Changes in carotenoid content and biosynthetic gene expression in juice sacs of four orange varieties (Citrus sinensis) differing in flesh fruit color.
The contribution of carotenoid composition to the color range of the fruit juice sacs of four orange varieties (Citrus sinensis) differing in flesh color, namely, Shamouti (normal orange color), Sanguinelli ("blood cultivar" purple color), Cara Cara navel (pink-reddish), and Huang pi Chen (yellowish color), was investigated. To this end, qualitative and quantitative analyses of carotenoid contents were first performed by high-performance liquid chromatography (HPLC) using a C30 column and a photodiode array detector in February, at a late developmental fruit stage. Concomitantly, transcript levels of Dxs, the gene controlling the first step of the MEP pathway, and six genes involved in beta,beta-xanthophyll biosynthesis (Psy, Pds, Zds, Lcy-b, Hy-b, and Zep) were determined in August, November, and February. Transcript level measurement was carried out by real-time RT-PCR on total RNA from juice sacs. The four orange varieties displayed different carotenoid profiles. Shamouti and Sanguinelli oranges accumulated mainly beta,beta-xanthophylls as expected in typically colored oranges, whereas Cara Cara navel orange accumulated linear carotenes in addition to cis-violaxanthin. Huang pi Chen fruit flesh orange was characterized by a strong reduction of total carotenoid content. Whereas gene expression was relatively low and similar in August (before color break) in all four varieties, in November (during color break), Dxs, Zds, Hy-b, and Zep expression was higher in Cara Cara and Huang pi Chen oranges. The beta,beta-xanthophyll accumulation observed in February in Shamouti and Sanguinelli oranges was apparently related to the increase of transcript levels of all measured genes (i.e., Dxs, Psy, Pds, Zds, Hy-b, and Zep) except Lcy-b. At this time, however, transcript levels in Cara Cara were rather similar to those found in Sanguinelli, although both showed different carotenoid compositions. The Huang pi Chen phenotype correlated with lower expression of Dxs and Psy genes. These results revealed a general pattern of transcript change in juice sacs of citrus fruit, characterized by an apparent coordination of Dxs and Psy expression and a general increase in mRNA levels of carotenoid biosynthetic genes. These transcript changes correlated well with the beta,beta-xanthophyll accumulation, the normal carotenoid set, observed in Shamouti and Sanguinelli oranges and suggest that the preferential accumulation of linear carotenes in Cara Cara navel and the practical absence of carotenoids in Huang pi Chen oranges were not predominantly due to changes in regulation of carotenoid biosynthetic genes at the transcriptional level.
Fanciullino AL
，Cerćos M
，Dhique-Mayer
，Froelicher Y
，Talón M
，Ollitrault P
，Morillon R
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《JOURNAL OF AGRICULTURAL AND FOOD CHEMISTRY》
Regulation of carotenoid biosynthesis during fruit maturation in the red-fleshed orange mutant Cara Cara.
Cara Cara is a spontaneous bud mutation of Navel orange (Citrus. sinensis L. Osbeck) characterized by developing fruits with a pulp of bright red coloration due to the presence of lycopene. Peel of mutant fruits is however orange and indistinguishable from its parental. To elucidate the basis of lycopene accumulation in Cara Cara, we analyzed carotenoid profile and expression of three isoprenoid and nine carotenoid genes in flavedo and pulp of Cara Cara and Navel fruits throughout development and maturation. The pulp of the mutant accumulated high amounts of lycopene, but also phytoene and phytofluene, from early developmental stages. The peel of Cara Cara also accumulated phytoene and phytofluene. The expression of isoprenoid genes and of carotenoid biosynthetic genes downstream PDS (phytoene desaturase) was higher in the pulp of Cara Cara than in Navel. Not important differences in the expression of these genes were observed between the peel of both oranges. Moreover, the content of the plant hormone ABA (abscisic acid) was lower in the pulp of Cara Cara, but the expression of two genes involved in its biosynthesis was higher. The results suggest that an altered carotenoid composition may conduct to a positive feedback regulatory mechanism of carotenoid biosynthesis in citrus fruits. Increased levels of isoprenoid precursors in the mutant that could be channeled to carotenoid biosynthesis may be related to the red-fleshed phenotype of Cara Cara.
Alquezar B
，Rodrigo MJ
，Zacarías L
《PHYTOCHEMISTRY》
Accumulation of carotenoids and expression of carotenoid biosynthetic genes during maturation in citrus fruit.
The relationship between carotenoid accumulation and the expression of carotenoid biosynthetic genes during fruit maturation was investigated in three citrus varieties, Satsuma mandarin (Citrus unshiu Marc.), Valencia orange (Citrus sinensis Osbeck), and Lisbon lemon (Citrus limon Burm.f.). We cloned the cDNAs for phytoene synthase (CitPSY), phytoene desaturase (CitPDS), zeta-carotene (car) desaturase (CitZDS), carotenoid isomerase (CitCRTISO), lycopene beta-cyclase (CitLCYb), beta-ring hydroxylase (CitHYb), zeaxanthin (zea) epoxidase (CitZEP), and lycopene epsilon-cyclase (CitLCYe) from Satsuma mandarin, which shared high identities in nucleotide sequences with Valencia orange, Lisbon lemon, and other plant species. With the transition of peel color from green to orange, the change from beta,epsilon-carotenoid (alpha-car and lutein) accumulation to beta,beta-carotenoid (beta-car, beta-cryptoxanthin, zea, and violaxanthin) accumulation was observed in the flavedos of Satsuma mandarin and Valencia orange, accompanying the disappearance of CitLCYe transcripts and the increase in CitLCYb transcripts. Even in green fruit, high levels of beta,epsilon-carotenoids and CitLCYe transcripts were not observed in the juice sacs. As fruit maturation progressed in Satsuma mandarin and Valencia orange, a simultaneous increase in the expression of genes (CitPSY, CitPDS, CitZDS, CitLCYb, CitHYb, and CitZEP) led to massive beta,beta-xanthophyll (beta-cryptoxanthin, zea, and violaxanthin) accumulation in both the flavedo and juice sacs. The gene expression of CitCRTISO was kept low or decreased in the flavedo during massive beta,beta-xanthophyll accumulation. In the flavedo of Lisbon lemon and Satsuma mandarin, massive accumulation of phytoene was observed with a decrease in the transcript level for CitPDS. Thus, the carotenoid accumulation during citrus fruit maturation was highly regulated by the coordination of the expression among carotenoid biosynthetic genes. In this paper, the mechanism leading to diversity in beta,beta-xanthophyll compositions between Satsuma mandarin and Valencia orange was also discussed on the basis of the substrate specificity of beta-ring hydroxylase and the balance of expression between upstream synthesis genes (CitPSY, CitPDS, CitZDS, and CitLCYb) and downstream synthesis genes (CitHYb and CitZEP).
Kato M
，Ikoma Y
，Matsumoto H
，Sugiura M
，Hyodo H
，Yano M
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