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Plant Science Today (PST)

Review Articles

Vol. 13 No. sp5 (2026): Recent Advances in Agriculture

Coloured maize as a functional crop: Integrating bioactive pigments, nutritional quality and genetic improvement

DOI
https://doi.org/10.14719/pst.15552
Submitted
14 May 2026
Published
28-07-2026

Abstract

Maize (Zea mays L.) is one of the world’s most important cereal crops, serving as a major source of calories, protein and industrial raw materials. However, conventional white and yellow cultivars are frequently deficient in essential micronutrients and health-promoting phytochemicals, contributing to widespread micronutrient malnutrition and hidden hunger in maize-dependent populations. Coloured maize genotypes are characterised by purple, blue, red and orange kernels. These genotypes represent an underutilised reservoir of nutritional and genetic diversity, enriched in anthocyanins, carotenoids, phenolic acids and minerals with strong antioxidant and functional properties. This review synthesises current knowledge on the nutritional composition, biochemical diversity and health relevance of coloured maize, emphasising the molecular regulation of flavonoid and carotenoid biosynthesis pathways and the roles of key structural and regulatory genes controlling pigment accumulation. The importance of traditional landraces and germplasm conservation in maintaining allelic richness for nutritional traits is discussed alongside advances in phenotyping, near-infrared spectroscopy, molecular markers and marker-assisted breeding. Progress in quality protein maize, provitamin A biofortification and the development of anthocyanin-rich sweet corn illustrate the integration of metabolic genetics with applied breeding strategies. Collectively, coloured maize is represented as a multifunctional crop capable of enhancing dietary quality, supporting functional food development and contributing to sustainable agricultural systems. Harnessing its genetic variability through genomics-assisted selection and targeted pathway engineering offers a promising route toward nutritionally superior, climate-resilient maize cultivars that can contribute to future food and nutritional security.

References

  1. 1. Matsuoka Y, Vigouroux Y, Goodman MM, Sanchez GJ, Buckler E, Doebley J. A single domestication for maize is shown by multilocus microsatellite genotyping. Proc Natl Acad Sci U S A. 2002;99(9):6080–4.https://doi.org/10.1073/pnas.052125199
  2. 2. Fang H, Fu X, Ge H, Zhang A, Shan T, Wang Y, et al. Genetic basis of maize kernel oil-related traits revealed by high-density SNP markers in a recombinant inbred line population. BMC Plant Biol. 2021;21:384. https://doi.org/10.1186/s12870-021-03089-0
  3. 3. Food and Agriculture Organization of the United Nations. World maize production statistics. Rome: FAO; 2025.
  4. 4. Food and Agriculture Organization of the United Nations. FAOSTAT statistical database. Rome: FAO; 2023.
  5. 5. Ministry of Agriculture and Farmers Welfare. Agricultural statistics at a glance 2023. New Delhi: Government of India; 2023.
  6. 6. United Nations. UN Comtrade: International trade statistics database. New York: United Nations; 2022.
  7. 7. Bouis HE, Saltzman A. Improving nutrition through biofortification: A review of evidence from HarvestPlus, 2003 through 2016. Glob Food Secur. 2017;12:49–58. https://doi.org/10.1016/j.gfs.2017.01.009
  8. 8. Ranum P, Peña-Rosas JP, García-Casal MN. Global maize production, utilization and consumption. Ann N Y Acad Sci. 2014;1312(1):105–12. https://doi.org/10.1111/nyas.12396
  9. 9. Niu L, Liu L, Zhang J, Scali M, Wang W, Hu X, et al. Genetic engineering of starch biosynthesis in maize seeds for efficient enzymatic digestion during bioethanol production. Int J Mol Sci. 2022;24(4):3927. https://doi.org/10.3390/ijms24043927
  10. 10. Di Y, Na R, Xia H, Wang Y, Li F. Irradiation effects on characteristics and ethanol fermentation of maize starch. Int J Biol Macromol. 2023;246:125602. https://doi.org/10.1016/j.ijbiomac.2023.125602
  11. 11. Zilic S, Serpen A, Akıllıoğlu G, Gökmen V, Vančetović J. Phenolic compounds, carotenoids, anthocyanins and antioxidant capacity of coloured maize (Zea mays L.) kernels. J Agric Food Chem. 2012;60(5):1224–31. https://doi.org/10.1021/jf204367z
  12. 12. Prasanna BM, Vasal SK, Kassahun B, Singh NN. Quality protein maize. Curr Sci. 2001;81(10):1308–19.
  13. 13. Giusti MM, Wrolstad RE. Acylated anthocyanins from edible sources and their applications in food systems. Biochem Eng J. 2003;14(3):217–25. https://doi.org/10.1016/S1369-703X(02)00221-8
  14. 14. Jing P, Noriega V, Schwartz SJ, Giusti MM. Effects of growing conditions on purple corncob (Zea mays L.) anthocyanins. J Agric Food Chem. 2007;55(21):8625–9. https://doi.org/10.1021/jf070755q
  15. 15. He J, Giusti MM. Anthocyanins: natural colourants with health-promoting properties. Annu Rev Food Sci Technol. 2010;1:163–87.https://doi.org/10.1146/annurev.food.080708.100754
  16. 16. Nishino H, Murakoshi M, Tokuda H, Satomi Y. Cancer prevention by carotenoids. Arch Biochem Biophys. 2009;483(2):165–8. https://doi.org/10.1016/j.abb.2008.09.011
  17. 17. Slavin J. Whole grains and human health. Nutr Res Rev. 2004;17(1):99–110. https://doi.org/10.1079/NRR200374
  18. 18. Traber MG, Stevens JF. Vitamins C and E: beneficial effects from a mechanistic perspective. Free Radic Biol Med. 2011;51(5):1000–13. https://doi.org/10.1016/j.freeradbiomed.2011.05.017
  19. 19. Shahidi F, Ambigaipalan P. Phenolics and polyphenolics in foods, beverages and spices: antioxidant activity and health effects—a review. J Funct Foods. 2015;18:820–97. https://doi.org/10.1016/j.jff.2015.06.018
  20. 20. Kasote DM, Katyare SS, Hegde MV, Bae H. Significance of antioxidant potential of plants and its relevance to therapeutic applications. Int J Biol Sci. 2015;11(8):982–91. https://doi.org/10.7150/ijbs.12096
  21. 21. Gogoi P, Bhuyan DJ, Bhattacharyya P, Choudhary B, Saikia S, Baruah DK, et al. Evaluating free and bound polyphenols, flavonoids and antioxidant properties of pigmented maize landraces. Food Chem Adv. 2025;6:100870. https://doi.org/10.1016/j.focha.2025.100870
  22. 22. Berardo N, Mazzinelli G, Valoti P, Laganà P, Redaelli R. Characterization of maize germplasm for the chemical composition of the grain. J Agric Food Chem. 2009;57(6):2378–84. https://doi.org/10.1021/jf803688t
  23. 23. Tenaillon MI, Charcosset A. A European perspective on maize history. C R Biol. 2011;334(3):221–8. https://doi.org/10.1016/j.crvi.2010.12.015
  24. 24. Smith JSC, Smith OS. The description and assessment of distances between inbred lines of maize: the utility of morphological, biochemical and genetic descriptors and a scheme for testing distinctiveness between inbred lines. Maydica. 1989;34:141–50.
  25. 25. Caporaso N, Whitworth MB, Fisk ID. Near-infrared spectroscopy and hyperspectral imaging for non-destructive quality assessment of cereal grains. Appl Spectrosc Rev. 2018;53(8):667–87. https://doi.org/10.1080/05704928.2018.1425214
  26. 26. Piperno DR, Flannery KV. The earliest archaeological maize (Zea mays L.) from highland Mexico: new accelerator mass spectrometry dates and their implications. Proc Natl Acad Sci U S A. 2001;98(4):2101–3. https://doi.org/10.1073/pnas.98.4.2101
  27. 27. Kim HY, Lee KY, Kim M, Hong M, Deepa P, Kim S. A review of the biological properties of purple corn (Zea mays L.). Sci Pharm. 2023;91(1):6. https://doi.org/10.3390/scipharm91010006
  28. 28. Allard RW. Principles of plant breeding. New York: John Wiley & Sons; 1960.
  29. 29. Falconer DS. Introduction to quantitative genetics. Scotts Valley (CA): CreateSpace Independent Publishing Platform; 2017.
  30. 30. Machikowa T, Saetang C. Correlation and path coefficient analysis on seed yield in sunflower (Helianthus annuus L.). Suranaree J Sci Technol. 2008;15(3):243–8.
  31. 31. Duvick DN. The use of cytoplasmic male sterility in hybrid seed production. Econ Bot. 1959;13(3):167–95. https://doi.org/10.1007/BF02860581
  32. 32. Hahlbrock K, Scheel D. Physiology and molecular biology of phenylpropanoid metabolism. Annu Rev Plant Physiol Plant Mol Biol. 1989;40:347–69. https://doi.org/10.1146/annurev.pp.40.060189.002023
  33. 33. Emerson RA. The genetic relations of plant colours in maize. Washington (DC): Carnegie Institution of Washington; 1921. https://doi.org/10.5962/bhl.title.16496
  34. 34. Dooner HK, Nelson OE. Controlling element-induced alterations in UDP-glucose: flavonoid glucosyltransferase. Proc Natl Acad Sci U S A. 1977;74(12):5623–7. https://doi.org/10.1073/pnas.74.12.5623
  35. 35. Ludwig SR, Habera LF, Dellaporta SL, Wessler SR. Lc, a member of the maize R gene family responsible for tissue-specific anthocyanin production, encodes a protein similar to transcriptional activators and contains the myc-homology region. Proc Natl Acad Sci U S A. 1989;86(18):7092–6. https://doi.org/10.1073/pnas.86.18.7092
  36. 36. Anirban A, Hayward A, Hong HT, Masouleh AK, Henry RJ, O'Hare TJ. Breaking the tight genetic linkage between the a1 and sh2 genes led to the development of anthocyanin-rich purple-pericarp super-sweetcorn. Sci Rep. 2023;13:1050. https://doi.org/10.1038/s41598-023-28083-4
  37. 37. Paulsmeyer M, Chatham L, Becker T, West M, West L, Juvik J. Survey of anthocyanin composition and concentration in diverse maize germplasms. J Agric Food Chem. 2017;65(21):4341–50. https://doi.org/10.1021/acs.jafc.7b00771
  38. 38. Iiyama K, Lam TB, Stone BA. Covalent cross-links in the cell wall. Plant Physiol. 1994;104(2):315–20. https://doi.org/10.1104/pp.104.2.315
  39. 39. Lopez-Martinez LX, Oliart-Ros RM, Valerio-Alfaro G, Lee CH, Parkin KL, Garcia HS. Antioxidant activity, phenolic compounds and anthocyanin content of eighteen strains of Mexican maize. LWT Food Sci Technol. 2009;42(6):1187–92. https://doi.org/10.1016/j.lwt.2008.10.010
  40. 40. Yan J, Kandianis CB, Harjes CE, Bai L, Kim EH, Yang X, et al. Rare genetic variation at Zea mays crtRB1 increases β-carotene in maize grain. Nat Genet. 2010;42(4):322–7. https://doi.org/10.1038/ng.551
  41. 41. Tohge T, Watanabe M, Hoefgen R, Fernie AR. Shikimate and phenylalanine biosynthesis in the green lineage. Front Plant Sci. 2013;4:62. https://doi.org/10.3389/fpls.2013.00062
  42. 42. Rodriguez MD, Monsierra L, Mansilla PS, Pérez GT, Pascual-Teresa S. Phenolic characterization of a purple maize (Zea mays cv. Moragro) by HPLC-QTOF-MS and study of its bioaccessibility using a simulated in vitro digestion/Caco-2 culture model. J Agric Food Chem. 2024;72(12):6327–38. https://doi.org/10.1021/acs.jafc.3c08960
  43. 43. Sekhon RS, Lin H, Childs KL, Hansey CN, Buell CR, de Leon N, et al. Genome-wide atlas of transcription during maize development. Plant J. 2011;66(4):553–63. https://doi.org/10.1111/j.1365-313X.2011.04527.x
  44. 44. Cocciolone SM, Chopra S, Flint-Garcia SA, McMullen MD, Peterson T. Tissue-specific patterns of a maize MYB transcription factor are epigenetically regulated. Plant J. 2001;27(5):467–78. https://doi.org/10.1046/j.1365-313X.2001.01124.x
  45. 45. Fedoroff NV, Furtek DB, Nelson OE. Cloning of the bronze locus in maize using the transposable controlling element Activator (Ac). Proc Natl Acad Sci U S A. 1984;81(12):3825–9. https://doi.org/10.1073/pnas.81.12.3825
  46. 46. Grotewold E, Athma P, Peterson T. Alternatively spliced products of the maize P gene encode proteins with homology to the DNA-binding domain of myb-like transcription factors. Proc Natl Acad Sci U S A. 1991;88(11):4587–91. https://doi.org/10.1073/pnas.88.11.4587
  47. 47. Cone KC, Burr FA, Burr B. Molecular analysis of the maize anthocyanin regulatory locus C1. Proc Natl Acad Sci U S A. 1986;83(24):9631–5. https://doi.org/10.1073/pnas.83.24.9631
  48. 48. Petroni K, Tonelli C. Recent advances on the regulation of anthocyanin synthesis in reproductive organs. Plant Sci. 2011;181(3):219–29. https://doi.org/10.1016/j.plantsci.2011.05.009
  49. 49. Buckner B, Kelson TL, Robertson DS. Cloning of the y1 locus of maize, a gene involved in the biosynthesis of carotenoids. Plant Cell. 1990;2(9):867–76. https://doi.org/10.1105/tpc.2.9.867
  50. 50. Radicella JP, Turks D, Chandler VL. Cloning and nucleotide sequence of a cDNA encoding B-Peru, a regulatory protein of the anthocyanin pathway in maize. Plant Mol Biol. 1991;17(1):127–30. https://doi.org/10.1007/BF00036813
  51. 51. Menssen A, Höhmann S, Martin W, Schnable PS, Peterson PA, Saedler H, et al. The En/Spm transposable element of Zea mays contains splice sites generating a novel intron from a dSpm element in the A2 gene. EMBO J. 1990;9(10):3051–7. https://doi.org/10.1002/j.1460-2075.1990.tb07501.x
  52. 52. Marrs KA, Alfenito MR, Lloyd AM, Walbot V. A glutathione S-transferase involved in vacuolar transfer encoded by the maize gene Bronze-2. Nature. 1995;375(6530):397–400. https://doi.org/10.1038/375397a0
  53. 53. Grotewold E, Chamberlin MA, Snook ME, Siame BA, Butler LG, Swenson J, et al. Engineering secondary metabolism in maize cells by ectopic expression of transcription factors. Plant Cell. 1998;10(5):721–40. https://doi.org/10.1105/tpc.10.5.721
  54. 54. Carey CC, Strahle JT, Selinger DA, Chandler VL. Mutations in the pale aleurone colour1 regulatory gene of Zea mays anthocyanin pathway have distinct phenotypes relative to the functionally similar TTG1 gene in Arabidopsis thaliana. Plant Cell. 2004;16(2):450–64. https://doi.org/10.1105/tpc.018796
  55. 55. Harjes CE, Rocheford TR, Bai L, Brutnell TP, Kandianis CB, Sowinski SG, et al. Natural genetic variation in lycopene epsilon cyclase tapped for maize biofortification. Science. 2008;319(5861):330–3. https://doi.org/10.1126/science.1150255
  56. 56. Sharma M, Chai C, Morohashi K, Grotewold E, Snook ME, Chopra S. Expression of flavonoid 3′-hydroxylase is controlled by P1, the regulator of 3-deoxyflavonoid biosynthesis in maize. BMC Plant Biol. 2012;12:196. https://doi.org/10.1186/1471-2229-12-196
  57. 57. Winkel-Shirley B. Flavonoid biosynthesis: a colourful model for genetics, biochemistry, cell biology and biotechnology. Plant Physiol. 2001;126(2):485–93. https://doi.org/10.1104/pp.126.2.485
  58. 58. Castellarin SD, Pfeiffer A, Sivilotti P, Degan M, Peterlunger E, Di Gaspero G. Transcriptional regulation of anthocyanin biosynthesis in ripening fruits of grapevine under seasonal water deficit. Plant Cell Environ. 2007;30(11):1381–99. https://doi.org/10.1111/j.1365-3040.2007.01716.x
  59. 59. Sun W, Liang L, Meng X, Li Y, Gao F, Liu X, et al. Biochemical and molecular characterization of a flavonoid 3-O-glycosyltransferase responsible for anthocyanin and flavonol biosynthesis in Freesia hybrida. Front Plant Sci. 2016;7:410. https://doi.org/10.3389/fpls.2016.00410
  60. 60. Enaru B, Drețcanu G, Pop TD, Stǎnilǎ A, Diaconeasa Z. Anthocyanins: factors affecting their stability and degradation. Antioxidants (Basel). 2021;10(12):1967. https://doi.org/10.3390/antiox10121967
  61. 61. Zhao YW, Wang CK, Huang XY, Hu DG. Anthocyanin stability and degradation in plants. Plant Signal Behav. 2021;16(12):1987767. https://doi.org/10.1080/15592324.2021.1987767
  62. 62. Qian Y, Zhang T, Yu Y, Gou L, Yang J, Xu J, et al. Regulatory mechanisms of bHLH transcription factors in plant adaptive responses to abiotic stresses. Front Plant Sci. 2021;12:677611. https://doi.org/10.3389/fpls.2021.677611
  63. 63. Li C, Yu W, Xu J, Lu X, Liu Y. Anthocyanin biosynthesis induced by MYB transcription factors in plants. Int J Mol Sci. 2022;23(19):11701. https://doi.org/10.3390/ijms231911701
  64. 64. Lu N, Jun JH, Li Y, Yang T, Liu Z, Grotewold E, et al. An unconventional proanthocyanidin pathway in maize. Nat Commun. 2023;14:4349. https://doi.org/10.1038/s41467-023-40014-5
  65. 65. Giménez P, Just-Borràs A, Gombau J, Canals JM, Zamora F. Effects of laccase from Botrytis cinerea on oxidative degradation of anthocyanins. OENO One. 2023;57(3):243–53. https://doi.org/10.20870/oeno-one.2023.57.3.7567
  66. 66. Wu X, Zhu S, He L, Cheng G, Li T, Meng W, et al. Phenylalanine ammonia-lyase: a core regulator of plant carbon metabolic flux redistribution—from molecular mechanisms to stress adaptability. Plants (Basel). 2025;14(24):3811. https://doi.org/10.3390/plants14243811
  67. 67. Sharma G, Muthusamy V, Mishra SJ, Kasana RK, Chhabra R, Sarma GR, et al. Composition of anthocyanins in kernels among sub-tropically adapted blue- and red-maize breeding lines. Eur Food Res Technol. 2026;252(3):699–712. https://doi.org/10.1007/s00217-026-05064-1
  68. 68. Betrán FJ, Beck D, Bänziger M, Edmeades GO. Secondary traits in parental inbreds and hybrids under stress and non-stress environments in tropical maize. Field Crops Res. 2003;83(1):51–65. https://doi.org/10.1016/S0378-4290(03)00061-3
  69. 69. Indian Agricultural Research Institute. Hybrid development and seed production of biofortified maize hybrids. New Delhi: Indian Agricultural Research Institute; 2025.
  70. 70. Muthusamy V, Hossain F, Thirunavukkarasu N, Choudhary M, Saha S, Bhat JS, et al. Development of beta-carotene-rich maize hybrids through marker-assisted introgression of beta-carotene hydroxylase allele. PLoS One. 2014;9(12):e113583. https://doi.org/10.1371/journal.pone.0113583
  71. 71. Saenz E, Abdala LJ, Borras L, Gerde JA. Maize kernel colour depends on the interaction between hardness and carotenoid concentration. J Cereal Sci. 2020;91:102901. https://doi.org/10.1016/j.jcs.2019.102901
  72. 72. Vazquez-Carrillo MG, Palos-Hernandez A, Gonzalez-Paramas AM, Santos-Buelga C, Garcia-Cruz L, Arellano-Vazquez JL, et al. Blue maize with pigmented germ: phytochemical compounds and tortilla colour. Food Chem. 2024;463:141109. https://doi.org/10.1016/j.foodchem.2024.141109
  73. 73. Vasal SK. Quality protein maize: overcoming the hurdles. J Crop Prod. 2002;6(1–2):193–227. https://doi.org/10.1300/J144v06n01_11
  74. 74. Mertz ET, Bates LS, Nelson OE. A mutant gene that changes protein composition and increases lysine content of maize endosperm. Science. 1964;145(3629):279–80. https://doi.org/10.1126/science.145.3629.279
  75. 75. Holding DR, Hunter BG, Chung T, Gibbon BC, Ford CF, Bharti AK, et al. Genetic analysis of opaque2 modifier loci in quality protein maize. Theor Appl Genet. 2008;117(2):157–70. https://doi.org/10.1007/s00122-008-0762-y
  76. 76. Twumasi-Afriyie S, Palacios-Rojas N, Friesen D, Teklewold A, Gissa DW, De Groote H, et al. Quality protein maize: development and dissemination for food security. Nutr Bull. 2016;41(1):38–45. https://doi.org/10.1111/nbu.12166
  77. 77. Hossain F, Jaiswal SK, Muthusamy V, Zunjare RU, Mishra SJ, Chand G, et al. Enhancement of nutritional quality in maize kernel through marker-assisted breeding for vte4, crtRB1 and opaque2 genes. J Appl Genet. 2023;64(3):431–43. https://doi.org/10.1007/s13353-023-00768-6
  78. 78. Indian Agricultural Research Institute. Annual report 2023–24. New Delhi: Indian Agricultural Research Institute; 2024.
  79. 79. Das AK, Gowda MM, Muthusamy V, Zunjare RU, Chauhan HS, Baveja A, Bhatt V, Chand G, Bhat JS, Guleria SK, et al. Development of maize hybrids with enhanced vitamin E, vitamin A, lysine and tryptophan through molecular breeding. Front Plant Sci. 2021; 12:659381. doi:10.3389/fpls.2021.659381. https://doi.org/10.3389/fpls.2021.659381
  80. 80. Senthil N, Thirusenduraselvi D, Bharathi P, Chandran S, Jagadeeshselvam N, Karthikeyan A, et al. Enhancing beta-carotene concentration in parental lines of CO6 maize hybrid through marker-assisted backcross breeding (MABB). Front Nutr. 2020;7:134. https://doi.org/10.3389/fnut.2020.00134
  81. 81. Kumari Vinodhana N, Ravikesavan R, Senthil N, Sivakumar S, Sathya Sheela KRV, Datta HM, et al. Enrichment of beta-carotene content in maize inbred UMI 1201 through marker-assisted backcross breeding. Electron J Plant Breed. 2025;16(3):323–30. https://doi.org/10.37992/2025.1603.036
  82. 82. Mondini L, Noorani A, Pagnotta MA. Assessing plant genetic diversity by molecular tools. Diversity. 2009;1(1):19–35. https://doi.org/10.3390/d1010019
  83. 83. Holly AA, Michael PN, John JA. Comparative transcriptomics of anthocyanin accumulation in the pericarp of pigmented purple corn. Theor Appl Genet. 2026;139(1):15. https://doi.org/10.1007/s00122-025-05137-x
  84. 84. Guo G, Xia H, Zhang A, Shan T, Wang Y, Li P, et al. Comparative transcriptomic and metabolomic analysis reveals mechanisms of selenium-regulated anthocyanin synthesis in waxy maize (Zea mays L.). Front Plant Sci. 2024;15:1466756. https://doi.org/10.3389/fpls.2024.1466756
  85. 85. Chachar Z, Lai R, Ahmed N, Lingling M, Chachar S, Paker NP, et al. Cloned genes and genetic regulation of anthocyanin biosynthesis in maize: a comparative review. Front Plant Sci. 2024;15:1310634. https://doi.org/10.3389/fpls.2024.1310634
  86. 86. Khusnutdinov E, Sukhareva A, Panfilova M, Mikhaylova E. Anthocyanin biosynthesis genes as model genes for genome editing in plants. Int J Mol Sci. 2021;22(16):8752. https://doi.org/10.3390/ijms22168752
  87. 87. He M, Li J, Jin W, Liu J. Unraveling the genetic and molecular mechanisms of anthocyanin biosynthesis and accumulation in maize kernels. Front Plant Sci. 2026;17:1771678. https://doi.org/10.3389/fpls.2026.1771678
  88. 88. Finkel ML, Sanchez S, Mak T, Granstein J, Lefkowitz A. Anthocyanin-rich purple corn extract and its effects on the blood pressure of adults. J Evid Based Complement Altern Med. 2013;18(4):237–42. https://doi.org/10.1177/2156587213482942

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