Aluminum (Al) toxicity is a major yield-limiting factor for crops in acidic soils. In this work, we have investigated the potential role of spermidine (Spd) on Al toxicity in rice chloroplasts. Exogenous Spd markedly reduced Al concentration and elevated other nutrient elements such as Mn, Mg, Fe, K, Ca, and Mo in chloroplasts of Al-treated plants. Meanwhile, Spd further activated arginine decarboxylase (ADC) activity of key enzyme in polyamine (PA) synthesis, and enhanced PA contents in chloroplasts. Spd application dramatically addressed Al-induced chlorophyll (Chl) losses, inhibited thylakoid membrane protein complexes degradation, especially photosystem II (PSII), and significantly depressed the accumulations of superoxide radical (O₂·^-), hydrogen peroxide (H₂O₂), and malondialdehyde (MDA) in chloroplasts. Spd addition activated antioxidant enzyme activities and decreased soluble sugar content in chloroplasts compared with Al treatment alone. Spd not only reversed the inhibition of photosynthesis-related gene transcript levels induced by Al toxicity, but diminished the increased expression of Chl catabolism-related genes. Furthermore, Chl fluorescence analysis showed that Spd protected PSII reaction centers and photosynthetic electron transport chain under Al stress, thus improving photosynthetic performance. These results suggest that PAs are involved in Al tolerance in rice chloroplasts and can effectively protect the integrity and function of photosynthetic apparatus, especially PSII, by mitigating oxidative damage induced by Al toxicity.

Rare earth elements (REEs) are emerging as an anticipated pollution in the environment due to their active use in many areas. However, the effects of REEs on the photosynthesis of rice have not been thoroughly explored. Therefore, this study emphasizes how high levels of La(III) affect the thylakoid membrane of rice seedlings, thereby inhibiting photosynthesis and growth. Here, we reported that rice plants treated with La(III) exhibited an increase in La accumulation in the leaves, accompanied by a decrease in chlorophyll content and photosynthetic capacity. La(III) exposure decreased Mg content in leaves, but possibly increased other nutrients including Cu, Mn, and Zn through systemic endocytosis. K-band and L-band appeared in the fluorescence OJIP transients, indicating La(III) stress destroyed the donor and receptor sides of photosystem II (PSII). Numerous reaction centers (RC/CS_m) were inactivated by La(III) treatment, which resulted in a reduction in electron transport capacity (decreased ET_o/RC and ET_o/CS_m) and an increase in the dissipation of the excess excitation energy by heat (increased DI_o/RC and DI_o/CS_m). The BN-PAGE analysis of thylakoid membrane protein complexes showed that La(III) induced the degradation of supercomplexes, PSII core, LHCII, PSI core, LHCI, and F1-ATPase binding Cyt b₆f complex. Collectively, this study revealed that La(III) causes significant degradation of thylakoid membrane proteins, thereby promoting the decomposition of photosynthetic complexes, ultimately destroying the chloroplast structure and reducing the photosynthetic performance of rice seedlings.