{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,22]],"date-time":"2026-09-22T20:42:27Z","timestamp":1790109747393,"version":"4.0.1"},"reference-count":121,"publisher":"Oxford University Press (OUP)","issue":"4","license":[{"start":{"date-parts":[[2020,4,2]],"date-time":"2020-04-02T00:00:00Z","timestamp":1585785600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/journals\/pages\/open_access\/funder_policies\/chorus\/standard_publication_model"}],"funder":[{"DOI":"10.13039\/100012359","name":"DBT","doi-asserted-by":"publisher","award":["BT\/04\/NE\/2009"],"award-info":[{"award-number":["BT\/04\/NE\/2009"]}],"id":[{"id":"10.13039\/100012359","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2020,7,29]]},"abstract":"<jats:title>Abstract<\/jats:title><jats:p>Carotenoids are mostly C40 terpenoids that participate in several important functions in plants including photosynthesis, responses to various forms of stress, signal transduction and photoprotection. While the antioxidant potential of carotenoids is of particular importance for human health, equally important is the role of \u03b2-carotene as the precursor for vitamin A in the human diet. Rice, which contributes upto 40% of dietary energy for mankind, contains very low level of \u03b2-carotene, thereby making it an important crop for enhancing \u03b2-carotene accumulation in its grains and consequently targeting vitamin A deficiency. Biosynthesis of carotenoids in the endosperm of white rice is blocked at the first enzymatic step wherein geranylgeranyl diphosphate is converted to phytoene by the action of phytoene synthase (PSY). Strategies aimed at enhancing \u03b2-carotene levels in the endosperm of white rice identified Narcissus pseudonarcissus (npPSY) and bacterial CRT1 as the regulators of the carotenoid biosynthetic pathway in rice. Besides transcriptional regulation of PSY, posttranscriptional regulation of PSY expression by OR gene, molecular synergism between \u03b5-LCY and \u03b2-LCY and epigenetic control of CRITSO through SET DOMAIN containing protein appear to be the other regulatory nodes which regulate carotenoid biosynthesis and accumulation in rice grains. In this review, we elucidate a comprehensive and deeper understanding of the regulatory mechanisms of carotenoid metabolism in crops that will enable us to identify an effective tool to alleviate carotenoid content in rice grains.<\/jats:p>","DOI":"10.1093\/bfgp\/elaa007","type":"journal-article","created":{"date-parts":[[2020,2,27]],"date-time":"2020-02-27T12:19:22Z","timestamp":1582805962000},"page":"324-335","source":"Crossref","is-referenced-by-count":8,"title":["A multifocal approach towards understanding the complexities of carotenoid biosynthesis and accumulation in rice grains"],"prefix":"10.1093","volume":"19","author":[{"given":"Upasna","family":"Chettry","sequence":"first","affiliation":[{"name":"Department of Botany, North-Eastern Hill University, Shillong 793022, India"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nikhil K","family":"Chrungoo","sequence":"additional","affiliation":[{"name":"Department of Botany, North-Eastern Hill University, Shillong 793022, India"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2020,4,2]]},"reference":[{"issue":"2","key":"2020072910370220200_ref1","doi-asserted-by":"crossref","first-page":"1710","DOI":"10.3390\/molecules16021710","article-title":"Carotenoids and their isomers: color pigments in fruits and vegetables","volume":"16","author":"Khoo","year":"2011","journal-title":"Molecules"},{"issue":"5","key":"2020072910370220200_ref2","doi-asserted-by":"crossref","first-page":"266","DOI":"10.1016\/j.tplants.2010.02.003","article-title":"Source to sink: regulation of carotenoid biosynthesis in plants","volume":"15","author":"Cazzonelli","year":"2010","journal-title":"Trends Plant Sci"},{"key":"2020072910370220200_ref3","doi-asserted-by":"crossref","first-page":"e0158","DOI":"10.1199\/tab.0158","article-title":"Carotenoid biosynthesis in Arabidopsis: a colorful pathway","volume":"10","author":"Ruiz-Sola","year":"2012","journal-title":"Arabidopsis Book"},{"issue":"4","key":"2020072910370220200_ref4","doi-asserted-by":"crossref","first-page":"597","DOI":"10.1111\/tpj.12386","article-title":"Carotenoid oxidation products as stress signals in plants","volume":"79","author":"Havaux","year":"2014","journal-title":"Plant J"},{"issue":"9","key":"2020072910370220200_ref5","doi-asserted-by":"crossref","first-page":"792","DOI":"10.1016\/j.tplants.2016.06.001","article-title":"Synthesis and function of apocarotenoid signals in plants","volume":"21","author":"Hou","year":"2016","journal-title":"Trends Plant Sci"},{"issue":"3","key":"2020072910370220200_ref6","doi-asserted-by":"crossref","first-page":"375","DOI":"10.1111\/tpj.13268","article-title":"Transient expression in Nicotiana benthamiana for rapid functional analysis of genes involved in non-photochemical quenching and carotenoid biosynthesis","volume":"88","author":"Leonelli","year":"2016","journal-title":"Plant J"},{"issue":"20","key":"2020072910370220200_ref7","doi-asserted-by":"crossref","first-page":"4921","DOI":"10.1093\/jxb\/ery242","article-title":"A role for \u03b2, \u03b2-xanthophylls in Arabidopsis UV-B photoprotection","volume":"69","author":"Emiliani","year":"2018","journal-title":"J Exp Bot"},{"key":"2020072910370220200_ref8","doi-asserted-by":"crossref","first-page":"466","DOI":"10.3390\/nu6020466","article-title":"Potential role of carotenoids as antioxidants in human health and disease","volume":"6","author":"Fiedor","year":"2014","journal-title":"Nutrients"},{"issue":"1","key":"2020072910370220200_ref9","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.bbabio.2005.05.008","article-title":"Cyclic endoperoxides of \u03b2-carotene, potential pro-oxidants, as products of chemical quenching of singlet oxygen","volume":"1709","author":"Fiedor","year":"2005","journal-title":"Biochim Biophys Acta Bioenerg"},{"key":"2020072910370220200_ref10","author":"FAOSTAT"},{"key":"2020072910370220200_ref11","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1016\/j.abb.2018.06.001","article-title":"Carotenoids in human nutrition and health","volume":"652","author":"Eggersdorfer","year":"2018","journal-title":"Arch Biochem Biophys"},{"issue":"4","key":"2020072910370220200_ref12","doi-asserted-by":"crossref","first-page":"482","DOI":"10.1038\/nbt1082","article-title":"Improving the nutritional value of Golden Rice through increased pro-vitamin A content","volume":"23","author":"Paine","year":"2005","journal-title":"Nat Biotechnol"},{"issue":"1","key":"2020072910370220200_ref13","doi-asserted-by":"crossref","first-page":"441","DOI":"10.1104\/pp.104.057927","article-title":"Why is Golden Rice golden (yellow) instead of red?","volume":"138","author":"Schaub","year":"2005","journal-title":"Plant Physiol"},{"issue":"33","key":"2020072910370220200_ref14","doi-asserted-by":"crossref","first-page":"464","DOI":"10.1111\/tpj.12384","article-title":"An in vitro system for the rapid functional characterization of genes involved in carotenoid biosynthesis and accumulation","volume":"77","author":"Bai","year":"2014","journal-title":"Plant J"},{"key":"2020072910370220200_ref15","doi-asserted-by":"crossref","first-page":"1622","DOI":"10.3389\/fpls.2016.01622","article-title":"Metabolic regulation of carotenoid-enriched Golden Rice line","volume":"7","author":"Gayen","year":"2016","journal-title":"Front Plant Sci"},{"issue":"12","key":"2020072910370220200_ref16","doi-asserted-by":"crossref","first-page":"1440","DOI":"10.1016\/j.molp.2018.09.007","article-title":"From Golden Rice to aSTARice: bioengineering astaxanthin biosynthesis in rice endosperm","volume":"11","author":"Zhu","year":"2018","journal-title":"Mol Plant"},{"issue":"1","key":"2020072910370220200_ref17","first-page":"96","article-title":"Plant wealth of Northeast India with reference to ethnobotany","volume":"8","author":"Mao","year":"2009","journal-title":"Indian J Tradit Know"},{"issue":"4","key":"2020072910370220200_ref18","doi-asserted-by":"crossref","first-page":"348","DOI":"10.3746\/pnf.2014.19.4.348","article-title":"Protective effects of black rice extracts on oxidative stress induced by tert-butyl hydroperoxide in HepG2 cells","volume":"19","author":"Lee","year":"2014","journal-title":"Prev Nutr Food Sci"},{"issue":"5","key":"2020072910370220200_ref19","doi-asserted-by":"crossref","first-page":"964","DOI":"10.1021\/jf00125a002","article-title":"Rice product volatiles: a review","volume":"32","author":"Maga","year":"1984","journal-title":"J Agric Food Chem"},{"issue":"2","key":"2020072910370220200_ref20","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1002\/(SICI)1097-0010(199602)70:2<151::AID-JSFA478>3.0.CO;2-U","article-title":"Comparative studies on volatile components of non-fragrant and fragrant rices","volume":"70","author":"Widjaja","year":"1996","journal-title":"J Sci Food Agric"},{"issue":"5","key":"2020072910370220200_ref21","doi-asserted-by":"crossref","first-page":"1101","DOI":"10.1021\/jf0108720","article-title":"Comparison of key aroma compounds in cooked brown rice varieties based on aroma extract dilution analyses","volume":"50","author":"Jezussek","year":"2002","journal-title":"J Agric Food Chem"},{"key":"2020072910370220200_ref22","first-page":"958","article-title":"2-Acetyl-1-pyrroline: an important aroma component of cooked rice","volume":"23","author":"Buttery","year":"1982","journal-title":"Chem Ind"},{"issue":"13","key":"2020072910370220200_ref23","doi-asserted-by":"crossref","first-page":"4696","DOI":"10.1021\/jf0606609","article-title":"Anthocyanin composition in black, blue, pink, purple, and red cereal grains","volume":"54","author":"Abdel-Aal","year":"2006","journal-title":"J Agric Food Chem"},{"issue":"3","key":"2020072910370220200_ref24","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1016\/j.tplants.2008.12.004","article-title":"Not just a grain of rice: the quest for quality","volume":"14","author":"Fitzgerald","year":"2009","journal-title":"Trends Plant Sci"},{"issue":"24","key":"2020072910370220200_ref25","doi-asserted-by":"crossref","first-page":"12804","DOI":"10.1021\/jf103277g","article-title":"Variation and correlation analysis of flavonoids and carotenoids in Korean pigmented rice (Oryza sativa L.) cultivars","volume":"58","author":"Kim","year":"2010","journal-title":"J Agric Food Chem"},{"issue":"4","key":"2020072910370220200_ref26","doi-asserted-by":"crossref","first-page":"1077","DOI":"10.1021\/jf0623992","article-title":"Rapid discrimination of scented rice by solid-phase microextraction, mass spectrometry, and multivariate analysis used as a mass sensor","volume":"55","author":"Laguerre","year":"2007","journal-title":"J Agric Food Chem"},{"issue":"8","key":"2020072910370220200_ref27","doi-asserted-by":"crossref","first-page":"1591","DOI":"10.1023\/B:BIOC.0000021330.81998.bb","article-title":"Agro-biodiversity in subsistence-oriented farming systems in a Philippine upland region: nutritional considerations","volume":"13","author":"Frei","year":"2004","journal-title":"Biodivers Conserv"},{"issue":"1","key":"2020072910370220200_ref28","doi-asserted-by":"crossref","first-page":"1631","DOI":"10.1038\/s41598-018-38233-8","article-title":"Comparative transcriptomics approach in elucidation of carotenoid biosynthesis regulation in grains of rice (Oryza sativa L.)","volume":"9","author":"Chettry","year":"2019","journal-title":"Sci Rep"},{"key":"2020072910370220200_ref29","first-page":"1","volume-title":"Microbial Carotenoids from Bacteria and Microalgae","author":"Paniagua-Michel","year":"2012"},{"issue":"1\u20132","key":"2020072910370220200_ref30","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1023\/B:PHYT.0000004180.25066.62","article-title":"Quantitative assessment of crosstalk between the two isoprenoid biosynthesis pathways in plants by NMR spectroscopy","volume":"2","author":"Schuhr","year":"2003","journal-title":"Phytochem Rev"},{"issue":"01","key":"2020072910370220200_ref31","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1055\/s-2004-830446","article-title":"Transport of isoprenoid intermediates across chloroplast envelope membranes","volume":"7","author":"Fl\u00fcgge","year":"2005","journal-title":"Plant Biol"},{"issue":"6","key":"2020072910370220200_ref32","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1016\/j.plipres.2005.09.003","article-title":"Biogenesis, molecular regulation and function of plant isoprenoids","volume":"44","author":"Bouvier","year":"2005","journal-title":"Prog Lipid Res"},{"issue":"2","key":"2020072910370220200_ref33","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1105\/tpc.010303","article-title":"Cloning of tangerine from tomato reveals a carotenoid isomerase essential for the production of \u03b2-carotene and xanthophylls in plants","volume":"14","author":"Isaacson","year":"2002","journal-title":"Plant Cell"},{"issue":"11\u201312","key":"2020072910370220200_ref34","doi-asserted-by":"crossref","first-page":"1045","DOI":"10.1515\/znc-1991-11-1219","article-title":"Functional complementation in Escherichia coli of different phytoene desaturase genes and analysis of accumulated carotenes","volume":"46","author":"Linden","year":"1991","journal-title":"Z Naturforsch C"},{"issue":"5","key":"2020072910370220200_ref35","doi-asserted-by":"crossref","first-page":"1071","DOI":"10.1046\/j.1365-313X.1997.11051071.x","article-title":"Transgenic rice (Oryza sativa) endosperm expressing daffodil (Narcissus pseudonarcissus) phytoene synthase accumulates phytoene, a key intermediate of provitamin A biosynthesis","volume":"11","author":"Burkhardt","year":"1997","journal-title":"Plant J"},{"issue":"5451","key":"2020072910370220200_ref36","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1126\/science.287.5451.303","article-title":"Engineering the provitamin A (\u03b2-carotene) biosynthetic pathway into (carotenoid-free) rice endosperm","volume":"287","author":"Ye","year":"2000","journal-title":"Science"},{"issue":"5","key":"2020072910370220200_ref37","doi-asserted-by":"crossref","first-page":"849","DOI":"10.1111\/pbi.13059","article-title":"Enhancing carotenoid biosynthesis in rice endosperm by metabolic engineering","volume":"17","author":"Tian","year":"2019","journal-title":"Plant Biotechnol J"},{"issue":"3","key":"2020072910370220200_ref38","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1007\/s11816-017-0440-0","article-title":"Effect of codon optimization on the enhancement of the \u03b2-carotene contents in rice endosperm","volume":"11","author":"Jeong","year":"2017","journal-title":"Plant Biotechnol Rep"},{"issue":"1","key":"2020072910370220200_ref39","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1007\/s11248-016-9977-x","article-title":"Reconstruction of the astaxanthin biosynthesis pathway in rice endosperm reveals a metabolic bottleneck at the level of endogenous \u03b2-carotene hydroxylase activity","volume":"26","author":"Bai","year":"2017","journal-title":"Transgenic Res"},{"issue":"1","key":"2020072910370220200_ref40","doi-asserted-by":"crossref","first-page":"6883","DOI":"10.1038\/s41598-017-07198-5","article-title":"Single genetic locus improvement of iron, zinc and \u03b2-carotene content in rice grains","volume":"7","author":"Singh","year":"2017","journal-title":"Sci Rep"},{"issue":"6","key":"2020072910370220200_ref41","doi-asserted-by":"crossref","first-page":"846","DOI":"10.1007\/s004250000352","article-title":"Regulation and activation of phytoene synthase, a key enzyme in carotenoid biosynthesis, during photomorphogenesis","volume":"211","author":"Welsch","year":"2000","journal-title":"Planta"},{"issue":"3","key":"2020072910370220200_ref42","doi-asserted-by":"crossref","first-page":"625","DOI":"10.1046\/j.1365-313X.1997.d01-16.x","article-title":"Light-dependent regulation of carotenoid biosynthesis occurs at the level of phytoene synthase expression and is mediated by phytochrome in Sinapis alba and Arabidopsis thaliana seedlings","volume":"12","author":"Von Lintig","year":"1997","journal-title":"Plant J"},{"issue":"377","key":"2020072910370220200_ref43","doi-asserted-by":"crossref","first-page":"2107","DOI":"10.1093\/jxb\/erf059","article-title":"Regulation of carotenoid formation during tomato fruit ripening and development","volume":"53","author":"Bramley","year":"2002","journal-title":"J Exp Bot"},{"issue":"25","key":"2020072910370220200_ref44","doi-asserted-by":"crossref","first-page":"11626","DOI":"10.1073\/pnas.0914428107","article-title":"Direct regulation of phytoene synthase gene expression and carotenoid biosynthesis by phytochrome-interacting factors","volume":"107","author":"Toledo-Ortiz","year":"2010","journal-title":"Proc Natl Acad Sci"},{"issue":"4","key":"2020072910370220200_ref45","doi-asserted-by":"crossref","first-page":"687","DOI":"10.1023\/A:1006256302570","article-title":"Phytoene synthase-2 enzyme activity in tomato does not contribute to carotenoid synthesis in ripening fruit","volume":"40","author":"Fraser","year":"1999","journal-title":"Plant Mol Biol"},{"issue":"1","key":"2020072910370220200_ref46","doi-asserted-by":"crossref","first-page":"367","DOI":"10.1104\/pp.108.117028","article-title":"A third phytoene synthase is devoted to abiotic stress-induced abscisic acid formation in rice and defines functional diversification of phytoene synthase genes","volume":"147","author":"Welsch","year":"2008","journal-title":"Plant Physiol"},{"issue":"9","key":"2020072910370220200_ref47","doi-asserted-by":"crossref","first-page":"3725","DOI":"10.1105\/tpc.112.104174","article-title":"Plastid localization of the key carotenoid enzyme phytoene synthase is altered by isozyme, allelic variation, and activity","volume":"24","author":"Shumskaya","year":"2012","journal-title":"Plant Cell"},{"issue":"4","key":"2020072910370220200_ref48","doi-asserted-by":"crossref","first-page":"588","DOI":"10.1111\/j.1365-313X.2009.03899.x","article-title":"Novel lycopene epsilon cyclase activities in maize revealed through perturbation of carotenoid biosynthesis","volume":"59","author":"Bai","year":"2009","journal-title":"Plant J"},{"issue":"5861","key":"2020072910370220200_ref49","doi-asserted-by":"crossref","first-page":"330","DOI":"10.1126\/science.1150255","article-title":"Natural genetic variation in lycopene epsilon cyclase tapped for maize biofortification","volume":"319","author":"Harjes","year":"2008","journal-title":"Science"},{"issue":"3","key":"2020072910370220200_ref50","doi-asserted-by":"crossref","first-page":"1568","DOI":"10.1104\/pp.111.181107","article-title":"The tomato MADS-box transcription factor RIPENING INHIBITOR interacts with promoters involved in numerous ripening processes in a COLORLESS NONRIPENING-dependent manner","volume":"157","author":"Martel","year":"2011","journal-title":"Plant Physiol"},{"issue":"3","key":"2020072910370220200_ref51","doi-asserted-by":"crossref","first-page":"1049","DOI":"10.1111\/nph.13647","article-title":"An R2R3-MYB transcription factor regulates carotenoid pigmentation in Mimulus lewisii flowers","volume":"209","author":"Sagawa","year":"2016","journal-title":"New Phytol"},{"key":"2020072910370220200_ref52","first-page":"1073","article-title":"Transcription factor RAP2.2 and its interacting partner SINAT2: stable elements in the carotenogenesis of Arabidopsis leaves","volume-title":"Plant Physiol","author":"Welsch","year":"2007"},{"issue":"4","key":"2020072910370220200_ref53","doi-asserted-by":"crossref","first-page":"2657","DOI":"10.1104\/pp.17.01830","article-title":"The citrus transcription factor CsMADS6 modulates carotenoid metabolism by directly regulating carotenogenic genes","volume":"176","author":"Lu","year":"2018","journal-title":"Plant Physiol"},{"issue":"11","key":"2020072910370220200_ref54","doi-asserted-by":"crossref","first-page":"3141","DOI":"10.1093\/jxb\/erv120","article-title":"Combined transcript, proteome, and metabolite analysis of transgenic maize seeds engineered for enhanced carotenoid synthesis reveals pleotropic effects in core metabolism","volume":"66","author":"Decourcelle","year":"2015","journal-title":"J Exp Bot"},{"issue":"3","key":"2020072910370220200_ref55","doi-asserted-by":"crossref","first-page":"830","DOI":"10.1104\/pp.18.01122","article-title":"Changing form and function through carotenoids and synthetic biology","volume":"79","author":"Wurtzel","year":"2019","journal-title":"Plant Physiol"},{"issue":"8","key":"2020072910370220200_ref56","doi-asserted-by":"crossref","first-page":"1795","DOI":"10.1105\/tpc.012526","article-title":"Contrasting effects of selection on sequence diversity and linkage disequilibrium at two phytoene synthase loci","volume":"15","author":"Palaisa","year":"2003","journal-title":"Plant Cell"},{"issue":"26","key":"2020072910370220200_ref57","doi-asserted-by":"crossref","first-page":"9885","DOI":"10.1073\/pnas.0307839101","article-title":"Long-range patterns of diversity and linkage disequilibrium surrounding the maize Y1 gene are indicative of an asymmetric selective sweep","volume":"101","author":"Palaisa","year":"2004","journal-title":"Proc Natl Acad Sci"},{"issue":"2","key":"2020072910370220200_ref58","doi-asserted-by":"crossref","first-page":"1181","DOI":"10.1104\/pp.107.098996","article-title":"Maize Y9 encodes a product essential for 15-cis-\u03b6-carotene isomerization","volume":"144","author":"Li","year":"2007","journal-title":"Plant Physiol"},{"key":"2020072910370220200_ref59","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1016\/j.pbi.2017.03.011","article-title":"Illuminating colors: regulation of carotenoid biosynthesis and accumulation by light","volume":"37","author":"Llorente","year":"2017","journal-title":"Curr Opin Plant Biol"},{"issue":"3","key":"2020072910370220200_ref60","doi-asserted-by":"crossref","first-page":"1333","DOI":"10.1104\/pp.107.111120","article-title":"PSY3, a new member of the phytoene synthase gene family conserved in the Poaceae and regulator of abiotic stress-induced root carotenogenesis","volume":"146","author":"Li","year":"2008","journal-title":"Plant Physiol"},{"issue":"3","key":"2020072910370220200_ref61","doi-asserted-by":"crossref","first-page":"1334","DOI":"10.1104\/pp.108.122119","article-title":"The maize phytoene synthase gene family: overlapping roles for carotenogenesis in endosperm, photomorphogenesis, and thermal stress tolerance","volume":"147","author":"Li","year":"2008","journal-title":"Plant Physiol"},{"key":"2020072910370220200_ref62","doi-asserted-by":"crossref","first-page":"253","DOI":"10.1146\/annurev-arplant-043015-111737","article-title":"Plastoglobuli: plastid microcompartments with integrated functions in metabolism, plastid developmental transitions, and environmental adaptation","volume":"68","author":"van Wijk","year":"2017","journal-title":"Annu Rev Plant Biol"},{"issue":"4","key":"2020072910370220200_ref63","doi-asserted-by":"crossref","first-page":"1550","DOI":"10.1104\/pp.15.00243","article-title":"Tissue-specific apocarotenoid glycosylation contributes to carotenoid homeostasis in Arabidopsis leaves","volume":"168","author":"L\u00e4tari","year":"2015","journal-title":"Plant Physiol"},{"issue":"11","key":"2020072910370220200_ref64","doi-asserted-by":"crossref","first-page":"3558","DOI":"10.1073\/pnas.1420831112","article-title":"Arabidopsis OR proteins are the major posttranscriptional regulators of phytoene synthase in controlling carotenoid biosynthesis","volume":"112","author":"Zhou","year":"2015","journal-title":"Proc Natl Acad Sci"},{"issue":"1","key":"2020072910370220200_ref65","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1016\/j.molp.2017.11.003","article-title":"Clp protease and OR directly control the proteostasis of phytoene synthase, the crucial enzyme for carotenoid biosynthesis in Arabidopsis","volume":"11","author":"Welsch","year":"2018","journal-title":"Mol Plant"},{"issue":"1","key":"2020072910370220200_ref66","doi-asserted-by":"crossref","first-page":"376","DOI":"10.1104\/pp.16.01256","article-title":"Distinct mechanisms of the ORANGE protein in controlling carotenoid flux","volume":"173","author":"Chayut","year":"2017","journal-title":"Plant Physiol"},{"issue":"10","key":"2020072910370220200_ref67","doi-asserted-by":"crossref","first-page":"4183","DOI":"10.1105\/tpc.113.113001","article-title":"Arabidopsis J-protein J20 delivers the first enzyme of the plastidial isoprenoid pathway to protein quality control","volume":"25","author":"Pulido","year":"2013","journal-title":"Plant Cell"},{"issue":"1","key":"2020072910370220200_ref68","doi-asserted-by":"crossref","first-page":"e1005824","DOI":"10.1371\/journal.pgen.1005824","article-title":"Specific Hsp100 chaperones determine the fate of the first enzyme of the plastidial isoprenoid pathway for either refolding or degradation by the stromal Clp protease in Arabidopsis","volume":"12","author":"Pulido","year":"2016","journal-title":"PLoS Genet"},{"issue":"1","key":"2020072910370220200_ref69","doi-asserted-by":"crossref","first-page":"252","DOI":"10.1111\/nph.13580","article-title":"Arabidopsis GERANYLGERANYL DIPHOSPHATE SYNTHASE 11 is a hub isozyme required for the production of most photosynthesis-related isoprenoids","volume":"209","author":"Ruiz-Sola","year":"2016","journal-title":"New Phytol"},{"issue":"6","key":"2020072910370220200_ref70","doi-asserted-by":"crossref","first-page":"3020","DOI":"10.1080\/15592324.2017.1331197","article-title":"Functions and substrates of plastoglobule-localized metallopeptidase PGM48","volume":"12","author":"Bhuiyan","year":"2017","journal-title":"Plant Signal Behav"},{"issue":"2","key":"2020072910370220200_ref71","doi-asserted-by":"crossref","first-page":"562","DOI":"10.1104\/pp.109.137042","article-title":"Timing and biosynthetic potential for carotenoid accumulation in genetically diverse germplasm of maize","volume":"150","author":"Vallabhaneni","year":"2009","journal-title":"Plant Physiol"},{"issue":"5","key":"2020072910370220200_ref72","doi-asserted-by":"crossref","first-page":"851","DOI":"10.1007\/s00122-015-2475-3","article-title":"Genome-wide association analysis reveals new targets for carotenoid biofortification in maize","volume":"128","author":"Suwarno","year":"2015","journal-title":"Theor Appl Genet"},{"issue":"26","key":"2020072910370220200_ref73","doi-asserted-by":"crossref","first-page":"6866","DOI":"10.1073\/pnas.1705689114","article-title":"A recruiting protein of geranylgeranyl diphosphate synthase controls metabolic flux toward chlorophyll biosynthesis in rice","volume":"114","author":"Zhou","year":"2017","journal-title":"Proc Natl Acad Sci"},{"issue":"11","key":"2020072910370220200_ref74","doi-asserted-by":"crossref","first-page":"4560","DOI":"10.1105\/tpc.113.116210","article-title":"Subchromoplast sequestration of carotenoids affects regulatory mechanisms in tomato lines expressing different carotenoid gene combinations","volume":"25","author":"Nogueira","year":"2013","journal-title":"Plant Cell"},{"issue":"1","key":"2020072910370220200_ref75","doi-asserted-by":"crossref","first-page":"405","DOI":"10.1104\/pp.105.1.405","article-title":"Carotenoid biosynthesis during tomato fruit development (evidence for tissue-specific gene expression)","volume":"105","author":"Fraser","year":"1994","journal-title":"Plant Physiol"},{"issue":"1","key":"2020072910370220200_ref76","doi-asserted-by":"crossref","first-page":"557","DOI":"10.1146\/annurev.arplant.49.1.557","article-title":"Genes and enzymes of carotenoid biosynthesis in plants","volume":"49","author":"Cunningham","year":"1998","journal-title":"Annu Rev Plant Biol"},{"issue":"1","key":"2020072910370220200_ref77","doi-asserted-by":"crossref","first-page":"204","DOI":"10.1104\/pp.112.198556","article-title":"Synergistic interactions between carotene ring hydroxylases drive lutein formation in plant carotenoid biosynthesis","volume":"160","author":"Quinlan","year":"2012","journal-title":"Plant Physiol"},{"issue":"3","key":"2020072910370220200_ref78","doi-asserted-by":"crossref","first-page":"435","DOI":"10.1111\/j.1365-3040.2005.01492.x","article-title":"Carotenoid accumulation and function in seeds and non-green tissues","volume":"29","author":"Howitt","year":"2006","journal-title":"Plant Cell Environ"},{"issue":"6","key":"2020072910370220200_ref79","doi-asserted-by":"crossref","first-page":"1020","DOI":"10.1111\/j.1365-3040.2011.02301.x","article-title":"Regulatory control of high levels of carotenoid accumulation in potato tubers","volume":"34","author":"Zhou","year":"2011","journal-title":"Plant Cell Environ"},{"key":"2020072910370220200_ref80","doi-asserted-by":"crossref","first-page":"29","DOI":"10.3389\/fpls.2012.00029","article-title":"Maize provitamin A carotenoids, current resources, and future metabolic engineering challenges","volume":"3","author":"Wurtzel","year":"2012","journal-title":"Front Plant Sci"},{"key":"2020072910370220200_ref81","doi-asserted-by":"crossref","first-page":"1197","DOI":"10.3389\/fpls.2016.01197","article-title":"Carotenoids in staple cereals: metabolism, regulation, and genetic manipulation","volume":"7","author":"Zhai","year":"2016","journal-title":"Front Plant Sci"},{"key":"2020072910370220200_ref82","doi-asserted-by":"crossref","first-page":"273","DOI":"10.1007\/978-3-319-39126-7_10","volume-title":"Carotenoids in Nature","author":"Li","year":"2016"},{"issue":"6","key":"2020072910370220200_ref83","doi-asserted-by":"crossref","first-page":"232","DOI":"10.1016\/S1360-1385(99)01414-4","article-title":"Carotenoid sequestration in plants: the role of carotenoid-associated proteins","volume":"4","author":"Vishnevetsky","year":"1999","journal-title":"Trends Plant Sci"},{"issue":"10","key":"2020072910370220200_ref84","doi-asserted-by":"crossref","first-page":"1601","DOI":"10.1093\/pcp\/pcq136","article-title":"Chromoplast differentiation: current status and perspectives","volume":"51","author":"Egea","year":"2010","journal-title":"Plant Cell Physiol"},{"issue":"2","key":"2020072910370220200_ref85","doi-asserted-by":"crossref","first-page":"102","DOI":"10.1016\/j.abb.2013.07.002","article-title":"Chromoplast biogenesis and carotenoid accumulation","volume":"539","author":"Li","year":"2013","journal-title":"Arch Biochem Biophys"},{"key":"2020072910370220200_ref86","doi-asserted-by":"crossref","first-page":"15036","DOI":"10.1038\/hortres.2015.36","article-title":"Carotenoid metabolism and regulation in horticultural crops","volume":"2","author":"Yuan","year":"2015","journal-title":"Horticulture research"},{"issue":"1","key":"2020072910370220200_ref87","doi-asserted-by":"crossref","first-page":"195","DOI":"10.1111\/pbi.12373","article-title":"Bottlenecks in carotenoid biosynthesis and accumulation in rice endosperm are influenced by the precursor\u2013product balance","volume":"14","author":"Bai","year":"2016","journal-title":"Plant Biotechnol J"},{"issue":"2","key":"2020072910370220200_ref88","doi-asserted-by":"crossref","first-page":"190","DOI":"10.1111\/j.1399-3054.2008.01058.x","article-title":"Phytoene desaturase is present in a large protein complex in the plastid membrane","volume":"133","author":"Lopez","year":"2008","journal-title":"Physiol Plant"},{"issue":"1","key":"2020072910370220200_ref89","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1186\/s12870-015-0426-4","article-title":"Carotenoid accumulation affects redox status, starch metabolism, and flavonoid\/anthocyanin accumulation in citrus","volume":"15","author":"Cao","year":"2015","journal-title":"BMC Plant Biol"},{"issue":"1","key":"2020072910370220200_ref90","doi-asserted-by":"crossref","first-page":"140","DOI":"10.1111\/pbi.12365","article-title":"Product stability and sequestration mechanisms in Solanum tuberosum engineered to biosynthesize high value ketocarotenoids","volume":"14","author":"Mortimer","year":"2016","journal-title":"Plant Biotechnol J"},{"issue":"1","key":"2020072910370220200_ref91","doi-asserted-by":"crossref","first-page":"12","DOI":"10.3732\/ajb.94.1.12","article-title":"Amyloplast to chromoplast conversion in developing ornamental tobacco floral nectaries provides sugar for nectar and antioxidants for protection","volume":"94","author":"Horner","year":"2007","journal-title":"Am J Bot"},{"issue":"13","key":"2020072910370220200_ref92","doi-asserted-by":"crossref","first-page":"2594","DOI":"10.1002\/jsfa.5767","article-title":"Physical barriers to carotenoid bioaccessibility. Ultrastructure survey of chromoplast and cell wall morphology in nine carotenoid-containing fruits and vegetables","volume":"92","author":"Jeffery","year":"2012","journal-title":"J Sci Food Agric"},{"issue":"5","key":"2020072910370220200_ref93","doi-asserted-by":"crossref","first-page":"1037","DOI":"10.1007\/s00425-014-2121-3","article-title":"Lipid-dissolved \u03b3-carotene, \u03b2-carotene, and lycopene in globular chromoplasts of peach palm (Bactris gasipaes Kunth) fruits","volume":"240","author":"Hempel","year":"2014","journal-title":"Planta"},{"issue":"5","key":"2020072910370220200_ref94","doi-asserted-by":"crossref","first-page":"1063","DOI":"10.1007\/s00425-014-2147-6","article-title":"Regulatory control of carotenoid accumulation in winter squash during storage","volume":"240","author":"Zhang","year":"2014","journal-title":"Planta"},{"issue":"1","key":"2020072910370220200_ref95","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"},{"issue":"1","key":"2020072910370220200_ref96","doi-asserted-by":"crossref","first-page":"59","DOI":"10.1046\/j.1365-313x.2001.01008.x","article-title":"A novel gene mutation that confers abnormal patterns of \u03b2-carotene accumulation in cauliflower (Brassica oleracea var. botrytis)","volume":"26","author":"Li","year":"2001","journal-title":"Plant J"},{"issue":"3\u20134","key":"2020072910370220200_ref97","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1007\/s00709-004-0059-1","article-title":"The chromoplasts of Or mutants of cauliflower (Brassica oleracea L. var. botrytis)","volume":"224","author":"Paolillo","year":"2004","journal-title":"Protoplasma"},{"issue":"6","key":"2020072910370220200_ref98","doi-asserted-by":"crossref","first-page":"1074","DOI":"10.1111\/tpj.12351","article-title":"Multiple microscopic approaches demonstrate linkage between chromoplast architecture and carotenoid composition in diverse Capsicum annuum fruit","volume":"76","author":"Kilcrease","year":"2013","journal-title":"Plant J"},{"issue":"12","key":"2020072910370220200_ref99","doi-asserted-by":"crossref","first-page":"3594","DOI":"10.1105\/tpc.106.046417","article-title":"The cauliflower Or gene encodes a DnaJ cysteine-rich domain-containing protein that mediates high levels of \u03b2-carotene accumulation","volume":"18","author":"Lu","year":"2006","journal-title":"Plant Cell"},{"key":"2020072910370220200_ref100","doi-asserted-by":"crossref","first-page":"33563","DOI":"10.1038\/srep33563","article-title":"Orange protein has a role in phytoene synthase stabilization in sweetpotato","volume":"6","author":"Park","year":"2016","journal-title":"Sci Rep"},{"issue":"1","key":"2020072910370220200_ref101","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1105\/tpc.108.063131","article-title":"Regulation of carotenoid composition and shoot branching in Arabidopsis by a chromatin modifying histone methyltransferase, SDG8","volume":"21","author":"Cazzonelli","year":"2009","journal-title":"Plant Cell"},{"key":"2020072910370220200_ref102","doi-asserted-by":"crossref","first-page":"1017","DOI":"10.3389\/fpls.2019.01017","article-title":"Transcriptional regulation of carotenoid biosynthesis in plants: so many regulators, so little consensus","volume":"10","author":"Stanley","year":"2019","journal-title":"Front Plant Sci"},{"issue":"2","key":"2020072910370220200_ref103","doi-asserted-by":"crossref","first-page":"154","DOI":"10.1038\/nbt.2462","article-title":"Single-base resolution methylomes of tomato fruit development reveal epigenome modifications associated with ripening","volume":"31","author":"Zhong","year":"2013","journal-title":"Nat Biotechnol"},{"issue":"4","key":"2020072910370220200_ref104","doi-asserted-by":"crossref","first-page":"488","DOI":"10.1111\/j.1745-4522.2008.00135.x","article-title":"Quantification of carotenoid and flavonoid components in brans of some Thai black rice cultivars using supercritical fluid extraction and high-performance liquid chromatography-mass spectrometry","volume":"15","author":"Nakornriab","year":"2008","journal-title":"J Food Lipid"},{"issue":"24","key":"2020072910370220200_ref105","doi-asserted-by":"crossref","first-page":"11914","DOI":"10.1021\/jf802613c","article-title":"Carotenoids in raw and parboiled brown and milled rice","volume":"56","author":"Lamberts","year":"2008","journal-title":"J Agric Food Chem"},{"issue":"33","key":"2020072910370220200_ref106","doi-asserted-by":"crossref","first-page":"7976","DOI":"10.1021\/jf401937b","article-title":"Phytochemical profiles of black, red, brown, and white rice from the Camargue region of France","volume":"61","author":"Pereira-Caro","year":"2013","journal-title":"J Agric Food Chem"},{"issue":"6","key":"2020072910370220200_ref107","doi-asserted-by":"crossref","first-page":"e1004416","DOI":"10.1371\/journal.pgen.1004416","article-title":"The HY5-PIF regulatory module coordinates light and temperature control of photosynthetic gene transcription","volume":"10","author":"Toledo-Ortiz","year":"2014","journal-title":"PLoS Genet"},{"issue":"6","key":"2020072910370220200_ref108","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"},{"issue":"3","key":"2020072910370220200_ref109","doi-asserted-by":"crossref","first-page":"923","DOI":"10.1105\/tpc.110.081273","article-title":"Transcriptome and metabolite profiling show that APETALA2a is a major regulator of tomato fruit ripening","volume":"23","author":"Karlova","year":"2011","journal-title":"Plant Cell"},{"issue":"5566","key":"2020072910370220200_ref110","doi-asserted-by":"crossref","first-page":"343","DOI":"10.1126\/science.1068181","article-title":"A MADS-box gene necessary for fruit ripening at the tomato ripening-inhibitor (rin) locus","volume":"296","author":"Vrebalov","year":"2002","journal-title":"Science"},{"issue":"2","key":"2020072910370220200_ref111","doi-asserted-by":"crossref","first-page":"212","DOI":"10.1111\/j.1365-313X.2008.03491.x","article-title":"DNA-binding specificity, transcriptional activation potential, and the rin mutation effect for the tomato fruit-ripening regulator RIN","volume":"55","author":"Ito","year":"2008","journal-title":"Plant J"},{"issue":"6","key":"2020072910370220200_ref112","doi-asserted-by":"crossref","first-page":"1081","DOI":"10.1111\/j.1365-313X.2009.04064.x","article-title":"TOMATO AGAMOUS-LIKE 1 is a component of the fruit ripening regulatory network","volume":"60","author":"Itkin","year":"2009","journal-title":"Plant J"},{"issue":"10","key":"2020072910370220200_ref113","doi-asserted-by":"crossref","first-page":"3041","DOI":"10.1105\/tpc.109.066936","article-title":"Fleshy fruit expansion and ripening are regulated by the tomato SHATTERPROOF gene TAGL1","volume":"21","author":"Vrebalov","year":"2009","journal-title":"Plant Cell"},{"issue":"12","key":"2020072910370220200_ref114","doi-asserted-by":"crossref","first-page":"e14427","DOI":"10.1371\/journal.pone.0014427","article-title":"Functional analysis of the Arlequin mutant corroborates the essential role of the Arlequin\/TAGL1 gene during reproductive development of tomato","volume":"5","author":"Gim\u00e9nez","year":"2010","journal-title":"PLoS One"},{"issue":"6","key":"2020072910370220200_ref115","doi-asserted-by":"crossref","first-page":"1107","DOI":"10.1007\/s00425-011-1561-2","article-title":"Direct targets of the tomato-ripening regulator RIN identified by transcriptome and chromatin immunoprecipitation analyses","volume":"235","author":"Fujisawa","year":"2012","journal-title":"Planta"},{"issue":"2","key":"2020072910370220200_ref116","doi-asserted-by":"crossref","first-page":"371","DOI":"10.1105\/tpc.112.108118","article-title":"A large-scale identification of direct targets of the tomato MADS box transcription factor RIPENING INHIBITOR reveals the regulation of fruit ripening","volume":"25","author":"Fujisawa","year":"2013","journal-title":"Plant Cell"},{"issue":"1","key":"2020072910370220200_ref117","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1105\/tpc.113.119453","article-title":"Transcriptional regulation of fruit ripening by tomato FRUITFULL homologs and associated MADS box proteins","volume":"26","author":"Fujisawa","year":"2014","journal-title":"Plant Cell"},{"issue":"1","key":"2020072910370220200_ref118","doi-asserted-by":"crossref","first-page":"22","DOI":"10.1134\/S1021443719010205","article-title":"TERF1 regulates nuclear gene expression through chloroplast retrograde signals","volume":"66","author":"Wu","year":"2019","journal-title":"Russ J Plant Physiol"},{"issue":"1","key":"2020072910370220200_ref119","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1093\/pcp\/pct162","article-title":"A new tomato NAC (N AM\/A TAF1\/2\/C UC2) transcription factor, SlNAC4, functions as a positive regulator of fruit ripening and carotenoid accumulation","volume":"55","author":"Zhu","year":"2013","journal-title":"Plant Cell Physiol"},{"issue":"6","key":"2020072910370220200_ref120","doi-asserted-by":"crossref","first-page":"3002","DOI":"10.1080\/15592324.2016.1184808","article-title":"Light-specific transcriptional regulation of the accumulation of carotenoids and phenolic compounds in rice leaves","volume":"11","author":"Mohanty","year":"2016","journal-title":"Plant Signal Behav"},{"issue":"1","key":"2020072910370220200_ref121","doi-asserted-by":"crossref","first-page":"41","DOI":"10.5114\/bta.2017.66616","article-title":"In silico identification of transcription factors associated with the biosynthesis of carotenoids in corn (Zea mays L.)","volume":"98","author":"Zinati","year":"2017","journal-title":"BioTechnologia"}],"container-title":["Briefings in Functional Genomics"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/academic.oup.com\/bfg\/article-pdf\/19\/4\/324\/33543106\/elaa007.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"http:\/\/academic.oup.com\/bfg\/article-pdf\/19\/4\/324\/33543106\/elaa007.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,10,16]],"date-time":"2022-10-16T19:11:16Z","timestamp":1665947476000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/bfg\/article\/19\/4\/324\/5814220"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,4,2]]},"references-count":121,"journal-issue":{"issue":"4","published-online":{"date-parts":[[2020,4,2]]},"published-print":{"date-parts":[[2020,7,29]]}},"URL":"https:\/\/doi.org\/10.1093\/bfgp\/elaa007","relation":{},"ISSN":["2041-2657"],"issn-type":[{"value":"2041-2657","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2020,7]]},"published":{"date-parts":[[2020,4,2]]}}}