Studies on carotenoids in watermelon flesh
Affiliation(s)
School of Chemical Engineering and Energy, Zhengzhou University, Zhengzhou, China;. Department of Bioengineering, Zhengzhou University, Zhengzhou, China.
School of Chemical Engineering and Energy, Zhengzhou University, Zhengzhou, China;. Department of Bioengineering, Zhengzhou University, Zhengzhou, China.
ABSTRACT
Carotenoids are responsible for the different flesh colors in watermelon fruit, such as white, salmon yellow, orange, pale yellow, canary yellow, crimson red, and scarlet red. In red-fleshed watermelons lycopene constitutes the major pigment and b-carotene the secondary. The predominant carotenoid in yellow-fleshed watermelon is neoxanthin. Lycopene content in watermelon is related to genotype and ploidy level, harvest maturity, and growth and development conditions. Watermelon flesh colors are controlled by several gene loci. There are two or three alleles identified at each gene locus. Up to now several full-length cDNAs or gene fragments encoding enzymes in the carotenoid metabolic pathway have been isolated and characterized from mature watermelon fruits. Differential expression of carotenogenic genes was examined in flesh, ovary, leaf, and root tissues across different colored fleshes (white, canary yellow, salmon yellow, orange, and red). Carotenogenic gene expression was also analyzed at three fruit developmental stages (10, 20, and 30 days postanthesis) in five flesh colors of watermelon cultivars (red, pink, orange, yellow, and white).
Cite this paper
Zhao, W. , Lv, P. and Gu, H. (2013) Studies on carotenoids in watermelon flesh. Agricultural Sciences, 4, 13-20. doi: 10.4236/as.2013.47A003.
Zhao, W. , Lv, P. and Gu, H. (2013) Studies on carotenoids in watermelon flesh. Agricultural Sciences, 4, 13-20. doi: 10.4236/as.2013.47A003.
References
[1] King, S.R., Davis, A.R. and Bang, H. (2009) New flesh colors in watermelon? HortScience, 44, 576. [2] Tadmor, Y., King, S., Levi, A., Davis, A., Meir, A., Wasserman, B., Hirschberg, J. and Lewinsohn, E. (2005) Comparative fruit colouration in watermelon and tomato. Food Research International, 38, 837-841. doi:10.1016/j.foodres.2004.07.011 [3] Perkins-Veazie, P., Collins, J.K., Davis, A.R. and Roberts, W. (2006) Carotenoid content of 50 watermelon cultivars. Journal of Agricultural and Food Chemistry, 54, 2593-2597. doi:10.1021/jf052066p [4] Jia, G., Zhao, W., Shen, H., Wang, B. and Ma, S. (2008) Analysis of carotenoids in watermelon by column chromatography separation. Journal of Anhui Agricultural Sciences, 36, 8416-8418. (in Chinese) [5] Kang, B., Zhao, W., Hou, Y. and Tian, P. (2010) Expression of carotenogenic genes during the development and ripening of watermelon fruit. Scientia Horticulturae, 124, 368-375. doi:10.1016/j.scienta.2010.01.027 [6] Hu, L. (2009) Separation and analysis of carotenoids in watermelon. Ms.D. Thesis, Zhengzhou University, Zhengzhou. (in Chinese) [7] Hu, L., Guo, X., Zhao, W., Wang, J. and Zhang, G. (2011) Separation and analysis of carotenoids in watermelon flesh. Journal of Henan Agricultural Sciences, 40, 150-156. (in Chinese) [8] Bang, H., Davis, A., Kim, S., Leskovar, D. and King, S. (2010) Flesh color inheritance and gene interactions among canary yellow, pale yellow, and red watermelon. Journal of the American Society for Horticultural Science, 135, 362-368. [9] Liu, C., Zhang, H., Dai, Z., Liu, X., Liu, Y., Deng, X., Chen, F. and Xu, J. (2012) Volatile chemical and carotenoid profiles in watermelons [Citrullus vulgaris (Thunb.) Schrad (Cucurbitaceae)] with different flesh colors. Food Science and Biotechnology, 21, 531-541. doi:10.1007/s10068-012-0068-3 [10] Hui, B. and Li, J. (2008) Content and composition comparisons of carotenoids between red-flesh watermelon and yellow-flesh watermelon. Food Science, 29, 587-591. (in Chinese) [11] Collins, J.K., Perkins-Veazie, P. and Roberts, W. (2006) Lycopene: From plants to humans. HortScience, 41, 1135-1144. [12] USDA-NCC (1998) Carotenoid database for US foods. http://www.nal.usda.gov/fnic/food [13] Heinonen, M.I., Ollilainen, V., Linkola, E.K., Varo Pertti, T. and Koivistoinen, P.E. (1989) Carotenoids in Finnish foods: vegetables, fruits and berries. Journal of Agricultural and Food Chemistry, 37, 655-659. doi:10.1021/jf00087a017 [14] Holden, J.M., Eldridge, A.L., Beecher, G.R., Buzzard, I.M., Bhagwat, S., Davis, C.S., Douglass, L.W., Gebhardt, E.S, Haytowitz, D. and Schakel, S. (1999) Carotenoid Content of U.S. foods: An update of the database. Journal of Food Composition and Analysis, 12, 169-196. doi:10.1006/jfca.1999.0827 [15] US Department of Agriculture, ARS (2005) USDA National Nutrient Database for Standard Reference, release 18. US Department of Agriculture, Washington DC. [16] Perkins-Veazie, P., Collins, J.K. and Pair, S.D. (2000) Watermelon lycopene content varies with cultivar and maturity stage. HortScience, 35, 412. [17] Perkins-Veazie, P., Collins, J.K., Pair, S.D. and Roberts, W. (2001) Lycopene content differs among red-fleshed watermelon cultivars. Journal of the Science of Food and Agriculture, 81, 983-987. doi:10.1002/jsfa.880 [18] Perkins-Veazie, P., Roberts, W., Collins, J.K. and Perez, K. (2003) Lycopene variation among watermelons: Culivars, potassium, and ripeness. HortScience, 38, 1295. [19] Perkins-Veazie, P. and Collins, J.K. (2004) Flesh quality and lycopene stability of fresh-cut watermelon. Postharvest Biology and Technology, 31, 159-166. doi:10.1016/j.postharvbio.2003.08.005 [20] Perkins-Veazie, P. and Collins, J.K. (2006) Carotenoid changes of intact watermelons after storage. Journal of Agricultural and Food Chemistry, 54, 5868-5874. doi:10.1021/jf0532664 [21] Perkins-Veazie, P. (2007) Ripening events in seeded watermelons. HortScience, 42, 927. [22] Perkins-Veazie, P. and Zhang, X. (2007) Grafting increases lycopene in seedless watermelon. HortScience, 42, 959. [23] Davis, A.R., Fish, W.W. and Perkins-Veazie, P. (2003) A rapid hexane-free method for analyzing lycopene content in watermelon. Journal of Food Science, 68, 326-332. doi:10.1111/j.1365-2621.2003.tb14160.x [24] Leskovar, D.I., Bang, H., Crosby, K.M., Maness, N., Franco, J.A. and Perkins-Veazie, P. (2004) Lycopene, carbohydrates, ascorbic acid and yield components of diploid and triploid watermelon cultivars and affected by deficit irrigation. Journal of Horticultural Science and Biotechnology, 79, 75-81. [25] Yuan, P., Liu, W., Lu, X., Zhao, S., Lu, J., Yan, Z., He, N., Zhu, H. and Guan, L. (2012) Change of lycopene contents during fruit development of different ploidy watermelon. Journal of Fruit Science, 29, 890-894. (in Chinese) [26] Jeffery, J., Davis, A. and King, S. (2012) Understanding the carotenoid biosynthetic pathway: Observation of four color variants of developing watermelon fruit. Israel Journal of Plant Sciences, 60, 425-434. [27] Sun, L., Ding, S., Li, J., Zhu, F. and Wang J. (2006) Comparative analysis of lycopene contents among 8 watermelon cultivars. Acta Agriculturae Shanghai, 22, 66-68. (in Chinese) [28] Wan, X., Liu, W., Yan, Z., Zhao, S., He, N., Liu, P. and Dai, J. (2011) Changes of the contents of functional substances including lycopene, citrulline and ascorbic acid during watermelon fruits development. Scientia Agricultura Sinica, 44, 2738-2747. (in Chinese) [29] Henderson, W.R. (1989) Inheritance of orange flesh color in watermelon. Cucurbit Genetics Cooperative Report, 12, 59-63. [30] Shimotsuma, M. (1963) Cytogenetical studies in the genus Citrullus. VII. Inheritance of several characters in wtaremelons. Japanese Journal of Breeding, 13, 235-240. [31] Navot, N., Sarfatti, M. and Zamir, D. (1990) Linkage relationships of genes affecting bitterness and flesh color in watermelon. Journal of Heredity, 81, 162-165. [32] Henderson, W.R., Scott, G.H. and Wehner, T.C. (1998) Interaction of flesh color genes in watermelon. Journal of Heredity, 89, 50-53. doi:10.1093/jhered/89.1.50 [33] Guner, N. and Wehner, T.C. (2004) The genes of watermelon. HortScience, 39, 1175-1182. [34] Wehner, T.C. (2008) Overview of the genes of watermelon. Proceedings of the IXth EUCARPIA Meeting on Genetics and Breeding of Cucurbitaceae, Avignon, 21-24 May 2008, 79-89. [35] Gusmini, G. and Wehner, T.C. (2006) Qualitative inheritance of rind pattern and flesh color in watermelon. Journal of Heredity, 97, 177-185. doi:10.1093/jhered/esj023 [36] Hirschberg, J. (2001) Carotenoid biosynthesis in flowering plants. Current Opinion in Plant Biology, 4, 210-218. doi:10.1016/S1369-5266(00)00163-1 [37] Fraser, P.D. and Bramley, P.M. (2004) The biosynthesis and nutritional uses of carotenoids. Progress in Lipid Research, 43, 228-265. doi:10.1016/j.plipres.2003.10.002 [38] Sadmann, G., Romer, S. and Fraser, P.D. (2006) Understanding carotenoid metabolism as a necessity for genetic engineering of crop plants. Metabolic Engineering, 8, 291-302. doi:10.1016/j.ymben.2006.01.005 [39] Li, F.Q., Murillo, C. and Wurtzel, E.T. (2007) Maize Y9 encodes a product essential for 15-cis-zeta-carotene isomerization. Plant Physiology, 144, 1181-1189. doi:10.1104/pp.107.098996 [40] Christopher, I.C. and Barry, J.P. (2010) Source to sink: Regulation of carotenoid biosynthesis in plants. Trends in Plant Science, 15, 266-274. doi:10.1016/j.tplants.2010.02.003 [41] Zhao, W. (2011) Natural pigments for use in foods. Encyclopedia of biotechnology in agriculture and food. Taylor & Francis, London. [42] Auldridge, M.E., McCarty, D.R. and Klee, H.J. (2006) Plant carotenoid cleavage oxygenases and their apocarotenoid products. Current Opinion in Plant Biology, 9, 315-321. doi:10.1016/j.pbi.2006.03.005 [43] Cunningham Jr., F.X. and Gannt, E. (1998) Genes and enzymes of carotenoid biosynthesis in plants. Plant Physiology and Plant Molecular Biology, 49, 557-583. doi:10.1146/annurev.arplant.49.1.557 [44] Sauret-Gueto, S., Botella-Pavia, P. and Rodríguez-Concepción, M. (2003) Molecular tools for the metabolic engineering of carotenoid biosynthesis in plants. Recent Research Developments in Plant Molecular Biology, 1, 339-363. [45] Kato, M., Ikoma, Y., Matsumoto, H., Sugiura, M., Hyodo, H. and Yano, M. (2004) Accumulation of carotenoids and expression of carotenoid biosynthetic genes during maturation in citrus fruit. Plant Physiology, 134, 824-837. doi:10.1104/pp.103.031104 [46] Bang, H. (2005) Environmental and genetic strategies to improve carotenoids and quality in watermelon. Ph.D. Thesis, Texas A&M University, College Station. [47] Bang, H., Kim, S., Leskovar, D. and King, S. (2007) Development of a codominant CAPS marker for allelic selection between canary yellow and red watermelon based on SNP in lycopene β-cyclase (LCYB) gene. Molecular Breeding, 20, 63-72. doi:10.1007/s11032-006-9076-4 [48] Guo, S., Liu, J., Zheng, Y., Huang, M., Zhang, H., Gong, G., He, H., Ren, Y., Zhong, S., Fei, Z. and Xu, Y. (2011) Characterization of transcriptome dynamics during watermelon fruit development: Sequencing, assembly, annotation and gene expression profiles. BMC Genomics, 12, 454. doi:10.1186/1471-2164-12-454 [49] Kang, B. and Zhao, W. (2007) Isolation and characterization of a watermelon partial cDNA ecoding phytoene synthase gene. Acta Botanica Boreali-Occidentalia Sinica, 27, 908-911. (in Chinese)
[1] King, S.R., Davis, A.R. and Bang, H. (2009) New flesh colors in watermelon? HortScience, 44, 576. [2] Tadmor, Y., King, S., Levi, A., Davis, A., Meir, A., Wasserman, B., Hirschberg, J. and Lewinsohn, E. (2005) Comparative fruit colouration in watermelon and tomato. Food Research International, 38, 837-841. doi:10.1016/j.foodres.2004.07.011 [3] Perkins-Veazie, P., Collins, J.K., Davis, A.R. and Roberts, W. (2006) Carotenoid content of 50 watermelon cultivars. Journal of Agricultural and Food Chemistry, 54, 2593-2597. doi:10.1021/jf052066p [4] Jia, G., Zhao, W., Shen, H., Wang, B. and Ma, S. (2008) Analysis of carotenoids in watermelon by column chromatography separation. Journal of Anhui Agricultural Sciences, 36, 8416-8418. (in Chinese) [5] Kang, B., Zhao, W., Hou, Y. and Tian, P. (2010) Expression of carotenogenic genes during the development and ripening of watermelon fruit. Scientia Horticulturae, 124, 368-375. doi:10.1016/j.scienta.2010.01.027 [6] Hu, L. (2009) Separation and analysis of carotenoids in watermelon. Ms.D. Thesis, Zhengzhou University, Zhengzhou. (in Chinese) [7] Hu, L., Guo, X., Zhao, W., Wang, J. and Zhang, G. (2011) Separation and analysis of carotenoids in watermelon flesh. Journal of Henan Agricultural Sciences, 40, 150-156. (in Chinese) [8] Bang, H., Davis, A., Kim, S., Leskovar, D. and King, S. (2010) Flesh color inheritance and gene interactions among canary yellow, pale yellow, and red watermelon. Journal of the American Society for Horticultural Science, 135, 362-368. [9] Liu, C., Zhang, H., Dai, Z., Liu, X., Liu, Y., Deng, X., Chen, F. and Xu, J. (2012) Volatile chemical and carotenoid profiles in watermelons [Citrullus vulgaris (Thunb.) Schrad (Cucurbitaceae)] with different flesh colors. Food Science and Biotechnology, 21, 531-541. doi:10.1007/s10068-012-0068-3 [10] Hui, B. and Li, J. (2008) Content and composition comparisons of carotenoids between red-flesh watermelon and yellow-flesh watermelon. Food Science, 29, 587-591. (in Chinese) [11] Collins, J.K., Perkins-Veazie, P. and Roberts, W. (2006) Lycopene: From plants to humans. HortScience, 41, 1135-1144. [12] USDA-NCC (1998) Carotenoid database for US foods. http://www.nal.usda.gov/fnic/food [13] Heinonen, M.I., Ollilainen, V., Linkola, E.K., Varo Pertti, T. and Koivistoinen, P.E. (1989) Carotenoids in Finnish foods: vegetables, fruits and berries. Journal of Agricultural and Food Chemistry, 37, 655-659. doi:10.1021/jf00087a017 [14] Holden, J.M., Eldridge, A.L., Beecher, G.R., Buzzard, I.M., Bhagwat, S., Davis, C.S., Douglass, L.W., Gebhardt, E.S, Haytowitz, D. and Schakel, S. (1999) Carotenoid Content of U.S. foods: An update of the database. Journal of Food Composition and Analysis, 12, 169-196. doi:10.1006/jfca.1999.0827 [15] US Department of Agriculture, ARS (2005) USDA National Nutrient Database for Standard Reference, release 18. US Department of Agriculture, Washington DC. [16] Perkins-Veazie, P., Collins, J.K. and Pair, S.D. (2000) Watermelon lycopene content varies with cultivar and maturity stage. HortScience, 35, 412. [17] Perkins-Veazie, P., Collins, J.K., Pair, S.D. and Roberts, W. (2001) Lycopene content differs among red-fleshed watermelon cultivars. Journal of the Science of Food and Agriculture, 81, 983-987. doi:10.1002/jsfa.880 [18] Perkins-Veazie, P., Roberts, W., Collins, J.K. and Perez, K. (2003) Lycopene variation among watermelons: Culivars, potassium, and ripeness. HortScience, 38, 1295. [19] Perkins-Veazie, P. and Collins, J.K. (2004) Flesh quality and lycopene stability of fresh-cut watermelon. Postharvest Biology and Technology, 31, 159-166. doi:10.1016/j.postharvbio.2003.08.005 [20] Perkins-Veazie, P. and Collins, J.K. (2006) Carotenoid changes of intact watermelons after storage. Journal of Agricultural and Food Chemistry, 54, 5868-5874. doi:10.1021/jf0532664 [21] Perkins-Veazie, P. (2007) Ripening events in seeded watermelons. HortScience, 42, 927. [22] Perkins-Veazie, P. and Zhang, X. (2007) Grafting increases lycopene in seedless watermelon. HortScience, 42, 959. [23] Davis, A.R., Fish, W.W. and Perkins-Veazie, P. (2003) A rapid hexane-free method for analyzing lycopene content in watermelon. Journal of Food Science, 68, 326-332. doi:10.1111/j.1365-2621.2003.tb14160.x [24] Leskovar, D.I., Bang, H., Crosby, K.M., Maness, N., Franco, J.A. and Perkins-Veazie, P. (2004) Lycopene, carbohydrates, ascorbic acid and yield components of diploid and triploid watermelon cultivars and affected by deficit irrigation. Journal of Horticultural Science and Biotechnology, 79, 75-81. [25] Yuan, P., Liu, W., Lu, X., Zhao, S., Lu, J., Yan, Z., He, N., Zhu, H. and Guan, L. (2012) Change of lycopene contents during fruit development of different ploidy watermelon. Journal of Fruit Science, 29, 890-894. (in Chinese) [26] Jeffery, J., Davis, A. and King, S. (2012) Understanding the carotenoid biosynthetic pathway: Observation of four color variants of developing watermelon fruit. Israel Journal of Plant Sciences, 60, 425-434. [27] Sun, L., Ding, S., Li, J., Zhu, F. and Wang J. (2006) Comparative analysis of lycopene contents among 8 watermelon cultivars. Acta Agriculturae Shanghai, 22, 66-68. (in Chinese) [28] Wan, X., Liu, W., Yan, Z., Zhao, S., He, N., Liu, P. and Dai, J. (2011) Changes of the contents of functional substances including lycopene, citrulline and ascorbic acid during watermelon fruits development. Scientia Agricultura Sinica, 44, 2738-2747. (in Chinese) [29] Henderson, W.R. (1989) Inheritance of orange flesh color in watermelon. Cucurbit Genetics Cooperative Report, 12, 59-63. [30] Shimotsuma, M. (1963) Cytogenetical studies in the genus Citrullus. VII. Inheritance of several characters in wtaremelons. Japanese Journal of Breeding, 13, 235-240. [31] Navot, N., Sarfatti, M. and Zamir, D. (1990) Linkage relationships of genes affecting bitterness and flesh color in watermelon. Journal of Heredity, 81, 162-165. [32] Henderson, W.R., Scott, G.H. and Wehner, T.C. (1998) Interaction of flesh color genes in watermelon. Journal of Heredity, 89, 50-53. doi:10.1093/jhered/89.1.50 [33] Guner, N. and Wehner, T.C. (2004) The genes of watermelon. HortScience, 39, 1175-1182. [34] Wehner, T.C. (2008) Overview of the genes of watermelon. Proceedings of the IXth EUCARPIA Meeting on Genetics and Breeding of Cucurbitaceae, Avignon, 21-24 May 2008, 79-89. [35] Gusmini, G. and Wehner, T.C. (2006) Qualitative inheritance of rind pattern and flesh color in watermelon. Journal of Heredity, 97, 177-185. doi:10.1093/jhered/esj023 [36] Hirschberg, J. (2001) Carotenoid biosynthesis in flowering plants. Current Opinion in Plant Biology, 4, 210-218. doi:10.1016/S1369-5266(00)00163-1 [37] Fraser, P.D. and Bramley, P.M. (2004) The biosynthesis and nutritional uses of carotenoids. Progress in Lipid Research, 43, 228-265. doi:10.1016/j.plipres.2003.10.002 [38] Sadmann, G., Romer, S. and Fraser, P.D. (2006) Understanding carotenoid metabolism as a necessity for genetic engineering of crop plants. Metabolic Engineering, 8, 291-302. doi:10.1016/j.ymben.2006.01.005 [39] Li, F.Q., Murillo, C. and Wurtzel, E.T. (2007) Maize Y9 encodes a product essential for 15-cis-zeta-carotene isomerization. Plant Physiology, 144, 1181-1189. doi:10.1104/pp.107.098996 [40] Christopher, I.C. and Barry, J.P. (2010) Source to sink: Regulation of carotenoid biosynthesis in plants. Trends in Plant Science, 15, 266-274. doi:10.1016/j.tplants.2010.02.003 [41] Zhao, W. (2011) Natural pigments for use in foods. Encyclopedia of biotechnology in agriculture and food. Taylor & Francis, London. [42] Auldridge, M.E., McCarty, D.R. and Klee, H.J. (2006) Plant carotenoid cleavage oxygenases and their apocarotenoid products. Current Opinion in Plant Biology, 9, 315-321. doi:10.1016/j.pbi.2006.03.005 [43] Cunningham Jr., F.X. and Gannt, E. (1998) Genes and enzymes of carotenoid biosynthesis in plants. Plant Physiology and Plant Molecular Biology, 49, 557-583. doi:10.1146/annurev.arplant.49.1.557 [44] Sauret-Gueto, S., Botella-Pavia, P. and Rodríguez-Concepción, M. (2003) Molecular tools for the metabolic engineering of carotenoid biosynthesis in plants. Recent Research Developments in Plant Molecular Biology, 1, 339-363. [45] Kato, M., Ikoma, Y., Matsumoto, H., Sugiura, M., Hyodo, H. and Yano, M. (2004) Accumulation of carotenoids and expression of carotenoid biosynthetic genes during maturation in citrus fruit. Plant Physiology, 134, 824-837. doi:10.1104/pp.103.031104 [46] Bang, H. (2005) Environmental and genetic strategies to improve carotenoids and quality in watermelon. Ph.D. Thesis, Texas A&M University, College Station. [47] Bang, H., Kim, S., Leskovar, D. and King, S. (2007) Development of a codominant CAPS marker for allelic selection between canary yellow and red watermelon based on SNP in lycopene β-cyclase (LCYB) gene. Molecular Breeding, 20, 63-72. doi:10.1007/s11032-006-9076-4 [48] Guo, S., Liu, J., Zheng, Y., Huang, M., Zhang, H., Gong, G., He, H., Ren, Y., Zhong, S., Fei, Z. and Xu, Y. (2011) Characterization of transcriptome dynamics during watermelon fruit development: Sequencing, assembly, annotation and gene expression profiles. BMC Genomics, 12, 454. doi:10.1186/1471-2164-12-454 [49] Kang, B. and Zhao, W. (2007) Isolation and characterization of a watermelon partial cDNA ecoding phytoene synthase gene. Acta Botanica Boreali-Occidentalia Sinica, 27, 908-911. (in Chinese)