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Biofortification of rice ( Oryza sativa  L.) with micronutrients is widely recognized as a sustainable strategy to alleviate human iron (Fe) and zinc (Zn) deficiencies in developing countries where rice is the staple food. Constitutive overexpression of the rice nicotianamine synthase ( OsNAS ) genes has been successfully implemented to increase Fe and Zn concentrations in unpolished and polished rice grain. Intensive research is now needed to couple this high-micronutrient trait with high grain yields. We investigated associations of increased grain Fe and Zn concentrations with agro-morphological traits of backcross twice second filial (BC 2 F 2 ) transgenic progeny carrying  OsNAS1  or  OsNAS2  overexpression constructs under  indica/japonica  and  japonica/japonica  genetic backgrounds. Thirteen agro-morphological traits were evaluated in BC 2 F 2  transgenic progeny grown under hydroponic conditions. Concentrations of eight mineral nutrients (Fe, Zn, copper, manganese, calcium, magnesium, potassium, and phosphorus) in roots, stems/sheaths, non-flag leaves, flag leaves, panicles, and grain were also determined. A distance-based linear model (DistLM) was utilized to extract plant tissue nutrient predictors accounting for the largest variation in agro-morphological traits differing between transgenic and non-transgenic progeny. Overall, the BC 2 F 2  transgenic progeny contained up to 148% higher Fe and 336% higher Zn concentrations in unpolished grain compared to non-transgenic progeny. However, unpolished grain concentrations surpassing 23 μg Fe g -1  and 40 μg Zn g -1  in BC 2 F 2   indica/japonica  progeny, and 36 μg Fe g -1  and 56 μg Zn g 1  in BC 2 F 2   japonica/japonica  progeny, were associated with significant reductions in grain yield. DistLM analyses identified grain-Zn and panicle-magnesium as the primary nutrient predictors associated with grain yield reductions in the  indica/japonica  and  japonica/japonica  background, respectively. We subsequently produced polished grain from high-yield BC 2 F 2  transgenic progeny carrying either the  OsNAS1  or  OsNAS2  overexpression constructs. The  OsNAS2  overexpressing progeny had higher percentages of Fe and Zn in polished rice grain compared to the  OsNAS1  overexpressing progeny. Results from this study demonstrate that genetic background has a major effect on the development of Fe and Zn biofortified rice. Moreover, our study shows that high-yielding rice lines with Fe and Zn biofortified polished grain can be developed by  OsNAS2  overexpression and monitoring for Zn overaccumulation in the grain.

Association of Increased Grain Iron and Zinc Concentrations with Agro-morphological Traits of Biofortified Rice