Influence of climate change on soil microbial community

Introduction

Climate change has emerged as a significant driver of environmental transformations in recent decades. It is characterized by rise in global temperatures, shifting precipitation patterns and increased frequency of extreme weather events. The present state of climate change is unprecedented, with global temperatures having already increased by about 1.2°C above preindustrial levels (IPCC, 2021) [24] . This warming trend is expected to persist, with forecasts indicating a possible temperature rise of 1.5°C to 4.5°C by the end of the 21st century, depending on future emission scenarios (IPCC, 2021; Gattuso et al., 2021) [24,19] . This increase in temperature is accompanied by prominent shifts in precipitation patterns, resulting in changes in soil moisture levels, which are crucial factors influencing the dynamics of soil microbial communities (Seneviratne et al., 2021) [39] . Soil microbial communities, which include fungi, protists, viruses, bacteria, and archaea, play crucial roles in determining the structure and function of terrestrial ecosystems. They do this by participating in biogeochemical cycles and establishing close associations with plant roots. A significant challenge in deepening our understanding of the functional role of soil microbial communities is connecting the changes in their structure and function in response to climate change. Tackling this challenge necessitates a new understanding of the mechanisms that enable soil microbial communities to resist and recover from climate extremes (Bardgett et al., 2020) [15] Historically, Earth's climate has undergone natural fluctuations, but the speed and magnitude of the current changes are unprecedented in modern times. In the pre-industrial era, soil microbial communities evolved under relatively stable climatic conditions, playing key roles in nutrient cycling, organic matter decomposition, and maintaining soil health (Delgado-Baquerizo et al., 2020) [41] . Soil microbial communities, comprising bacteria, fungi, archaea, and viruses, are vital for nutrient cycling, organic matter breakdown, and soil structure formation (Delgado-Baquerizo et al., 2020; Singh et al., 2020) [41] . The rapid and ongoing changes in climate are now exerting unprecedented pressure on these communities, potentially threatening these essential ecosystem services (Wagg et al., 2021) [47] . Over million years, these communities have adapted to natural cycles of warming and cooling. However, the current rate of climate change is much faster, providing these communities with less time to adapt, which could lead to disruptions in critical ecosystem functions. The future impact of climate change on soil microbial communities raises significant concerns.

As global temperatures rise and changes in precipitation patterns and extreme weather events become more frequent, soil moisture and temperature conditions are likely to shift. These environmental changes can alter the composition, diversity, and function of soil microbial communities, which are crucial for maintaining soil fertility and the capacity for carbon sequestration (Schmidt et al., 2019; Trivedi et al., 2020) [38,45] . Such shifts in microbial communities may disrupt essential ecosystem functions, leading to decreased soil health and productivity. Additionally, the interaction between climate change and soil microbes could create feedback loops that further exacerbate climate change. For instance, changes in microbial activity could result in higher greenhouse gas emissions from soil, such as increased carbon dioxide and methane release, thereby accelerating global warming (Bradford et al., 2020; Crowther et al., 2019) [7,12] . This potential for a feedback loop where soil microbes contribute to the intensification of climate change highlights the importance of understanding and mitigating these impacts to preserve ecosystem stability and function. Addressing these challenges is critical for ensuring the resilience of ecosystems in the face of ongoing climate change.

Climatic factors affecting soil microbes

Since microorganisms are essential in soil organic matter decomposition and other carbon cycling processes, understanding how they respond to global changes can significantly enhance our knowledge of carbon cycling and its feedback mechanisms related to climate change (Yang et al., 2021) [52] . When ecosystems experience disturbances like increased dryness or changes from agricultural practices, the composition and activity of these microbial communities can change significantly. Such shifts may lead to major changes in how soil organic carbon (SOC) is stored or released. These changes in SOC dynamics are crucial because they directly impact the soil's ability to remain productive and sustainable over time (Ramirez et al., 2020). The characteristics of the environment where soil microbes live, such as the long-term climate and the soil's physical and chemical properties, can change the natural traits of soil microbial communities. These traits affect how well these communities can withstand and recover from extreme climate conditions (Bardgett et al., 2020) [15] .

Temperature change

Soil is the largest terrestrial reservoir of carbon (C). Warmer temperatures accelerate organic matter decomposition, potentially reducing soil organic carbon (SOC) levels, which are essential for maintaining soil aggregate stability (Six et al., 2000). The decline in SOC content leads to weakened binding agents between soil particles, decreasing the structural integrity of aggregates (Bronick & Lal, 2005) [8] . Soil microorganisms drive carbon losses to the atmosphere, but their response to temperature change due to climate warming is often short-lived and unpredictable. Long-term geothermal studies show that microbial activity is inherently sensitive to temperature increases and does not acclimate to warming, causing an initial acceleration in carbon loss. However, this effect is temporary, as substrate depletion reduces microbial biomass, limiting their influence on the ecosystem. A biogeochemical model confirms that warming accelerates microbial processes, but substrate availability ultimately constrains carbon loss from soils (Walker et al., 2018) [48] . A global meta-analysis showed that elevated temperature decreases soil microbial biomass nitrogen (N) while increasing N mineralization, nitrification, and denitrification rates, leading to a significant rise in N2O emissions. These changes suggest a shift in microbial N cycling from anabolic to catabolic processes, driven by higher temperatures. Increased N cycling processes were closely linked to enhanced microbial carbon metabolism, highlighting the interconnectedness of soil C and N cycles (Dai et al., 2020) [13] .

Change in rainfall pattern

Variations in rainfall patterns affect soil water conditions and microbial activity at different depths, influencing plant-soil interactions. An experiment was conducted in which 18 O stable isotope probing (SIP) to examine bacterial and fungal communities at three depths in soil mesocosms exposed to frequent or infrequent watering for 12 weeks. Results revealed that soil depth had a greater impact on microbial response to rewetting than the rainfall pattern (Engelhardt et al., 2018) [17] . Reduced rainfall can significantly impact soil microbial communities and affect overall ecosystem processes. This metaanalysis found that less rainfall led to a decrease in total soil microbial biomass and bacterial abundance, while fungal communities remained mostly unchanged. Interestingly, only certain bacterial groups, like Proteobacteria, were affected, indicating that while the overall microbial biomass declined, the composition of microbial communities didn't change much. The study also showed that the size of rainfall reductions mattered more than the duration, with larger reductions in biomass seen in wetter areas. In regions with high rainfall, reductions in soil organic carbon and respiration were closely linked to decreases in microbial biomass, suggesting that these ecosystems may be particularly vulnerable to future droughts (Ren et al., 2018) [18] .

Elevation in co2 levels

Prior to the industrial era, atmospheric CO₂ levels were stable at around 280 parts per million (ppm) due to balanced natural processes within the carbon cycle. For nearly 6,000 years, CO₂ levels hovered around this range without significant variation (Nunes et al., 2023) [54] . Following the onset of industrialization in the late 18th century, CO₂ levels began a notable rise as fossil fuel combustion, deforestation, and industrial processes released large amounts of CO₂ into the atmosphere. This upward trend continued through the 20th century and has sharply accelerated in recent decades, with current levels exceeding 420 ppm-over 50% higher than pre-industrial levels (Nunes et al., 2023) [54] . Higher levels of atmospheric CO₂ can change the way microbial networks are structured. While this hasn't been directly tested, it is possible that these changes could affect how vulnerable microbial communities are to shifting into different states when faced with extreme climate events. This means that increased CO₂ levels might make soil microbes either more or less stable in their response to sudden environmental changes, depending on how their internal networks are altered (Bardgett et al., 2020) [15] . Before industrialization, atmospheric CO2 levels were much lower, but burning fossil fuels and deforestation have significantly raised them. CO2 capture and storage is a strategy to reduce these levels and fight climate change, but leaks from storage sites can cause rapid acidification, potentially harming soil microbes. On a deeper level, CO2 influences microbes by altering enzyme activity, electron transfer, and gene expression (Yu et al., 2019) [53] .

Soil moisture

Long-term drought experiments revealed that soil microbial communities develop increased resilience to fluctuations in soil moisture during drying-rewetting (D/RW) cycles, though not resistance to drought itself. Rewetting events, rather than low soil moisture during drought, were key in shaping microbial responses. These cycles promoted efficient recovery, with microbes reducing respiration relative to bacterial growth. Such adaptations to soil moisture changes could help slow down the predicted loss of soil carbon under future drought conditions (EA de et al., 2019) [14] 1.5 UV radiation Ultraviolet (UV) radiation impacts both plants and soil microorganisms, altering their interactions. UV-A (320-400 nm) and UV-B (280-320 nm) influence plant morphogenesis and phototropism, while UV-B and UV-C stimulate secondary metabolite production. Short-wave UV radiation (<350 nm) directly damages pathogens through DNA disruption and increased membrane permeability, with UV-C (100-280 nm) being the most effective but also harmful to plants. UV-B and UV-C also indirectly affect microbes by enhancing plant defence responses, reactive oxygen species (ROS) accumulation, and secondary metabolite production (Vanhaelewyn et al., 2020) [46] . Changes in climate and stratospheric ozone dynamics are altering exposure to UV-B radiation, affecting both plants and soil microbes. While ozone recovery may reduce future UV-B levels, climate change introduces new combinations of UV-B with other factors like temperature, water availability, and elevated CO2 (Bornman et al., 2019) [6] . These interactions impact key ecosystem processes such as microbial-driven litter decomposition, pest-pathogen interactions, and species competition. Ozone depletion, particularly in the southern hemisphere, is also contributing to shifts in precipitation, temperature, and fire regimes, influencing microbial activity in terrestrial ecosystems (Bornman et al., 2019) [6] .

Microbial response to climate change 2.1 Shift in microbial habitat

Microbial traits are essential for soil carbon cycling, influencing how microorganisms respond to environmental changes. Climate change, by altering factors like moisture, temperature, and salinity, can push microbes to prioritize maintenance activities such as enzyme production, cellular repair, and defence overgrowth. This shift in microbial focus under stressful conditions affects the balance between carbon decomposition and stabilization in soils. As environmental pressures increase, microbial activity changes, potentially altering soil carbon stocks and contributing to climate change feedback. Understanding these microbial responses is crucial for predicting future ecosystem changes (Malik et al., 2020) [28] . Forests play a critical role in combating climate change by storing carbon in their soils, but they are increasingly threatened by rising carbon dioxide levels, warming temperatures, droughts, and nitrogen pollution. These environmental shifts lead to a microbial shift, where the balance of fungi and bacteria that help forests adapt is altered, impacting their ability to support carbon storage and nutrient cycling. Droughts weaken forest resilience by triggering pathogen outbreaks and wildfires (Baldrian et al., 2023) [3] .

Change in enzyme activities

Global environmental changes are altering soil extracellular enzyme activities (EEAs), which are critical for carbon and nutrient cycling. Nitrogen addition generally stimulates carbon and phosphorus-acquisition EEAs but suppresses oxidase activity, while phosphorus addition reduces P-acquisition enzymes. Changes in precipitation and warming also affect enzyme activities, with drought dramatically reducing oxidase activity. EEAs are more sensitive to nutrient additions than to climate change, with significant implications for soil carbon storage and plant productivity (Xiao et al., 2018) [51] . Climate change-induced thawing of permafrost leads to a shift in microbial communities, where fungal species like ericoid mycorrhizal fungi are replaced by bacteria and archaea that thrive in nitrogen-rich environments, such as ammonia-oxidizing microbes. This transition alters the nitrogen cycle, reducing microbes that support plant growth and potentially accelerating permafrost degradation (Seitz et al., 2022) [55] . Changes in soil moisture due to climate shifts, such as more frequent droughts, lead to microbial community changes, favouring species that can tolerate desiccation and produce protective substances like trehalose. On the other hand, wetter conditions promote anaerobic microbes, including methane-producing archaea, which can alter greenhouse gas emissions and soil health (Vicuña et al., 2021) [56] . Climate warming significantly increases soil ligninase activity by 21.4%, which is closely linked to higher soil respiration (SR), while cellulase activity remains unaffected (Chen et al., 2018) [35] . This increase in ligninase activity suggests that microorganisms are enhancing the breakdown of recalcitrant carbon in response to warming, sustaining long-term rises in SR. The effect of warming on ligninase activity grows over time, indicating that climate change is likely to accelerate carbon decomposition in soils (Chen et al., 2018) [35] . Drought, a major consequence of climate change, significantly impacts soil microorganisms by reducing their abundance, altering their structure, and decreasing enzyme production critical for nutrient cycling, such as oxidoreductases and phosphatases. This decline in microbial activity lowers soil fertility, negatively affecting plant productivity. However, organic amendments like compost can enhance soil enzyme activity, while plant growth-promoting microbes (Plant growth promoting Bacteria and plant growth promoting fungi) can help plants cope with drought (Bogati et al., 2022) [5] . Climate warming influences soil carbon dynamics by increasing the ratio of ligninase to cellulase activity, which reflects a shift in microbial preference toward decomposing more resistant carbon sources. This enzymatic shift is associated with a reduction in soil carbon stocks, particularly with long-term warming, which decreases the recalcitrant carbon pool by 14% (Chen et al., 2020) [13] . [57] Drought Reduction in microbial biomass Dehydrogenase, Phosphatase 21% reduction in heterotrophic respiration.

Chen et al., (2018) [35] Precipitation increase.

Increase in enzyme production and microbial biomass. Phosphatase 15-25% increase in activity. [59] 2.3 Impact on respiration Repeated cycles of drying and wetting can cause long-lasting changes in the functional state of soils, which can be measured through soil respiration -the process by which microbes break down organic matter and release carbon dioxide (CO₂). The way soil respiration responds to these cycles depends on how the land has been used in the past. Soils that were previously used for farming (arable agriculture) are more likely to undergo a sudden change in their respiration rates compared to soils that have a history of being grasslands, which tend to be more stable in their respiration response. This suggests that land use history plays a significant role in determining how soil respiration and overall soil function respond to environmental stress (Bardgett et al., 2020) [15] . The temperature response of soil microbial respiration (Rh) is crucial for understanding how these microorganisms will react to climate change, especially as temperatures rise. A recent study that analysed 25 soil samples from various forests, ranging from tropical to cold-temperate regions, found that the optimum temperature (for Rh is much higher than previously thought between 38.5 and 46.0 °C, compared to the old benchmark of 35 °C. This discovery highlights that varies significantly across different climates, with northern regions potentially releasing more CO2 as temperatures rise (Liu et al., 2018) [27] . [44] 3. Impact on soil health and functions

Soil structure & aggregate stability

The breakdown of soil structure and weakening of aggregate stability due to climate change has significant consequences for both soil health and agricultural productivity. When soil aggregates weaken, they easily break apart, which increases the risk of erosion, strips away valuable topsoil, and reduces the soil's ability to absorb water. As a result, soils become less effective at holding onto essential nutrients and moisture, directly impacting crop yields. This issue is particularly pressing in arid and semi-arid regions, where water is already scarce. In these areas, declining soil quality can further threaten food production, compounding the challenges faced by farmers (Schlesinger & Andrews, 2000) [37] . Soil microbiomes play a crucial role in agroecosystems, driving soil fertility, crop productivity, and stress resilience. Their functions are closely tied to soil structure, including aggregation and pore connectivity, which regulate water, oxygen, and nutrient flow. Agricultural practices can alter soil structure, impacting microbial processes and leading to issues like soil erosion and reduced fertility. Sustainable practices like integrated soil fertility management aim to improve soil structure and enhance microbial biodiversity (Hartmann et al., 2023) [21] . [26] Soil organic carbon Carbon within soil aggregates, related to stability. Dry combustion Zhang et al., 2023 [26] Bulk Density Indicates Soil compaction. Core sampling.

Phefadu & Munjonji, [33] 2024

Aggregate size distribution Distribution of aggregate sizes which shows stability across different sizes. Image analysis and sieving.

Phefadu & Munjonji, 2024 [33]

Soil erosion & compaction

Soils are essential for food security and providing various ecosystem services, but climate change poses significant challenges to their health. It affects soils directly through shifts in temperature, rainfall, and moisture levels, and indirectly through adaptations like irrigation and changes in crop rotation. A review of 20 case studies across Europe revealed that while many adaptation strategies can help reduce soil erosion and boost soil organic carbon, they might also lead to increased compaction in some areas (Hamidov et al., 2018) [20] . Climate and land use changes significantly impact soil erosion and land degradation. A study focused on a critical dam watershed near the Strait of Hormuz examined how these factors affect soil erosion now and, in the future (Azimi et al., 2019) [2] . Using advanced modelling techniques, the study found that rainfall erosivity is expected to increase under various climate scenarios, with the highest erosivity projected for 2030. Future land use changes are likely to convert rangelands into agricultural areas, resulting in decreased vegetation cover and increased soil erosion, which could reach up to 14.5/ha/year. Among the various factors contributing to soil erosion, the soil cover (C factor) is one that watershed managers can actively manage to help reduce soil loss and mitigate the adverse effects of climate change (Azimi et al., 2019) [2] .

Rate of decomposition

Microbial decomposition of soil organic matter (SOM) plays a significant role in the carbon cycle by releasing up to 60 peta-grams (Pg) of carbon as CO₂ from soils into the atmosphere each year. This amount is about six times greater than the current annual rate of human-caused (anthropogenic) carbon emissions, making microbial decomposition a crucial part of the global carbon cycle. Studies have shown that microbial decomposition of SOM generally increases exponentially with temperature (Moinet et al., 2021) [31] .

Nutrient cycling

Soil microbiomes consist of a wide variety of microorganisms, including bacteria, fungi, archaea, viruses, and protozoa. These microbes are crucial for cycling nutrients in the soil, which means they help break down and recycle essential elements. This process is important for keeping the soil healthy, supporting plant growth, and even influencing the climate. As microbes cycle nutrients, they can either release or absorb greenhouse gases like carbon dioxide (CO2), methane (CH4), and nitrous oxide (N2O). This happens because different microbes have different roles and capabilities. Some microbes release these gases into the atmosphere, while others might help reduce their levels. This diversity in microbial functions helps regulate not only soil health and plant growth but also impacts the climate by controlling the levels of greenhouse gases (B Mohanty et al., 2021) [30] 3.5 Impact on chemical properties Climate change significantly impacts carbon sequestration by altering key factors such as temperature, CO2 levels, and precipitation, which influence nutrient cycles like mineralization, decomposition, and leaching in soils. These changes affect soil fertility and plant nutrient availability. Enhancing soil carbon sequestration through effective management practices is critical not only for mitigating climate change but also for improving plant nutrition and boosting agricultural productivity, particularly in developing regions facing food security challenges (Elbasiouny et al., 2022) [16] . [15] . This increased tolerance mainly happens because the community reorganizes itself, favouring microbial groups that are better able to survive during droughts. As the environment changes, these droughtresistant microbes become more dominant, making the entire community more resilient to similar stress in the future. They can also shape the inherent characteristics of soil microbial communities. A global meta-analysis found that soils with high organic carbon content and a balanced pH had less variation or instability in microbial biomass over time, suggesting that these factors help reduce stress and enhance the stability of soil microbial communities (Bardgett et al., 2020) [15] .

Role of mutualistic relationships.

In a microbial community, the overall abilities and functions of the microbiome go beyond just the individual microbes present. This is because different microbial species interact with each other in complex ways and with the host plants in various types of relationships: mutualistic, synergistic, commensalistic, amensalistic, or parasitic. These interactions affect each microbe's survival, growth, and reproduction. When focusing mutualistic relationships, where both the microbes and plants benefit from each other, these interactions become especially important in resisting climate change. For example, mutualistic soil microbes can help plants better absorb nutrients and water, making them more resilient to environmental stresses like drought or extreme temperatures. This improved plant health supports soil fertility, which in turn creates a healthier environment for the microbes, reinforcing the entire system's ability to cope with climate change impacts (Suman et al., 2022).

Conclusion

The impact of climate change on soil microbial communities is complex and transformative, affecting both soil health and its role in supporting ecosystems. As temperatures rise and rainfall patterns shift, microbial habitats change, altering the balance of species and the activity of key enzymes. Elevated CO₂ and fluctuations in soil moisture also influence microbial respiration, nutrient cycles, and decomposition rates. Increased UV exposure further disrupts microbial functions, while soil structure and stability weaken, leading to erosion and compaction issues. Climate-driven stresses like drought test microbes' resilience, affecting their ability to support plant growth and maintain essential mutualistic relationships with plants. Together, these changes reshape the intricate web of life belowground, ultimately impacting soil fertility, carbon storage, and agricultural productivity.

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