In humans, iron (Fe) and zinc (Zn) deficiencies result in major worldwide health problems. Transgenic technologies to produce Fe-and Zn-biofortified rice varieties offer a promising potential solution. Nicotianamine, the precursor of phytosiderophores, chelates Fe 2+ and Zn 2+ and plays an important role in transporting these metals to both vegetative and reproductive organs within the plant. Our objective was to increase Fe and Zn contents in rice grains by overexpressing the barley nicotianamine synthase gene HvNAS1. HvNAS1-overexpressing transgenic rice showed increased HvNAS1 expression and subsequent increases in endogenous nicotianamine and phytosiderophore content in shoots, roots, and seeds. Fe and Zn concentrations in polished T 1 seeds from transgenic plants increased more than three and twofold, respectively; Fe and Zn concentrations also increased in both polished and brown T 2 seeds. These results suggest that the overproduction of nicotianamine enhances the translocation of Fe and Zn into rice grains.

Iron uptake and translocation in plants are important processes for both plant and human nutrition, whereas relatively little is known about the molecular mechanisms of iron transport within the plant body. Several reports have shown that yellow stripe 1 (YS1) and YS1-like (YSL) transporters mediate metal-phytosiderophore uptake and/or metal-nicotianamine translocation. Among the 18 YSL genes in rice (OsYSLs), OsYSL18 is predicted to encode a polypeptide of 679 amino acids containing 13 putative transmembrane domains. An OsYSL18-green fluorescent protein (GFP) fusion was localized to the plasma membrane when transiently expressed in onion epidermal cells.Electrophysiological measurements using Xenopus laevis oocytes showed that OsYSL18 transports iron(III)-deoxymugineic acid, but not iron(II)-nicotianamine, zinc(II)-deoxymugineic acid, or zinc(II)-nicotianamine. Reverse transcriptase PCR analysis revealed more OsYSL18 transcripts in flowers than in shoots or roots. OsYSL18 promoterb-glucuronidase (GUS) analysis revealed that OsYSL18 was expressed in reproductive organs including the pollen tube. In vegetative organs, OsYSL18 was specifically expressed in lamina joints, the inner cortex of crown roots, and phloem parenchyma and companion cells at the basal part of every leaf sheath. These results suggest that OsYSL18 is an ironphytosiderophore transporter involved in the translocation of iron in reproductive organs and phloem in joints.

SummaryNicotianamine (NA), a metal chelator ubiquitous in higher plants, serves as an antihypertensive substance in humans. To engineer a novel antihypertensive rice that contains larger amounts of NA, the barley NA synthase gene, HvNAS1 , was introduced into rice via Agrobacterium -mediated transformation. The introduced HvNAS1 was driven by pGluB-1 , which induces strong gene expression in the endosperm of rice seeds. The NA content in transgenic rice seeds was up to fourfold greater than that in non-transgenic rice

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