The road toward Cd-safe rice: From mass selection to marker-assisted selection and genetic manipulation

Rice is an important dietary source of the toxic mineral cadmium (Cd) for populations in which rice is the main staple food. When grown in agricultural soils that are contaminated with Cd, rice often accumulates excessive Cd into the grains, which is a serious threat to agricultural sustainability and human health. To limit Cd accumulation in rice grains, studies on the genetic basis of Cd accumulation in rice have been carried out extensively, and some low-Cd rice varieties have also been developed in recent years. However, the challenges in low-Cd rice breeding still exist because the outcomes of the current genetic improvements for low-Cd rice cannot fully meet the requirements for the development of Cd-safe rice at present. In this review, we outline the progress in understanding the physiological mechanisms and the genetic nature of Cd accumulation in rice and summarize the strategies and outcomes of low-Cd rice breeding over the past decade. By graphing the physiological mechanism of Cd transport in the rice plant, three key steps and some underlying genes are summarized and discussed. Also, two genetic features of the natural variation in rice grain-Cd accumulation, the phenotypic plasticity and subspecies divergence, and the potential genetic explanations for these features are also discussed. Finally, we summarize and discuss current progress and the potential issues in low-Cd rice breeding using different breeding strategies. We hope to propose strategies for future success in the breeding of low-Cd rice varieties over the next decade.