{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T14:21:55Z","timestamp":1788272515792,"version":"build-2803163510"},"reference-count":54,"publisher":"MDPI AG","issue":"11","license":[{"start":{"date-parts":[[2026,5,22]],"date-time":"2026-05-22T00:00:00Z","timestamp":1779408000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Natural Science Foundation of Guangdong Province, China","award":["2025A1515012326"],"award-info":[{"award-number":["2025A1515012326"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Fruit coloration is a key determinant of tomato quality, yet how light and temperature interact to regulate pigmentation during ripening remains unclear. Using a semi-in-fruit experimental system, we demonstrate that while high light accelerates chlorophyll degradation and lycopene accumulation at 25 \u00b0C, supra-optimal temperature (40 \u00b0C) completely abolishes lycopene biosynthesis irrespective of light conditions, primarily through transcriptional suppression of SlPSY1 and SlGGPS2. Elevated postharvest temperatures (\u226530 \u00b0C) not only change the carotenoid composition but also reduce the antioxidant capacity and vitamin C content in fruit. Temperature-switch experiments revealed a critical developmental window, days 2\u20134 after ethylene treatment, during which temperature exerts dominant control over carotenoid metabolism. Exposure to high temperature within this window irreversibly shifts pigment accumulation from lycopene to yellow\/orange carotenoids. These findings identify a temporally precise regulatory nexus integrating environmental signals with the ripening program, offering a framework for targeted temperature management to optimize tomato color and nutritional quality.<\/jats:p>","DOI":"10.3390\/ijms27114690","type":"journal-article","created":{"date-parts":[[2026,5,22]],"date-time":"2026-05-22T16:06:08Z","timestamp":1779465968000},"page":"4690","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":2,"title":["Temperature Dominates Light in Regulating Lycopene During a Critical Period in Postharvest Tomato Fruit"],"prefix":"10.3390","volume":"27","author":[{"given":"Jinyan","family":"Chen","sequence":"first","affiliation":[{"name":"College of Life Sciences, South China Agricultural University, Guangzhou 510642, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chenyang","family":"He","sequence":"additional","affiliation":[{"name":"College of Life Sciences, South China Agricultural University, Guangzhou 510642, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qu","family":"Luo","sequence":"additional","affiliation":[{"name":"College of Life Sciences, South China Agricultural University, Guangzhou 510642, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yujuan","family":"Zhong","sequence":"additional","affiliation":[{"name":"Guangdong Key Laboratory for New Technology Research of Vegetables, Vegetable Research Institute, Guangdong Academy of Agricultural Sciences, Guangzhou 510640, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yingchao","family":"Xu","sequence":"additional","affiliation":[{"name":"Guangdong Key Laboratory for New Technology Research of Vegetables, Vegetable Research Institute, Guangdong Academy of Agricultural Sciences, Guangzhou 510640, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jiayu","family":"Luo","sequence":"additional","affiliation":[{"name":"College of Life Sciences, South China Agricultural University, Guangzhou 510642, China"},{"name":"Guangdong Key Laboratory for New Technology Research of Vegetables, Vegetable Research Institute, Guangdong Academy of Agricultural Sciences, Guangzhou 510640, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Huaiyuan","family":"Li","sequence":"additional","affiliation":[{"name":"National Demonstration Center for Experimental Essentials of Plant Biology Education (SCAU)\/Basic Experiment and Practice Training Center, South China Agricultural University, Guangzhou 510642, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xuelian","family":"Zhang","sequence":"additional","affiliation":[{"name":"College of Life Sciences, South China Agricultural University, Guangzhou 510642, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,5,22]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"76","DOI":"10.1007\/s00425-023-04108-0","article-title":"Differential defensive and nutritional traits among cultivated tomato and its wild relatives shape their interactions with a specialist herbivore","volume":"257","author":"Bento","year":"2023","journal-title":"Planta"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1007\/s10142-023-01162-5","article-title":"Beyond green and red: Unlocking the genetic orchestration of tomato fruit color and pigmentation","volume":"23","author":"Naeem","year":"2023","journal-title":"Funct. Integr. Genom."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"977","DOI":"10.1016\/j.bbabio.2010.12.007","article-title":"Chlorophyll breakdown in higher plants","volume":"1807","year":"2011","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_4","first-page":"e158","article-title":"Carotenoid biosynthesis in Arabidopsis: A colorful pathway","volume":"10","year":"2012","journal-title":"Arab. Book"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"4","DOI":"10.1007\/s00344-013-9392-y","article-title":"Mechanism and significance of chlorophyll breakdown","volume":"33","author":"Christ","year":"2014","journal-title":"J. Plant Growth Regul."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"68","DOI":"10.1016\/j.molp.2014.12.007","article-title":"Carotenoid metabolism in plants","volume":"8","author":"Nisar","year":"2015","journal-title":"Mol. Plant"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"2147","DOI":"10.1105\/tpc.16.00428","article-title":"Arabidopsis STAY-GREEN, mendel\u2019s green cotyledon gene, encodes magnesium-dechelatase","volume":"28","author":"Shimoda","year":"2016","journal-title":"Plant Cell"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"940","DOI":"10.1111\/j.1365-313X.2009.03919.x","article-title":"Defect in non-yellow coloring 3, an alpha\/beta hydrolase-fold family protein, causes a stay-green phenotype during leaf senescence in rice","volume":"59","author":"Morita","year":"2009","journal-title":"Plant J."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"767","DOI":"10.1105\/tpc.108.064089","article-title":"Pheophytin pheophorbide hydrolase (pheophytinase) is involved in chlorophyll breakdown during leaf senescence in Arabidopsis","volume":"21","author":"Schelbert","year":"2009","journal-title":"Plant Cell"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"935","DOI":"10.1093\/jxb\/erv509","article-title":"Functional characterization and hormonal regulation of the PHEOPHYTINASE gene LpPPH controlling leaf senescence in perennial ryegrass","volume":"67","author":"Zhang","year":"2016","journal-title":"J. Exp. Bot."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"15259","DOI":"10.1073\/pnas.2036571100","article-title":"Chlorophyll breakdown: Pheophorbide a oxygenase is a Rieske-type iron-sulfur protein, encoded by the accelerated cell death 1 gene","volume":"100","author":"Tanner","year":"2003","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1105\/tpc.106.044404","article-title":"In vivo participation of red chlorophyll catabolite reductase in chlorophyll breakdown","volume":"19","author":"Anders","year":"2007","journal-title":"Plant Cell"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"3322","DOI":"10.1093\/jxb\/erae095","article-title":"Defective mutations in STAY-GREEN 1, PHYTOENE SYNTHASE 1, and MYB12 genes lead to formation of green ripe fruit in tomato","volume":"75","author":"Cui","year":"2024","journal-title":"J. Exp. Bot."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"1472","DOI":"10.1111\/nph.15973","article-title":"Advances and future directions in betalain metabolic engineering","volume":"224","author":"Polturak","year":"2019","journal-title":"New Phytol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"257","DOI":"10.1007\/s12038-016-9602-4","article-title":"Differential expression of carotenoid biosynthetic pathway genes in two contrasting tomato genotypes for lycopene content","volume":"41","author":"Pandurangaiah","year":"2016","journal-title":"J. Biosci."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"58","DOI":"10.1016\/j.molp.2017.09.010","article-title":"Carotenoid metabolism in plants: The role of plastids","volume":"11","author":"Sun","year":"2018","journal-title":"Mol. Plant"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"3663","DOI":"10.1111\/pce.14684","article-title":"The role of environmental stress in fruit pigmentation","volume":"46","author":"Espley","year":"2023","journal-title":"Plant Cell Environ."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"108","DOI":"10.1016\/j.biotechadv.2009.10.003","article-title":"Light signaling genes and their manipulation towards modulation of phytonutrient content in tomato fruits","volume":"28","author":"Azari","year":"2010","journal-title":"Biotechnol. Adv."},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Llorente, B., D\u2019Andrea, L., and Rodriguez-Concepcion, M. (2016). Evolutionary recycling of light signaling components in fleshy fruits: New insights on the role of pigments to monitor ripening. Front. Plant Sci., 7, Erratum in Front Plant Sci.\u00a02022, 13, 900067.","DOI":"10.3389\/fpls.2022.900067"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"135690","DOI":"10.1016\/j.foodchem.2023.135690","article-title":"Unravelling effects of red\/far-red light on nutritional quality and the role and mechanism in regulating lycopene synthesis in postharvest cherry tomatoes","volume":"414","author":"Song","year":"2023","journal-title":"Food Chem."},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Toledo-Ortiz, G., Johansson, H., Lee, K.P., Bou-Torrent, J., Stewart, K., Steel, G., Rodr\u00edguez-Concepci\u00f3n, M., and Halliday, K.J. (2014). The HY5-PIF regulatory module coordinates light and temperature control of photosynthetic gene transcription. PLoS Genet., 10.","DOI":"10.1371\/journal.pgen.1004416"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"e148","DOI":"10.1199\/tab.0148","article-title":"Phytochrome signaling mechanisms","volume":"9","author":"Li","year":"2011","journal-title":"Arab. Book"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1002\/jsfa.1370","article-title":"Effects of environmental factors and agricultural techniques on antioxidant content of tomatoes","volume":"83","author":"Dumas","year":"2003","journal-title":"J. Sci. Food Agric."},{"key":"ref_24","doi-asserted-by":"crossref","unstructured":"Wang, Y., Zhang, C., Xu, B., Fu, J., Du, Y., Fang, Q., Dong, B., and Zhao, H. (2022). Temperature regulation of carotenoid accumulation in the petals of sweet osmanthus via modulating expression of carotenoid biosynthesis and degradation genes. BMC Genom., 23.","DOI":"10.1186\/s12864-022-08643-0"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"6930","DOI":"10.1002\/jsfa.12054","article-title":"Low and high storage temperature inhibited the coloration of mandarin fruit (Citrus unshiu Marc.) with different mechanism","volume":"102","author":"Ge","year":"2022","journal-title":"J. Sci. Food Agric."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"325","DOI":"10.1093\/nar\/30.1.325","article-title":"PlantCARE, a database of plant cis-acting regulatory elements and a portal to tools for in silico analysis of promoter sequences","volume":"30","author":"Lescot","year":"2002","journal-title":"Nucleic Acids Res."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1016\/j.scienta.2019.03.019","article-title":"Optimal red light irradiation time to increase health-promoting compounds in tomato fruit postharvest","volume":"251","author":"Panjai","year":"2019","journal-title":"Sci. Hortic."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"2211","DOI":"10.1111\/pce.13854","article-title":"A transcriptomic, metabolomic and cellular approach to the physiological adaptation of tomato fruit to high temperature","volume":"44","author":"Almeida","year":"2021","journal-title":"Plant Cell Environ."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"590","DOI":"10.1016\/S2095-3119(18)62062-3","article-title":"Supplemental blue and red light promote lycopene synthesis in tomato fruits","volume":"18","author":"Xie","year":"2019","journal-title":"J. Integr. Agric."},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Pola, W., Sugaya, S., and Photchanachai, S. (2020). Influence of postharvest temperatures on carotenoid biosynthesis and phytochemicals in mature green chili (Capsicum annuum L.). Antioxidants, 9.","DOI":"10.3390\/antiox9030203"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"135602","DOI":"10.1016\/j.foodchem.2023.135602","article-title":"High temperature elevates carotenoid accumulation of banana fruit via upregulation of MaEIL9 module","volume":"412","author":"Zhu","year":"2023","journal-title":"Food Chem."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"869","DOI":"10.1104\/pp.20.00019","article-title":"Phytochrome-dependent temperature perception modulates isoprenoid metabolism","volume":"183","author":"Bianchetti","year":"2020","journal-title":"Plant Physiol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"34","DOI":"10.1016\/j.micres.2019.05.003","article-title":"Effect of growth temperature on biosynthesis and accumulation of carotenoids in cyanobacterium Anabaena sp. PCC 7120 under diazotrophic conditions","volume":"226","author":"Bujas","year":"2019","journal-title":"Microbiol. Res."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.plantsci.2016.04.016","article-title":"Chlorophyll loss associated with heat-induced senescence in bentgrass","volume":"249","author":"Jespersen","year":"2016","journal-title":"Plant Sci."},{"key":"ref_35","doi-asserted-by":"crossref","unstructured":"Li, Z., Tang, M., Hassan, M.J., Zhang, Y., Han, L., and Peng, Y. (2021). Adaptability to high temperature and stay-green genotypes associated with variations in antioxidant, chlorophyll metabolism, and \u03b3-aminobutyric acid accumulation in creeping bentgrass species. Front. Plant Sci., 12.","DOI":"10.3389\/fpls.2021.750728"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"4051","DOI":"10.1093\/jxb\/erp238","article-title":"Accumulation of soluble sugars in peel at high temperature leads to stay-green ripe banana fruit","volume":"60","author":"Yang","year":"2009","journal-title":"J. Exp. Bot."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1016\/j.scienta.2014.10.026","article-title":"Characterization of the stage dependency of high temperature on green ripening reveals a distinct chlorophyll degradation regulation in banana fruit","volume":"180","author":"Du","year":"2014","journal-title":"Sci. Hortic."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"110980","DOI":"10.1016\/j.postharvbio.2019.110980","article-title":"Imbalanced expression of stay-green 1 alleles in banana AAB\/ABB cultivars prevents high-temperature-induced green ripening as in AAA Cavendish fruit","volume":"158","author":"Wu","year":"2019","journal-title":"Postharvest Biol. Technol."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Su, L.Y., Diretto, E., Purgatto, E., Danoun, S., Zouine, M., Li, Z.G., Roustan, J.P., Bouzayen, M., Giuliano, G., and Chervin, C. (2015). Carotenoid accumulation during tomato fruit ripening is modulated by the auxin-ethylene balance. BMC Plant Biol., 15.","DOI":"10.1186\/s12870-015-0495-4"},{"key":"ref_40","unstructured":"Gross, K.C., Wang, C.Y., and Saltveit, M.E. (2016). Ethylene effects, The Commercial Storage of Fruits, Vegetables, and Florist and Nursery Stocks."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"773","DOI":"10.21273\/HORTTECH04668-20","article-title":"Physiological response of mature green tomatoes to treatment with ethylene at high temperature","volume":"30","author":"Loayza","year":"2020","journal-title":"HortTechnology"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"368","DOI":"10.1016\/S0176-1617(11)80064-7","article-title":"The effect of high temperature and ethylene treatment on the ripening of tomatoes","volume":"136","author":"Yang","year":"1990","journal-title":"J. Plant Physiol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"16674","DOI":"10.1038\/s41598-017-16851-y","article-title":"Regulation of ethylene-responsive SlWRKYs involved in color change during tomato fruit ripening","volume":"7","author":"Wang","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"3337","DOI":"10.1093\/jxb\/erae121","article-title":"Galactinol synthase 2 influences the metabolism of chlorophyll, carotenoid, and ethylene in tomato fruits","volume":"75","author":"Zhang","year":"2024","journal-title":"J. Exp. Bot."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"936","DOI":"10.1111\/j.1365-313X.2010.04384.x","article-title":"A tomato (Solanum lycopersicum) APETALA2\/ERF gene, SlAP2a, is a negative regulator of fruit ripening","volume":"64","author":"Chung","year":"2010","journal-title":"Plant J."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"1408","DOI":"10.1093\/plcell\/koad030","article-title":"Proteasomal degradation of MaMYB60 mediated by the E3 ligase MaBAH1 causes high temperature-induced repression of chlorophyll catabolism and green ripening in banana","volume":"35","author":"Wei","year":"2023","journal-title":"Plant Cell"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"32","DOI":"10.1016\/j.foodres.2012.11.024","article-title":"Processing tomato pulp in the presence of lipids: The impact on lycopene bioaccessibility","volume":"51","author":"Colle","year":"2013","journal-title":"Food Res. Int."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"992","DOI":"10.1105\/tpc.010405","article-title":"Zeaxanthin accumulation in the absence of a functional xanthophyll cycle protects Chlamydomonas reinhardtii from photooxidative stress","volume":"15","author":"Baroli","year":"2003","journal-title":"Plant Cell"},{"key":"ref_49","doi-asserted-by":"crossref","unstructured":"Guo, F., Zhou, W., Zhang, J.C., Xu, Q., and Deng, X.X. (2012). Effect of the citrus lycopene \u03b2-cyclase transgene on carotenoid metabolism in transgenic tomato fruits. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0032221"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1038\/s41438-020-00431-9","article-title":"SlGRAS4 accelerates fruit ripening by regulating ethylene biosynthesis genes and SlMADS1 in tomato","volume":"8","author":"Liu","year":"2021","journal-title":"Hortic. Res."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"72","DOI":"10.1016\/j.phytochem.2014.11.007","article-title":"Fruits from ripening impaired, chlorophyll degraded and jasmonate insensitive tomato mutants have altered tocopherol content and composition","volume":"111","author":"Almeida","year":"2015","journal-title":"Phytochemistry"},{"key":"ref_52","doi-asserted-by":"crossref","unstructured":"Ponchel, F., Toomes, C., Bransfield, K., Leong, F.T., Douglas, S.H., Field, S.L., Bell, S.M., Combaret, V., Puisieux, A., and Mighell, A.J. (2003). Real-time PCR based on SYBR-Green I fluorescence: An alternative to the TaqMan assay for a relative quantification of gene rearrangements, gene amplifications and micro gene deletions. BMC Biotechnol., 3.","DOI":"10.1186\/1472-6750-3-18"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"129","DOI":"10.1016\/j.foodchem.2011.03.006","article-title":"Purification and characterisation of aquamarine blue pigment from the shells of abalone (Haliotis discus hannai Ino)","volume":"128","author":"Cai","year":"2011","journal-title":"Food Chem."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"1751","DOI":"10.1111\/1750-3841.15682","article-title":"Carotenoid composition and antioxidant activities of Chinese orange-colored tomato cultivars and the effects of thermal processing on the bioactive components","volume":"86","author":"Yang","year":"2021","journal-title":"J. Food Sci."}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/11\/4690\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,1]],"date-time":"2026-06-01T04:10:31Z","timestamp":1780287031000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/11\/4690"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,5,22]]},"references-count":54,"journal-issue":{"issue":"11","published-online":{"date-parts":[[2026,6]]}},"alternative-id":["ijms27114690"],"URL":"https:\/\/doi.org\/10.3390\/ijms27114690","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,5,22]]}}}