Rice is one of the most important staple crops,feeding more than half of the population on Earth.How to steadily increase yield has been a major challenge in the context of an ever-growing popula-tion and climate change.Over the past decades,breeding prac-tices have greatly increased cereal grain yield mainly owing to the adoption of semidwarf alleles and the development of hybrid crop varieties.However,this achievement relies heavily on fertilizer application,and over-fertilization results in massive energy con-sumption,widespread water eutrophication,significant acidifica-tion of croplands,and a heavy economic burden on farmers(Zhang,2007).