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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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