4. 初级代谢调控Regulation of carotenoid biosynthesis.pdfVIP

4. 初级代谢调控Regulation of carotenoid biosynthesis.pdf

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4. 初级代谢调控Regulation of carotenoid biosynthesis

Regulation of carotenoid biosynthesis in plants: evidence for a key role of hydroxymethylbutenyl diphosphate reductase in controlling the supply of plastidial isoprenoid precursors Patricia Botella-Pav??a, O?scar Besumbes, Michael A. Phillips, Lorenzo Carretero-Paulet, Albert Boronat and Manuel Rodr??guez-Concepcio?n* Departament de Bioqu??mica i Biologia Molecular, Facultat de Qu??mica, Universitat de Barcelona, Mart?? i Franque?s 1-7, 08028 Barcelona, Spain Received 14 May 2004; revised 9 July 2004; accepted 20 July 2004. *For correspondence (fax t34 93 402 1219; e-mail mrodrigu@sun.bq.ub.es). Summary Carotenoids are isoprenoid pigments that function as photoprotectors, precursors of the hormone abscisic acid (ABA), colorants and nutraceuticals. A major problem for the metabolic engineering of high carotenoid levels in plants is the limited supply of their isoprenoid precursor geranylgeranyl diphosphate (GGPP), formed by condensation of isopentenyl diphosphate (IPP) and dimethylallyl diphosphate (DMAPP) units usually synthesized by the methylerythritol phosphate (MEP) pathway in plastids. Our earlier work with three of the seven MEP pathway enzymes suggested that the first reaction of the pathway catalyzed by deoxyxylulose 5-phosphate synthase (DXS) is limiting for carotenoid biosynthesis during tomato (Lycopersicon esculentum) fruit ripening. Here we investigate the contribution of the enzyme hydroxymethylbutenyl diphoshate reductase (HDR), which simultaneously synthesizes IPP and DMAPP in the last step of the pathway. A strong upregulation of HDR gene expression was observed in correlation with carotenoid production during both tomato fruit ripening and Arabidopsis thaliana seedling deetiolation. Constitutive overexpression of the tomato cDNA encoding HDR in Arabidopsis did not increase carotenoid levels in etioplasts. By contrast, light- grown transgenic plants showed higher carotenoid levels and an enhanced seed dormancy phenotype suggestive of increased ABA l

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