1 Introduction

Forest ecosystems are among the most productive and complex biomes on Earth, playing a crucial role in global carbon cycling and serving as significant carbon sinks (Baldrian, 2016; Lladó et al., 2017). These ecosystems are characterized by a high level of spatial heterogeneity and dynamic processes that range from short-term events to long-term ecological developments. The primary producers in these ecosystems, mainly trees, contribute to the structure and functioning of forests by providing habitats and resources for a diverse array of organisms, including fungi, bacteria, and archaea (Baldrian, 2016). The intricate interactions among these organisms and their environment are essential for maintaining ecosystem health and resilience, particularly in the face of global changes such as climate warming and increased carbon dioxide levels (Lladó et al., 2017).

Leaf litter, the layer of fallen leaves and other organic material on the forest floor, is a critical component of forest ecosystems. It plays a dual role in nutrient and carbon cycling and in regulating microclimatic conditions (Sayer, 2005). The decomposition of leaf litter is a key process in biogeochemical cycles, driven primarily by microbial communities, including fungi and bacteria (Baldrian, 2017; Bani et al., 2018). Fungi are particularly important due to their ability to produce specific enzymes that break down complex plant biomass, while bacteria contribute significantly to the decomposition of fungal mycelia and nitrogen cycling processes (Baldrian, 2017). The quality and quantity of leaf litter can influence microbial community composition and activity, thereby affecting soil fertility and ecosystem functioning (Bani et al., 2018; Feng et al., 2022).

This study aims to synthesize current knowledge on the role of leaf litter in forest soil fertility and microbial diversity. Specifically, it will examine the processes of leaf litter decomposition and its impact on nutrient cycling and soil organic carbon pools. Explore the interactions between leaf litter and microbial communities, including the effects of litter diversity on microbial activity and community structure. Assess the influence of environmental factors, such as climate and soil properties, on the decomposition process and microbial dynamics. And identify gaps in the current understanding and suggest directions for future research to enhance our knowledge of leaf litter's role in forest ecosystems. By integrating findings from various studies, this study will provide a comprehensive understanding of how leaf litter contributes to forest soil fertility and microbial diversity, highlighting its importance in maintaining ecosystem health and resilience.

2 Composition and Decomposition of Leaf Litter

2.1 Chemical composition of leaf litter

Leaf litter is composed of various chemical constituents, including lignin, cellulose, hemicellulose, tannins, and nutrients such as nitrogen (N), phosphorus (P), and calcium (Ca). These components play crucial roles in determining the decomposition rate and nutrient cycling within forest ecosystems. For instance, cellulose is a dominant polysaccharide in plant litter and significantly influences the decomposition process due to its abundance and the specific microbial communities it attracts (Štursová et al., 2012). Lignin, another major component, is more resistant to decomposition and requires specialized enzymes produced primarily by fungi (Bani et al., 2018). The nutrient content, such as N and P, also affects the microbial activity and decomposition rates, with higher nutrient levels generally promoting faster decomposition (Luo et al., 2017).

2.2 Decomposition processes and factors influencing decomposition rates

The decomposition of leaf litter is a complex process influenced by both abiotic and biotic factors. This process is essential for nutrient cycling and maintaining soil fertility in forest ecosystems.

2.2.1 Abiotic factors (temperature, moisture)

Temperature and moisture are critical abiotic factors that significantly influence the decomposition rates of leaf litter. Higher temperatures generally accelerate microbial activity, leading to faster decomposition rates. Moisture availability is equally important, as it affects the microbial community structure and their enzymatic activities. For example, in a study on macrophyte leaf litter, it was found that decomposition rates varied significantly with temperature, with higher rates observed at elevated temperatures (Zhao et al., 2020). Similarly, soil moisture levels can impact the microbial community composition and activity, thereby influencing the decomposition process (Dong et al., 2021).

2.2.2 Biotic factors (microbial activity, fauna)

Microbial communities, including bacteria and fungi, are the primary agents of leaf litter decomposition. Fungi, in particular, play a dominant role due to their ability to produce specific enzymes that break down complex organic compounds like lignin and cellulose (Bani et al., 2018). Bacteria also contribute significantly, especially in the later stages of decomposition when simpler compounds are more prevalent (Štursová et al., 2012). Soil fauna, such as earthworms and insects, also play a crucial role by physically breaking down litter and enhancing microbial access to organic matter. A meta-analysis revealed that soil fauna could contribute up to 30.9% to the forest litter decomposition rate, with their impact being more pronounced in warmer and moister climates (Xu et al., 2020).

2.3 Role of leaf litter in nutrient cycling

Leaf litter plays a pivotal role in nutrient cycling within forest ecosystems. As litter decomposes, it releases essential nutrients back into the soil, which are then available for plant uptake. This process is crucial for maintaining soil fertility and supporting plant growth. The decomposition of leaf litter contributes to the formation of soil organic matter (SOM), which is vital for soil structure and nutrient retention (Prescott and Vesterdal, 2021). The nutrient content of the litter, such as N and P, influences the rate of nutrient release and the overall nutrient dynamics in the soil. For instance, mixed leaf litter has been shown to enhance microbial diversity and nutrient release compared to single-species litter, thereby promoting more efficient nutrient cycling (Liu et al., 2022).

3 Impact of Leaf Litter on Soil Fertility

3.1 Contribution of decomposed leaf litter to soil organic matter

Decomposed leaf litter plays a crucial role in contributing to soil organic matter (SOM). The decomposition process, primarily driven by microbial communities, breaks down complex organic compounds in leaf litter into simpler forms, which are then incorporated into the soil. This process not only enriches the soil with organic carbon but also enhances its overall fertility. For instance, the addition of leaf litter has been shown to increase soil organic carbon mineralization, thereby contributing to the soil's organic matter content (Wang et al., 2014). Furthermore, volatile organic compounds (VOCs) released during leaf litter decomposition can diffuse into the soil matrix, contributing to various soil carbon pools and altering soil microbial communities (McBride et al., 2020).

3.2 Nutrient release and availability (nitrogen, phosphorus, potassium)

The decomposition of leaf litter is a critical process for the release and availability of essential nutrients such as nitrogen (N), phosphorus (P), and potassium (K). During decomposition, microorganisms, particularly fungi and bacteria, break down the organic matter, releasing these nutrients back into the soil (Figure 1). Studies have shown that the nutrient content and rate of decomposition of mixed litters are significantly different from those of single species, with mixed litters promoting higher microbial diversity and nutrient release (Liu et al., 2022). Additionally, the enrichment of ecosystems by nutrients like N and P has been found to have significant effects on fungal decomposers, which play a pivotal role in nutrient cycling (Jabiol et al., 2018). The addition of leaf litter has also been shown to increase soil microbial biomass carbon and nitrogen, further enhancing nutrient availability (Zhang et al., 2022).

3.3 Enhancement of soil structure and water retention

Leaf litter decomposition not only contributes to soil fertility but also enhances soil structure and water retention. The organic matter from decomposed leaf litter improves soil aggregation, which in turn enhances soil porosity and water-holding capacity. This is particularly important in maintaining soil health and preventing erosion. The presence of diverse microbial communities during litter decomposition has been linked to improved soil structure and stability (Pei et al., 2017). Moreover, the decomposition process can influence soil microbial community structure, which plays a role in maintaining soil physical properties. The addition of leaf litter has been shown to stimulate soil microbial activity, which can lead to better soil structure and increased water retention (Dong et al., 2021).

4 Influence of Leaf Litter on Microbial Diversity

4.1 Microbial communities in leaf litter (bacteria, fungi, actinomycetes)

Leaf litter serves as a critical habitat for diverse microbial communities, including bacteria, fungi, and actinomycetes. Studies have shown that the composition and diversity of these microbial communities are influenced by the type and diversity of leaf litter. For instance, in a Schrenk’s Spruce forest, the dominant bacterial phyla in leaf litter were Proteobacteria, Acidobacteria, and Actinomycetes, while the dominant fungal phyla were Basidiomycota, Ascomycota, and Mortierellomycota (Zhu et al., 2022). Similarly, in a Mediterranean oak forest, bacterial and fungal communities in leaf litter varied significantly during decomposition, with distinct succession patterns observed for each community (Santonja et al., 201). The presence of high-tannin leaf litter from transgenic poplars also influenced microbial communities, with notable changes in the abundance of Actinobacteria and various fungal classes (Winder et al., 2013).

4.2 Interactions between leaf litter and soil microbiota

The interactions between leaf litter and soil microbiota are complex and dynamic, significantly affecting soil microbial community structure and function. Leaf litter inputs can alter soil microbial activities and nutrient cycling, as observed in various forest ecosystems. For example, litter and root manipulations in a Schrenk’s Spruce forest led to changes in soil bacterial and fungal communities, with litter removal decreasing the diversity of these communities (Zhu et al., 2022). In a subtropical forest ecosystem, increased leaf litter diversity was positively correlated with the abundance of mycorrhizal fungi and actinomycetes, indicating strong interactions between aboveground litter and belowground microbial communities (Pei et al., 2017). Additionally, the composition of microbial communities in litter and soil is influenced by tree species, with distinct microbial communities developing on decomposing leaf litters of different tree species (Prescott and Grayston, 2013).

4.3 Impact of leaf litter on microbial diversity and abundance

Leaf litter has a profound impact on microbial diversity and abundance in forest soils. The diversity of leaf litter inputs can lead to cascading effects on microbial communities. In a Mediterranean oak forest, increased litter species diversity was associated with higher fungal diversity but lower bacterial diversity, highlighting the differential impact of litter diversity on microbial communities (Santonja et al., 2018). In a mixed Quercus acutissima and Robinia pseudoacacia forest, the decomposition rate of litter and the stability of microbial alpha diversity were higher compared to a pure Quercus acutissima forest, suggesting that mixed forests support more stable microbial communities (Dong et al., 2021). Furthermore, changes in litter input, such as litter removal or addition, can significantly affect soil microbial biomass and enzyme activity, with varying effects depending on forest stand density and soil quality (Wang et al., 2022). Overall, leaf litter plays a crucial role in shaping the diversity and abundance of microbial communities in forest soils, thereby influencing ecosystem functions such as nutrient cycling and decomposition.

5 Leaf Litter and Soil Health

5.1 Indicators of soil health influenced by leaf litter

Leaf litter plays a crucial role in influencing various indicators of soil health, including soil physicochemical properties, microbial biomass, and nutrient cycling. The addition of leaf litter from different tree species significantly affects soil total nitrogen, available NPK, and soil microbial biomass, which are critical indicators of soil health (Sun et al., 2017). For instance, the decomposition of mixed coniferous and broadleaf litters has been shown to enhance the carbon metabolic intensity, richness, and diversity of soil microbial communities, thereby improving soil health (Naimei, 2011). Additionally, the quality and quantity of leaf litter can influence the rate of litter decomposition and nutrient release, which are essential for maintaining soil fertility and microbial diversity (Silva et al., 2018).

The microbial community structure is also a vital indicator of soil health influenced by leaf litter. Studies have shown that the addition of leaf litter can alter the composition and functional characteristics of soil microbial communities. For example, the decomposition of leaf litter from dominant tree species enhances the metabolic capacity and functional diversity of soil microbes, which is crucial for soil health and forest succession (Liang et al., 2015). Moreover, the presence of specific microbial taxa, such as nitrifying and nitrogen-fixing bacteria, can be indicative of the nutrient status and overall health of the soil (Tanikawa et al., 2022).

5.2 Role in disease suppression and plant health

Leaf litter not only contributes to soil health but also plays a significant role in disease suppression and plant health. The decomposition of leaf litter can lead to the release of various organic compounds and nutrients that enhance soil microbial activity and diversity, which in turn can suppress soil-borne pathogens. For instance, the addition of leaf litter has been shown to increase the ATP-to-microbial biomass C ratio and the ergosterol-to-microbial biomass C ratio, indicating a more active and diverse microbial community capable of outcompeting pathogenic organisms (Salamanca et al., 2006).

Furthermore, the presence of a diverse microbial community in the soil, fostered by leaf litter decomposition, can enhance plant health by promoting beneficial symbiotic relationships and improving nutrient availability. The increased microbial diversity and activity can lead to better nutrient cycling and availability, which supports plant growth and resilience against diseases (Dong et al., 2021). Additionally, the specific composition of leaf litter, such as the carbon-to-nitrogen ratio and lignin content, can influence the microbial community structure and its ability to suppress plant pathogens (Naimei, 2011).

In summary, leaf litter significantly influences soil health indicators and plays a vital role in disease suppression and plant health by enhancing soil microbial diversity and activity. The decomposition of leaf litter from various tree species contributes to nutrient cycling, improves soil physicochemical properties, and fosters a diverse microbial community that supports healthy plant growth and resilience against diseases (Liang et al., 2015; Dong et al., 2021).

6 Case Studies

6.1 Effects of leaf litter on soil fertility in temperate forests

Leaf litter plays a crucial role in maintaining soil fertility in temperate forests. The decomposition of leaf litter releases essential nutrients back into the soil, which are vital for plant growth. For instance, the leachates from broadleaf litter in temperate forests contain higher amounts of carbon and nitrogen, which significantly enhance soil microbial activity and respiration, leading to improved soil fertility (Joly et al., 2016). Additionally, the diversity of leaf litter can influence the microbial community structure and nutrient cycling, as observed in mixed Quercus acutissima and Robinia pseudoacacia forests in Northern China, where mixed forests showed more stable microbial diversity and better nutrient cycling compared to pure forests (Dong et al., 2021).

6.2 Leaf litter's role in microbial diversity in tropical rainforests

In tropical rainforests, leaf litter significantly impacts microbial diversity and function. The identity of leaf litter species, rather than their diversity, has been shown to shape microbial functions and the abundance of soil organisms. For example, in Ecuadorian tropical montane rainforests, leaf litter from species with low carbon-to-nitrogen ratios, such as Cecropia andina, improved microbial activity and increased the abundance of microarthropods (Sánchez-Galindo et al., 2021). Furthermore, the addition of different leaf litter types to tropical forest soils in the Philippines altered soil microbial biomass and energy charge, indicating that leaf litter quality directly influences microbial community dynamics (Salamanca et al., 2006).

6.3 Leaf litter's impact on soil health in boreal forests

In boreal forests, leaf litter contributes to soil health by influencing microbial community structure and nutrient availability. The decomposition of leaf litter and deadwood is primarily driven by fungi, which play a significant role in nutrient cycling and maintaining soil health (Bani et al., 2018). However, the impact of leaf litter on microbial biomass and community structure can vary. For instance, a global meta-analysis revealed that aboveground litter removal decreased soil microbial biomass and altered the fungi-to-bacteria ratio, highlighting the importance of leaf litter in maintaining microbial balance and soil health in forest ecosystems (Jing et al., 2021).

6.4 Best practices for leaf litter management in mixed forests

Effective leaf litter management in mixed forests involves maintaining a balance between litter diversity and quality to support soil microbial communities and nutrient cycling. Studies have shown that mixed-species leaf litter can lead to non-additive effects on soil microbial activity, suggesting that the combination of different litter types can provide complementary resources for microorganisms (Joly et al., 2016). In subtropical forests, higher litter species diversity was associated with increased microbial biomass and enzyme activity, indicating that diverse litter inputs can enhance soil health and ecosystem functions (Pei et al., 2017). Therefore, promoting mixed-species forests and managing leaf litter to ensure a variety of litter types can be beneficial for maintaining soil fertility and microbial diversity.

7 Management Practices for Enhancing Leaf Litter Benefits

7.1 Sustainable forest management practices

Sustainable forest management practices are essential for maintaining the ecological balance and enhancing the benefits of leaf litter in forest ecosystems. One of the key practices involves the strategic management of both aboveground and belowground litter inputs. Research indicates that root litter inputs exert a larger control on microbial biomass than aboveground litter inputs, suggesting that forest management should consider the balance between these two types of litter to optimize soil microbial communities and carbon stability (Jing et al., 2021). Additionally, the decomposition of leaf litter and deadwood plays a crucial role in nutrient cycling and soil fertility, with fungi being major contributors due to their enzymatic capabilities (Bani et al., 2018). Therefore, promoting the growth of diverse fungal communities through sustainable practices can enhance the decomposition process and improve soil health.

Another important aspect is the management of litter diversity and identity. Studies have shown that leaf litter identity significantly influences microbial functions and microarthropod abundance, with high-quality species (low C-to-N ratio) improving resource quality and microbial activity (Sánchez-Galindo et al., 2021). This highlights the importance of maintaining a diverse range of tree species in forests to ensure a balanced and effective decomposition process. Moreover, the use of mixed litters, such as coniferous and broadleaf combinations, has been found to increase the carbon metabolic function of soil microbial communities, further supporting the need for diverse litter inputs (Naimei, 2011).

7.2 Leaf litter management in forestry and agriculture

In forestry and agriculture, effective leaf litter management can significantly enhance soil fertility and microbial diversity. One approach is the use of vermicomposting, which involves the recycling of nutrients from leaf litter waste using earthworms like Eisenia fetida. This method has been shown to improve soil physicochemical properties and increase microbial populations, making it a valuable practice for sustainable soil fertility management (Suthar and Gairola, 2014). Additionally, the incorporation of different leaf litters into soil can alter soil properties and enhance microbial community functions. For instance, the addition of leaf litters from various tree species to Panax ginseng-growing soil significantly affected soil nitrogen, phosphorus, and potassium levels, as well as microbial biomass (Sun et al., 2017).

Furthermore, the use of leaf litter extraction fluids from dominant tree species can enhance the metabolic capacity and functional diversity of soil microbes, contributing to forest succession and soil quality improvement (Liang et al., 2015). This practice can be particularly useful in abandoned lands or areas undergoing reforestation, where enhancing soil microbial activity is crucial for ecosystem recovery.

8 Challenges and Future Directions

8.1 Challenges in studying leaf litter decomposition and its effects

Studying leaf litter decomposition in forest ecosystems presents several challenges. One significant challenge is the complexity of interactions between microbial communities and environmental variables such as temperature and moisture, which can influence decomposition rates and nutrient cycling (Bani et al., 2018). Additionally, the variability in litter quality, including differences in chemical composition and physical structure, further complicates the study of decomposition processes (Elias et al., 2020; Sánchez-Galindo et al., 2021). The presence of diverse microbial communities, including fungi and bacteria, and their succession over time adds another layer of complexity, making it difficult to isolate specific factors that drive decomposition (He et al., 2016; Bani et al., 2018). Moreover, the effects of forest management practices, such as logging and the creation of forest gaps, on litter decomposition and soil fauna diversity are context-dependent and require long-term studies to fully understand (Huang et al., 2020; Dong et al., 2021).

8.2 Knowledge gaps and research needs

Despite significant progress in understanding leaf litter decomposition, several knowledge gaps remain. One major gap is the limited understanding of the role of bacteria in decomposition, which has historically been overshadowed by the focus on fungi (Bani et al., 2018). Additionally, the interactions between microbial community composition, litter chemistry, and decomposition rates are not fully understood, particularly in different forest types and under varying environmental conditions (He et al., 2016; Luo et al., 2017). There is also a need for more research on the effects of litter diversity versus litter identity on microbial functions and soil fauna abundance, as current studies provide mixed results (Santonja et al., 2017; Sánchez-Galindo et al., 2021). Furthermore, the impact of climate change, particularly increased drought, on litter decomposition and soil biota requires further investigation to predict future ecosystem responses (Santonja et al., 2017).

8.3 Future prospects for leaf litter research in forest ecosystems

Future research on leaf litter decomposition should focus on several key areas. First, there is a need for more comprehensive studies that integrate the roles of both fungi and bacteria in decomposition processes, considering their interactions and contributions to nutrient cycling (Luo et al., 2017; Bani et al., 2018). Long-term experiments that manipulate environmental variables, such as temperature and moisture, will help elucidate the effects of climate change on decomposition rates and microbial community dynamics (Santonja et al., 2017). Additionally, studies should explore the effects of different forest management practices, including the creation of forest gaps and mixed-species plantations, on litter decomposition and soil biodiversity (Huang et al., 2020; Dong et al., 2021). Advances in molecular techniques, such as metatranscriptomics, can provide deeper insights into the diversity and functional roles of protists and other microorganisms in leaf litter (Voß et al., 2019). Finally, research should aim to develop predictive models that incorporate microbial community composition, litter quality, and environmental factors to better understand and manage forest ecosystems in the face of global change (He et al., 2016; Luo et al., 2017).

9 Concluding Remarks

The role of leaf litter in forest soil fertility and microbial diversity is multifaceted and significant. Research indicates that both the diversity and identity of leaf litter influence microbial functions and soil properties. For instance, leaf litter diversity can enhance microbial activity and nutrient cycling, although the specific effects can vary over time and with different litter compositions. High-quality leaf litter, characterized by a low carbon-to-nitrogen ratio, tends to improve resource quality and increase the abundance of soil microarthropods. Additionally, the chemical composition of leaf litter leachates significantly affects soil microbial activity, with broadleaf litter generally providing more labile carbon and nutrients compared to coniferous litter. The diversity of microbial communities in the litter layer is also influenced by the diversity and identity of the leaf litter, with distinct succession patterns observed for bacterial and fungal communities.

The findings underscore the importance of maintaining tree species diversity in forest ecosystems to support soil fertility and microbial diversity. Forest management practices should consider the composition and diversity of leaf litter to promote healthy soil microbial communities and efficient nutrient cycling. For example, incorporating a mix of high-quality leaf litter species can enhance microbial activity and improve soil health. Additionally, understanding the specific effects of different tree species on soil properties can inform reforestation and conservation strategies, ensuring that the selected species contribute positively to soil microbial diversity and ecosystem functions. The role of leaf litter in shaping soil microbial communities also highlights the need for preserving native tree species and preventing the loss of biodiversity, which could have cascading effects on soil health and forest ecosystem stability.

Leaf litter plays a crucial role in forest ecosystems by influencing soil fertility, microbial diversity, and nutrient cycling. The interactions between leaf litter and soil microorganisms are complex and driven by both the diversity and identity of the litter. These interactions are essential for maintaining the health and productivity of forest soils. As such, leaf litter should be recognized as a vital component of forest ecosystems, with significant implications for forest management and conservation efforts. By fostering diverse and high-quality leaf litter inputs, we can support robust soil microbial communities and ensure the long-term sustainability of forest ecosystems.

Acknowledgments

The author sincerely appreciates the valuable opinions and suggestions provided by the three anonymous reviewers, whose meticulous review greatly helped us improve the quality of this article.

Conflict of Interest Disclosure

The author affirms that this research was conducted without any commercial or financial relationships that could be construed as a potential conflict of interest.

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