":"2006","journal-title":"Plant Sci."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1111\/j.1751-1097.1999.tb01944.x","article-title":"Environmental significance of anthocyanins in plant stress responses","volume":"70","year":"1999","journal-title":"Photochem. Photobiol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1085","DOI":"10.2307\/3870059","article-title":"Stress-induced phenylpropanoid metabolism","volume":"7","author":"Dixon","year":"1995","journal-title":"Plant Cell."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"1176","DOI":"10.1111\/j.1365-3040.2011.02316.x","article-title":"High temperature reduces apple fruit color via modulation of the anthocyanin regulatory complex","volume":"34","author":"Wang","year":"2011","journal-title":"Plant Cell Environ."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"187","DOI":"10.1080\/14620316.2005.11511915","article-title":"Color accumulation patterns and the anthocyanin biosynthetic pathway in \u2018Red Delicious\u2019 apple variants","volume":"80","author":"Korchinsky","year":"2005","journal-title":"J. Hortic. Sci. Biotech."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"20","DOI":"10.21273\/JASHS.132.1.20","article-title":"Endogenous ethylene and color development in the skin of \u2018Pink Lady\u2019 apple","volume":"132","author":"Whale","year":"2007","journal-title":"J. Am. Soc. Hortic. Sci."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"1138","DOI":"10.1021\/jf072540m","article-title":"Prestorage ultraviolet-white light irradiation alters apple peel metabolome","volume":"56","author":"Rudell","year":"2008","journal-title":"J. Agric. Food Chem."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"718","DOI":"10.1080\/14620316.2002.11511562","article-title":"Expression of anthocyanin biosynthetic genes in Malus sylvestris L. \u2018Mutsu\u2019 non-red apples","volume":"77","author":"Kondo","year":"2002","journal-title":"J. Hortic. Sci. Biotech."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1016\/S0304-4238(98)00161-7","article-title":"Friut bagging, a useful method for studying anthocyanin synthesis and gene m expression in apple","volume":"77","author":"Ju","year":"1998","journal-title":"Sci. Hortic."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"474","DOI":"10.21273\/HORTSCI.32.3.474C","article-title":"Bag and liner color greatly affect apple temperature under full sunlight","volume":"32","author":"Ritenour","year":"1997","journal-title":"HortScience"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"403","DOI":"10.1016\/S0168-9452(03)00201-2","article-title":"Molecular cloning and analysis of anthocyanin biosynthesis genes preferentially expressed in apple skin","volume":"165","author":"Kim","year":"2003","journal-title":"Plant Sci."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"743","DOI":"10.1007\/BF00029856","article-title":"Cloning and molecular analysis of structural genes involved in flavonoid and stilbene biosynthesis in grape (Vitis vinifera L.)","volume":"24","author":"Sparvoli","year":"1994","journal-title":"Plant Mol. Biol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"865","DOI":"10.21273\/JASHS.116.5.865","article-title":"Activity of phenylalanine ammonia-lyase (PAL) and concentrations of anthocyanins and phenolics in developing strawberry fruit","volume":"116","author":"Cheng","year":"1991","journal-title":"J. Am. Soc. Hortic. Sci."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1016\/0304-4238(94)00739-3","article-title":"Relationships among phenylalanine ammonia lyase activity, simple phenol concentration and anthocyanin accumulation in apple","volume":"61","author":"Ju","year":"1995","journal-title":"Sci. Hortic."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"191","DOI":"10.2503\/jjshs.57.191","article-title":"Color development of 4 apple cultivars grown in the Southeast of Japan, with special reference for bagging","volume":"57","author":"Kubo","year":"1998","journal-title":"J. Jpn. Soc. Hortic. Sci."},{"key":"ref_35","unstructured":"Stafford, H.A. (1990). Flavonoid Metabolism, CRC Press."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"1652","DOI":"10.1104\/pp.102.018242","article-title":"Biochemical and molecular characterization of a novel UDP-glucose: Anthocyanin 3'-O-Glucosyltransferase, a key enzyme for blue anthocyanin biosynthesis, from Gentian","volume":"132","author":"Okuhara","year":"2003","journal-title":"Plant Physiol."},{"key":"ref_37","first-page":"1318","article-title":"Relationship between anthocyanin biosynthesis and related enzymes activity in Pyrus pyrifolia mantianhong and its bud sports aoguan","volume":"7","author":"Feng","year":"2008","journal-title":"Sci. Agric. Sin."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"2437","DOI":"10.1093\/jxb\/err415","article-title":"Introduction of apple ANR genes into tobacco inhibits expression of both CHI and DFR genes in flowers, leading to loss of anthocyanin","volume":"63","author":"Han","year":"2012","journal-title":"J. Exp. Bot."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"1254","DOI":"10.21273\/HORTSCI.27.12.1254","article-title":"Reporting of objective color measurements","volume":"27","author":"McGuire","year":"1992","journal-title":"HortScience"}],"container-title":["Molecules"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1420-3049\/18\/2\/1549\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T21:44:33Z","timestamp":1760219073000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1420-3049\/18\/2\/1549"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2013,1,25]]},"references-count":39,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2013,2]]}},"alternative-id":["molecules18021549"],"URL":"https:\/\/doi.org\/10.3390\/molecules18021549","relation":{},"ISSN":["1420-3049"],"issn-type":[{"value":"1420-3049","type":"electronic"}],"subject":[],"published":{"date-parts":[[2013,1,25]]}}}