Boron contributes to excessive aluminum tolerance in trifoliate orange (Poncirus trifoliata (L.) Raf.) by inhibiting cell wall deposition and promoting vacuole compartmentation

Boron (B) is an indispensable micronutrient for plant growth that can also alleviate aluminum (Al) toxicity. However, limited data are available on the underlying mechanisms behind this phenomenon. Here, we found that a certain range of B application could alleviate the inhibitory effects of Al toxicity on citrus. Transcriptome analysis revealed that several Al stress-responsive genes and pathways were differentially affected and enriched, such as coding for the secretion of organic acid and the distribution of Al in subcellular components after B addition. Specifically, B application enhanced rhizosphere pH and induced malate exudation by expressing PtALMT4 and PtALMT9 genes occurred in Al-treated root, which ultimately reduced the absorption of Al and coincided with down-regulated the expression of PtNrat1. Moreover, B supply suppressed the pectin methyl-esterase (PME) activity and displayed a lower level of PtPME2 expression, while enhanced the PtSTAR1 expression, which is responsible for reducing cell wall (CW) Al deposition. Boron addition enhanced the PtALS1 and PtALS3 expression, accompanied by a higher proportion of vacuolar Al compartmentation during Al exposure. Collectively, the protective effects of B on root injury induced by Al is mainly by subsiding the Al uptake in the root apoplast and compartmentalizing Al into vacuole.