Arid and semiarid apple producing areas suffer from severe alkalinity of soil, which strongly affects the yield and quality of apples. Dopamine (DA) is involved in metabolic activities in response to abiotic stress in plants. To detect the effects of exogenous DA application on the adaption of apple (Malus hupehensis) seedlings to alkali stress and as a protection from oxidative stress, 0.1 mM DA was identified as the most suitable concentration by hydroponic culture. Further experimentation showed that the growth and photosynthesis of apple seedlings were significantly inhibited under alkali stress, and more reactive oxygen species accumulated, compared with control. However, exogenous DA application suppressed the loss of the plant height, root length, chlorophyll levels, and photosynthetic capacity of apple seedlings that were caused by alkali stress. In the leaves of alkali stressed seedlings, the catalase, superoxide dismutase, and peroxidase activities were lower and hydrogen peroxide and malondialdehyde levels were higher than in the untreated plants. The presence of DA significantly alleviated such effects of alkali stress. In addition, exogenous DA application increased the antioxidant capacity of apple seedlings under alkali stress by increasing the level of chlorogenic acid. These results are significant for improving the alkali tolerance of apple in apple-producing areas with alkalized soil.

Drought seriously affects the yield and quality of apples. γ-aminobutyric acid (GABA) plays an important role in the responses of plants to various stresses. However, the role and possible mechanism of GABA in the drought response of apple seedlings remain unknown. To explore the effect of GABA on apple seedlings under drought stress, seedlings of Malus hupehensis were treated with seven concentrations of GABA, and the response of seedlings under 15-day drought stress was observed. The results showed that 0.5 mM GABA was the most effective at relieving drought stress. Treatment with GABA reduced the relative electrical conductivity and MDA content of leaves induced by drought stress and significantly increased the relative water content of leaves. Exogenous GABA significantly decreased the stomatal conductance and intercellular carbon dioxide concentration and transpiration rate, and it significantly increased the photosynthetic rate under drought. GABA also reduced the accumulation of superoxide anions and hydrogen peroxide in leaf tissues under drought and increased the activities of POD, SOD, and CAT and the content of GABA. Exogenous treatment with GABA acted through the accumulation of abscisic acid (ABA) in the leaves to significantly decrease stomatal conductance and increase the stomatal closure rate, and the levels of expression of ABA-related genes PYL4, ABI1, ABI2, HAB1, ABF3, and OST1 changed in response to drought. Taken together, exogenous GABA can enhance the drought tolerance of apple seedlings.

Drought stress is an environmental factor that seriously threatens plant growth, development and yield. VQ proteins are transcriptional regulators that have been reported to be involved in plant growth, development and the responses to biotic and abiotic stressors. However, the relationship between VQ proteins and drought stress has not been well documented in plants. In this study, overexpressing the apple VQ motif-containing protein (MdVQ37) gene in apple plants markedly reduced the tolerance to drought. Physiological and biochemical studies further demonstrated lower enzymatic activities and decreased photosynthetic capacity in transgenic lines compared with wild-type (WT) plants under drought stress. Ultrastructural analysis of leaves showed that the leaves and palisade tissues from the transgenic lines were significantly thinner than those from WT plants. Salicylic acid (SA) analysis indicated that overexpression of MdVQ37 increased the accumulation of 2,5-DHBA by up-regulating the expression of the SA catabolic gene, which ultimately resulted to a significant reduction in endogenous SA content and the disruption of the SA-dependent signaling pathway under drought stress. Applying SA partially increased the survival rate of the transgenic lines under drought stress. These results demonstrate that the regulatory function of apple MdVQ37 is implicated in drought stress, through a change in leaf development and SA homeostasis. This study provides novel insight into understanding the multiple functions of VQ proteins.

Soil alkalization affects apple production in northwest China. Autophagy is a highly conserved degradative protein pathway in eukaryotes. Autophagy in plants can be activated by various abiotic factors. We previously identified the positive role of the autophagy-related gene MdATG18a in drought, nitrogen deficiency and resistance to Diplocarpon mali infection in apple. However, it is still unclear whether ATG18a is related to alkaline stress. In this study, we used hydroponic culture to simulate alkaline stress and found that the overexpression of MdATG18a significantly improved the tolerance of apple to alkaline stress. The overexpression of MdATG18a increased biomass, photosynthetic rate and antioxidant capacity of transgenic plants compared with wild-type plants under alkaline stress. The overexpression of MdATG18a promoted γ-aminobutyric acid (GABA) shunt via an increase in glutamate (GABA precursor) and GABA contents and upregulation of GABA shunt-related genes. In addition, the overexpression of MdATG18a significantly upregulated the expression of other core ATG genes and increased the formation of autophagosomes under alkaline stress. In conclusion, these results suggest that the overexpression of MdATG18a in apple enhances alkaline tolerance and the GABA shunt, which may be owing to the increase in autophagic activity.