An overview of the direct and indirect effects of acid rain on plants: Relationships among acid rain, soil, microorganisms, and plants

Acid rain (AR) causes numerous environmental problems and complex negative effects on plants globally. Many studies have previously reported on direct effects of AR or its depositional substances on plant injury and performance. However, few studies have addressed the indirect effects of AR on plants as mediated by soil microorganisms and the abiotic environment of the soil rhizosphere. The indirect effects (e.g., AR → soil microorganisms→plants) need greater attention, because acidic deposition not only affects the distribution, composition, abundance, function, and activity of plant-associated microorganisms, but also influences the dynamics of some substances in the soil in a way that may be harmful to plants. Therefore, this review not only focused on the direct effects of AR on plant performance, growth, and biomass allocations from a whole-plant perspective, but also addressed the pathway of AR–soil chemical characteristics-plants, which explains how soil solute leaching and acidification by AR will reduce the availability of essential nutrients and increase the availability of heavy metals for plants, affecting carbon and nitrogen cycles. Mainly, we evaluated the AR-soil microorganisms-plants pathway by: 1) synthesizing the potential roles of soil microbes in alleviating soil acidic stress on plants and the adverse effects of AR on plant-associated soil microorganisms; 2) exploring how plant mycorrhizal types affect the detection of AR effect on plants. The meta-analysis showed that the effects of AR-induced pH on leaf chlorophyll content, plant height, and plant root biomass were dependent on plant mycorrhizal types. Some possible reasons for different synergy between mycorrhizal symbiotic types and plants were discussed. Future research relating to the effects of AR on plants should focus on the combined direct and indirect effects to evaluate how AR affects plant performance comprehensively.