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This study aimed to elucidate the metabolic regulatory mechanisms underlying volatile aroma biosynthesis in Malus domestica \u201cNingqiu\u201d apples, thereby providing theoretical support for the comprehensive utilization of aroma resources. Our methodological framework integrated headspace solid-phase microextraction gas chromatography\u2013mass spectrometry (HS-SPME-GC-MS), ultra-high-performance liquid chromatography\u2013orbitrap mass spectrometry (UHPLC-OE-MS), and Illumina high-throughput sequencing to generate comprehensive metabolomic and transcriptomic profiles of peel and pulp tissues. Critical differential aroma compound classes were identified, including esters, aldehydes, alcohols, terpenoids, and ketones, with their metabolic pathways systematically mapped through KEGG functional annotation. Our findings revealed substantial transcriptomic and metabolomic divergence across carotenoid, terpenoid, and fatty acid metabolic pathways. Integrative analysis of multi-omics data revealed 26 and 31 putative biologically significant hub genes in peel and pulp tissues, respectively, putatively associated with the observed metabolic signatures. Among these, five core genes\u2014farnesyl diphosphate synthase (FDPS1.X1), alcohol acyltransferases (AAT1 and AAT3), alcohol dehydrogenase (ADH3), and carotenoid cleavage dioxygenase (CCD3)\u2014were recognized as shared regulatory determinants between both tissue types. Furthermore, terpene synthase (TPS7) emerged as a peel-specific regulatory factor, while hydroperoxide lyase (HPL2), alcohol dehydrogenases (ADH2 and ADH4), and alcohol acyltransferase (AAT2) were identified as pulp-exclusive modulators of metabolic differentiation. The experimental findings provide foundational insights into the molecular basis of aroma profile variation in Malus domestica \u201cNingqiu\u201d and establish a functional genomics framework for precision breeding initiatives targeting fruit quality optimization through transcriptional regulatory network manipulation.<\/jats:p>","DOI":"10.3390\/plants14081165","type":"journal-article","created":{"date-parts":[[2025,4,9]],"date-time":"2025-04-09T07:49:02Z","timestamp":1744184942000},"page":"1165","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":10,"title":["Integrated Analysis of the Metabolome and Transcriptome During Apple Ripening to Highlight Aroma Determinants in Ningqiu Apples"],"prefix":"10.3390","volume":"14","author":[{"given":"Jun","family":"Ma","sequence":"first","affiliation":[{"name":"Horticultural Research Institute, Ningxia Academy of Agriculture and Forestry, Yinchuan 750002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Guangzong","family":"Li","sequence":"additional","affiliation":[{"name":"Farmland Irrigation Research Institute, Chinese Academy of Agricultural Sciences, Xinxiang 453002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yannan","family":"Chu","sequence":"additional","affiliation":[{"name":"Horticultural Research Institute, Ningxia Academy of Agriculture and Forestry, Yinchuan 750002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Haiying","family":"Yue","sequence":"additional","affiliation":[{"name":"Horticultural Research Institute, Ningxia Academy of Agriculture and Forestry, Yinchuan 750002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zehua","family":"Xu","sequence":"additional","affiliation":[{"name":"Horticultural Research Institute, Ningxia Academy of Agriculture and Forestry, Yinchuan 750002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jiaqi","family":"Wu","sequence":"additional","affiliation":[{"name":"Horticultural Research Institute, Ningxia Academy of Agriculture and Forestry, Yinchuan 750002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaolong","family":"Li","sequence":"additional","affiliation":[{"name":"Horticultural Research Institute, Ningxia Academy of Agriculture and Forestry, Yinchuan 750002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yonghua","family":"Jia","sequence":"additional","affiliation":[{"name":"Horticultural Research Institute, Ningxia Academy of Agriculture and Forestry, Yinchuan 750002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,4,9]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","unstructured":"Christodoulou, M.D., and Culham, A. (2021). When do apples stop growing, and why does it matter?. PLoS ONE, 16.","DOI":"10.1371\/journal.pone.0252288"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"2376","DOI":"10.1104\/pp.17.00311","article-title":"Ripening transcriptomic program in red and white grapevine varieties correlates with apple skin anthocyanin accumulation","volume":"174","author":"Massonnet","year":"2017","journal-title":"Plant Physiol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"1026","DOI":"10.21273\/HORTSCI.35.6.1026","article-title":"Factors that influence biosynthesis of volatile flavor compounds in apple fruits","volume":"35","author":"Fellman","year":"2000","journal-title":"HortScience"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"327","DOI":"10.1016\/0308-8146(95)00100-X","article-title":"Unsaturated fatty acid bioconversion by apple pomace enzyme system. Factors influencing the production of aroma compounds","volume":"55","author":"Almosnino","year":"1996","journal-title":"Food Chem."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"6157","DOI":"10.1021\/jf100172e","article-title":"Expression of genes associated with aroma formation derived from the fatty acid pathway during peach fruit ripening","volume":"58","author":"Zhang","year":"2010","journal-title":"J. Agric. Food Chem."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"221","DOI":"10.1002\/ffj.3739","article-title":"Enzymatic reactions towards aldehydes, An overview","volume":"38","author":"Schober","year":"2023","journal-title":"Flavour Fragr. J."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Liu, Y., Zhou, J., Yi, C., Chen, F., Liu, Y., Liao, Y., Zhang, Z., Liu, W., and Lv, J. (2023). Integrative analysis of non-targeted metabolome and transcriptome reveals the mechanism of volatile formation in pepper fruit. Front. Genet., 14.","DOI":"10.3389\/fgene.2023.1290492"},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Lu, H., Zhao, H., Zhong, T., Chen, D., Wu, Y., and Xie, Z. (2024). Molecular regulatory mechanisms affecting fruit aroma. Foods, 13.","DOI":"10.3390\/foods13121870"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"4497","DOI":"10.1093\/jxb\/ers134","article-title":"Metabolic and gene expression analysis of apple (Malus \u00d7 domestica) carotenogenesis","volume":"63","author":"Dejnoprat","year":"2012","journal-title":"J. Exp. Bot."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"425","DOI":"10.1080\/15538362.2015.1031432","article-title":"Post-harvest ripening changes in wood apple (Feronia elephantum Corr), an underutilized fruit","volume":"15","author":"Lakshmi","year":"2015","journal-title":"Int. J. Fruit Sci."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"9052","DOI":"10.1080\/10408398.2022.2064422","article-title":"Free and glycosidically bound aroma compounds in fruit: Biosynthesis, transformation, and practical control","volume":"63","author":"Chen","year":"2023","journal-title":"Crit. Rev. Food Sci. Nutr."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1016\/j.cofs.2018.03.005","article-title":"Biogeneration of aroma compounds","volume":"19","author":"Sales","year":"2018","journal-title":"Curr. Opin. Food Sci."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1016\/S0925-5214(98)00087-8","article-title":"Preharvest factors influencing flavor of fresh fruit and vegetables","volume":"15","author":"Mattheis","year":"1999","journal-title":"Postharvest Biol. Technol."},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Ma, J., Li, X., Chu, Y., Yue, H., Xu, Z., Li, B., Wu, X., Gan, J., and Jia, Y. (2024). Characterization of Changes in Ripening Process of Volatile Apple Compounds Based on HS-SPME-GC-MS Analysis. Agriculture, 14.","DOI":"10.3390\/agriculture14101787"},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Li, G., Liu, J., Zhang, H., Jia, L., Liu, Y., Li, J., Zhou, S., Wang, P., Tan, M., and Shao, J. (2023). Volatile metabolome and floral transcriptome analyses reveal the volatile components of strongly fragrant progeny of Malus \u00d7 robusta. Front. Plant Sci., 14.","DOI":"10.3389\/fpls.2023.1065219"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"7904","DOI":"10.1021\/acs.jafc.2c07719","article-title":"Transcriptome and metabolomics integrated analysis reveals MdMYB94 associated with esters biosynthesis in apple (Malus \u00d7 Domestica)","volume":"71","author":"Li","year":"2023","journal-title":"J. Agric. Food Chem."},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Yang, S., Li, D., Li, S., Yang, H., and Zhao, Z. (2022). GC-MS metabolite and transcriptome analyses reveal the differences of volatile synthesis and gene expression profiling between two apple varieties. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23062939"},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Savoi, S., Santiago, A., Ordu\u00f1a, L., and Matus, J.T. (2022). Transcriptomic and metabolomic integration as a resource in grapevine to study fruit metabolite quality traits. Front. Plant Sci., 13.","DOI":"10.3389\/fpls.2022.937927"},{"key":"ref_19","unstructured":"Blanpied, G.D., and Silsby, K.J. (1992). Predicting Harvest Date Windows for Apples, Cornell Cooperative Extension."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"1296","DOI":"10.1021\/acs.jchemed.6b00891","article-title":"Determination of titratable acidity in wine using potentiometric, conductometric, and photometric methods","volume":"94","author":"Volmer","year":"2017","journal-title":"J. Chem. Educ."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"116019","DOI":"10.1016\/j.lwt.2024.116019","article-title":"Aroma formation mechanism by the drying step during Congou black tea processing: Analyses by HP-SPME and SAFE with GC-MS","volume":"198","author":"Ma","year":"2024","journal-title":"LWT"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"907","DOI":"10.1038\/s41587-019-0201-4","article-title":"Graph-based genome alignment and genotyping with HISAT2 and HISAT-genotype","volume":"37","author":"Kim","year":"2019","journal-title":"Nat. Biotechnol."},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Doppler, M., Kluger, B., Bueschl, C., Schneider, C., Krska, R., Delcambre, S., Hiller, K., Lemmens, M., and Schuhmacher, R. (2016). Stable Isotope-Assisted Evaluation of Different Extraction Solvents for Untargeted Metabolomics of Plants. Int. J. Mol. Sci., 17.","DOI":"10.3390\/ijms17071017"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1038\/nmeth.3317","article-title":"HISAT: A fast spliced aligner with low memory requirements","volume":"12","author":"Kim","year":"2015","journal-title":"Nat. Methods"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1650","DOI":"10.1038\/nprot.2016.095","article-title":"Transcript-level expression analysis of RNA-seq experiments with HISAT, StringTie and Ballgown","volume":"11","author":"Pertea","year":"2016","journal-title":"Nat. Protoc."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"290","DOI":"10.1038\/nbt.3122","article-title":"StringTie enables improved reconstruction of a transcriptome from RNA-seq reads","volume":"33","author":"Pertea","year":"2015","journal-title":"Nat. Biotechnol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1111\/j.2517-6161.1995.tb02031.x","article-title":"Controlling the false discovery rate: A practical and powerful approach to multiple testing","volume":"57","author":"Benjamini","year":"1995","journal-title":"J. R. Stat. Soc. Ser. B Methodol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1093\/bioinformatics\/btp616","article-title":"edgeR: A Bioconductor package for differential expression analysis of digital gene expression data","volume":"26","author":"Robinson","year":"2010","journal-title":"Bioinformatics"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"D545","DOI":"10.1093\/nar\/gkaa970","article-title":"KEGG: Integrating viruses and cellular organisms","volume":"49","author":"Kanehisa","year":"2021","journal-title":"Nucleic Acids Res."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"1060","DOI":"10.1038\/nprot.2011.335","article-title":"Procedures for large-scale metabolic profiling of serumand plasma using gas chromatography and liquid chromatography coupled to massspectrometry","volume":"6","author":"Dunn","year":"2011","journal-title":"Nat. Protoc."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1005","DOI":"10.1038\/nprot.2010.50","article-title":"Global metabolic profiling procedures for urine using UPLC-MS","volume":"5","author":"Want","year":"2010","journal-title":"Nat. Protoc."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"1575","DOI":"10.1007\/s11306-015-0809-4","article-title":"An integrated targeted metabolomic platform for high-throughput metabolite profiling and automated data processing","volume":"11","author":"Cai","year":"2015","journal-title":"Metabolomics"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"1469","DOI":"10.1093\/plphys\/kiab583","article-title":"Heterologous expression of Bixa orellana cleavage dioxygenase 4\u20133 drives crocin but not bixin biosynthesis","volume":"188","author":"Frusciante","year":"2022","journal-title":"Plant Physiol."},{"key":"ref_34","first-page":"498","article-title":"Analysis of metabolomic data: Tools, current strategies and future challenges for omics data integration","volume":"18","author":"Cambiaghi","year":"2017","journal-title":"Brief. Bioinform."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"6656","DOI":"10.1038\/s41467-022-34537-6","article-title":"Metabolite annotation from knowns to unknowns through knowledge-guided multi-layer metabolic networking","volume":"13","author":"Zhou","year":"2022","journal-title":"Nat. Commun."},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Langfelder, P., and Horvath, S. (2008). WGCNA: An R package for weighted correlation network analysis. BMC Bioinform., 9.","DOI":"10.1186\/1471-2105-9-559"},{"key":"ref_37","unstructured":"Langfelder, P., and Horvath, S. (2014). Tutorials for the WGCNA Package, UCLA."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"3538","DOI":"10.1002\/csc2.20473","article-title":"Co-expression gene network analysis of cold-tolerant Solanum commersonii reveals new insights in response to low temperatures","volume":"61","author":"Esposito","year":"2021","journal-title":"Crop Sci."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Iannaccone, M., Ianni, A., Contaldi, F., Esposito, S., Martino, C., Bennato, F., De Angelis, E., Grotta, L., Pomilio, F., and Giansante, D. (2019). Whole blood transcriptome analysis in ewes fed with hemp seed supplemented diet. Sci. Rep., 9.","DOI":"10.1038\/s41598-019-52712-6"},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Horvath, S., and Dong, J. (2008). Geometric interpretation of gene co-expression network analysis. PLoS Comput. Biol., 4.","DOI":"10.1371\/journal.pcbi.1000117"},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Vernocchi, P., Gili, T., Conte, F., Del Chierico, F., Conta, G., and Miccheli APutignani, L. (2020). Network analysis of gut microbiome and metabolome to discover microbiota-linked biomarkers in patients affected by non-small cell lung cancer. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21228730"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"1195","DOI":"10.1111\/hepr.13386","article-title":"Weighted gene coexpression network analysis reveals hub genes involved in cholangiocarcinoma progression and prognosis","volume":"49","author":"Tian","year":"2019","journal-title":"Hepatol. Res."},{"key":"ref_43","first-page":"1","article-title":"Molecular physiology of fruit growth in apple","volume":"47","author":"Malladi","year":"2020","journal-title":"Hortic. Rev."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"127074","DOI":"10.1016\/j.foodchem.2020.127074","article-title":"Insights into the aroma profiles and characteristic aroma of \u2018Honeycrisp\u2019 apple (Malus \u00d7 domestica)","volume":"327","author":"Yan","year":"2020","journal-title":"Food Chem."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"84","DOI":"10.1007\/s00425-003-1191-4","article-title":"Cloning and functional expression of an (E, E)-\u03b1-farnesene synthase cDNA from peel tissue of apple fruit","volume":"219","author":"Pechous","year":"2004","journal-title":"Planta"},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Zhang, H., Chen, M., Wen, H., Wang, Z., Chen, J., Fang, L., Zhang, H., Xie, Z., Jiang, D., and Cheng, Y. (2020). Transcriptomic and metabolomic analyses provide insight into the volatile compounds of citrus leaves and flowers. BMC Plant Biol., 20.","DOI":"10.1186\/s12870-019-2222-z"},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Espino-D\u00edaz, M., Sep\u00falveda, D.R., Gonz\u00e1lez-Aguilar, G., and I Olivas, G. (2016). Biochemistry of apple aroma, A review. Food Technol. Biotechnol., 54.","DOI":"10.17113\/ftb.54.04.16.4248"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/S0065-2296(08)00801-X","article-title":"Aroma volatiles: Biosynthesis and mechanisms of modulation during fruit ripening","volume":"50","author":"Defilippi","year":"2009","journal-title":"Adv. Bot. Res."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1016\/0925-5214(96)00020-8","article-title":"The effect of harvest date on aroma compound production from \u2018Golden Delicious\u2019 apple fruit and relationship to respiration and ethylene production","volume":"8","author":"Song","year":"1996","journal-title":"Postharvest Biol. Technol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1016\/j.postharvbio.2013.02.006","article-title":"Lipoxygenase-associated apple volatiles and their relationship with aroma perception during ripening","volume":"82","author":"Contreras","year":"2013","journal-title":"Postharvest Biol. Technol."},{"key":"ref_51","doi-asserted-by":"crossref","unstructured":"Song, J. (2010). Major enzymes of flavor volatiles production and regulation infresh fruits and vegetables. Enzymes in Fruit and Vegetable Processing, CRC Press.","DOI":"10.1201\/9781420094343-c3"},{"key":"ref_52","doi-asserted-by":"crossref","unstructured":"Liu, X., Hao, N., Feng, R., Meng, Z., Li, Y., and Zhao, Z. (2021). Transcriptome and metabolite profiling analyses provide insight into volatile aroma compounds of the apple cultivar \u2018Ruixue\u2019 and its parents during fruit development. BMC Plant Biol., 21.","DOI":"10.1186\/s12870-021-03032-3"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"491","DOI":"10.1016\/j.enzmictec.2006.11.025","article-title":"Lipoxygenase in fruits and vegetables: A review","volume":"40","author":"Baysal","year":"2007","journal-title":"Enzym. Microb. Technol."},{"key":"ref_54","doi-asserted-by":"crossref","unstructured":"Jesenko, E., Vidrih, R., and Zlati\u0107, E. (2025). Effects of Hexanal Supplementation on Volatile Compound Profiles and Quality Parameters of \u2018Fuji Kiku\u2019 Apples During Cold Storage and Shelf Life. Agronomy, 15.","DOI":"10.3390\/agronomy15020292"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"2627","DOI":"10.1021\/jf011152w","article-title":"Investigations of aroma volatile biosynthesis under anoxic conditions and in different tissues of \u201cRedchief Delicious\u201d apple fruit (Malus domestica Borkh.)","volume":"50","author":"Rudell","year":"2002","journal-title":"J. Agric. Food Chem."},{"key":"ref_56","unstructured":"Chan, H.W.S. (1987). Autoxidation of Unsaturated Lipids, Academic Press."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"511","DOI":"10.21273\/HORTSCI12631-17","article-title":"Effects of MAP treatment on aroma compounds and enzyme activities in flat peach during storage and shelf life","volume":"53","author":"Zhou","year":"2018","journal-title":"HortScience"},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Han, Y. (2015). Effect of Fruit Types and Temperature on Formation of Volatiles in the Lipoxygenase (LOX) Pathway, The Ohio State University.","DOI":"10.4172\/2472-0542.1000102"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"289","DOI":"10.47836\/pjtas.47.1.20","article-title":"Hexanal Treatment for Improving the Shelf-life and Quality of Fruits: A Review","volume":"47","author":"Azhar","year":"2024","journal-title":"Pertanika J. Trop. Agric. Sci."},{"key":"ref_60","doi-asserted-by":"crossref","unstructured":"Liu, G., Huang, L., and Lian, J. (2023). Alcohol acyltransferases for the biosynthesis of esters. Biotechnol. Biofuels Bioprod., 16.","DOI":"10.1186\/s13068-023-02343-x"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"286","DOI":"10.1016\/j.postharvbio.2007.07.003","article-title":"Changes in biosynthesis of aroma volatile compounds during on-tree maturation of \u2018Pink Lady\u00ae\u2019 apples","volume":"47","author":"Villatoro","year":"2008","journal-title":"Postharvest Biol. Technol."},{"key":"ref_62","doi-asserted-by":"crossref","unstructured":"Rajendran, S., Silcock, P., and Bremer, P. (2023). Flavour volatiles of fermented vegetable and fruit substrates: A review. Molecules, 28.","DOI":"10.3390\/molecules28073236"},{"key":"ref_63","doi-asserted-by":"crossref","unstructured":"Villano, C., Demurtas, O.C., Esposito, S., Granell, A., Rambla, J.L., Piombino, P., Frusciante, L., Carputo, D., Diretto, G., and Aversano, R. (2023). Integrative analysis of metabolome and transcriptome profiles to highlight aroma determinants in Aglianico and Falanghina grape berries. BMC Plant Biol., 23.","DOI":"10.1186\/s12870-023-04280-1"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"442","DOI":"10.1111\/nph.12175","article-title":"A STAY-GREEN protein S l SGR 1 regulates lycopene and \u03b2-carotene accumulation by interacting directly with S l PSY 1 during ripening processes in tomato","volume":"198","author":"Luo","year":"2013","journal-title":"New Phytol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"609","DOI":"10.1007\/s11032-011-9577-7","article-title":"Functional allelic diversity of the apple alcohol acyl-transferase gene MdAAT1 associated with fruit ester volatile contents in apple cultivars","volume":"29","author":"Dunemann","year":"2012","journal-title":"Mol. Breed."},{"key":"ref_66","unstructured":"Rapp, A., and Marais, J. (1993). Shelf life of wine: Changes in aroma substances during storage and ageing of white wines. Developments in Food Science, Elsevier."},{"key":"ref_67","doi-asserted-by":"crossref","unstructured":"Ferreira, V., de la Fuente, A., and S\u00e1enz-Navajas, M.P. (2022). Wine aroma vectors and sensory attributes. Managing Wine Quality, Woodhead Publishing.","DOI":"10.1016\/B978-0-08-102067-8.00008-7"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1016\/j.plaphy.2017.12.001","article-title":"Genome-wide analysis of carotenoid cleavage oxygenase genes and their responses to various phytohormones and abiotic stresses in apple (Malus domestica)","volume":"123","author":"Chen","year":"2018","journal-title":"Plant Physiol. Biochem."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"1215","DOI":"10.1002\/mnfr.200900608","article-title":"A review on the beneficial aspects of food processing","volume":"54","author":"Fogliano","year":"2010","journal-title":"Mol. Nutr. Food Res."}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/14\/8\/1165\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,9]],"date-time":"2025-10-09T17:13:10Z","timestamp":1760029990000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/14\/8\/1165"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,4,9]]},"references-count":69,"journal-issue":{"issue":"8","published-online":{"date-parts":[[2025,4]]}},"alternative-id":["plants14081165"],"URL":"https:\/\/doi.org\/10.3390\/plants14081165","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,4,9]]}}}