Nitrogen limits zinc-mediated stimulation of tillering in rice by modifying phytohormone balance under low-temperature stress

Zinc (Zn) can improve resistance of rice plants to abiotic stress and promote the recovery of rice tillers by influencing auxin (IAA) synthesis, Zn absorption is closely related to nitrogen (N) concentrations; however, the mechanisms by which Zn regulates rice tiller recovery at low temperatures under different N levels are rarely studied. In this study, a solution culture experiment was conducted at two temperatures, three different Zn concentrations, and two N concentrations. Low temperature during the vegetative stage damaged the rice antioxidant system, suppressed nutrient uptake, and decreased the number of tillers. Increasing Zn application improved the resistance of rice plants to low-temperature stress under normal N levels, enhanced nutrient absorption, improved tiller bud cytokinin (CTK) concentrations, and the CTK/IAA ratio, which accelerated tiller recovery relative to plants receiving standard Zn treatment. Shoot Zn concentrations were positively correlated with tiller numbers at normal N levels. High N attenuated the positive effect of Zn on rice tillering at low temperatures due to higher shoot Zn and IAA accumulation, and lower tiller buds CTK/IAA ratios. However, appropriate Zn application under high N levels was conducive to breaking tiller dormancy and promoting tillering growth spurts when recovering to normal temperature, which was related to the synergistic regulation of hormone balance and nutrient absorption by N and Zn. In conclusion, Zn could enhance rice low-temperature stress resilience and facilitate tiller recovery via balancing the hormones in tiller buds. However, higher environmental N limited the stimulation of Zn on rice tillering under low temperature.