A global perspective on carotenoids: Metabolism, biotechnology, and benefits for nutrition and health

Progress in Lipid Research

Volumn 70, Issue , 2018, Pages 62-93

  Rodriguez Concepcion, Manuel ^a   Avalos, Javier ^b   Bonet, M Luisa ^c,j,k   Boronat, Albert ^a,d   Gomez Gomez, Lourdes ^e   Hornero Mendez, Damaso ^f   Limon, M Carmen ^b   Meléndez Martínez, Antonio J ^b   Olmedilla Alonso, Begoña ^g   Palou, Andreu ^c,j,k   Ribot, Joan ^c,j,k   Rodrigo, Maria J ^h   Zacarias, Lorenzo ^h   Zhu, Changfu ⁱ  

^a UNIVERSITAT AUTÒNOMA DE BARCELONA   (Spain)

^b UNIVERSITY OF SEVILLE   (Spain)

^c UNIVERSITAT DE LES ILLES BALEARS   (Spain)

^d UNIVERSITY OF BARCELONA   (Spain)

^e UNIVERSITY OF CASTILLA LA MANCHA   (Spain)

^f INSTITUTO DE LA GRASA   (Spain)

^g INSTITUTE OF FOOD SCIENCE   (Spain)

^h INSTITUTO DE AGROQUÍMICA Y TECNOLOGÍA DE ALIMENTOS CSIC   (Spain)

ⁱ UNIVERSITY OF LLEIDA   (Spain)

^j Instituto de Salud Carlos III   (Spain)

^k HEALTH RESEARCH INSTITUTE OF THE BALEARIC ISLANDS IDISBA   (Spain)

more..

Author keywords

Biotechnology; Carotenoid; Health; Metabolism; Nutrition; Pigment

Indexed keywords

BETA CAROTENE; CANTHAXANTHIN; CAPSANTHIN; CAROTENOID; CHYLOMICRON; CROCETIN; DECAPRENOXANTHIN; ISOPRENOID; LYCOPENE; PERIDININ; PHYTOENE; PHYTOFLUENE; RHODOXANTHIN; SEMI BETA CAROTENONE; TRANSCRIPTION FACTOR; TRANSCRIPTION FACTOR NRF2; UNCLASSIFIED DRUG; VIOLAXANTHIN; XANTHOPHYLL; ZEAXANTHIN; ZEAXANTHIN EPOXIDASE;

ABSORPTION; ANIMAL; ARCHAEON; BIOAVAILABILITY; BIOSYNTHESIS; BIOTECHNOLOGY; CELL DIFFERENTIATION; CHEMICAL STRUCTURE; CHLOROPLAST; CHROMOPLAST; CLASSIFICATION; CYCLIZATION; DIET; DIETARY INTAKE; ENZYME ACTIVE SITE; ENZYME LOCALIZATION; ENZYME MECHANISM; FOOD; FOOD COMPOSITION; FUNGUS; HUMAN; IN VITRO STUDY; IN VIVO STUDY; ISOMER; KNOWLEDGE; LIVER; MAIZE; MEDICAL RESEARCH; MICROALGA; NICOTIANA TABACUM; NON PHOTOSYNTHETIC BACTERIA; NONHUMAN; NUTRITION; OILSEED; OXIDATIVE STRESS; PHOTOTROPHIC BACTERIUM; PHYSICAL CHEMISTRY; PLANT CELL; PLANT LEAF; PLANT ROOT; PLASTID; POTATO; PRIORITY JOURNAL; PROTEIN CLEAVAGE; PROTEIN INTERACTION; REVIEW; RICE; SCIENTIST; SORGHUM; SOYBEAN; STORAGE; SWEET ORANGE; SWEET POTATO; TOMATO; VITAMIN METABOLISM; CROP; HEALTH; METABOLISM; NUTRITIONAL SCIENCE;

ANIMALS; BIOTECHNOLOGY; CAROTENOIDS; CROPS, AGRICULTURAL; HEALTH; HUMANS; NUTRITIONAL SCIENCES;

EID: 85046458195     PISSN: 01637827     EISSN: 18732194     Source Type: Journal    
DOI: 10.1016/j.plipres.2018.04.004     Document Type: Review

References (471)

1. Yabuzaki, J., Carotenoids Database: structures, chemical fingerprints and distribution among organisms. Database (Oxford), 2017, 2017.
2. Britton, G., Structure and properties of carotenoids in relation to function. FASEB J 9 (1995), 1551–1558.
3. Britton, G., Liaaen-Jensen, S., Pfander, H., Carotenoids Birkhäuser, Basel, Switzerland. 2004.
4. Weedon, B.C.L., Moss, G.P., Structure and nomenclature. Britton, G., Liaaen-Jensen, S., Pfander, H., (eds.) Carotenoids, 1995, Birkhauser, Basel, Switzerland, 27–44.
5. Melendez-Martinez, A.J., Paulino, M., Stinco, C.M., Mapelli-Brahm, P., Wang, X.D., Study of the time-course of cis/trans (Z/E) isomerization of lycopene, phytoene, and phytofluene from tomato. J Agric Food Chem 62 (2014), 12399–12406.
6. Chasse, G.A., Mak, M.L., Deretey, E., Farkas, I., Torday, L.L., Papp, J.G., et al. An ab initio computational study on selected lycopene isomers. J Mol Struct 571 (2001), 27–37.
7. Rodriguez-Amaya, D.B., Changes in carotenoids during processing and storage of foods. Arch Latinoam Nutr 49 (1999), 38S–47S.
8. Schieber, A., Carle, R., Occurrence of carotenoid cis-isomers in food: Technological, analytical, and nutritional implications. Trends Food Sci Technol 16 (2005), 416–422.
9. Than, A., Bramley, P.M., Davies, B.H., Rees, A.F., Stereochemistry of phytoene. Phytochemistry 11 (1972), 3187–3192.
10. Isaacson, T., Ronen, G., Zamir, D., Hirschberg, J., Cloning of tangerine from tomato reveals a carotenoid isomerase essential for the production of beta-carotene and xanthophylls in plants. Plant Cell 14 (2002), 333–342.
11. Schmitz, H.H., Emenhiser, C., Schwartz, S.J., HPLC separation of geometric carotene isomers using a calcium hydroxide stationary phase. J Agric Food Chem 43 (1995), 1212–1218.
12. Xu, C.J., Fraser, P.D., Wang, W.J., Bramley, P.M., Differences in the carotenoid content of ordinary citrus and lycopene-accumulating mutants. J Agric Food Chem 54 (2006), 5474–5481.
13. Meléndez-Martínez, A.J., Vicario, I.M., Heredia, F.J., Geometrical isomers of violaxanthin in orange juice. Food Chem 104 (2007), 169–175.
14. Meléndez-Martínez, A.J., Britton, G., Vicario, I.M., Heredia, F.J., Identification of isolutein (lutein epoxide) as cis-antheraxanthin in orange juice. J Agric Food Chem 53 (2005), 9369–9373.
15. Bohm, V., Puspitasari-Nienaber, N.L., Ferruzzi, M.G., Schwartz, S.J., Trolox equivalent antioxidant capacity of different geometrical isomers of alpha-carotene, beta-carotene, lycopene, and zeaxanthin. J Agric Food Chem 50 (2002), 221–226.
16. Stahl, W., Schwarz, W., Sundquist, A.R., Sies, H., cis-trans isomers of lycopene and beta-carotene in human serum and tissues. Arch Biochem Biophys 294 (1992), 173–177.
17. Meléndez-Martínez, A.J., Britton, G., Vicario, I.M., Heredia, F.J., The complex carotenoid pattern of orange juices from concentrate. Food Chem 109 (2008), 546–553.
18. Bone, R.A., Landrum, J.T., Hime, G.W., Cains, A., Zamor, J., Stereochemistry of the human macular carotenoids. Invest Ophthalmol Vis Sci 34 (1993), 2033–2040.
19. Rasmussen, H.M., Muzhingi, T., Eggert, E.M.R., Johnson, E.J., Lutein, zeaxanthin, meso-zeaxanthin content in egg yolk and their absence in fish and seafood. J Food Compos Anal 27 (2012), 139–144.
20. Johnson, E.J., Neuringer, M., Russell, R.M., Schalch, W., Snodderly, D.M., Nutritional manipulation of primate retinas, III: Effects of lutein or zeaxanthin supplementation on adipose tissue and retina of xanthophyll-free monkeys. Invest Ophthalmol Vis Sci 46 (2005), 692–702.
21. Mínguez-Mosquera, M.I., Hornero-Méndez, D., Changes in carotenoid esterification during the fruit ripening of capsicum annuum Cv. Bola. J Agric Food Chem 42 (1994), 640–644.
22. Delgado-Pelayo, R., Hornero-Méndez, D., Identification and quantitative analysis of carotenoids and their esters from sarsaparilla (Smilax aspera L.) berries. J Agric Food Chem 60 (2012), 8225–8232.
23. Murillo, E., Giuffrida, D., Menchaca, D., Dugo, P., Torre, G., Meléndez-Martinez, A.J., et al. Native carotenoids composition of some tropical fruits. Food Chem 140 (2013), 825–836.
24. Mercadante, A.Z., Rodrigues, D.B., Petry, F.C., Mariutti, L.R.B., Carotenoid esters in foods - A review and practical directions on analysis and occurrence. Food Res Int 99 ((2017), 830–850.
25. Breithaupt, D.E., Wirt, U., Bamedi, A., Differentiation between lutein monoester regioisomers and detection of lutein diesters from marigold flowers (Tagetes erecta L.) and Several fruits by liquid chromatography-mass spectrometry. J Agric Food Chem 50 (2002), 66–70.
26. Breitbaupt, D.E., Bamedi, A., Carotenoids and carotenoid esters in potatoes (Solanum tuberosum L.): New insights into an ancient vegetable. J Agric Food Chem 50 (2002), 7175–7181.
27. Pfister, S., Meyer, P., Steck, A., Pfander, H., Isolation and structure elucidation of carotenoid-glycosyl esters in gardenia fruits (Gardenia jasminoides Ellis) and saffron (Crocus sativus Linne). J Agric Food Chem 44 (1996), 2612–2615.
28. Bhosale, P., Bernstein, P.S., Vertebrate and invertebrate carotenoid-binding proteins. 2007, 121–127.
29. Shahidi, F., Metusalach, J.A., Brown, Carotenoid pigments in seafoods and aquaculture. Crit Rev Food Sci Nutr 38 (1998), 1–67.
30. Hoischen, D., Colmenares, L.U., Liu, J., Simmons, C.J., Britton, G., Liu, R.S.H., Fluorinated analogs of the carotenoprotein, α-crustacyanin. Bioorg Chem 26 (1998), 365–374.
31. Melendez-Martinez, A.J., Mapelli-Brahm, P., Benitez-Gonzalez, A., Stinco, C.M., A comprehensive review on the colorless carotenoids phytoene and phytofluene. Arch Biochem Biophys 572 (2015), 188–200.
32. Meléndez-Martínez, A.J., Paulino, M., Stinco, C.M., Mapelli-Brahm, P., Wang, X.D., Study of the time-course of cis/trans (Z/E) isomerization of lycopene, phytoene, and phytofluene from tomato. J Agric Food Chem 62 (2014), 12399–12406.
33. Martínez, A., Stinco, C.M., Meléndez-Martinez, A.J., Free radical scavenging properties of phytofluene and phytoene isomers as compared to Lycopene: a combined experimental and theoretical study. J Phys Chem B 118 (2014), 9819–9825.
34. Pokhilko, A., Bou-Torrent, J., Pulido, P., Rodriguez-Concepcion, M., Ebenhoh, O., Mathematical modelling of the diurnal regulation of the MEP pathway in Arabidopsis. New Phytol 206 (2015), 1075–1085.
35. Penuelas, J., Munne-Bosch, S., Isoprenoids: an evolutionary pool for photoprotection. Trends Plant Sci 10 (2005), 166–169.
36. Leonelli, L., Brooks, M.D., Niyogi, K.K., Engineering the lutein epoxide cycle into Arabidopsis thaliana. Proc Natl Acad Sci U S A 114 (2017), E7002–E7008.
37. Garcia-Plazaola, J.I., Esteban, R., Fernandez-Marin, B., Kranner, I., Porcar-Castell, A., Thermal energy dissipation and xanthophyll cycles beyond the Arabidopsis model. Photosynth Res 113 (2012), 89–103.
38. Esteban, R., Becerril, J.M., Garcia-Plazaola, J.I., Lutein epoxide cycle, more than just a forest tale. Plant Signal Behav 4 (2009), 342–344.
39. Hashimoto, H., Uragami, C., Cogdell, R.J., Carotenoids and photosynthesis. Subcell Biochem 79 (2016), 111–139.
40. Hou, X., Rivers, J., Leon, P., McQuinn, R.P., Pogson, B.J., Synthesis and function of apocarotenoid signals in plants. Trends Plant Sci 21 (2016), 792–803.
41. Llorente, B., Martinez-Garcia, J.F., Stange, C., Rodriguez-Concepcion, M., Illuminating colors: regulation of carotenoid biosynthesis and accumulation by light. Curr Opin Plant Biol 37 (2017), 49–55.
42. Britton, G., Functions of intact carotenoids. Britton, G., Liaaen-Jensen, S., Pfander, H., (eds.) Carotenoids, vol 4: natural functions, 2008, Birkhäuser Verlag, Basel, Switzerland, 189–212.
43. M.A. Ruiz-Sola, M. Rodriguez-Concepcion, Carotenoid biosynthesis in Arabidopsis: a colorful pathway, Arabidopsis book 10 (2012) e0158.
44. Nisar, N., Li, L., Lu, S., Khin, N.C., Pogson, B.J., Carotenoid metabolism in plants. Mol Plant 8 (2015), 68–82.
45. Moise, A.R., Al-Babili, S., Wurtzel, E.T., Mechanistic aspects of carotenoid biosynthesis. Chem Rev 114 (2014), 164–193.
46. Rodriguez-Concepcion, M., Boronat, A., Breaking new ground in the regulation of the early steps of plant isoprenoid biosynthesis. Curr Opin Plant Biol 25 (2015), 17–22.
47. Wright, L.P., Rohwer, J.M., Ghirardo, A., Hammerbacher, A., Ortiz-Alcaide, M., Raguschke, B., et al. Deoxyxylulose 5-phosphate synthase controls flux through the methylerythritol 4-phosphate pathway in arabidopsis. Plant Physiol 165 (2014), 1488–1504.
48. Hemmerlin, A., Post-translational events and modifications regulating plant enzymes involved in isoprenoid precursor biosynthesis. Plant Sci 203–204 (2013), 41–54.
49. Flores-Pérez, U., Sauret-Güeto, S., Gas, E., Jarvis, P., Rodríguez-Concepción, M., A mutant impaired in the production of plastome-encoded proteins uncovers a mechanism for the homeostasis of isoprenoid biosynthetic enzymes in arabidopsis plastids. Plant Cell 20 (2008), 1303–1315.
50. Pulido, P., Llamas, E., Llorente, B., Ventura, S., Wright, L.P., Rodriguez-Concepcion, M., 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. PLoS Genet, 12, 2016, e1005824.
51. Pulido, P., Toledo-Ortiz, G., Phillips, M.A., Wright, L.P., Rodriguez-Concepcion, M., Arabidopsis J-protein J20 delivers the first enzyme of the plastidial isoprenoid pathway to protein quality control. Plant Cell 25 (2013), 4183–4194.
52. Llamas, E., Pulido, P., Rodriguez-Concepcion, M., Interference with plastome gene expression and Clp protease activity in Arabidopsis triggers a chloroplast unfolded protein response to restore protein homeostasis. PLoS Genet, 13, 2017, e1007022.
53. Ghirardo, A., Wright, L.P., Bi, Z., Rosenkranz, M., Pulido, P., Rodriguez-Concepcion, M., et al. Metabolic flux analysis of plastidic isoprenoid biosynthesis in poplar leaves emitting and nonemitting isoprene. Plant Physiol 165 (2014), 37–51.
54. Sauret-Gueto, S., Botella-Pavia, P., Flores-Perez, U., Martinez-Garcia, J.F., San Roman, C., Leon, P., et al. Plastid cues posttranscriptionally regulate the accumulation of key enzymes of the methylerythritol phosphate pathway in Arabidopsis. Plant Physiol 141 (2006), 75–84.
55. Ruiz-Sola, M.A., Coman, D., Beck, G., Barja, M.V., Colinas, M., Graf, A., et al. Arabidopsis geranylgeranyl diphosphate synthase 11 is a hub isozyme required for the production of most photosynthesis-related isoprenoids. New Phytol 209 (2016), 252–264.
56. Ruiz-Sola, M.A., Barja, M.V., Manzano, D., Llorente, B., Schipper, B., Beekwilder, J., et al. A single arabidopsis gene encodes two differentially targeted geranylgeranyl diphosphate synthase isoforms. Plant Physiol 172 (2016), 1393–1402.
57. Zhou, F., Wang, C.Y., Gutensohn, M., Jiang, L., Zhang, P., Zhang, D., et al. A recruiting protein of geranylgeranyl diphosphate synthase controls metabolic flux toward chlorophyll biosynthesis in rice. Proc Natl Acad Sci U S A 114 (2017), 6866–6871.
58. Ji, J., Wang, G., Wang, J., Wang, P., Functional analysis of multiple carotenogenic genes from Lycium barbarum and Gentiana lutea L. for their effects on beta-carotene production in transgenic tobacco. Biotechnol Lett 31 (2009), 305–312.
59. Chen, W., He, S., Liu, D., Patil, G.B., Zhai, H., Wang, F., et al. a sweetpotato geranylgeranyl pyrophosphate synthase gene, ibggps, increases carotenoid content and enhances osmotic stress tolerance in Arabidopsis thaliana. PLoS One, 10, 2015, e0137623.
60. Tata, S.K., Jung, J., Kim, Y.H., Choi, J.Y., Jung, J.Y., Lee, I.J., et al. Heterologous expression of chloroplast-localized geranylgeranyl pyrophosphate synthase confers fast plant growth, early flowering and increased seed yield. Plant Biotechnol J 14 (2016), 29–39.
61. Fraser, P.D., Romer, S., Shipton, C.A., Mills, P.B., Kiano, J.W., Misawa, N., et al. Evaluation of transgenic tomato plants expressing an additional phytoene synthase in a fruit-specific manner. Proc Natl Acad Sci U S A 99 (2002), 1092–1097.
62. Liu, L., Shao, Z., Zhang, M., Wang, Q., Regulation of carotenoid metabolism in tomato. Mol Plant 8 (2015), 28–39.
63. Shumskaya, M., Wurtzel, E.T., The carotenoid biosynthetic pathway: thinking in all dimensions. Plant Sci 208 (2013), 58–63.
64. Shumskaya, M., Bradbury, L.M., Monaco, R.R., Wurtzel, E.T., Plastid localization of the key carotenoid enzyme phytoene synthase is altered by isozyme, allelic variation, and activity. Plant Cell 24 (2012), 3725–3741.
65. Welsch, R., Zhou, X., Yuan, H., Alvarez, D., Sun, T., Schlossarek, D., et al. Clp protease and OR directly control the proteostasis of phytoene synthase, the crucial enzyme for carotenoid biosynthesis in arabidopsis. Mol Plant 11 (2018), 149–162.
66. D'Andrea, L., Simon-Moya, M., Llorente, B., Llamas, E., Marro, M., Loza-Alvarez, P., et al. Interference with Clp protease impairs carotenoid accumulation during tomato fruit ripening. J Exp Bot 69 (2018), 1557–1568.
67. Zhou, X., Welsch, R., Yang, Y., Alvarez, D., Riediger, M., Yuan, H., et al. Arabidopsis OR proteins are the major posttranscriptional regulators of phytoene synthase in controlling carotenoid biosynthesis. Proc Natl Acad Sci U S A 112 (2015), 3558–3563.
68. Park, S., Kim, H.S., Jung, Y.J., Kim, S.H., Ji, C.Y., Wang, Z., et al. Orange protein has a role in phytoene synthase stabilization in sweetpotato. Sci Rep, 6, 2016, 33563.
69. Luo, Z., Zhang, J., Li, J., Yang, C., Wang, T., Ouyang, B., et al. A STAY-GREEN protein SlSGR1 regulates lycopene and beta-carotene accumulation by interacting directly with SlPSY1 during ripening processes in tomato. New Phytol 198 (2013), 442–452.
70. Arango, J., Jourdan, M., Geoffriau, E., Beyer, P., Welsch, R., Carotene hydroxylase activity determines the levels of both alpha-carotene and total carotenoids in orange carrots. Plant Cell 26 (2014), 2223–2233.
71. Kachanovsky, D.E., Filler, S., Isaacson, T., Hirschberg, J., Epistasis in tomato color mutations involves regulation of phytoene synthase 1 expression by cis-carotenoids. Proc Natl Acad Sci U S A 109 (2012), 19021–19026.
72. Chen, Y., Li, F., Wurtzel, E.T., Isolation and characterization of the Z-ISO gene encoding a missing component of carotenoid biosynthesis in plants. Plant Physiol 153 (2010), 66–79.
73. Li, R., Kang, C., Song, X., Yu, L., Liu, D., He, S., et al. A zeta-carotene desaturase gene, IbZDS, increases beta-carotene and lutein contents and enhances salt tolerance in transgenic sweetpotato. Plant Sci 262 (2017), 39–51.
74. McQuinn, R.P., Wong, B., Giovannoni, J.J., AtPDS overexpression in tomato: exposing unique patterns of carotenoid self-regulation and an alternative strategy for the enhancement of fruit carotenoid content. Plant Biotechnol J 16 (2018), 482–494.
75. Renato, M., Pateraki, I., Boronat, A., Azcon-Bieto, J., Tomato fruit chromoplasts behave as respiratory bioenergetic organelles during ripening. Plant Physiol 166 (2014), 920–933.
76. Nawrocki, W.J., Tourasse, N.J., Taly, A., Rappaport, F., Wollman, F.A., The plastid terminal oxidase: its elusive function points to multiple contributions to plastid physiology. Annu Rev Plant Biol 66 (2015), 49–74.
77. Renato, M., Boronat, A., Azcon-Bieto, J., Respiratory processes in non-photosynthetic plastids. Front Plant Sci, 6, 2015, 496.
78. Beltran, J., Kloss, B., Hosler, J.P., Geng, J., Liu, A., Modi, A., et al. Control of carotenoid biosynthesis through a heme-based cis-trans isomerase. Nat Chem Biol 11 (2015), 598–605.
79. Cazzonelli, C.I., Pogson, B.J., Source to sink: regulation of carotenoid biosynthesis in plants. Trends Plant Sci 15 (2010), 266–274.
80. Kim, J., Smith, J.J., Tian, L., Dellapenna, D., The evolution and function of carotenoid hydroxylases in Arabidopsis. Plant Cell Physiol 50 (2009), 463–479.
81. Mellado-Ortega, E., Hornero-Mendez, D., Isolation and identification of lutein esters, including their regioisomers, in tritordeum (xTritordeum Ascherson et Graebner) grains: Evidence for a preferential xanthophyll acyltransferase activity. Food Chem 135 (2012), 1344–1352.
82. Fernandez-Orozco, R., Gallardo-Guerrero, L., Hornero-Mendez, D., Carotenoid profiling in tubers of different potato (Solanum sp) cultivars: accumulation of carotenoids mediated by xanthophyll esterification. Food Chem 141 (2013), 2864–2872.
83. van Wijk, K.J., Kessler, F., Plastoglobuli: plastid microcompartments with integrated functions in metabolism, plastid developmental transitions, and environmental adaptation. Annu Rev Plant Biol 68 (2017), 253–289.
84. Ariizumi, T., Kishimoto, S., Kakami, R., Maoka, T., Hirakawa, H., Suzuki, Y., et al. Identification of the carotenoid modifying gene PALE YELLOW PETAL 1 as an essential factor in xanthophyll esterification and yellow flower pigmentation in tomato (Solanum lycopersicum). Plant J 79 (2014), 453–465.
85. Cunningham, F.X., Gantt, E., Genes and enzymes of carotenoid biosynthesis in plants. Ann Rev Plant Physiol Plant Mol Biol 49 (1998), 557–583.
86. Lopez, A.B., Yang, Y., Thannhauser, T.W., Li, L., Phytoene desaturase is present in a large protein complex in the plastid membrane. Physiol Plant 133 (2008), 190–198.
87. Ytterberg, A.J., Peltier, J.B., van Wijk, K.J., Protein profiling of plastoglobules in chloroplasts and chromoplasts. A surprising site for differential accumulation of metabolic enzymes. Plant Physiol 140 (2006), 984–997.
88. Barsan, C., Zouine, M., Maza, E., Bian, W., Egea, I., Rossignol, M., et al. Proteomic analysis of chloroplast-to-chromoplast transition in tomato reveals metabolic shifts coupled with disrupted thylakoid biogenesis machinery and elevated energy-production components. Plant Physiol 160 (2012), 708–725.
89. Hallin, E.I., Hasan, M., Guo, K., Akerlund, H.E., Molecular studies on structural changes and oligomerisation of violaxanthin de-epoxidase associated with the pH-dependent activation. Photosynth Res 129 (2016), 29–41.
90. Joyard, J., Ferro, M., Masselon, C., Seigneurin-Berny, D., Salvi, D., Garin, J., et al. Chloroplast proteomics and the compartmentation of plastidial isoprenoid biosynthetic pathways. Mol Plant 2 (2009), 1154–1180.
91. Welsch, R., Beyer, P., Hugueney, P., Kleinig, H., von Lintig, J., Regulation and activation of phytoene synthase, a key enzyme in carotenoid biosynthesis, during photomorphogenesis. Planta 211 (2000), 846–854.
92. Galpaz, N., Ronen, G., Khalfa, Z., Zamir, D., Hirschberg, J., A chromoplast-specific carotenoid biosynthesis pathway is revealed by cloning of the tomato white-flower locus. Plant Cell 18 (2006), 1947–1960.
93. Fraser, P.D., Schuch, W., Bramley, P.M., Phytoene synthase from tomato (Lycopersicon esculentum) chloroplasts—partial purification and biochemical properties. Planta 211 (2000), 361–369.
94. Nogueira, M., Mora, L., Enfissi, E.M., Bramley, P.M., Fraser, P.D., Subchromoplast sequestration of carotenoids affects regulatory mechanisms in tomato lines expressing different carotenoid gene combinations. Plant Cell 25 (2013), 4560–4579.
95. Li, L., Yuan, H., Zeng, Y., Xu, Q., Plastids and carotenoid accumulation. Subcell Biochem 79 (2016), 273–293.
96. Liebers, M., Grubler, B., Chevalier, F., Lerbs-Mache, S., Merendino, L., Blanvillain, R., et al. Regulatory shifts in plastid transcription play a key role in morphological conversions of plastids during plant development. Front Plant Sci, 8, 2017, 23.
97. Howitt, C.A., Pogson, B.J., Carotenoid accumulation and function in seeds and non-green tissues. Plant Cell Environ 29 (2006), 435–445.
98. Solymosi, K., Schoefs, B., Etioplast and etio-chloroplast formation under natural conditions: the dark side of chlorophyll biosynthesis in angiosperms. Photosynth Res 105 (2010), 143–166.
99. Rodriguez-Villalon, A., Gas, E., Rodriguez-Concepcion, M., Phytoene synthase activity controls the biosynthesis of carotenoids and the supply of their metabolic precursors in dark-grown Arabidopsis seedlings. Plant J 60 (2009), 424–435.
100. Yuan, H., Owsiany, K., Sheeja, T.E., Zhou, X., Rodriguez, C., Li, Y., et al. A Single amino acid substitution in an orange protein promotes carotenoid overaccumulation in arabidopsis. Plant Physiol 169 (2015), 421–431.
101. Egea, I., Barsan, C., Bian, W., Purgatto, E., Latche, A., Chervin, C., et al. Chromoplast differentiation: current status and perspectives. Plant Cell Physiol 51 (2010), 1601–1611.
102. Sun, T., Yuan, H., Cao, H., Yazdani, M., Tadmor, Y., Li, L., Carotenoid metabolism in plants: The role of plastids. Mol Plant 11 (2018), 58–74.
103. Schweiggert, R.M., Carle, R., Carotenoid deposition in plant and animal foods and its impact on bioavailability. Crit Rev Food Sci Nutr 57 (2017), 1807–1830.
104. Lado, J., Zacarias, L., Gurrea, A., Page, A., Stead, A., Rodrigo, M.J., Exploring the diversity in Citrus fruit colouration to decipher the relationship between plastid ultrastructure and carotenoid composition. Planta 242 (2015), 645–661.
105. Hempel, J., Amrehn, E., Quesada, S., Esquivel, P., Jimenez, V.M., Heller, A., et al. Lipid-dissolved gamma-carotene, beta-carotene, and lycopene in globular chromoplasts of peach palm (Bactris gasipaes Kunth) fruits. Planta 240 (2014), 1037–1050.
106. Sitte, P., Falk, H., Liedvogel, B., Chromoplasts. Czygan, F.G.C., (eds.) Pigments in plants, 1980, G Fischer Verl, Stuttgart, New York, 117–148.
107. Gross, J., Pigments in fruits. Schweigert, B.S., (eds.) Food science and technology: a series of monographs, 1987, London Academic Press.
108. Li, L., Paolillo, D.J., Parthasarathy, M.V., Dimuzio, E.M., Garvin, D.F., A novel gene mutation that confers abnormal patterns of beta-carotene accumulation in cauliflower (Brassica oleracea var. botrytis). Plant J 26 (2001), 59–67.
109. Lu, S., Van Eck, J., Zhou, X., Lopez, A.B., O'Halloran, D.M., Cosman, K.M., et al. The cauliflower Or gene encodes a DnaJ cysteine-rich domain-containing protein that mediates high levels of beta-carotene accumulation. Plant Cell 18 (2006), 3594–3605.
110. Lopez, A.B., Van Eck, J., Conlin, B.J., Paolillo, D.J., O'Neill, J., Li, L., Effect of the cauliflower Or transgene on carotenoid accumulation and chromoplast formation in transgenic potato tubers. J Exp Bot 59 (2008), 213–223.
111. Bai, C., Rivera, S.M., Medina, V., Alves, R., Vilaprinyo, E., Sorribas, A., et al. An in vitro system for the rapid functional characterization of genes involved in carotenoid biosynthesis and accumulation. Plant J 77 (2014), 464–475.
112. Bai, C., Capell, T., Berman, J., Medina, V., Sandmann, G., Christou, P., et al. Bottlenecks in carotenoid biosynthesis and accumulation in rice endosperm are influenced by the precursor-product balance. Plant Biotechnol J 14 (2016), 195–205.
113. Berman, J., Zorrilla-Lopez, U., Medina, V., Farre, G., Sandmann, G., Capell, T., et al. The Arabidopsis ORANGE (AtOR) gene promotes carotenoid accumulation in transgenic corn hybrids derived from parental lines with limited carotenoid pools. Plant Cell Rep 36 (2017), 933–945.
114. Chayut, N., Yuan, H., Ohali, S., Meir, A., Sa'ar, U., Tzuri, G., et al. Distinct Mechanisms of the ORANGE Protein in Controlling Carotenoid Flux. Plant Physiol 173 (2017), 376–389.
115. Kilambi, H.V., Kumar, R., Sharma, R., Sreelakshmi, Y., Chromoplast-specific carotenoid-associated protein appears to be important for enhanced accumulation of carotenoids in hp1 tomato fruits. Plant Physiol 161 (2013), 2085–2101.
116. Kilcrease, J., Rodriguez-Uribe, L., Richins, R.D., Arcos, J.M., Victorino, J., O'Connell, M.A., Correlations of carotenoid content and transcript abundances for fibrillin and carotenogenic enzymes in Capsicum annum fruit pericarp. Plant Sci 232 (2015), 57–66.
117. Nupur, L.N., Vats, A., Dhanda, S.K., Raghava, G.P., Pinnaka, A.K., Kumar, A., ProCarDB: A database of bacterial carotenoids. BMC Microbiol, 16, 2016, 96.
118. Chen, C.W., Hsu, S.H., Lin, M.T., Hsu, Y.H., Mass production of C50 carotenoids by Haloferax mediterranei in using extruded rice bran and starch under optimal conductivity of brined medium. Bioprocess Biosyst Eng 38 (2015), 2361–2367.
119. Rodrigo-Banos, M., Garbayo, I., Vilchez, C., Bonete, M.J., Martinez-Espinosa, R.M., Carotenoids from haloarchaea and their potential in biotechnology. Mar Drugs 13 (2015), 5508–5532.
120. Kelly, M., Norgard, S., Liaaen-Jensen, S., Bacterial carotenoids. 31. C50-carotenoids 5. Carotenoids of Halobacterium salinarium, especially bacterioruberin. Acta Chem Scand 24 (1970), 2169–2182.
121. Mandelli, F., Miranda, V.S., Rodrigues, E., Mercadante, A.Z., Identification of carotenoids with high antioxidant capacity produced by extremophile microorganisms. World J Microbiol Biotechnol 28 (2012), 1781–1790.
122. Calegari-Santos, R., Diogo, R.A., Fontana, J.D., Bonfim, T.M., Carotenoid production by halophilic archaea under different culture conditions. Curr Microbiol 72 (2016), 641–651.
123. Boronat, A., Rodriguez-Concepcion, M., Terpenoid biosynthesis in prokaryotes. Adv Biochem Eng Biotechnol 148 (2015), 3–18.
124. Hundle, B.S., Beyer, P., Kleinig, H., Englert, G., Hearst, J.E., Carotenoids of Erwinia herbicola and an Escherichia coli HB101 strain carrying the Erwinia herbicola carotenoid gene cluster. Photochem Photobiol 54 (1991), 89–93.
125. Armstrong, G.A., Eubacteria show their true colors: Genetics of carotenoid pigment biosynthesis from microbes to plants. J Bacteriol 176 (1994), 4795–4802.
126. Armstrong, G.A., Hearst, J.E., Carotenoids 2: Genetics and molecular biology of carotenoid pigment biosynthesis. FASEB J 10 (1996), 228–237.
127. Schweiggert, R.M., Carle, R., Carotenoid production by bacteria, microalgae, and fungi. Kaczor, A., Baranska, M., (eds.) Carotenoids: Nutrition, analysis and technology, 2016, John Wiley & Sons, Chichester, UK.
128. Misawa, N., Nakagawa, M., Kobayashi, K., Yamano, S., Izawa, Y., Nakamura, K., et al. Elucidation of the Erwinia uredovora carotenoid biosynthetic pathway by functional analysis of gene products expressed in Escherichia coli. J Bacteriol 172 (1990), 6704–6712.
129. Takano, H., The regulatory mechanism underlying light-inducible production of carotenoids in nonphototrophic bacteria. Biosci Biotechnol Biochem 80 (2016), 1264–1273.
130. Pasamontes, L., Hug, D., Tessier, M., Hohmann, H.P., Schierle, J., van Loon, A.P., Isolation and characterization of the carotenoid biosynthesis genes of Flavobacterium sp. strain R1534. Gene 185 (1997), 35–41.
131. Elias-Arnanz, M., Padmanabhan, S., Murillo, F.J., Light-dependent gene regulation in nonphototrophic bacteria. Curr Opin Microbiol 14 (2011), 128–135.
132. Galbis-Martinez, M., Padmanabhan, S., Murillo, F.J., Elias-Arnanz, M., CarF mediates signaling by singlet oxygen, generated via photoexcited protoporphyrin IX, in Myxococcus xanthus light-induced carotenogenesis. J Bacteriol 194 (2012), 1427–1436.
133. Armstrong, G.A., Genetics of eubacterial carotenoid biosynthesis: A colorful tale. Annu Rev Microbiol 51 (1997), 629–659.
134. Tian, B., Xu, Z., Sun, Z., Lin, J., Hua, Y., Evaluation of the antioxidant effects of carotenoids from Deinococcus radiodurans through targeted mutagenesis, chemiluminescence, and DNA damage analyses. Biochim Biophys Acta 1770 (2007), 902–911.
135. Zhang, L., Yang, Q., Luo, X., Fang, C., Zhang, Q., Tang, Y., Knockout of crtB or crtI gene blocks the carotenoid biosynthetic pathway in Deinococcus radiodurans R1 and influences its resistance to oxidative DNA-damaging agents due to change of free radicals scavenging ability. Arch Microbiol 188 (2007), 411–419.
136. Gao, P., Davies, J., Kao, R.Y.T., Dehydrosqualene desaturase as a novel target for anti-virulence therapy against Staphylococcus aureus. MBio, 8, 2017.
137. Liu, G.Y., Essex, A., Buchanan, J.T., Datta, V., Hoffman, H.M., Bastian, J.F., et al. Staphylococcus aureus golden pigment impairs neutrophil killing and promotes virulence through its antioxidant activity. J Exp Med 202 (2005), 209–215.
138. Takaichi, S., Carotenoids and carotenogenesis in anoxygenic photosynthetic bacteria. Frank, H.A., Young, A.J., Britton, G., Cogdell, R.J., (eds.) The photochemistry of carotenoids: Advances in photosynthesis, 1999, Kluwer Academic, Dordrecht, 39–69.
139. Takaichi, S., Distribution and biosynthesis of carotenoids. The purple phototrophic bacteria. Hunter, C.N., Daldal, F., Thurnauer, M.C., Beaty, J.T., (eds.) Adv photosynthesis respiration, 2009, 97–117.
140. Fraser, N.J., Hashimoto, H., Cogdell, R.J., Carotenoids and bacterial photosynthesis: The story so far. Photosynth Res 70 (2001), 249–256.
141. Toth, T.N., Chukhutsina, V., Domonkos, I., Knoppova, J., Komenda, J., Kis, M., et al. Carotenoids are essential for the assembly of cyanobacterial photosynthetic complexes. Biochim Biophys Acta 1847 (2015), 1153–1165.
142. Zakar, T., Laczko-Dobos, H., Toth, T.N., Gombos, Z., Carotenoids assist in cyanobacterial photosystem II assembly and function. Front Plant Sci, 7, 2016, 295.
143. Kirilovsky, D., Kerfeld, C.A., Cyanobacterial photoprotection by the orange carotenoid protein. Nat Plants, 2, 2016, 16180.
144. Goodwin, T.W., The nature and distribution of carotenoids in some blue-green algae. J Gen Microbiol 17 (1957), 467–473.
145. Takaichi, S., Carotenoids in algae: distributions, biosyntheses and functions. Mar Drugs 9 (2011), 1101–1118.
146. Takaichi, S., Mochimaru, M., Carotenoids and carotenogenesis in cyanobacteria: unique ketocarotenoids and carotenoid glycosides. Cell Mol Life Sci 64 (2007), 2607–2619.
147. Sandmann, G., Mautz, J., Breitenbach, J., Control of light-dependent keto carotenoid biosynthesis in Nostoc 7120 by the transcription factor NtcA. Z Naturforsch C 71 (2016), 303–311.
148. Mulders, K.J., Lamers, P.P., Martens, D.E., Wijffels, R.H., Phototrophic pigment production with microalgae: Biological constraints and opportunities. J Phycol 50 (2014), 229–242.
149. Raposo, M.F., de Morais, A.M., de Morais, R.M., Carotenoids from marine microalgae: A valuable natural source for the prevention of chronic diseases. Mar Drugs 13 (2015), 5128–5155.
150. Henriquez, V., Escobar, C., Galarza, J., Gimpel, J., Carotenoids in Microalgae. Subcell Biochem 79 (2016), 219–237.
151. Wichuk, K., Brynjolfsson, S., Fu, W., Biotechnological production of value-added carotenoids from microalgae: Emerging technology and prospects. Bioengineered 5 (2014), 204–208.
152. Solovchenko, A., Neverov, K., Carotenogenic response in photosynthetic organisms: a colorful story. Photosynth Res 133 (2017), 31–47.
153. Solovchenko, A.E., Physiological role of neutral lipid accumulation in eukaryotic microalgae under stresses. Russ J Plant Physiol 59 (2012), 167–176.
154. Del Campo, J.A., Garcia-Gonzalez, M., Guerrero, M.G., Outdoor cultivation of microalgae for carotenoid production: Current state and perspectives. Appl Microbiol Biotechnol 74 (2007), 1163–1174.
155. Guedes, A.C., Amaro, H.M., Malcata, F.X., Microalgae as sources of carotenoids. Mar Drugs 9 (2011), 625–644.
156. Lamers, P.P., Janssen, M., De Vos, R.C., Bino, R.J., Wijffels, R.H., Exploring and exploiting carotenoid accumulation in Dunaliella salina for cell-factory applications. Trends Biotechnol 26 (2008), 631–638.
157. Oren, A., A hundred years of Dunaliella research: 1905–2005. Saline Syst, 1, 2005, 2.
158. Shah, M.M., Liang, Y., Cheng, J.J., Daroch, M., Astaxanthin-producing green microalga Haematococcus pluvialis: From single cell to high value commercial products. Front Plant Sci, 7, 2016, 531.
159. Avalos, J., Díaz-Sánchez, V., García-Martínez, J., Castrillo, M., Ruger-Herreros, M., Limón, M.C., Carotenoids. Martín, J.F., García-Estrada, C., Zeilinger, S., (eds.) Biosynthesis and molecular genetics of fungal secondary metabolites, 2014, Springer, 149–185.
160. Avalos, J., Nordzieke, S., Parra, O., Pardo-Medina, J., Limón, M.C., Carotenoid production by filamentous fungi and yeasts. Sibirny, A.A., (eds.) Biotechnology of yeasts and filamentous fungi, 2017.
161. Estrada, A.F., Brefort, T., Mengel, C., Diaz-Sanchez, V., Alder, A., Al-Babili, S., et al. Ustilago maydis accumulates beta-carotene at levels determined by a retinal-forming carotenoid oxygenase. Fungal Genet Biol 46 (2009), 803–813.
162. Daub, M.E., Payne, G.A., The role of carotenoids in resistance of fungi to cercosporin. Phytopathology 79 (1989), 180–185.
163. Prado-Cabrero, A., Estrada, A.F., Al-Babili, S., Avalos, J., Identification and biochemical characterization of a novel carotenoid oxygenase: Elucidation of the cleavage step in the Fusarium carotenoid pathway. Mol Microbiol 64 (2007), 448–460.
164. Saelices, L., Youssar, L., Holdermann, I., Al-Babili, S., Avalos, J., Identification of the gene responsible for torulene cleavage in the Neurospora carotenoid pathway. Mol Genet Genomics 278 (2007), 527–537.
165. Estrada, A.F., Youssar, L., Scherzinger, D., Al-Babili, S., Avalos, J., The ylo-1 gene encodes an aldehyde dehydrogenase responsible for the last reaction in the Neurospora carotenoid pathway. Mol Microbiol 69 (2008), 1207–1220.
166. Diaz-Sanchez, V., Estrada, A.F., Trautmann, D., Al-Babili, S., Avalos, J., The gene carD encodes the aldehyde dehydrogenase responsible for neurosporaxanthin biosynthesis in Fusarium fujikuroi. FEBS J 278 (2011), 3164–3176.
167. Prado-Cabrero, A., Scherzinger, D., Avalos, J., Al-Babili, S., Retinal biosynthesis in fungi: Characterization of the carotenoid oxygenase CarX from Fusarium fujikuroi. Eukaryot Cell 6 (2007), 650–657.
168. Avalos, J., Cerdá-Olmedo, E., Fungal carotenoid production. Arora, D.K., (eds.) Handbook of fungal biotechnology, 2004, Marcel Dekker, Inc., New York, 367–378.
169. Pollmann, H., Breitenbach, J., Sandmann, G., Engineering of the carotenoid pathway in Xanthophyllomyces dendrorhous leading to the synthesis of zeaxanthin. Appl Microbiol Biotechnol 101 (2017), 103–111.
170. Ledetzky, N., Osawa, A., Iki, K., Pollmann, H., Gassel, S., Breitenbach, J., et al. Multiple transformation with the crtYB gene of the limiting enzyme increased carotenoid synthesis and generated novel derivatives in Xanthophyllomyces dendrorhous. Arch Biochem Biophys 545 (2014), 141–147.
171. Avalos, J., Limon, M.C., Biological roles of fungal carotenoids. Curr Genet 61 (2015), 309–324.
172. Prado, M.M., Prado-Cabrero, A., Fernandez-Martin, R., Avalos, J., A gene of the opsin family in the carotenoid gene cluster of Fusarium fujikuroi. Curr Genet 46 (2004), 47–58.
173. Estrada, A.F., Avalos, J., Regulation and targeted mutation of opsA, coding for the NOP-1 opsin orthologue in Fusarium fujikuroi. J Mol Biol 387 (2009), 59–73.
174. Sutter, R.P., Mutations affecting sexual development in Phycomyces blakesleeanus. Proc Natl Acad Sci U S A 72 (1975), 127–130.
175. Vershinin, A., Biological functions of carotenoids—diversity and evolution. Biofactors 10 (1999), 99–104.
176. Moran, N.A., Jarvik, T., Lateral transfer of genes from fungi underlies carotenoid production in aphids. Science 328 (2010), 624–627.
177. Altincicek, B., Kovacs, J.L., Gerardo, N.M., Horizontally transferred fungal carotenoid genes in the two-spotted spider mite Tetranychus urticae. Biol Lett 8 (2012), 253–257.
178. Cobbs, C., Heath, J., Stireman, J.O., 3rd, P. Abbot, Carotenoids in unexpected places: Gall midges, lateral gene transfer, and carotenoid biosynthesis in animals. Mol Phylogenet Evol 68 (2013), 221–228.
179. Novakova, E., Moran, N.A., Diversification of genes for carotenoid biosynthesis in aphids following an ancient transfer from a fungus. Mol Biol Evol 29 (2012), 313–323.
180. Zhao, C., Nabity, P.D., Phylloxerids share ancestral carotenoid biosynthesis genes of fungal origin with aphids and adelgids. PLoS One, 12, 2017, e0185484.
181. Grbic, M., Van Leeuwen, T., Clark, R.M., Rombauts, S., Rouze, P., Grbic, V., et al. The genome of Tetranychus urticae reveals herbivorous pest adaptations. Nature 479 (2011), 487–492.
182. Schweigert, F.J., Metabolism of carotenoids in mammals. Britton, G., Liaaen-Jensen, S., Pfander, H., (eds.) Carotenoids vol. 3: Biosynthesis and metabolism, 1998, Birkhäuser Verlag, Basel, Switzerland, 249–284.
183. Nozière, P., Graulet, B., Lucas, A., Martin, B., Grolier, P., Doreau, M., Carotenoids for ruminants: From forages to dairy products. 2006, 418–450.
184. Khachik, F., Spangler, C.J., Smith, J.C., Canfield, L.M., Steck, A., Pfander, H., Identification, quantification, and relative concentrations of carotenoids and their metabolites in human milk and serum. Anal Chem 69 (1997), 1873–1881.
185. Johnson, E.J., Role of lutein and zeaxanthin in visual and cognitive function throughout the lifespan. Nutr Rev 72 (2014), 605–612.
186. Perry, A., Rasmussen, H., Johnson, E.J., Xanthophyll (lutein, zeaxanthin) content in fruits, vegetables and corn and egg products. J Food Compos Anal 22 (2009), 9–15.
187. LaFountain, A.M., Prum, R.O., Frank, H.A., Diversity, physiology, and evolution of avian plumage carotenoids and the role of carotenoid-protein interactions in plumage color appearance. Arch Biochem Biophys 572 (2015), 201–212.
188. Maoka, T., Carotenoids in marine animals. 2011, 278–293.
189. Breithaupt, D.E., Modern application of xanthophylls in animal feeding - a review. 2007, 501–506.
190. Havaux, M., Carotenoid oxidation products as stress signals in plants. Plant J 79 (2014), 597–606.
191. Sui, X., Kiser, P.D., Lintig, J., Palczewski, K., Structural basis of carotenoid cleavage: From bacteria to mammals. Arch Biochem Biophys 539 (2013), 203–213.
192. White, M.D., Flashman, E., Catalytic strategies of the non-heme iron dependent oxygenases and their roles in plant biology. Curr Opin Chem Biol 31 (2016), 126–135.
193. Caceres, L.A., Lakshminarayan, S., Yeung, K.K., McGarvey, B.D., Hannoufa, A., Sumarah, M.W., et al. Repellent and attractive effects of alpha-, beta-, and dihydro-beta- ionone to generalist and specialist herbivores. J Chem Ecol 42 (2016), 107–117.
194. Rodriguez, A., Alquezar, B., Pena, L., Fruit aromas in mature fleshy fruits as signals of readiness for predation and seed dispersal. New Phytol 197 (2013), 36–48.
195. Macias, F.A., Lopez, A., Varela, R.M., Torres, A., Molinillo, J.M., Bioactive apocarotenoids annuionones F and G: Structural revision of annuionones A, B and E. Phytochemistry 65 (2004), 3057–3063.
196. Akiyama, K., Matsuzaki, K., Hayashi, H., Plant sesquiterpenes induce hyphal branching in arbuscular mycorrhizal fungi. Nature 435 (2005), 824–827.
197. Walter, M.H., Floss, D.S., Strack, D., Apocarotenoids: Hormones, mycorrhizal metabolites and aroma volatiles. Planta 232 (2010), 1–17.
198. McQuinn, R.P., Giovannoni, J.J., Pogson, B.J., More than meets the eye: From carotenoid biosynthesis, to new insights into apocarotenoid signaling. Curr Opin Plant Biol 27 (2015), 172–179.
199. Al-Babili, S., Bouwmeester, H.J., Strigolactones, a novel carotenoid-derived plant hormone. Annu Rev Plant Biol 66 (2015), 161–186.
200. Adhikari, E., Lee, D.K., Giavalisco, P., Sieburth, L.E., Long-distance signaling in bypass1 mutants: Bioassay development reveals the bps signal to be a metabolite. Mol Plant 6 (2013), 164–173.
201. Van Norman, J.M., Sieburth, L.E., Dissecting the biosynthetic pathway for the bypass1 root-derived signal. Plant J 49 (2007), 619–628.
202. Avendano-Vazquez, A.O., Cordoba, E., Llamas, E., San Roman, C., Nisar, N., De la Torre, S., et al. An uncharacterized apocarotenoid-derived signal generated in zeta-carotene desaturase mutants regulates leaf development and the expression of chloroplast and nuclear genes in arabidopsis. Plant Cell 26 (2014), 2524–2537.
203. Van Norman, J.M., Zhang, J., Cazzonelli, C.I., Pogson, B.J., Harrison, P.J., Bugg, T.D., et al. Periodic root branching in Arabidopsis requires synthesis of an uncharacterized carotenoid derivative. Proc Natl Acad Sci U S A 111 (2014), E1300–9.
204. Christodoulou, E., Kadoglou, N.P., Kostomitsopoulos, N., Valsami, G., Saffron: A natural product with potential pharmaceutical applications. J Pharm Pharmacol 67 (2015), 1634–1649.
205. Rivera-Madrid, R., Aguilar-Espinosa, M., Cardenas-Conejo, Y., Garza-Caligaris, L.E., Carotenoid derivates in achiote (Bixa orellana) seeds: Synthesis and health promoting properties. Front Plant Sci, 7, 2016, 1406.
206. Li, H.H., Hao, R.L., Wu, S.S., Guo, P.C., Chen, C.J., Pan, L.P., et al. Occurrence, function and potential medicinal applications of the phytohormone abscisic acid in animals and humans. Biochem Pharmacol 82 (2011), 701–712.
207. Fridlender, M., Kapulnik, Y., Koltai, H., Plant derived substances with anti-cancer activity: From folklore to practice. Front Plant Sci, 6, 2015, 799.
208. Croglio, M.P., Haake, J.M., Ryan, C.P., Wang, V.S., Lapier, J., Schlarbaum, J.P., et al. Analogs of the novel phytohormone, strigolactone, trigger apoptosis and synergize with PARP inhibitors by inducing DNA damage and inhibiting DNA repair. Oncotarget 7 (2016), 13984–14001.
209. Ansari, M., Emami, S., beta-Ionone and its analogs as promising anticancer agents. Eur J Med Chem 123 (2016), 141–154.
210. Kloer, D.P., Schulz, G.E., Structural and biological aspects of carotenoid cleavage. Cell Mol Life Sci 63 (2006), 2291–2303.
211. dela Sena, C., Riedl, K.M., Narayanasamy, S., Curley, R.W. Jr., Schwartz, S.J., Harrison, E.H., The human enzyme that converts dietary provitamin A carotenoids to vitamin A is a dioxygenase. J Biol Chem 289 (2014), 13661–13666.
212. Schmidt, H., Kurtzer, R., Eisenreich, W., Schwab, W., The carotenase AtCCD1 from Arabidopsis thaliana is a dioxygenase. J Biol Chem 281 (2006), 9845–9851.
213. Ahrazem, O., Gomez-Gomez, L., Rodrigo, M.J., Avalos, J., Limon, M.C., Carotenoid cleavage oxygenases from microbes and photosynthetic organisms: Features and functions. Int J Mol Sci, 17, 2016.
214. Schwartz, S.H., Tan, B.C., Gage, D.A., Zeevaart, J.A., McCarty, D.R., Specific oxidative cleavage of carotenoids by VP14 of maize. Science 276 (1997), 1872–1874.
215. Nambara, E., Marion-Poll, A., Abscisic acid biosynthesis and catabolism. Annu Rev Plant Biol 56 (2005), 165–185.
216. Alder, A., Jamil, M., Marzorati, M., Bruno, M., Vermathen, M., Bigler, P., et al. The path from beta-carotene to carlactone, a strigolactone-like plant hormone. Science 335 (2012), 1348–1351.
217. Lin, H., Wang, R., Qian, Q., Yan, M., Meng, X., Fu, Z., et al. DWARF27, an iron-containing protein required for the biosynthesis of strigolactones, regulates rice tiller bud outgrowth. Plant Cell 21 (2009), 1512–1525.
218. Waters, M.T., Brewer, P.B., Bussell, J.D., Smith, S.M., Beveridge, C.A., The Arabidopsis ortholog of rice DWARF27 acts upstream of MAX1 in the control of plant development by strigolactones. Plant Physiol 159 (2012), 1073–1085.
219. Schwartz, S.H., Qin, X., Zeevaart, J.A., Characterization of a novel carotenoid cleavage dioxygenase from plants. J Biol Chem 276 (2001), 25208–25211.
220. Simkin, A.J., Schwartz, S.H., Auldridge, M., Taylor, M.G., Klee, H.J., The tomato carotenoid cleavage dioxygenase 1 genes contribute to the formation of the flavor volatiles beta-ionone, pseudoionone, and geranylacetone. Plant J 40 (2004), 882–892.
221. Vallabhaneni, R., Bradbury, L.M., Wurtzel, E.T., The carotenoid dioxygenase gene family in maize, sorghum, and rice. Arch Biochem Biophys 504 (2010), 104–111.
222. da Silva Messias, R., Galli, V., Dos Anjos, E.S.S.D., Rombaldi, C.V., Carotenoid biosynthetic and catabolic pathways: Gene expression and carotenoid content in grains of maize landraces. Nutrients 6 (2014), 546–563.
223. Tan, B.C., Guan, J.C., Ding, S., Wu, S., Saunders, J.W., Koch, K.E., et al. Structure and origin of the white cap locus and its role in evolution of grain color in maize. Genetics 206 (2017), 135–150.
224. Floss, D.S., Walter, M.H., Role of carotenoid cleavage dioxygenase 1 (CCD1) in apocarotenoid biogenesis revisited. Plant Signal Behav 4 (2009), 172–175.
225. Ilg, A., Beyer, P., Al-Babili, S., Characterization of the rice carotenoid cleavage dioxygenase 1 reveals a novel route for geranial biosynthesis. FEBS J 276 (2009), 736–747.
226. Ilg, A., Bruno, M., Beyer, P., Al-Babili, S., Tomato carotenoid cleavage dioxygenases 1A and 1B: Relaxed double bond specificity leads to a plenitude of dialdehydes, mono-apocarotenoids and isoprenoid volatiles. FEBS Open Biol 4 (2014), 584–593.
227. Frusciante, S., Diretto, G., Bruno, M., Ferrante, P., Pietrella, M., Prado-Cabrero, A., et al. Novel carotenoid cleavage dioxygenase catalyzes the first dedicated step in saffron crocin biosynthesis. Proc Natl Acad Sci U S A 111 (2014), 12246–12251.
228. Ahrazem, O., Rubio-Moraga, A., Berman, J., Capell, T., Christou, P., Zhu, C., et al. The carotenoid cleavage dioxygenase CCD2 catalysing the synthesis of crocetin in spring crocuses and saffron is a plastidial enzyme. New Phytol 209 (2016), 650–663.
229. Rubio, A., Rambla, J.L., Santaella, M., Gomez, M.D., Orzaez, D., Granell, A., et al. Cytosolic and plastoglobule-targeted carotenoid dioxygenases from Crocus sativus are both involved in beta-ionone release. J Biol Chem 283 (2008), 24816–24825.
230. Huang, F.C., Molnar, P., Schwab, W., Cloning and functional characterization of carotenoid cleavage dioxygenase 4 genes. J Exp Bot 60 (2009), 3011–3022.
231. Bruno, M., Beyer, P., Al-Babili, S., The potato carotenoid cleavage dioxygenase 4 catalyzes a single cleavage of beta-ionone ring-containing carotenes and non-epoxidated xanthophylls. Arch Biochem Biophys 572 (2015), 126–133.
232. Rodrigo, M.J., Alquezar, B., Alos, E., Medina, V., Carmona, L., Bruno, M., et al. A novel carotenoid cleavage activity involved in the biosynthesis of Citrus fruit-specific apocarotenoid pigments. J Exp Bot 64 (2013), 4461–4478.
233. Bruno, M., Koschmieder, J., Wuest, F., Schaub, P., Fehling-Kaschek, M., Timmer, J., et al. Enzymatic study on AtCCD4 and AtCCD7 and their potential to form acyclic regulatory metabolites. J Exp Bot 67 (2016), 5993–6005.
234. Ma, G., Zhang, L., Matsuta, A., Matsutani, K., Yamawaki, K., Yahata, M., et al. Enzymatic formation of beta-citraurin from beta-cryptoxanthin and Zeaxanthin by carotenoid cleavage dioxygenase4 in the flavedo of citrus fruit. Plant Physiol 163 (2013), 682–695.
235. Ahrazem, O., Diretto, G., Argandona, J., Rubio-Moraga, A., Julve, J.M., Orzaez, D., et al. Evolutionarily distinct carotenoid cleavage dioxygenases are responsible for crocetin production in Buddleja davidii. J Exp Bot 68 (2017), 4663–4677.
236. Gonzalez-Jorge, S., Ha, S.H., Magallanes-Lundback, M., Gilliland, L.U., Zhou, A., Lipka, A.E., et al. Carotenoid cleavage dioxygenase4 is a negative regulator of beta-carotene content in Arabidopsis seeds. Plant Cell 25 (2013), 4812–4826.
237. Campbell, R., Ducreux, L.J., Morris, W.L., Morris, J.A., Suttle, J.C., Ramsay, G., et al. The metabolic and developmental roles of carotenoid cleavage dioxygenase 4 from potato. Plant Physiol 154 (2010), 656–664.
238. Farre, G., Sanahuja, G., Naqvi, S., Bai, C., Capell, T., Zhu, C., et al. Travel advice on the road to carotenoids in plants. Plant Sci 179 (2010), 28–48.
239. Giuliano, G., Provitamin A biofortification of crop plants: A gold rush with many miners. Curr Opin Biotechnol 44 (2017), 169–180.
240. Mann, V., Harker, M., Pecker, I., Hirschberg, J., Metabolic engineering of astaxanthin production in tobacco flowers. Nat Biotechnol 18 (2000), 888–892.
241. Zhu, C., Gerjets, T., Sandmann, G., Nicotiana glauca engineered for the production of ketocarotenoids in flowers and leaves by expressing the cyanobacterial crtO ketolase gene. Transgenic Res 16 (2007), 813–821.
242. Gerjets, T., Sandmann, M., Zhu, C., Sandmann, G., Metabolic engineering of ketocarotenoid biosynthesis in leaves and flowers of tobacco species. Biotechnol J 2 (2007), 1263–1269.
243. Hasunuma, T., Miyazawa, S., Yoshimura, S., Shinzaki, Y., Tomizawa, K., Shindo, K., et al. Biosynthesis of astaxanthin in tobacco leaves by transplastomic engineering. Plant J 55 (2008), 857–868.
244. Lu, Y., Stegemann, S., Agrawal, S., Karcher, D., Ruf, S., Bock, R., Horizontal transfer of a synthetic metabolic pathway between plant species. Curr Biol 27 (2017), 3034–3041 (e3).
245. Mortimer, C.L., Misawa, N., Perez-Fons, L., Robertson, F.P., Harada, H., Bramley, P.M., et al. The formation and sequestration of nonendogenous ketocarotenoids in transgenic Nicotiana glauca. Plant Physiol 173 (2017), 1617–1635.
246. Mortimer, C.L., Misawa, N., Ducreux, L., Campbell, R., Bramley, P.M., Taylor, M., et al. Product stability and sequestration mechanisms in Solanum tuberosum engineered to biosynthesize high value ketocarotenoids. Plant Biotechnol J 14 (2016), 140–152.
247. Harada, H., Maoka, T., Osawa, A., Hattan, J., Kanamoto, H., Shindo, K., et al. Construction of transplastomic lettuce (Lactuca sativa) dominantly producing astaxanthin fatty acid esters and detailed chemical analysis of generated carotenoids. Transgenic Res 23 (2014), 303–315.
248. Simpson, K., Quiroz, L.F., Rodriguez-Concepcion, M., Stange, C.R., Differential contribution of the first two enzymes of the mep pathway to the supply of metabolic precursors for carotenoid and chlorophyll biosynthesis in carrot (Daucus carota). Front Plant Sci, 7, 2016, 1344.
249. Maass, D., Arango, J., Wust, F., Beyer, P., Welsch, R., Carotenoid crystal formation in Arabidopsis and carrot roots caused by increased phytoene synthase protein levels. PLoS One, 4, 2009, e6373.
250. Moreno, J.C., Pizarro, L., Fuentes, P., Handford, M., Cifuentes, V., Stange, C., Levels of lycopene beta-cyclase 1 modulate carotenoid gene expression and accumulation in Daucus carota. PLoS One, 8, 2013, e58144.
251. Moreno, J.C., Cerda, A., Simpson, K., Lopez-Diaz, I., Carrera, E., Handford, M., et al. Increased Nicotiana tabacum fitness through positive regulation of carotenoid, gibberellin and chlorophyll pathways promoted by Daucus carota lycopene beta-cyclase (Dclcyb1) expression. J Exp Bot 67 (2016), 2325–2338.
252. Jayaraj, J., Devlin, R., Punja, Z., Metabolic engineering of novel ketocarotenoid production in carrot plants. Transgenic Res 17 (2008), 489–501.
253. Fray, R.G., Wallace, A., Fraser, P.D., Valero, D., Hedden, P., Bramley, P.M., et al. Constitutive expression of a fruit phytoene synthase gene in transgenic tomatoes causes dwarfism by redirecting metabolites from the gibberellin pathway. Plant J 8 (1995), 693–701.
254. Lois, L.M., Rodríguez-Concepción, M., Gallego, F., Campos, N., Boronat, A., Carotenoid biosynthesis during tomato fruit development: Regulatory role of 1-deoxy-D-xylulose 5-phosphate synthase. Plant J 22 (2000), 503–513.
255. Enfissi, E.M.A., Fraser, P.D., Lois, L.M., Boronat, A., Schuch, W., Bramley, P.M., Metabolic engineering of the mevalonate and non-mevalonate isopentenyl diphosphate-forming pathways for the production of health-promoting isoprenoids in tomato. Plant Biotech J 3 (2005), 17–27.
256. Romer, S., Fraser, P.D., Kiano, J.W., Shipton, C.A., Misawa, N., Schuch, W., et al. Elevation of the provitamin A content of transgenic tomato plants. Nat Biotechnol 18 (2000), 666–669.
257. Enfissi, E.M., Nogueira, M., Bramley, P.M., Fraser, P.D., The regulation of carotenoid formation in tomato fruit. Plant J 89 (2017), 774–788.
258. D'Ambrosio, C., Giorio, G., Marino, I., Merendino, A., Petrozza, A., Salfi, L., et al. Virtually complete conversion of lycopene into β-carotene in fruits of tomato plants transformed with the tomato lycopene β-cyclase (tlcy-b) cDNA. Plant Sci 166 (2004), 207–214.
259. Ralley, L., Schuch, W., Fraser, P.D., Bramley, P.M., Genetic modification of tomato with the tobacco lycopene beta-cyclase gene produces high beta-carotene and lycopene fruit. Z Naturforsch C 71 (2016), 295–301.
260. Ralley, L., Enfissi, E.M., Misawa, N., Schuch, W., Bramley, P.M., Fraser, P.D., Metabolic engineering of ketocarotenoid formation in higher plants. Plant J 39 (2004), 477–486.
261. Huang, J.C., Zhong, Y.J., Liu, J., Sandmann, G., Chen, F., Metabolic engineering of tomato for high-yield production of astaxanthin. Metab Eng 17 (2013), 59–67.
262. Nogueira, M., Enfissi, E.M.A., Martinez Valenzuela, M.E., Menard, G.N., Driller, R.L., Eastmond, P.J., et al. Engineering of tomato for the sustainable production of ketocarotenoids and its evaluation in aquaculture feed. Proc Natl Acad Sci U S A 114 (2017), 10876–10881.
263. Pons, E., Alquezar, B., Rodriguez, A., Martorell, P., Genoves, S., Ramon, D., et al. Metabolic engineering of beta-carotene in orange fruit increases its in vivo antioxidant properties. Plant Biotechnol J 12 (2014), 17–27.
264. Ducreux, L.J.M., Morris, W.L., Hedley, P.E., Shepherd, T., Davies, H.V., Millam, S., et al. Metabolic engineering of high carotenoid potato tubers containing enhanced levels of beta-carotene and lutein. J Exp Bot 56 (2005), 81–89.
265. Morris, W.L., Ducreux, L.J., Hedden, P., Millam, S., Taylor, M.A., Overexpression of a bacterial 1-deoxy-D-xylulose 5-phosphate synthase gene in potato tubers perturbs the isoprenoid metabolic network: implications for the control of the tuber life cycle. J Exp Bot 57 (2006), 3007–3018.
266. Diretto, G., Al-Babili, S., Tavazza, R., Papacchioli, V., Beyer, P., Giuliano, G., Metabolic engineering of potato carotenoid content through tuber-specific overexpression of a bacterial mini-pathway. PLoS One, 2, 2007, e350.
267. Diretto, G., Tavazza, R., Welsch, R., Pizzichini, D., Mourgues, F., Papacchioli, V., et al. Metabolic engineering of potato tuber carotenoids through tuber-specific silencing of lycopene epsilon cyclase. BMC Plant Biol, 6, 2006, 13.
268. Diretto, G., Welsch, R., Tavazza, R., Mourgues, F., Pizzichini, D., Beyer, P., et al. Silencing of beta-carotene hydroxylase increases total carotenoid and beta-carotene levels in potato tubers. BMC Plant Biol, 7, 2007, 11.
269. Romer, S., Lubeck, J., Kauder, F., Steiger, S., Adomat, C., Sandmann, G., Genetic engineering of a zeaxanthin-rich potato by antisense inactivation and co-suppression of carotenoid epoxidation. Metab Eng 4 (2002), 263–272.
270. Gerjets, T., Sandmann, G., Ketocarotenoid formation in transgenic potato. J Exp Bot 57 (2006), 3639–3645.
271. Morris, W.L., Ducreux, L.J., Fraser, P.D., Millam, S., Taylor, M.A., Engineering ketocarotenoid biosynthesis in potato tubers. Metab Eng 8 (2006), 253–263.
272. Li, L., Yang, Y., Xu, Q., Owsiany, K., Welsch, R., Chitchumroonchokchai, C., et al. The Or gene enhances carotenoid accumulation and stability during post-harvest storage of potato tubers. Mol Plant 5 (2012), 339–352.
273. Campbell, R., Morris, W.L., Mortimer, C.L., Misawa, N., Ducreux, L.J., Morris, J.A., et al. Optimising ketocarotenoid production in potato tubers: Effect of genetic background, transgene combinations and environment. Plant Sci 234 (2015), 27–37.
274. Park, S.C., Kim, S.H., Park, S., Lee, H.U., Lee, J.S., Park, W.S., et al. Enhanced accumulation of carotenoids in sweetpotato plants overexpressing IbOr-Ins gene in purple-fleshed sweetpotato cultivar. Plant Physiol Biochem 86 (2015), 82–90.
275. Shewmaker, C.K., Sheehy, J.A., Daley, M., Colburn, S., Ke, D.Y., Seed-specific overexpression of phytoene synthase: Increase in carotenoids and other metabolic effects. Plant J 20 (1999), 401–412.
276. Ravanello, M.P., Ke, D., Alvarez, J., Huang, B., Shewmaker, C.K., Coordinate expression of multiple bacterial carotenoid genes in canola leading to altered carotenoid production. Metab Eng 5 (2003), 255–263.
277. Fujisawa, M., Takita, E., Harada, H., Sakurai, N., Suzuki, H., Ohyama, K., et al. Pathway engineering of Brassica napus seeds using multiple key enzyme genes involved in ketocarotenoid formation. J Exp Bot 60 (2009), 1319–1332.
278. Yu, B., Lydiate, D.J., Young, L.W., Schafer, U.A., Hannoufa, A., Enhancing the carotenoid content of Brassica napus seeds by downregulating lycopene epsilon cyclase. Transgenic Res 17 (2008), 573–585.
279. Kim, M.J., Kim, J.K., Kim, H.J., Pak, J.H., Lee, J.H., Kim, D.H., et al. Genetic modification of the soybean to enhance the beta-carotene content through seed-specific expression. PLoS One, 7, 2012, e48287.
280. Schmidt, M.A., Parrott, W.A., Hildebrand, D.F., Berg, R.H., Cooksey, A., Pendarvis, K., et al. Transgenic soya bean seeds accumulating beta-carotene exhibit the collateral enhancements of oleate and protein content traits. Plant Biotechnol J 13 (2015), 590–600.
281. Pierce, E.C., LaFayette, P.R., Ortega, M.A., Joyce, B.L., Kopsell, D.A., Parrott, W.A., Ketocarotenoid production in soybean seeds through metabolic engineering. PLoS One, 10, 2015, e0138196.
282. Ye, X., Al-Babili, S., Kloti, A., Zhang, J., Lucca, P., Beyer, P., et al. Engineering the provitamin A (beta-carotene) biosynthetic pathway into (carotenoid-free) rice endosperm. Science 287 (2000), 303–305.
283. Paine, J.A., Shipton, C.A., Chaggar, S., Howells, R.M., Kennedy, M.J., Vernon, G., et al. Improving the nutritional value of Golden Rice through increased pro-vitamin A content. Nat Biotechnol 23 (2005), 482–487.
284. Pham, T.N., Dong, T.L., Tran, V.H., Tran, T.C.H., Effect of storage conditions on total carotenoid content in golden rice grains. Omonrice 14 (2006), 18–27.
285. Gayen, D., Ali, N., Sarkar, S.N., Datta, S.K., Datta, K., Down-regulation of lipoxygenase gene reduces degradation of carotenoids of golden rice during storage. Planta 242 (2015), 353–363.
286. Bai, C., Berman, J., Farre, G., Capell, T., Sandmann, G., Christou, P., et al. Reconstruction of the astaxanthin biosynthesis pathway in rice endosperm reveals a metabolic bottleneck at the level of endogenous beta-carotene hydroxylase activity. Transgenic Res 26 (2017), 13–23.
287. Aluru, M., Xu, Y., Guo, R., Wang, Z., Li, S., White, W., et al. Generation of transgenic maize with enhanced provitamin A content. J Exp Bot 59 (2008), 3551–3562.
288. Zhu, C., Naqvi, S., Breitenbach, J., Sandmann, G., Christou, P., Capell, T., Combinatorial genetic transformation generates a library of metabolic phenotypes for the carotenoid pathway in maize. Proc Natl Acad Sci U S A 105 (2008), 18232–18237.
289. Naqvi, S., Zhu, C., Farre, G., Sandmann, G., Capell, T., Christou, P., Synergistic metabolism in hybrid corn indicates bottlenecks in the carotenoid pathway and leads to the accumulation of extraordinary levels of the nutritionally important carotenoid zeaxanthin. Plant Biotechnol J 9 (2011), 384–393.
290. Berman, J., Zorrilla-Lopez, U., Sandmann, G., Capell, T., Christou, P., Zhu, C., The silencing of carotenoid beta-hydroxylases by rna interference in different maize genetic backgrounds increases the beta-carotene content of the endosperm. Int J Mol Sci, 18, 2017.
291. Farre, G., Perez-Fons, L., Decourcelle, M., Breitenbach, J., Hem, S., Zhu, C., et al. Metabolic engineering of astaxanthin biosynthesis in maize endosperm and characterization of a prototype high oil hybrid. Transgenic Res 25 (2016), 477–489.
292. Che, P., Zhao, Z.Y., Glassman, K., Dolde, D., Hu, T.X., Jones, T.J., et al. Elevated vitamin E content improves all-trans beta-carotene accumulation and stability in biofortified sorghum. Proc Natl Acad Sci U S A 113 (2016), 11040–11045.
293. Cong, L., Wang, C., Chen, L., Liu, H., Yang, G., He, G., Expression of phytoene synthase1 and carotene desaturase crtI genes result in an increase in the total carotenoids content in transgenic elite wheat (Triticum aestivum L.). J Agric Food Chem 57 (2009), 8652–8660.
294. Wang, C., Zeng, J., Li, Y., Hu, W., Chen, L., Miao, Y., et al. Enrichment of provitamin A content in wheat (Triticum aestivum L.) by introduction of the bacterial carotenoid biosynthetic genes CrtB and CrtI. J Exp Bot 65 (2014), 2545–2556.
295. Zeng, J., Wang, X., Miao, Y., Wang, C., Zang, M., Chen, X., et al. Metabolic engineering of wheat provitamin a by simultaneously overexpressing crtb and silencing carotenoid hydroxylase (TaHYD). J Agric Food Chem 63 (2015), 9083–9092.
296. Grune, T., Lietz, G., Palou, A., Ross, A.C., Stahl, W., Tang, G., et al. Beta-carotene is an important vitamin A source for humans. J Nutr 140 (2010), 2268S–2285S.
297. Maiani, G., Caston, M.J., Catasta, G., Toti, E., Cambrodon, I.G., Bysted, A., et al. Carotenoids: actual knowledge on food sources, intakes, stability and bioavailability and their protective role in humans. Mol Nutr Food Res 53:Suppl. 2 (2009), S194–218.
298. Howard, A.N., Thurnham, D.I., Lutein and atherosclerosis: Belfast versus Toulouse revisited. Med Hypotheses 98 (2017), 63–68.
299. Wang, X.D., Lycopene metabolism and its biological significance. Am J Clin Nutr, 96, 2012 (1214S-22S).
300. Yamaguchi, M., Role of carotenoid beta-cryptoxanthin in bone homeostasis. J Biomed Sci, 19, 2012, 36.
301. Cooper, D.A., Carotenoids in health and disease: recent scientific evaluations, research recommendations and the consumer. J Nutr 134 (2004), 221S–224S.
302. Fernández-García, E., Carvajal-Lérida, I., Jarén-Galán, M., Garrido-Fernández, J., Pérez-Gálvez, A., Hornero-Méndez, D., Carotenoids bioavailability from foods: From plant pigments to efficient biological activities. Food Res Int 46 (2012), 438–450.
303. Landrum, J.T., Bone, R.A., Lutein, zeaxanthin, and the macular pigment. Arch Biochem Biophys 385 (2001), 28–40.
304. Bonet, M.L., Canas, J.A., Ribot, J., Palou, A., Carotenoids and their conversion products in the control of adipocyte function, adiposity and obesity. Arch Biochem Biophys 572 (2015), 112–125.
305. Hamer, M., Chida, Y., Intake of fruit, vegetables, and antioxidants and risk of type 2 diabetes: Systematic review and meta-analysis. J Hypertens 25 (2007), 2361–2369.
306. Leermakers, E.T., Darweesh, S.K., Baena, C.P., Moreira, E.M., Melo van Lent, D., Tielemans, M.J., et al. The effects of lutein on cardiometabolic health across the life course: A systematic review and meta-analysis. Am J Clin Nutr 103 (2016), 481–494.
307. Milani, A., Basirnejad, M., Shahbazi, S., Bolhassani, A., Carotenoids: biochemistry, pharmacology and treatment. Br J Pharmacol 174 (2017), 1290–1324.
308. Britton, G., Khachik, F., Carotenoids in food. Britton, G., Liaaen-Jensen, S., Pfander, H., (eds.) Carotenoids vol. 5: Nutrition and health, 2009, Birkhauser, Basel, Switzerland, 45–66.
309. Lindbergh, C.A., Renzi-Hammond, L.M., Hammond, B.R., Terry, D.P., Mewborn, C.M., Puente, A.N., et al. Lutein and Zeaxanthin influence brain function in older adults: A randomized controlled trial. J Int Neuropsychol Soc 24 (2018), 77–90.
310. Hammond, B.R., Dietary carotenoids and the nervous system. Foods 4 (2015), 698–701.
311. Johnson, E.J., A possible role for lutein and zeaxanthin in cognitive function in the elderly. Am J Clin Nutr, 96, 2012 (1161S-5S).
312. Meléndez-Martínez, A.J., Mapelli-Brahm, P., Stinco, C.M., The colourless carotenoids phytoene and phytofluene: From dietary sources to their usefulness for the functional foods and nutricosmetics industries. J Food Compos Anal 67 (2018), 91–103.
313. Bjelakovic, G., Nikolova, D., Gluud, L.L., Simonetti, R.G., Gluud, C., Antioxidant supplements for prevention of mortality in healthy participants and patients with various diseases. Cochrane Database Syst Rev, 3, 2012, CD007176.
314. TATBCCPG, The alpha-tocopherol, beta-carotene lung cancer prevention study: Design, methods, participant characteristics, and compliance. The ATBC Cancer Prevention Study Group. Ann Epidemiol 4 (1994), 1–10.
315. Omenn, G.S., Goodman, G.E., Thornquist, M.D., Balmes, J., Cullen, M.R., Glass, A., et al. Effects of a combination of beta carotene and vitamin A on lung cancer and cardiovascular disease. N Engl J Med 334 (1996), 1150–1155.
316. Albanes, D., Heinonen, O.P., Taylor, P.R., Virtamo, J., Edwards, B.K., Rautalahti, M., et al. Alpha-Tocopherol and beta-carotene supplements and lung cancer incidence in the alpha-tocopherol, beta-carotene cancer prevention study: effects of base-line characteristics and study compliance. J Natl Cancer Inst 88 (1996), 1560–1570.
317. Monsen, E.R., Dietary reference intakes for the antioxidant nutrients: Vitamin C, vitamin E, selenium, and carotenoids. J Am Diet Assoc 100 (2000), 637–640.
318. Hennekens, C.H., Buring, J.E., Manson, J.E., Stampfer, M., Rosner, B., Cook, N.R., et al. Lack of effect of long-term supplementation with beta carotene on the incidence of malignant neoplasms and cardiovascular disease. N Engl J Med 334 (1996), 1145–1149.
319. Blot, W.J., Li, J.Y., Taylor, P.R., Guo, W., Dawsey, S.M., Li, B., The Linxian trials: Mortality rates by vitamin-mineral intervention group. Am J Clin Nutr, 62, 1995 (1424S-1426S).
320. Tang, G., Bioconversion of dietary provitamin A carotenoids to vitamin A in humans. Am J Clin Nutr, 91, 2010 (1468S-1473S).
321. Bonet, M.L., Ribot, J., Palou, A., Lipid metabolism in mammalian tissues and its control by retinoic acid. Biochim Biophys Acta 1821 (2012), 177–189.
322. Brun, P.J., Yang, K.J., Lee, S.A., Yuen, J.J., Blaner, W.S., Retinoids: Potent regulators of metabolism. Biofactors 39 (2013), 151–163.
323. Liu, Y., Chen, H., Mu, D., Fan, J., Song, J., Zhong, Y., et al. Circulating retinoic acid levels and the development of metabolic syndrome. J Clin Endocrinol Metab 101 (2016), 1686–1692.
324. Serra, F., Bonet, M.L., Puigserver, P., Oliver, J., Palou, A., Stimulation of uncoupling protein 1 expression in brown adipocytes by naturally occurring carotenoids. Int J Obes Relat Metab Disord 23 (1999), 650–655.
325. Amengual, J., Gouranton, E., van Helden, Y.G., Hessel, S., Ribot, J., Kramer, E., et al. Beta-carotene reduces body adiposity of mice via BCMO1. PLoS One, 6, 2011, e20644.
326. Carpentier, S., Knaus, M., Suh, M., Associations between lutein, zeaxanthin, and age-related macular degeneration: An overview. Crit Rev Food Sci Nutr 49 (2009), 313–326.
327. Sabour-Pickett, S., Nolan, J.M., Loughman, J., Beatty, S., A review of the evidence germane to the putative protective role of the macular carotenoids for age-related macular degeneration. Mol Nutr Food Res 56 (2012), 270–286.
328. Bernstein, P.S., Li, B., Vachali, P.P., Gorusupudi, A., Shyam, R., Henriksen, B.S., et al. Lutein, zeaxanthin, and meso-zeaxanthin: The basic and clinical science underlying carotenoid-based nutritional interventions against ocular disease. Prog Retin Eye Res 50 (2016), 34–66.
329. Nolan, J.M., Meagher, K., Kashani, S., Beatty, S., What is meso-zeaxanthin, and where does it come from?. Eye (Lond) 27 (2013), 899–905.
330. Krinsky, N.I., Landrum, J.T., Bone, R.A., Biologic mechanisms of the protective role of lutein and zeaxanthin in the eye. Annu Rev Nutr 23 (2003), 171–201.
331. Chew, E.Y., Clemons, T.E., Sangiovanni, J.P., Danis, R.P., Ferris, F.L., 3rd, M.J. Elman, et al. Secondary analyses of the effects of lutein/zeaxanthin on age-related macular degeneration progression: AREDS2 report No. 3. JAMA Ophthalmol 132 (2014), 142–149.
332. Mares, J., Lutein and zeaxanthin isomers in eye health and disease. Annu Rev Nutr 36 (2016), 571–602.
333. Rao, A.V., Rao, L.G., Carotenoids and human health. Pharmacol Res 55 (2007), 207–216.
334. Melendez-Martinez, A.J., Stinco, C.M., Liu, C., Wang, X.D., A simple HPLC method for the comprehensive analysis of cis/trans (Z/E) geometrical isomers of carotenoids for nutritional studies. Food Chem 138 (2013), 1341–1350.
335. Dias, M.G., Olmedilla-Alonso, B., Hornero-Méndez, D., Mercadante, A.Z., Osorio, C., Vargas-Murga, L., et al. Tabla de contenido en carotenoides de alimentos iberoamericanos. Meléndez-Martínez, A.J., (eds.) Carotenoides en agroalimentación y Salud, Terracota, 2017, Ciudad de México, México, 354–429.
336. Hornero-Mendez, D., Costa-Garcia, J., Minguez-Mosquera, M.I., Characterization of carotenoid high-producing Capsicum annuum cultivars selected for paprika production. J Agric Food Chem 50 (2002), 5711–5716.
337. Rodrigo, M.J., Marcos, J.F., Zacarias, L., Biochemical and molecular analysis of carotenoid biosynthesis in flavedo of orange (Citrus sinensis L.) during fruit development and maturation. J Agric Food Chem 52 (2004), 6724–6731.
338. Breithaupt, D.E., Bamedi, A., Carotenoid esters in vegetables and fruits: A screening with emphasis on beta-cryptoxanthin esters. J Agric Food Chem 49 (2001), 2064–2070.
339. Burri, B.J., Beta-cryptoxanthin as a source of vitamin A. J Sci Food Agric 95 (2015), 1786–1794.
340. Sajilata, M.G., Singhal, R.S., Kamat, M.Y., The carotenoid pigment zeaxanthin-A review. Comp Rev Food Sci Food Safety 7 (2008), 29–49.
341. Calvo, M.M., Lutein: A valuable ingredient of fruit and vegetables. Crit Rev Food Sci Nutr 45 (2005), 671–696.
342. Granado, F., Olmedilla, B., Blanco, I., Rojas Hidalgo, E., Carotenoid composition in raw and cooked Spanish vegetables. J Agric Food Chem, 40, 1992, 2135 2140.
343. Shi, J., Le Maguer, M., Lycopene in tomatoes: Chemical and physical properties affected by food processing. Crit Rev Biotechnol 20 (2000), 293–334.
344. Delgado-Pelayo, R., Gallardo-Guerrero, L., Hornero-Mendez, D., Carotenoid composition of strawberry tree (Arbutus unedo L.) fruits. Food Chem 199 (2016), 165–175.
345. Morales, P., Ferreira, I.C.F.R., Carvalho, A.M., Fernández-Ruiz, V., Sánchez-Mata, M.C., Cámara, M., et al. Wild edible fruits as a potential source of phytochemicals with capacity to inhibit lipid peroxidation. Eur J Lipid Sci Technol 115 (2013), 176–185.
346. de Rosso, V.V., Mercadante, A.Z., Identification and quantification of carotenoids, by HPLC-PDA-MS/MS, from Amazonian fruits. J Agric Food Chem 55 (2007), 5062–5072.
347. Sánchez-Mata, M.C., Cabrera Loera, R.D., Morales, P., Fernández-Ruiz, V., Cámara, M., Díez Marqués, C., et al. Wild vegetables of the Mediterranean area as valuable sources of bioactive compounds. Genet. Res. Crop Evol. 59 (2012), 431–443.
348. Murillo, E., Meléndez-Martínez, A.J., Portugal, F., Screening of vegetables and fruits from Panama for rich sources of lutein and zeaxanthin. Food Chem 122 (2010), 167–172.
349. Shewry, P.R., Hey, S., Do “ancient” wheat species differ from modern bread wheat in their contents of bioactive components?. J Cereal Sci 65 (2015), 236–243.
350. Atienza, S.G., Ballesteros, J., Martin, A., Hornero-Mendez, D., Genetic variability of carotenoid concentration and degree of esterification among tritordeum (xTritordeum Ascherson et Graebner) and durum wheat accessions. J Agric Food Chem 55 (2007), 4244–4251.
351. Brown, C.R., Breeding for phytonutrient enhancement of potato. Am J Potato Res 85 (2008), 298–307.
352. Britton, G., Overview on carotenoid biosynthesis. Britton, G., Liaaen-Jensen, S., Pfander, H., (eds.) Carotenoids, vol. 3: Biosynthesis, 1998, Birkhäuser Verlag, Basel, Switzerland, 13–147.
353. Camara, B., Brangeon, J., Carotenoid metabolism during chloroplast to chromoplast transformation in Capsicum annuum fruit. Planta 151 (1981), 359–364.
354. Hornero-Mendez, D., Minguez-Mosquera, M.I., Xanthophyll esterification accompanying carotenoid overaccumulation in chromoplast of Capsicum annuum ripening fruits is a constitutive process and useful for ripeness index. J Agric Food Chem 48 (2000), 1617–1622.
355. Granado, F., Olmedilla, B., Blanco, I., Gil-Martinez, E., Rojas-Hidalgo, E., Variability in the intercomparison of food carotenoid content data: a user's point of view. Crit Rev Food Sci Nutr 37 (1997), 621–633.
356. Mellado-Ortega, E., Hornero-Mendez, D., Lutein esterification in wheat flour increases the carotenoid retention and is induced by storage temperatures. Foods, 6, 2017.
357. Pérez-Gálvez, A., Minguez-Mosquera, M.I., Esterification of xanthophylls and its effect on chemical behavior and bioavailability of carotenoids in the human. Nutr Res 25 (2005), 631–640.
358. Rodriguez-Amaya, D.B., A guide to carotenoid analysis in foods. 2001.
359. Knockaert, G., Lemmens, L., Van Buggenhout, S., Hendrickx, M., Van Loey, A., Changes in β-carotene bioaccessibility and concentration during processing of carrot puree. Food Chem 133 (2012), 60–67.
360. Updike, A.A., Schwartz, S.J., Thermal processing of vegetables increases cis isomers of lutein and zeaxanthin. J Agric Food Chem 51 (2003), 6184–6190.
361. Mercadante, A.Z., Rodriguez-Amaya, D.B., Effects of ripening, cultivar differences, and processing on the carotenoid composition of mango. J Agric Food Chem 46 (1998), 128–130.
362. O'Neill, M.E., Carroll, Y., Corridan, B., Olmedilla, B., Granado, F., Blanco, I., et al. A European carotenoid database to assess carotenoid intakes and its use in a five-country comparative study. Br J Nutr 85 (2001), 499–507.
363. Biehler, E., Alkerwi, A.a., Hoffmann, L., Krause, E., Guillaume, M., Lair, M.L., et al. Contribution of violaxanthin, neoxanthin, phytoene and phytofluene to total carotenoid intake: Assessment in Luxembourg. J Food Compos Anal 25 (2012), 56–65.
364. Rodriguez-Amaya, D.B., Kimura, M., Godoy, H.T., Amaya-Farfan, J., Updated Brazilian database on food carotenoids: Factors affecting carotenoid composition. J Food Compos Anal 21 (2008), 445–463.
365. Beltran, B., Estevez, R., Cuadrado, C., Jimenez, S., Olmedilla-Alonso, B., Base de datos de carotenoides para valoracion de la ingesta dietetica de carotenos, xantofilas y de vitamina A; Utilizacion en un estudio comparativo del estado nutricional en vitamina A de adultos jovenes. Nutr Hosp 27 (2012), 1334–1343.
366. Bramley, P.M., Carotenoid biosynthesis and chlorophyll degradation. Seymour, G.B., Poole, M., Giovannoni, J.J., Tucker, G.A., (eds.) The molecular biology and biochemistry of fruit ripening, 2013, Wiley, 76–112.
367. Jenab, M., Salvini, S., van Gils, C.H., Brustad, M., Shakya-Shrestha, S., Buijsse, B., et al. Dietary intakes of retinol, beta-carotene, vitamin D and vitamin E in the European Prospective Investigation into Cancer and Nutrition cohort. Eur J Clin Nutr 63:Suppl. 4 (2009), S150–78.
368. Bohn, T., McDougall, G.J., Alegria, A., Alminger, M., Arrigoni, E., Aura, A.M., et al. Mind the gap-deficits in our knowledge of aspects impacting the bioavailability of phytochemicals and their metabolites—a position paper focusing on carotenoids and polyphenols. Mol Nutr Food Res 59 (2015), 1307–1323.
369. Bohn, T., Desmarchelier, C., Dragsted, L.O., Nielsen, C.S., Stahl, W., Ruhl, R., et al. Host-related factors explaining interindividual variability of carotenoid bioavailability and tissue concentrations in humans. Mol Nutr Food Res, 61, 2017.
370. Ranard, K.M., Jeon, S., Mohn, E.S., Griffiths, J.C., Johnson, E.J., Erdman, J.W. Jr., Dietary guidance for lutein: consideration for intake recommendations is scientifically supported. Eur J Nutr 56 (2017), 37–42.
371. Huang, Y.M., Dou, H.L., Huang, F.F., Xu, X.R., Zou, Z.Y., Lin, X.M., Effect of supplemental lutein and zeaxanthin on serum, macular pigmentation, and visual performance in patients with early age-related macular degeneration. Biomed Res Int, 2015, 2015, 564738.
372. Aggett, P.J., Population reference intakes and micronutrient bioavailability: A European perspective. Am J Clin Nutr, 91, 2010 (1433S-1437S).
373. Rodriguez-Amaya, D., Status of carotenoid analytical methods and in vitro assays for the assessment of food quality and health effects. Curr. Opin Food Sci 1 (2015), 56–63.
374. Bohn, T., Bioavailability of non-provitamin A carotenoids. Curr Nutr Food Sci 4 (2008), 240–258.
375. Castenmiller, J.J., West, C.E., Bioavailability and bioconversion of carotenoids. Annu Rev Nutr 18 (1998), 19–38.
376. Lemmens, L., Colle, I., Van Buggenhout, S., Palmero, P., Van Loey, A., Hendrickx, M., Carotenoid bioaccesibility in fruit- and vegetable-based food products as affected by product (micro) structural characteristics and the presente of lipids: A review. Trends Food Sci Technol 38 (2014), 125–135.
377. van Lieshout, M., West, C.E., van Breemen, R.B., Isotopic tracer techniques for studying the bioavailability and bioefficacy of dietary carotenoids, particularly beta-carotene, in humans: a review. Am J Clin Nutr 77 (2003), 12–28.
378. Carbonell-Capella, J.M., Buniowska, M., Barba, F.J., Esteve, M.J., Frígola, A., Analytical methods for determining bioavailability and bioaccesibility of bioactive compounds from fruits and vegetables: A review. Compr. Rev. Food Sci. Food Safety 13 (2014), 155–171.
379. Saini, R.K., Nile, S.H., Park, S.W., Carotenoids from fruits and vegetables: Chemistry, analysis, occurrence, bioavailability and biological activities. Food Res Int 76 (2015), 735–750.
380. Venema, K., Van Nuenen, M., Smeets-Peeters, M., Minekus, M., Havenaar, R., TNO's in vitro large intestinal model: An excellent screening tool for functional food and pharmaceutical research. Nutr Res 24 (2000), 558–564.
381. Minekus, M., Alminger, M., Alvito, P., Ballance, S., Bohn, T., Bourlieu, C., et al. A standardised static in vitro digestion method suitable for food - an international consensus. Food Funct 5 (2014), 1113–1124.
382. Estevez-Santiago, R., Olmedilla-Alonso, B., Fernandez-Jalao, I., Bioaccessibility of provitamin A carotenoids from fruits: Application of a standardised static in vitro digestion method. Food Funct 7 (2016), 1354–1366.
383. Granado-Lorencio, F., Blanco-Navarro, I., Perez-Sacristan, B., Hernandez-Alvarez, E., Biomarkers of carotenoid bioavailability. Food Res Int 99 (2017), 902–916.
384. Trumbo, P., Yates, A.A., Schlicker, S., Poos, M., Dietary reference intakes: Vitamin A, vitamin K, arsenic, boron, chromium, copper, iodine, iron, manganese, molybdenum, nickel, silicon, vanadium, and zinc. J Am Diet Assoc 101 (2001), 294–301.
385. Burri, B.J., Chang, J.S., Neidlinger, T.R., beta-Cryptoxanthin- and alpha-carotene-rich foods have greater apparent bioavailability than beta-carotene-rich foods in Western diets. Br J Nutr 105 (2011), 212–219.
386. Harrison, E.H., Mechanisms involved in the intestinal absorption of dietary vitamin A and provitamin A carotenoids. Biochim Biophys Acta 1821 (2012), 70–77.
387. Eroglu, A., Harrison, E.H., Carotenoid metabolism in mammals, including man: formation, occurrence, and function of apocarotenoids. J Lipid Res 54 (2013), 1719–1730.
388. Reboul, E., Absorption of vitamin A and carotenoids by the enterocyte: Focus on transport proteins. Nutrients 5 (2013), 3563–3581.
389. Shete, V., Quadro, L., Mammalian metabolism of beta-carotene: gaps in knowledge. Nutrients 5 (2013), 4849–4868.
390. von Lintig, J., Provitamin A metabolism and functions in mammalian biology. Am J Clin Nutr, 96, 2012 (1234S-44S).
391. Yeum, K.J., Russell, R.M., Carotenoid bioavailability and bioconversion. Annu Rev Nutr 22 (2002), 483–504.
392. Mapelli-Brahm, P., Corte-Real, J., Melendez-Martinez, A.J., Bohn, T., Bioaccessibility of phytoene and phytofluene is superior to other carotenoids from selected fruit and vegetable juices. Food Chem 229 (2017), 304–311.
393. Chitchumroonchokchai, C., Failla, M.L., Hydrolysis of zeaxanthin esters by carboxyl ester lipase during digestion facilitates micellarization and uptake of the xanthophyll by Caco-2 human intestinal cells. J Nutr 136 (2006), 588–594.
394. Tyssandier, V., Cardinault, N., Caris-Veyrat, C., Amiot, M.J., Grolier, P., Bouteloup, C., et al. Vegetable-borne lutein, lycopene, and beta-carotene compete for incorporation into chylomicrons, with no adverse effect on the medium-term (3-wk) plasma status of carotenoids in humans. Am J Clin Nutr 75 (2002), 526–534.
395. Lobo, G.P., Amengual, J., Li, H.N., Golczak, M., Bonet, M.L., Palczewski, K., et al. Beta,beta-carotene decreases peroxisome proliferator receptor gamma activity and reduces lipid storage capacity of adipocytes in a beta,beta-carotene oxygenase 1-dependent manner. J Biol Chem 285 (2010), 27891–27899.
396. Musinovic, H., Bonet, M.L., Granados, N., Amengual, J., von Lintig, J., Ribot, J., et al. beta-Carotene during the suckling period is absorbed intact and induces retinoic acid dependent responses similar to preformed vitamin A in intestine and liver, but not adipose tissue of young rats. Mol Nutr Food Res 58 (2014), 2157–2165.
397. Lobo, G.P., Hessel, S., Eichinger, A., Noy, N., Moise, A.R., Wyss, A., et al. ISX is a retinoic acid-sensitive gatekeeper that controls intestinal beta,beta-carotene absorption and vitamin A production. FASEB J 24 (2010), 1656–1666.
398. van den Berg, H., Faulks, R., Granado, H.F., Hirschberg, J., Olmedilla, B., Sandmann, G., et al. The potential for the improvement of carotenoid levels in foods and the likely systemic effects. J Sci Food Agric 80 (2000), 880–912.
399. Moussa, M., Gouranton, E., Gleize, B., Yazidi, C.E., Niot, I., Besnard, P., et al. CD36 is involved in lycopene and lutein uptake by adipocytes and adipose tissue cultures. Mol Nutr Food Res 55 (2011), 578–584.
400. Li, B., Vachali, P., Bernstein, P.S., Human ocular carotenoid-binding proteins. Photochem Photobiol Sci 9 (2010), 1418–1425.
401. Kawaguchi, R., Yu, J., Honda, J., Hu, J., Whitelegge, J., Ping, P., et al. A membrane receptor for retinol binding protein mediates cellular uptake of vitamin A. Science 315 (2007), 820–825.
402. Berry, D.C., Jacobs, H., Marwarha, G., Gely-Pernot, A., O'Byrne, S.M., DeSantis, D., et al. The STRA6 receptor is essential for retinol-binding protein-induced insulin resistance but not for maintaining vitamin A homeostasis in tissues other than the eye. J Biol Chem 288 (2013), 24528–24539.
403. Berry, D.C., Jin, H., Majumdar, A., Noy, N., Signaling by vitamin A and retinol-binding protein regulates gene expression to inhibit insulin responses. Proc Natl Acad Sci U S A 108 (2011), 4340–4345.
404. Alapatt, P., Guo, F., Komanetsky, S.M., Wang, S., Cai, J., Sargsyan, A., et al. Liver retinol transporter and receptor for serum retinol-binding protein (RBP4). J Biol Chem 288 (2013), 1250–1265.
405. Amengual, J., Widjaja-Adhi, M.A., Rodriguez-Santiago, S., Hessel, S., Golczak, M., Palczewski, K., et al. Two carotenoid oxygenases contribute to mammalian provitamin A metabolism. J Biol Chem 288 (2013), 34081–34096.
406. Hessel, S., Eichinger, A., Isken, A., Amengual, J., Hunzelmann, S., Hoeller, U., et al. CMO1 deficiency abolishes vitamin A production from beta-carotene and alters lipid metabolism in mice. J Biol Chem 282 (2007), 33553–33561.
407. Amengual, J., Lobo, G.P., Golczak, M., Li, H.N., Klimova, T., Hoppel, C.L., et al. A mitochondrial enzyme degrades carotenoids and protects against oxidative stress. FASEB J 25 (2011), 948–959.
408. Lindqvist, A., Andersson, S., Biochemical properties of purified recombinant human beta-carotene 15,15'-monooxygenase. J Biol Chem 277 (2002), 23942–23948.
409. Lobo, G.P., Isken, A., Hoff, S., Babino, D., von Lintig, J., BCDO2 acts as a carotenoid scavenger and gatekeeper for the mitochondrial apoptotic pathway. Development 139 (2012), 2966–2977.
410. Napoli, J.L., Cellular retinoid binding-proteins, CRBP, CRABP, FABP5: Effects on retinoid metabolism, function and related diseases. Pharmacol Ther 173 (2017), 19–33.
411. Krinsky, N.I., Yeum, K.J., Carotenoid-radical interactions. Biochem Biophys Res Commun 305 (2003), 754–760.
412. Ben-Dor, A., Steiner, M., Gheber, L., Danilenko, M., Dubi, N., Linnewiel, K., et al. Carotenoids activate the antioxidant response element transcription system. Mol Cancer Ther 4 (2005), 177–186.
413. Fiedor, J., Burda, K., Potential role of carotenoids as antioxidants in human health and disease. Nutrients 6 (2014), 466–488.
414. Johnson, E.J., Role of lutein and zeaxanthin in visual and cognitive function throughout the lifespan. Nutr Rev 72 (2014), 605–612.
415. Kaulmann, A., Bohn, T., Carotenoids, inflammation, and oxidative stress—implications of cellular signaling pathways and relation to chronic disease prevention. Nutr Res 34 (2014), 907–929.
416. Sosa, V., Moline, T., Somoza, R., Paciucci, R., Kondoh, H., ME, L.L., Oxidative stress and cancer: an overview. Ageing Res Rev 12 (2013), 376–390.
417. Bohm, F., Edge, R., Truscott, G., Interactions of dietary carotenoids with activated (singlet) oxygen and free radicals: Potential effects for human health. Mol Nutr Food Res 56 (2012), 205–216.
418. Palozza, P., Serini, S., Di Nicuolo, F., Piccioni, E., Calviello, G., Prooxidant effects of beta-carotene in cultured cells. Mol Aspects Med 24 (2003), 353–362.
419. Siems, W., Wiswedel, I., Salerno, C., Crifo, C., Augustin, W., Schild, L., et al. Beta-carotene breakdown products may impair mitochondrial functions—potential side effects of high-dose beta-carotene supplementation. J Nutr Biochem 16 (2005), 385–397.
420. Young, A.J., Lowe, G.M., Antioxidant and prooxidant properties of carotenoids. Arch Biochem Biophys 385 (2001), 20–27.
421. van Helden, Y.G., Keijer, J., Heil, S.G., Pico, C., Palou, A., Oliver, P., et al. Beta-carotene affects oxidative stress-related DNA damage in lung epithelial cells and in ferret lung. Carcinogenesis 30 (2009), 2070–2076.
422. Palozza, P., Serini, S., Di Nicuolo, F., Calviello, G., Modulation of apoptotic signalling by carotenoids in cancer cells. Arch Biochem Biophys 430 (2004), 104–109.
423. Schafer, Z.T., Grassian, A.R., Song, L., Jiang, Z., Gerhart-Hines, Z., Irie, H.Y., et al. Antioxidant and oncogene rescue of metabolic defects caused by loss of matrix attachment. Nature 461 (2009), 109–113.
424. Watson, J., Oxidants, antioxidants and the current incurability of metastatic cancers. Open Biol, 3, 2013, 120144.
425. Gregor, M.F., Hotamisligil, G.S., Inflammatory mechanisms in obesity. Annu Rev Immunol 29 (2011), 415–445.
426. Marseglia, L., Manti, S., D'Angelo, G., Nicotera, A., Parisi, E., Di Rosa, G., et al. Oxidative stress in obesity: A critical component in human diseases. Int J Mol Sci 16 (2014), 378–400.
427. Makki, K., Froguel, P., Wolowczuk, I., Adipose tissue in obesity-related inflammation and insulin resistance: Cells, cytokines, and chemokines. ISRN Inflamm, 2013, 2013, 139239.
428. Wisse, B.E., The inflammatory syndrome: the role of adipose tissue cytokines in metabolic disorders linked to obesity. J Am Soc Nephrol 15 (2004), 2792–2800.
429. Karkeni, E., Bonnet, L., Astier, J., Couturier, C., Dalifard, J., Tourniaire, F., et al. All-trans-retinoic acid represses chemokine expression in adipocytes and adipose tissue by inhibiting NF-kappaB signaling. J Nutr Biochem 42 (2017), 101–107.
430. Fenni, S., Hammou, H., Astier, J., Bonnet, L., Karkeni, E., Couturier, C., et al. Lycopene and tomato powder supplementation similarly inhibit high-fat diet induced obesity, inflammatory response, and associated metabolic disorders. Mol Nutr Food Res, 61, 2017.
431. Bhuvaneswari, V., Velmurugan, B., Balasenthil, S., Ramachandran, C.R., Nagini, S., Chemopreventive efficacy of lycopene on 7,12-dimethylbenz[a]anthracene-induced hamster buccal pouch carcinogenesis. Fitoterapia 72 (2001), 865–874.
432. Sharoni, Y., Linnewiel-Hermoni, K., Khanin, M., Salman, H., Veprik, A., Danilenko, M., et al. Carotenoids and apocarotenoids in cellular signaling related to cancer: A review. Mol Nutr Food Res 56 (2012), 259–269.
433. Zhang, X., Zhao, W.E., Hu, L., Zhao, L., Huang, J., Carotenoids inhibit proliferation and regulate expression of peroxisome proliferators-activated receptor gamma (PPARgamma) in K562 cancer cells. Arch Biochem Biophys 512 (2011), 96–106.
434. Lian, F., Wang, X.D., Enzymatic metabolites of lycopene induce Nrf2-mediated expression of phase II detoxifying/antioxidant enzymes in human bronchial epithelial cells. Int J Cancer 123 (2008), 1262–1268.
435. Xia, Y., Shen, S., Verma, I.M., NF-kappaB, an active player in human cancers. Cancer Immunol Res 2 (2014), 823–830.
436. Bastien, J., Rochette-Egly, C., Nuclear retinoid receptors and the transcription of retinoid-target genes. Gene 328 (2004), 1–16.
437. Aranda, A., Pascual, A., Nuclear hormone receptors and gene expression. Physiol Rev 81 (2001), 1269–1304.
438. Shaw, N., Elholm, M., Noy, N., Retinoic acid is a high affinity selective ligand for the peroxisome proliferator-activated receptor beta/delta. J Biol Chem 278 (2003), 41589–41592.
439. Rieck, M., Meissner, W., Ries, S., Muller-Brusselbach, S., Muller, R., Ligand-mediated regulation of peroxisome proliferator-activated receptor (PPAR) beta/delta: A comparative analysis of PPAR-selective agonists and all-trans retinoic acid. Mol Pharmacol 74 (2008), 1269–1277.
440. Schug, T.T., Berry, D.C., Shaw, N.S., Travis, S.N., Noy, N., Opposing effects of retinoic acid on cell growth result from alternate activation of two different nuclear receptors. Cell 129 (2007), 723–733.
441. Bushue, N., Wan, Y.J., Retinoid pathway and cancer therapeutics. Adv Drug Deliv Rev 62 (2010), 1285–1298.
442. Lian, F., Smith, D.E., Ernst, H., Russell, R.M., Wang, X.D., Apo-10'-lycopenoic acid inhibits lung cancer cell growth in vitro, and suppresses lung tumorigenesis in the A/J mouse model in vivo. Carcinogenesis 28 (2007), 1567–1574.
443. Gouranton, E., Aydemir, G., Reynaud, E., Marcotorchino, J., Malezet, C., Caris-Veyrat, C., et al. Apo-10'-lycopenoic acid impacts adipose tissue biology via the retinoic acid receptors. Biochim Biophys Acta 1811 (2011), 1105–1114.
444. Eroglu, A., Hruszkewycz, D.P., dela Sena, C., Narayanasamy, S., Riedl, K.M., Kopec, R.E., et al. Naturally occurring eccentric cleavage products of provitamin A beta-carotene function as antagonists of retinoic acid receptors. J Biol Chem 287 (2012), 15886–15895.
445. Ziouzenkova, O., Orasanu, G., Sukhova, G., Lau, E., Berger, J.P., Tang, G., et al. Asymmetric cleavage of beta-carotene yields a transcriptional repressor of retinoid X receptor and peroxisome proliferator-activated receptor responses. Mol Endocrinol 21 (2007), 77–88.
446. Shirakura, Y., Takayanagi, K., Mukai, K., Tanabe, H., Inoue, M., beta-cryptoxanthin suppresses the adipogenesis of 3T3-L1 cells via RAR activation. J Nutr Sci Vitaminol (Tokyo) 57 (2011), 426–431.
447. Inoue, M., Tanabe, H., Matsumoto, A., Takagi, M., Umegaki, K., Amagaya, S., et al. Astaxanthin functions differently as a selective peroxisome proliferator-activated receptor gamma modulator in adipocytes and macrophages. Biochem Pharmacol 84 (2012), 692–700.
448. Breitman, T.R., Takahashi, N., Retinoylation of proteins in mammalian cells. Biochem Soc Trans 24 (1996), 723–727.
449. Blomhoff, R., Blomhoff, H.K., Overview of retinoid metabolism and function. J Neurobiol 66 (2006), 606–630.
450. Al Tanoury, Z., Piskunov, A., Rochette-Egly, C., Vitamin A and retinoid signaling: Genomic and nongenomic effects. J Lipid Res 54 (2013), 1761–1775.
451. Yong, L.C., Forman, M.R., Beecher, G.R., Graubard, B.I., Campbell, W.S., Reichman, M.E., et al. Relationship between dietary intake and plasma concentrations of carotenoids in premenopausal women: Application of the USDA-NCI carotenoid food-composition database. Am J Clin Nutr 60 (1994), 223–230.
452. Tucker, K.L., Chen, H., Vogel, S., Wilson, P.W., Schaefer, E.J., Lammi-Keefe, C.J., Carotenoid intakes, assessed by dietary questionnaire, are associated with plasma carotenoid concentrations in an elderly population. J Nutr 129 (1999), 438–445.
453. Carroll, Y.L., Corridan, B.M., Morrissey, P.A., Carotenoids in young and elderly healthy humans: Dietary intakes, biochemical status and diet-plasma relationships. Eur J Clin Nutr 53 (1999), 644–653.
454. Curran-Celentano, J., Hammond, B.R. Jr., Ciulla, T.A., Cooper, D.A., Pratt, L.M., Danis, R.B., Relation between dietary intake, serum concentrations, and retinal concentrations of lutein and zeaxanthin in adults in a Midwest population. Am J Clin Nutr 74 (2001), 796–802.
455. Olmedilla, B., Granado, F., Southon, S., Wright, A.J., Blanco, I., Gil-Martinez, E., et al. Serum concentrations of carotenoids and vitamins A, E, and C in control subjects from five European countries. Br J Nutr 85 (2001), 227–238.
456. El-Sohemy, A., Baylin, A., Kabagambe, E., Ascherio, A., Spiegelman, D., Campos, H., Individual carotenoid concentrations in adipose tissue and plasma as biomarkers of dietary intake. Am J Clin Nutr 76 (2002), 172–179.
457. Tangney, C.C., Bienias, J.L., Evans, D.A., Morris, M.C., Reasonable estimates of serum vitamin E, vitamin C, and beta-cryptoxanthin are obtained with a food frequency questionnaire in older black and white adults. J Nutr 134 (2004), 927–934.
458. Burke, J.D., Curran-Celentano, J., Wenzel, A.J., Diet and serum carotenoid concentrations affect macular pigment optical density in adults 45 years and older. J Nutr 135 (2005), 1208–1214.
459. Faure, H., Preziosi, P., Roussel, A.M., Bertrais, S., Galan, P., Hercberg, S., et al. Factors influencing blood concentration of retinol, alpha-tocopherol, vitamin C, and beta-carotene in the French participants of the SU.VI.MAX trial. Eur J Clin Nutr 60 (2006), 706–717.
460. Talegawkar, S.A., Johnson, E.J., Carithers, T.C., Taylor, H.A., Bogle, M.L., Tucker, K.L., Carotenoid intakes, assessed by food-frequency questionnaires (FFQs), are associated with serum carotenoid concentrations in the Jackson Heart Study: Validation of the Jackson Heart Study Delta NIRI Adult FFQs. Public Health Nutr 11 (2008), 989–997.
461. Arab, L., Cambou, M.C., Craft, N., Wesseling-Perry, K., Jardack, P., Ang, A., Racial differences in correlations between reported dietary intakes of carotenoids and their concentration biomarkers. Am J Clin Nutr 93 (2011), 1102–1108.
462. George, S.M., Thompson, F.E., Midthune, D., Subar, A.F., Berrigan, D., Schatzkin, A., et al. Strength of the relationships between three self-reported dietary intake instruments and serum carotenoids: The Observing Energy and Protein Nutrition (OPEN) Study. Public Health Nutr 15 (2012), 1000–1007.
463. Wawrzyniak, A., Hamulka, J., Friberg, E., Wolk, A., Dietary, anthropometric, and lifestyle correlates of serum carotenoids in postmenopausal women. Eur J Nutr 52 (2013), 1919–1926.
464. Olmedilla-Alonso, B., Beltran-de-Miguel, B., Estevez-Santiago, R., Cuadrado-Vives, C., Markers of lutein and zeaxanthin status in two age groups of men and women: Dietary intake, serum concentrations, lipid profile and macular pigment optical density. Nutr J, 13, 2014, 52.
465. Duran-Cabral, M., Fernandez-Jalao, I., Estevez-Santiago, R., Olmedilla Alonso, B., Assessment of individual carotenoid and vitamin A dietary intake in overweight and obese Dominican subjects. Nutr Hosp 34 (2017), 407–415.
466. Pezdirc, K., Hutchesson, M.J., Whitehead, R., Ozakinci, G., Perrett, D., Collins, C.E., Fruit, vegetable and dietary carotenoid intakes explain variation in skin-color in young caucasian women: A cross-sectional study. Nutrients 7 (2015), 5800–5815.
467. Fraser, G.E., Jaceldo-Siegl, K., Henning, S.M., Fan, J., Knutsen, S.F., Haddad, E.H., et al. Biomarkers of dietary intake are correlated with corresponding measures from repeated dietary recalls and food-frequency questionnaires in the adventist health study-2. J Nutr 146 (2016), 586–594.
468. Estevez-Santiago, R., Beltran-de-Miguel, B., Olmedilla-Alonso, B., Assessment of dietary lutein, zeaxanthin and lycopene intakes and sources in the Spanish survey of dietary intake (2009–2010). Int J Food Sci Nutr 67 (2016), 305–313.
469. Beltran-de-Miguel, B., Estevez-Santiago, R., Olmedilla-Alonso, B., Assessment of dietary vitamin A intake (retinol, alpha-carotene, beta-carotene, beta-cryptoxanthin) and its sources in the National Survey of Dietary Intake in Spain (2009–2010). Int J Food Sci Nutr 66 (2015), 706–712.
470. Vargas-Murga, L., de Rosso, V.V., Mercadante, A.Z., AB, O., Fruits and vegetables in the Brazilian Household Budget Survey (2008–2009): carotenoid content and assessment of individual carotenoid intake. J Food Comp Anal 50 (2016), 88–96.
471. NHANES, What we eat in America. https://www.ars.usda.gov/ARSUserFiles/80400530/pdf/0910/tables_1-40_2009-2010.pdf, 2009–2010.