Ionomic and metabolomic analyses reveal the resistance response mechanism to saline-alkali stress in Malus halliana seedlings.

Saline-alkali stress is a major abiotic stress limiting plant growth. The selection of saline-alkali-tolerant rootstock is an effective strategy to reduce salinization-alkalization influence in apple production. M. halliana is a highly saline-alkali-resistant apple rootstock in northwestern China. However, few metabolic response studies have been conducted on this species. In plants under saline-alkali stress, the uptake of K, Mg and Zn in M. halliana leaves were inhibited, whereas the absorption of Fe, Cu or Mn were increased. Metabolic analysis revealed 140 differentially expressed metabolites, which were mainly involved in alkaloid biosynthesis, phenylalanine biosynthesis, ATP-binding cassette (ABC) transporters, and mineral absorption. Especially, the expression of sucrose, amino acids, alkaloids, flavonoids and carotenoids were significantly upregulated under saline-alkali stress. qRT-PCR analysis demonstrated that NHX8 and ZTP1 involved in Na and Fe transport were upregulated, while AKT1, MRS2-4 and ZTP29 involved in K, Mg and Zn transport were downregulated, respectively. ANT, ATP2A, CALM and SOS2 are involved in Ca signal transduction, and ABCB1, ABCC10 and NatA are key transporters that maintain ionic homeostasis. M. halliana regulates Na/K homeostasis by mediating Ca signalling and ABC transporters. The accumulation of metabolites contributes to improving the saline-alkali resistance of M. halliana because of the scavenging of ROS. An increase in pheophorbide a content in porphyrin and chlorophyll metabolism leads to leaf senescence in M. halliana leaves, which contributes to a reduction in stress-induced injury. These findings provide important insights into the saline-alkali tolerance mechanism in apple, which also provides an important starting point for future research.