Simple Summary Microorganisms and their enzymes are crucial to ensuring soil quality, health, and carbon sequestration. Their numerous reactions are essential for biogeochemical cycles. However, a comprehensive review is lacking to summarize the latest findings in agricultural and enzymatic research. Although the relationships between soil enzyme activities and different soil ecosystems, such as arctic and permafrost regions, tropics and subtropics, tundra, steppes, etc., have been intensively investigated, particularly in the context of climate changes, only a few reviews summarize the impact of climate change on soil enzyme activity. This review aims to highlight the main groups of microbial enzymes found in soil (such as alpha-glucosidases and beta-glucosidases, phosphatases, ureases, N-acetyl-glucosaminidases, peptidases, etc.), their role in the global nutrient cycles of carbon, nitrogen, phosphorus, sulfur, carbon sequestration, and the influence of intensive agriculture on microbial enzymatic activity, and the variations in enzyme activity across different climate zones and soil ecosystems. Furthermore, the review will emphasize the importance of microbial enzymes for soil fertility and present both current challenges and future perspectives.Abstract The extracellular enzymes secreted by soil microorganisms play a pivotal role in the decomposition of organic matter and the global cycles of carbon (C), phosphorus (P), and nitrogen (N), also serving as indicators of soil health and fertility. Current research is extensively analyzing these microbial populations and enzyme activities in diverse soil ecosystems and climatic regions, such as forests, grasslands, tropics, arctic regions and deserts. Climate change, global warming, and intensive agriculture are altering soil enzyme activities. Yet, few reviews have thoroughly explored the key enzymes required for soil fertility and the effects of abiotic factors on their functionality. A comprehensive review is thus essential to better understand the role of soil microbial enzymes in C, P, and N cycles, and their response to climate changes, soil ecosystems, organic farming, and fertilization. Studies indicate that the soil temperature, moisture, water content, pH, substrate availability, and average annual temperature and precipitation significantly impact enzyme activities. Additionally, climate change has shown ambiguous effects on these activities, causing both reductions and enhancements in enzyme catalytic functions.

Changes in soil CO2 and N2O emissions due to climate change and nitrogen input will result in increased levels of atmospheric CO2 and N2O, thereby feeding back into Earth's climate. Understanding the responses of soil carbon and nitrogen emissions mediated by microbe from permafrost peatland to temperature rising is important for modeling the regional carbon and nitrogen balance. This study conducted a laboratory incubation experiment at 15 and 20 degrees C to observe the impact of increasing temperature on soil CO2 and N2O emissions and soil microbial abundances in permafrost peatland. An NH4NO3 solution was added to soil at a concentration of 50 mg N kg(-1) to investigate the effect of nitrogen addition. The results indicated that elevated temperature, available nitrogen, and their combined effects significantly increased CO2 and N2O emissions in permafrost peatland. However, the temperature sensitivities of soil CO2 and N2O emissions were not affected by nitrogen addition. Warming significantly increased the abundances of methanogens, methanotrophs, and nirK-type denitrifiers, and the contents of soil dissolved organic carbon (DOC) and ammonia nitrogen, whereas nirS-type denitrifiers, beta-1,4-glucosidase (beta G), cellobiohydrolase (CBH), and acid phosphatase (AP) activities significantly decreased. Nitrogen addition significantly increased soil nirS-type denitrifiers abundances, beta-1,4-N- acetylglucosaminidase (NAG) activities, and ammonia nitrogen and nitrate nitrogen contents, but significantly reduced bacterial, methanogen abundances, CBH, and AP activities. A rising temperature and nitrogen addition had synergistic effects on soil fungal and methanotroph abundances, NAG activities, and DOC and DON contents. Soil CO2 emissions showed a significantly positive correlation with soil fungal abundances, NAG activities, and ammonia nitrogen and nitrate nitrogen contents. Soil N2O emissions showed positive correlations with soil fungal, methanotroph, and nirK-type denitrifiers abundances, and DOC, ammonia nitrogen, and nitrate contents. These results demonstrate the importance of soil microbes, labile carbon, and nitrogen for regulating soil carbon and nitrogen emissions. The results of this study can assist simulating the effects of global climate change on carbon and nitrogen cycling in permafrost peatlands.

In Arctic soils, warming accelerates decomposition of organic matter and increases emission of greenhouse gases (GHGs), contributing to a positive feedback to climate change. Although microorganisms play a key role in the processes between decomposition of organic matter and GHGs emission, the effects of warming on temporal responses of microbial activity are still elusive. In this study, treatments of warming and precipitation were conducted from 2012 to 2018 in Cambridge Bay, Canada. Soils of organic and mineral layers were collected monthly from June to September in 2018 and analyzed for extracellular enzyme activities and bacterial community structures. The activity of hydrolases was the highest in June and decreased thereafter over summer in both organic and mineral layers. Bacterial community structures changed gradually over summer, and the responses were distinct depending on soil layers and environmental factors; water content and soil temperature affected the shift of bacterial community structures in both layers, whereas bacterial abundance, dissolved organic carbon, and inorganic nitrogen did so in the organic layer only. The activity of hydrolases and bacterial community structures did not differ significantly among treatments but among months. Our results demonstrate that temporal variations may control extracellular enzyme activities and microbial community structure rather than the small effect of warming over a long period in high Arctic soil. Although the effects of the treatments on microbial activity were minor, our study provides insight that microbial activity may increase due to an increase in carbon availability, if the growing season is prolonged in the Arctic.

Microbial processes, including extracellular enzyme (exoenzyme) production, are a major driver of decomposition and a current topic of interest in Arctic soils due to the effects of climate warming. While enzyme activity levels are often assessed, we lack information on the specific location of these exoenzymes within the soil matrix. Identifying the locations of different soil enzymes is needed to improve our understanding of microbial and overall ecosystem function. Using soil obtained from Utqiagvik, Alaska, our objectives in the study are (1) to measure the activity of enzymes in soil pore water, (2) to examine the distribution of activity among soil particle size fractions using filtration, and (3) to cross these particle size fraction analyses with disruption techniques (blending to shred and sonication to further separate clumped/ aggregated soil materials) to assess how tightly bound the enzymes are to the particles. The results of the soil pore water assays showed little to no enzyme activity (<0.05 nmol g soil(-1) h(-1)), suggesting that enzymes are not abundant in soil pore water. In the soil cores, we detected activity for most of the hydrolytic enzymes, and there were clear differences among the particle size and disruption treatments. Higher activities in unfiltered and 50-mu m filters relative to much finer 2-mu m filters suggested that the enzymes were preferentially associated with larger particles in the soil, likely the organic material that makes up the bulk of these Arctic soils. Furthermore, in the sonication + blending treatment with no filter, 5 of 6 hydrolytic enzymes showed higher activity compared to blending only (and much higher than sonication only), further indicating that enzyme-substrate complexes throughout the organic matter component of the soil matrix are the sites of hydrolytic enzyme activity. These results suggest that the enzymes are likely bound to either the producing microbes, which are bound to the substrates, or directly to the larger organic substrates they are decomposing. This close-proximity binding may potentially minimize the transport of decomposition products away from the microbes that produce them.

Information about the seasonal changes in soil enzyme activity along altitudinal gradients is key for understanding the responses of soil biochemical processes to ongoing climate change. Therefore, a 3-year in situ soil core incubation experiment was conducted along a 2431-m altitudinal gradient in an alpine-gorge region on the eastern Qinghai-Tibet Plateau, which includes dry valley shrubland, valley-mountain ecotone forest, subalpine coniferous forest, alpine coniferous forest and alpine meadow. The invertase, urease, and acid phosphatase activities in the soil organic layer (OL) and mineral layer (ML) were measured at five critical periods from 2013 to 2017. Higher soil enzyme activity was observed in the subalpine coniferous forest and alpine meadow, whereas lower activity was observed in the dry valley shrubland and valley-mountain ecotone forest. Overall, the seasonal sensitivity of soil enzyme activity decreased with increases in altitude. In the subalpine and alpine coniferous forests, the seasonal sensitivity of soil enzyme activity was significantly higher in the ML than in the OL. The key drivers of the seasonal variation in soil enzyme activity exhibited marked variation with changes in altitude, and these variations depended on the enzyme type and soil layer. Significant effects of temperature and precipitation on seasonal fluctuations in enzyme activities were observed at lower altitudes, whereas soil freeze-thaw cycles significantly affected the seasonal changes in acid phosphatase activity at higher altitudes. In addition, the seasonal variations in specific enzyme activities differed from those found for enzyme activities, which indicated that microorganisms might have mediated the effects of environmental factors on soil enzyme activity by altering the enzyme production efficiency. Briefly, the seasonal fluctuations in soil enzyme activity and their dominant drivers varied greatly among different altitudes in the alpine-gorge region, implying that soil biochemical processes will exhibit differential responses to ongoing climate change at different altitudes.

Peatland is a key component of terrestrial ecosystems in permafrost regions and have important effects on climate warming. Soil enzymes are involved in biogeochemical cycle of soil carbon (C), nitrogen (N) and phosphorus (P), which can be used as early sensitive indicators of soil nutrient changes caused by climate change. To predict the possible effects of permafrost degradation on soil enzymes in peatlands, ten peatlands from three types of permafrost regions along the permafrost degradation sequence (predominantly continuous permafrost region-predominantly continuous and island permafrost region-sparsely island permafrost region) in northeast China were selected to examine the activities of soil invertase, beta-glucosidase, urease and acid phosphatase and their relationships with soil physicochemical properties. The results demonstrated that permafrost type had significant effect on soil enzyme activities. Soil enzyme activities in predominantly continuous and island permafrost region were significantly higher than those in sparsely island permafrost region and predominantly continuous permafrost region. The activities of four soil enzymes were higher in 0-15 cm than 15-30 cm soil layer. Soil enzymes activities were positively correlated with soil ammonia nitrogen (NH4+-N), soil moisture content (SMC), total phosphorus (TP) and total nitrogen (TN), but negatively correlated with soil nitrate nitrogen (NO3--N). Soil inorganic nitrogen and moisture contents were the main factors affecting soil enzyme activities, with NH4+-N accounted for 41.6% of the variance, SMC 29.6%, and NO3--N 11.0%. These results suggested that permafrost degradation may change soil enzyme activities by changing soil physicochemical properties. In this study, only 0-30 cm peat soil in permafrost regions was collected during the complete thawing period of permafrost active layer, further studies should be placed on the change of soil enzyme activities in active layer and permafrost layer during freezing and thawing process in the southernmost location of northeast China in the Eurasia permafrost body and boreal forest belt.

Sustained climate warming increases the frequency and strength of soil freeze-thaw (FT) events, which strongly affect the properties of soil microbial communities. To explore the responses and mechanisms of the frequency and strength of freeze-thaw events on soil microbial communities, a lab-scale FT test was conducted on forest soil in permafrost region from the Daxing'an Mountains, China. The number of FT cycles (FTN) had a greater effect on microbial communities than FT temperature fluctuation (FTF). The FTN and FTF explained 20.9 and 10.8% of the variation in microbial community structure, respectively, and 22.9 and 11.6% of the variation in enzyme activities, respectively. The total and subgroup microbial biomass, the ratio of fungi to bacteria (F/B), and C- and N-hydrolyzing enzyme activities all decreased with an increase in FTN. Among microbial groups, arbuscular mycorrhizal fungi (AMF) were the most sensitive to FT events. Based on the changes of F/B and AMF, the reduction in soil carbon sequestration caused by frequent FT events can be explained from a perspective of microorganisms. Based on redundancy analysis and Mental Test, soil moisture, total organic carbon, and total nitrogen were the major factors affecting microorganisms in FT events. In the forest ecosystem, soil water and fertilizer were important factors to resist the damage of FT to microorganism, and sufficient water and fertilizer can lighten the damage of FT events to microorganisms. As a result of this study, the understanding of the responses of soil microorganisms to the variation in FT patterns caused by climate changes has increased, which will lead to better predictions of the effects of likely climate change on soil microorganisms.

Fire frequency and severity are increasing in tundra and boreal regions as climate warms, which can directly affect climate feedbacks by increasing carbon (C) emissions from combustion of the large soil C pool and indirectly via changes in vegetation, permafrost thaw, hydrology, and nutrient availability. To better understand the direct and indirect effects of changing fire regimes in northern ecosystems, we examined how differences in soil burn severity (i.e., extent of soil organic matter combustion) affect soil C, nitrogen (N), and phosphorus (P) availability and microbial processes over time. We created experimental burns of three fire severities (low, moderate, and high) in a larch forest in the northeastern Siberian Arctic and analyzed soils at 1, 8 days, and 1 year postfire. Labile dissolved C and N increased with increasing soil burn severity immediately (1 day) postfire by up to an order of magnitude, but declined significantly 1 week later; both variables were comparable or lower than unburned soils by 1 year postfire. Soil burn severity had no effect on P in the organic layer, but P increased with increasing severity in mineral soil horizons. Most extracellular enzyme activities decreased by up to 70% with increasing soil burn severity. Increasing soil burn severity reduced soil respiration 1 year postfire by 50%. However, increasing soil burn severity increased net N mineralization rates 1 year postfire, which were 10-fold higher in the highest burn severity. While fires of high severity consumed approximately five times more soil C than those of low severity, soil C pools will also be driven by indirect effects of fire on soil processes. Our data suggest that despite an initial increase in labile C and nutrients with soil burn severity, soil respiration and extracellular activities related to the turnover of organic matter were greatly reduced, which may mitigate future C losses following fire.

Geomorphic disturbances to surrounding terrain induced by thermal degradation of permafrost often lead to surface ponding or soil saturation. However, interactions between soil moisture and temperature on belowground carbon processes are not fully understood. We conducted batch incubation for three temperature treatments [constant freezing (CF), constant thawing (CT), and fluctuating temperatures (FTC)] and two soil moisture conditions (ponded and unsaturated). Extracellular enzyme activity was higher under ponded conditions than under unsaturated conditions, resulting in higher dissolved organic carbon (DOC) levels for ponded conditions. More CO2 and less CH4 were emitted under unsaturated conditions than under ponded conditions. Carbon dioxide emission was similar for CT and FTC treatments regardless of moisture conditions. However, CH4 emission was higher under ponded conditions than under unsaturated conditions for CT treatments, but was very low for FTC treatments regardless of moisture conditions. Little CO2 and CH4 were produced in CF treatments. Despite similar CO2 and CH4 emission levels for CT and FTC treatments, lower DOC levels were observed in the latter, indicating slower soil organic carbon (SOC) decomposition. Similar DOC variation patterns between CT and CF treatments indicated that SOC decomposition was considerable and further degradation to CO2 or CH4 was negligible even for CF treatments. The SOC decomposition and CO2 and CH4 emissions were considerable for FTC treatments. Our results suggest that labile-C produced during SOC decomposition in seasonally frozen soils and permafrost may provide supplemental substrates that would enhance the positive feedback to climate change with rising temperatures and wetter conditions.

The lack of snow cover due to winter climate change has great potential to impact winter soil nitrogen cycling in boreal forests. A snow manipulation was conducted in a Tibetan spruce forest to explore the effects of snow absence on winter soil nitrogen dynamics by shelter method. Snow absence on average reduced soil temperatures at the depths of 0 cm and 5 cm by 1.44 degrees C and 0.33 degrees C, respectively, throughout the winter. Moreover, snow absence increased soil frost and freeze-thaw cycles. Soil net nitrogen mineralization and labile nitrogen pools (ammonium, nitrate and dissolved organic nitrogen) were higher in the snow absence plots compared to control plots. Snow absence increased soil microbial biomass carbon but did not affect microbial biomass nitrogen. Nevertheless, soil enzyme activities involved in nitrogen cycles were often lowered by snow absence over the winter. The results noted in this study suggest that warming-induced absence in seasonal snowpack may stimulate winter soil nitrogen availabilities by changing soil microhabitats, which has important implications for soil biogeochemical cycles in the subalpine forest ecosystems on the eastern Tibetan Plateau.