Recent research progress on soil microbial responses to drying–rewetting cycles

Abstract

Climatic change, such as increases in extreme drought and rainfall events and changes in rainfall intensity and pattern, has been strongly influencing soil moisture. The climatic change impact is particularly common in arid, semi-arid and Mediterranean regions, which is causing dramatic changes in the intensity and frequency of soil drying–rewetting cycles. The soil drying–rewetting cycle is a natural phenomenon that the soil experiences drying, then wetting, and then drying and rewetting again and again. When a dry soil is being rewetted, the amount of soil microbial biomass and its activity can be sharply increasing in a short time period, and then a large amount of gaseous carbon (C) and nitrogen (N) erupts from the soil. The sudden release of gaseous C and N is caused by the stimulation of the soil microbes. Such a phenomenon is called “Birch effect”. The drying–rewetting cycles have direct and indirect effects on soil microbes, and soil microbial responses to the drying and rewetting events play an important role in the feedbacks of terrestrial ecosystems. From aspects of soil microbial biomass, microbial activities and microbial structure, we review recent advances on studies regarding microbial responses to soil drying–rewetting cycles. We interpret the microbial responses using five different types of mechanisms: (1) Microbial stress mechanism: when a soil becomes dry, microorganisms must accumulate compatible solutes such as carbohydrates and aminoacids so that the soil microbes can equilibrate with their environment in order to avoid dehydrating and being killed. When the soil is rewetted, soil microbes must dispose of those osmolytes rapidly by transforming them into carbon dioxide (CO2), dissolved organic carbon (DOC) and nutrients in order to prevent water from being flowing into the cells. (2) Substrate supply mechanism: low soil moisture may result in the physical disruption of soil aggregates which leads to the exposure of new soil surfaces and of previously protected organic matter. When the soil is rewetted, its physical structure is further disrupted by swelling. The increased new soil surfaces and previously protected organic matter will improve the microorganism’s nutrient availability. (3) Soil hydrophobicity mechanism: soil hydrophobicity can cause the reduction of soil moisture and nutrient availability and inhibition of microbial decomposition of soil organic matter. Therefore, soil hydrophobicity is an important factor of explaining the activity of microorganism in drying and rewetting events. (4) Diffusive limitations mechanism: transportation of the soil microbe is limited in a dry soil. When soil moisture is increasing, soil microbial activity is enhanced along with the increased availability of substrate nutrients. (5) Predation mechanism: a moist soil is usually conducive to the increase of bacteria and fungi populations. In response, protozoa and nematodes also increase, leading to the fluctuation of the soil microbial community structure. On the basis of the literature review, we propose five important aspects to be considered in the future: (1) assessing soil microbes’ concrete adapting ways to the drying–rewetting cycles, (2) evaluating the microbial responses to the drying–rewetting cycles based on suitable indicators, (3) interpreting microbial responses to the drying–rewetting cycles by combining field investigation and laboratory controlling experiment, (4) investigating the microbial responses to the drying–rewetting cycles at different temporal and spatial scales.