Regulation of Tocopherol Biosynthesis During Fruit Maturation of Different Citrus Species

Tocopherols are plant-derived isoprenoids with vitamin E activity, which are involved in diverse physiological processes in plants. Although their biosynthesis has been extensively investigated in model plants, their synthesis in important fruit crops as Citrus has scarcely been studied. Therefore, the aim of this work was to initiate a physiological and molecular characterization of tocopherol synthesis and accumulation in Citrus fruits during maturation. For that purpose, we selected fruit of the four main commercial species: grapefruit (Citrus paradisi), lemon (Citrus limon), sweet orange (Citrus sinensis), and mandarin (Citrus clementina), and analyzed tocopherol content and the expression profile of 14 genes involved in tocopherol synthesis during fruit maturation in both the flavedo and pulp. The selected genes covered the pathways supplying the tocopherol precursors homogentisate (HGA) (TAT1 and HPPD) and phytyl pyrophosphate (PPP) (VTE5, VTE6, DXS1 and 2, GGPPS1 and 6, and GGDR) and the tocopherol-core pathway (VTE2, VTE3a, VTE3b, VTE1, and VTE4). Tocopherols accumulated mainly as α- and γ-tocopherol, and α-tocopherol was the predominant form in both tissues. Moreover, differences were detected between tissues, among maturation stages and genotypes. Contents were higher in the flavedo than in the pulp during maturation, and while they increased in the flavedo they decreased or were maintained in the pulp. Among genotypes, mature fruit of lemon accumulated the highest tocopherol content in both the flavedo and the pulp, whereas mandarin fruit accumulated the lowest concentrations, and grapefruit and orange had intermediate levels. Higher concentrations in the flavedo were associated with a higher expression of all the genes evaluated, and different genes are suitable candidates to explain the temporal changes in each tissue: (1) in the flavedo, the increase in tocopherols was concomitant with the up-regulation of TAT1 and VTE4, involved in the supply of HGA and the shift of γ- into α-tocopherol, respectively; and (2) in the pulp, changes paralleled the expression of VTE6, DXS2, and GGDR, which regulate PPP availability. Also, certain genes (i.e., VTE6, DXS2, and GGDR) were co-regulated and shared a similar pattern during maturation in both tissues, suggesting they are developmentally modulated.

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Main Authors: Rey, Florencia, Zacarías, Lorenzo, Rodrigo Esteve, María Jesús
Other Authors: CSIC - Unidad de Recursos de Información Científica para la Investigación (URICI)
Format: artículo biblioteca
Language:English
Published: Frontiers Media 2021-10-06
Subjects:Tocopherol, Vitamin E, Citrus, Fruit, Ripening, Gene expression,
Online Access:http://hdl.handle.net/10261/253596
http://dx.doi.org/10.13039/501100004837
http://dx.doi.org/10.13039/501100003359
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spelling dig-iata-es-10261-2535962021-11-05T02:20:58Z Regulation of Tocopherol Biosynthesis During Fruit Maturation of Different Citrus Species Rey, Florencia Zacarías, Lorenzo Rodrigo Esteve, María Jesús CSIC - Unidad de Recursos de Información Científica para la Investigación (URICI) Generalitat Valenciana Ministerio de Ciencia e Innovación (España) Tocopherol Vitamin E Citrus Fruit Ripening Gene expression Tocopherols are plant-derived isoprenoids with vitamin E activity, which are involved in diverse physiological processes in plants. Although their biosynthesis has been extensively investigated in model plants, their synthesis in important fruit crops as Citrus has scarcely been studied. Therefore, the aim of this work was to initiate a physiological and molecular characterization of tocopherol synthesis and accumulation in Citrus fruits during maturation. For that purpose, we selected fruit of the four main commercial species: grapefruit (Citrus paradisi), lemon (Citrus limon), sweet orange (Citrus sinensis), and mandarin (Citrus clementina), and analyzed tocopherol content and the expression profile of 14 genes involved in tocopherol synthesis during fruit maturation in both the flavedo and pulp. The selected genes covered the pathways supplying the tocopherol precursors homogentisate (HGA) (TAT1 and HPPD) and phytyl pyrophosphate (PPP) (VTE5, VTE6, DXS1 and 2, GGPPS1 and 6, and GGDR) and the tocopherol-core pathway (VTE2, VTE3a, VTE3b, VTE1, and VTE4). Tocopherols accumulated mainly as α- and γ-tocopherol, and α-tocopherol was the predominant form in both tissues. Moreover, differences were detected between tissues, among maturation stages and genotypes. Contents were higher in the flavedo than in the pulp during maturation, and while they increased in the flavedo they decreased or were maintained in the pulp. Among genotypes, mature fruit of lemon accumulated the highest tocopherol content in both the flavedo and the pulp, whereas mandarin fruit accumulated the lowest concentrations, and grapefruit and orange had intermediate levels. Higher concentrations in the flavedo were associated with a higher expression of all the genes evaluated, and different genes are suitable candidates to explain the temporal changes in each tissue: (1) in the flavedo, the increase in tocopherols was concomitant with the up-regulation of TAT1 and VTE4, involved in the supply of HGA and the shift of γ- into α-tocopherol, respectively; and (2) in the pulp, changes paralleled the expression of VTE6, DXS2, and GGDR, which regulate PPP availability. Also, certain genes (i.e., VTE6, DXS2, and GGDR) were co-regulated and shared a similar pattern during maturation in both tissues, suggesting they are developmentally modulated. This work was funded by the research grant RTI2018-095131-B-I00 of the Ministry of Science and Innovation/Agencia Española de Investigación/FEDER (Spanish Government) and PROMETEO/2020/027 (Generalitat Valenciana). FR was the recipient of a Ph.D. scholarship (POS_EXT_2016_1_133720) from ANII (Uruguay). The authors acknowledge support of the publication fee by the CSIC Open Access Publication Support Initiative through its Unit of Information Resources for Research (URICI). Peer reviewed 2021-11-04T06:41:12Z 2021-11-04T06:41:12Z 2021-10-06 artículo http://purl.org/coar/resource_type/c_6501 Frontiers in Plant Science 12: 743993 (2021) http://hdl.handle.net/10261/253596 10.3389/fpls.2021.743993 1664-462X http://dx.doi.org/10.13039/501100004837 http://dx.doi.org/10.13039/501100003359 en #PLACEHOLDER_PARENT_METADATA_VALUE# info:eu-repo/grantAgreement/AEI/Plan Estatal de Investigación Científica y Técnica y de Innovación 2017-2020/RTI2018-095131-B-I00 Publisher's version https://doi.org/10.3389/fpls.2021.743993 Sí open Frontiers Media
institution IATA ES
collection DSpace
country España
countrycode ES
component Bibliográfico
access En linea
databasecode dig-iata-es
tag biblioteca
region Europa del Sur
libraryname Biblioteca del IATA España
language English
topic Tocopherol
Vitamin E
Citrus
Fruit
Ripening
Gene expression
Tocopherol
Vitamin E
Citrus
Fruit
Ripening
Gene expression
spellingShingle Tocopherol
Vitamin E
Citrus
Fruit
Ripening
Gene expression
Tocopherol
Vitamin E
Citrus
Fruit
Ripening
Gene expression
Rey, Florencia
Zacarías, Lorenzo
Rodrigo Esteve, María Jesús
Regulation of Tocopherol Biosynthesis During Fruit Maturation of Different Citrus Species
description Tocopherols are plant-derived isoprenoids with vitamin E activity, which are involved in diverse physiological processes in plants. Although their biosynthesis has been extensively investigated in model plants, their synthesis in important fruit crops as Citrus has scarcely been studied. Therefore, the aim of this work was to initiate a physiological and molecular characterization of tocopherol synthesis and accumulation in Citrus fruits during maturation. For that purpose, we selected fruit of the four main commercial species: grapefruit (Citrus paradisi), lemon (Citrus limon), sweet orange (Citrus sinensis), and mandarin (Citrus clementina), and analyzed tocopherol content and the expression profile of 14 genes involved in tocopherol synthesis during fruit maturation in both the flavedo and pulp. The selected genes covered the pathways supplying the tocopherol precursors homogentisate (HGA) (TAT1 and HPPD) and phytyl pyrophosphate (PPP) (VTE5, VTE6, DXS1 and 2, GGPPS1 and 6, and GGDR) and the tocopherol-core pathway (VTE2, VTE3a, VTE3b, VTE1, and VTE4). Tocopherols accumulated mainly as α- and γ-tocopherol, and α-tocopherol was the predominant form in both tissues. Moreover, differences were detected between tissues, among maturation stages and genotypes. Contents were higher in the flavedo than in the pulp during maturation, and while they increased in the flavedo they decreased or were maintained in the pulp. Among genotypes, mature fruit of lemon accumulated the highest tocopherol content in both the flavedo and the pulp, whereas mandarin fruit accumulated the lowest concentrations, and grapefruit and orange had intermediate levels. Higher concentrations in the flavedo were associated with a higher expression of all the genes evaluated, and different genes are suitable candidates to explain the temporal changes in each tissue: (1) in the flavedo, the increase in tocopherols was concomitant with the up-regulation of TAT1 and VTE4, involved in the supply of HGA and the shift of γ- into α-tocopherol, respectively; and (2) in the pulp, changes paralleled the expression of VTE6, DXS2, and GGDR, which regulate PPP availability. Also, certain genes (i.e., VTE6, DXS2, and GGDR) were co-regulated and shared a similar pattern during maturation in both tissues, suggesting they are developmentally modulated.
author2 CSIC - Unidad de Recursos de Información Científica para la Investigación (URICI)
author_facet CSIC - Unidad de Recursos de Información Científica para la Investigación (URICI)
Rey, Florencia
Zacarías, Lorenzo
Rodrigo Esteve, María Jesús
format artículo
topic_facet Tocopherol
Vitamin E
Citrus
Fruit
Ripening
Gene expression
author Rey, Florencia
Zacarías, Lorenzo
Rodrigo Esteve, María Jesús
author_sort Rey, Florencia
title Regulation of Tocopherol Biosynthesis During Fruit Maturation of Different Citrus Species
title_short Regulation of Tocopherol Biosynthesis During Fruit Maturation of Different Citrus Species
title_full Regulation of Tocopherol Biosynthesis During Fruit Maturation of Different Citrus Species
title_fullStr Regulation of Tocopherol Biosynthesis During Fruit Maturation of Different Citrus Species
title_full_unstemmed Regulation of Tocopherol Biosynthesis During Fruit Maturation of Different Citrus Species
title_sort regulation of tocopherol biosynthesis during fruit maturation of different citrus species
publisher Frontiers Media
publishDate 2021-10-06
url http://hdl.handle.net/10261/253596
http://dx.doi.org/10.13039/501100004837
http://dx.doi.org/10.13039/501100003359
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