Leguminous plants are integral to both natural ecosystems and agricultural systems. They are renowned for their ability to fix atmospheric nitrogen through symbiotic relationships with rhizobial bacteria, which significantly enhances soil fertility and reduces the need for synthetic nitrogen fertilizers. This nitrogen-fixing capability not only supports the growth of leguminous plants themselves but also benefits subsequent crops in crop rotation systems, thereby promoting sustainable agricultural practices (Zhang et al., 2019; Stavridou et al., 2022). Additionally, legumes contribute to biodiversity and ecosystem stability by providing habitat and food for various organisms.

Soil microbial communities, comprising bacteria, fungi, archaea, and other microorganisms, play a crucial role in maintaining soil health and ecosystem functioning. These microbes are involved in nutrient cycling, organic matter decomposition, and the formation of soil structure. They also engage in complex interactions with plants, influencing plant growth, health, and productivity. For instance, mycorrhizal fungi enhance plant nutrient uptake, while other soil microbes can protect plants from pathogens. The diversity and composition of soil microbial communities are influenced by various factors, including plant species, soil type, and environmental conditions (Schlatter et al., 2015; Romdhane et al., 2021).

Understanding the interactions between leguminous plants and soil microbial communities is vital .These interactions can significantly impact soil nutrient dynamics and plant health, thereby influencing agricultural productivity and sustainability (Zhang et al., 2019; Stavridou et al., 2022). Leguminous plants can alter the composition and function of soil microbial communities, which in turn can affect soil health and ecosystem services. For example, the presence of leguminous plants has been shown to enhance microbial diversity and resilience, which are critical for soil ecosystem stability and function (Jiao et al., 2019; Zhang et al., 2019). Moreover, studying these interactions can provide insights into the mechanisms underlying plant-microbe symbioses and help develop strategies for improving crop performance and soil management practices (Lau and Lennon, 2011; Thrall et al., 2011).

This review provides an overview of the current status of the interaction between leguminous plants and soil microorganisms, and discusses in detail the biological basis of the interaction and its variation under different environmental and management conditions through case analysis. Through a comprehensive review and analysis, this review aims to understand the practical significance of these interactions on ecosystems and agricultural production, and optimize their application in agriculture, providing support for future research and practice.

2 Leguminous Plants and Their Role in Nitrogen Fixation

2.1 Leguminous plants and their unique ability to fix atmospheric nitrogen

Leguminous plants, belonging to the family Leguminosae (Fabaceae), are renowned for their unique ability to fix atmospheric nitrogen through a symbiotic relationship with nitrogen-fixing bacteria, primarily from the Rhizobium genus (Raza et al., 2020). This family is the third largest among angiosperms and includes a wide variety of forms ranging from annual herbs to large trees, distributed from tropical to arctic regions. The symbiotic relationship is facilitated by the formation of specialized structures called root nodules, where the bacteria reside and perform nitrogen fixation (Franche et al., 2009; Raza et al., 2020).

2.2 Mechanism of nitrogen fixation in leguminous plants

The process of nitrogen fixation in leguminous plants involves a complex interaction between the plant roots and nitrogen-fixing bacteria. The bacteria infect the root hairs of the host plant, leading to the formation of root nodules. Inside these nodules, the bacteria differentiate into a form known as bacteroids, which are capable of converting atmospheric nitrogen (N₂) into ammonia (NH₃) through the action of the enzyme nitrogenase (Liu et al., 2018; Raza et al., 2020). The plant supplies the bacteria with carbohydrates derived from photosynthesis, which the bacteria use as an energy source for the nitrogen fixation process. This mutualistic relationship ensures that the fixed nitrogen is made available to the plant in a form that can be assimilated for growth and development (Liu et al., 2018).

2.3 Importance of nitrogen fixation for plant growth and soil fertility

Nitrogen is a critical nutrient for plant growth, as it is a fundamental component of amino acids, proteins, and nucleic acids. However, atmospheric nitrogen is not directly accessible to most plants. The ability of leguminous plants to fix atmospheric nitrogen provides a significant advantage, allowing them to thrive in nitrogen-poor soils (Raza et al., 2020). This biological nitrogen fixation (BNF) not only supports the growth of the leguminous plants themselves but also enhances soil fertility by increasing the nitrogen content in the soil, benefiting subsequent crops planted in the same soil (Bhattacharjee et al., 2008; Patel, 2018; Raza et al., 2020). This process is environmentally friendly and sustainable, offering an alternative to chemical fertilizers, which can have detrimental effects on the environment.

3 Soil Microbial Communities

3.1 Types and functions of soil microbial communities

Soil microbial communities are composed of a diverse array of microorganisms, including bacteria, fungi, archaea, and protozoa. These microorganisms play crucial roles in various soil processes. Bacteria and fungi are the most abundant and functionally diverse groups, with bacteria such as Proteobacteria, Actinobacteria, and Bacteroidetes being dominant in many soil environments (Zhou et al., 2020a). Fungi, particularly mycorrhizal fungi, are essential for plant nutrient uptake and soil structure formation (Schweitzer et al., 2008; Zhang et al., 2019).

3.2 Role of soil microbes in nutrient cycling, organic matter decomposition, and soil structure

Soil microbes are integral to nutrient cycling, organic matter decomposition, and the maintenance of soil structure. They facilitate the breakdown of organic matter, releasing essential nutrients such as nitrogen, phosphorus, and potassium back into the soil, which are then available for plant uptake (Schlatter et al., 2015). Microbial mediation of niche differentiation in resource use is a key process, where different microbes access various nutrient pools, thus supporting plant nutrient partitioning. Additionally, soil microbes contribute to the formation and stabilization of soil aggregates, enhancing soil structure and porosity.

3.3 Factors influencing the composition and activity of soil microbial communities

Several factors influence the composition and activity of soil microbial communities, including plant species, soil type, and environmental conditions. Plant species and their associated root exudates significantly shape the microbial communities in the rhizosphere, the soil region influenced by root secretions. Soil type, including its pH, texture, and nutrient content, also plays a critical role in determining microbial community structure (Zhou et al., 2020a). Environmental factors such as moisture, temperature, and land management practices further modulate microbial activity and diversity (Paredes and Lebeis, 2016; Zhang et al., 2019).

4 Symbiotic Interactions Between Leguminous Plants and Soil Microbes

4.1 Types of symbiotic relationships between leguminous plants and soil microbes

Leguminous plants engage in various symbiotic relationships with soil microbes, primarily with rhizobia and arbuscular mycorrhizal fungi (AMF). The rhizobia-legume symbiosis involves rhizobial bacteria forming nodules on the roots of leguminous plants, where they fix atmospheric nitrogen into a form usable by the plant (Oldroyd et al., 2011). Another significant symbiotic relationship is the tripartite symbiosis, where both AMF and rhizobia simultaneously associate with the same leguminous host, enhancing nutrient acquisition and plant growth in nutrient-deficient soils (Chang et al., 2017; Lace and Ott, 2018).

4.2 Mechanisms of symbiosis formation and maintenance

The formation and maintenance of these symbiotic relationships involve complex molecular signaling and recognition processes. For rhizobia-legume symbiosis, the plant secretes flavonoids that trigger the production of Nod factors by rhizobia, which in turn initiate nodule formation on the plant roots (Oldroyd et al., 2011). Similarly, in AMF symbiosis, plants secrete strigolactones that activate mycorrhizal factors, facilitating the association with plant root hairs. The tripartite symbiosis requires coordinated gene regulation and mutual exchange of diffusible signal molecules to induce the expression of genes involved in the common symbiotic pathway (Chang et al., 2017; Lace and Ott, 2018).

4.3 Benefits of symbiosis for both plants and microbes

The symbiotic relationships between leguminous plants and soil microbes are mutually beneficial. For the plants, these symbioses enhance nutrient acquisition, particularly nitrogen and phosphorus, which are critical for plant growth and productivity (Oldroyd et al., 2011; Shtark et al., 2012; Chang et al., 2017). Rhizobia fix atmospheric nitrogen into ammonia, which the plant can use, while AMF improve phosphorus uptake through their extensive hyphal networks (Shtark et al., 2012; Chang et al., 2017). In return, the microbes receive carbohydrates and other organic compounds from the plant, which serve as energy sources (Oldroyd et al., 2011; Chang et al., 2017).

4.4 Examples of legume-microbe symbioses

The symbiotic relationship between leguminous plants and soil microorganisms is an important research field in plant ecology and agricultural production. These symbiotic relationships not only enhance plant nutrient absorption, but also play a crucial role in the global nitrogen cycle.

The symbiotic relationship between rhizobia and soybean (Glycine max) is the most typical example. Rhizobia can form nodules in the roots of soybeans, fixing nitrogen in the atmosphere and converting it into a form that plants can utilize, thereby increasing soil fertility and reducing dependence on chemical nitrogen fertilizer (Ikeda et al., 2010). This symbiotic relationship is of great significance for improving agricultural productivity.

The interaction between arbuscular mycorrhizal fungi (AMF) and peas (Pisum sativum) is another research focus. AMF helps plants more effectively absorb water and minerals, especially phosphorus, through symbiosis with pea roots. This symbiotic relationship not only enhances the growth and yield of peas, but also serves as an ideal model system for studying the mechanism of mycorrhizal symbiosis (Chang et al., 2017).

The tripartite symbiotic system of lotus (not specifically referring to a particular species) provides a unique research platform, involving the interaction between AMF, rhizobia, and host plants. There is a mutually beneficial relationship between wild leguminous plant Lotus japonicus and arbuscular mycorrhizal fungi (AMF) and nitrogen fixing rhizobia. The Lotus symbiotic genes play a wide-ranging role in building the root microbiota and highlight unexpected microbial interactions between root symbiotes and symbiotic communities. The bacterial root microbial community generally exhibits strong resistance to significant changes in the composition of root associated fungal communities, but simultaneously disrupting the symbiosis between AM and rhizobia increases the connectivity between bacterial root microbial communities (Figure 1) (Chang et al., 2017). This tripartite symbiotic relationship helps us gain a deeper understanding of how microorganisms collaborate with each other in complex environments, supporting plant growth, and also provides an important perspective for deciphering the genetic and molecular basis of symbiotic relationships.

5 Impact of Leguminous Plants on Soil Microbial Communities

5.1 Influence of leguminous plants on the diversity and composition of soil microbial communities

Leguminous plants significantly influence the diversity and composition of soil microbial communities. The cultivation of runner beans, for instance, has been shown to differentially shape the soil microbial community structure by recruiting specific bacteria and excluding others, thereby affecting the overall microbial diversity (Stavridou et al., 2022). Similarly, the presence of legumes in plant communities has been found to maintain higher microbial biomass compared to non-leguminous plants, indicating their crucial role in sustaining microbial diversity. In alpine grasslands, leguminous plant coverage, along with soil total nitrogen, explained a substantial portion of the variation in soil microbial communities, highlighting the importance of legumes in shaping microbial diversity (Cui et al., 2016). Furthermore, the diversity of soil bacterial communities under desert leguminous plants was significantly influenced by the type of rhizocompartment, with different bacterial communities being dominant in the root, rhizosphere, and bulk soil (Zhou et al., 2020a).

5.2 Changes in soil microbial activity and function in the presence of leguminous plants

The presence of leguminous plants also impacts soil microbial activity and function. For example, the cultivation of Bt maize, a genetically modified leguminous plant, resulted in increased microbial activity and nitrogen mineralization in the rhizosphere, demonstrating the functional changes induced by leguminous plants (Velasco et al., 2013). In experimental grassland ecosystems, the absence of legumes led to a decrease in microbial biomass and slower decomposition of organic material, underscoring the role of legumes in enhancing microbial activity. Additionally, leguminous plants like Hedysarum species in desert environments were found to enrich soil nutrients and support diverse microbial communities, which in turn influenced soil microbial functions such as nitrogen fixation and organic matter decomposition (Zhou et al., 2020a).

5.3 Long-term effects of leguminous plant cultivation on soil health

The long-term cultivation of leguminous plants has profound effects on soil health. Studies have shown that leguminous plants contribute to the maintenance of soil organic matter and nutrient cycling, which are critical for long-term soil fertility and health (Stavridou et al., 2022). The interactions between leguminous plants and soil microbes, particularly nitrogen-fixing bacteria, play a vital role in sustaining soil nitrogen levels, thereby enhancing soil fertility over time. Moreover, the hierarchical filtering and enriching effect of leguminous plants on beneficial microbes through their rhizocompartments further supports soil health by promoting a balanced and diverse microbial community (Zhou et al., 2020a). These long-term benefits underscore the importance of leguminous plants in sustainable agricultural practices and soil conservation efforts.

6 Case Study: Effects of Hedysarum Leguminous Plants on Soil Bacterial Communities in the Mu Us Desert, Northwest China

6.1 Study area and methods

The Mu Us Desert, located in northwest China, serves as the study area for examining the interactions between leguminous plants and soil microbial communities. This region is characterized by arid conditions and sandy soils, making it an ideal location to study desert leguminous plants such as Hedysarum mongolicum and Hedysarum scoparium. Zhou et al. (2020a) used high-throughput 16S rRNA genome sequencing and conventional soil physicochemical index measurements to analyze bacterial diversity and soil properties in different rhizosphere regions, including roots, rhizosphere soil, rhizosphere soil, and shrub soil (Figure 2).

The study revealed the diversity and interaction of bacterial communities in different rhizosphere environments by analyzing the roots and soil samples of two drought tolerant leguminous shrubs in the desert region of Northwest China. The aim was to understand the adaptability of these plants to desert environments and their ecological restoration effects.

6.2 Key findings and implications

The study revealed significant differences in bacterial community diversity across the different rhizocompartments. The root compartment was predominantly composed of Proteobacteria, Actinobacteria, Bacteroidetes, Tenericutes, and Chloroflexi, with Proteobacteria being the dominant genus. Soil nutrients were found to be higher in the rhizosphere soil compared to root-zone soil and intershrub bulk soil, except for total phosphorus and available phosphorus (Figure 3) (Zhou et al., 2020a). These findings suggest that desert leguminous plants have a hierarchical filtering and enriching effect on beneficial microbes through their rhizocompartments. Additionally, soil physicochemical factors such as pH and NH₄+-N significantly influenced the structure and composition of microbial communities in different rhizocompartments (Zhou et al., 2020a; 2020b).

This study revealed the complex interactions of bacterial communities in different soil and root environments through co-occurrence network analysis, and found that Proteobacteria dominated in the roots, while Actinobacteria was more significant in the soil under and between shrubs. This provides insights into microbial ecology in different soil and root environments, highlighting the main bacterial populations and their interactions, which are crucial for understanding soil health and plant microbial relationships.

6.3 Research significance

The implications of this case study extend beyond the Mu Us Desert, offering insights into the interactions between leguminous plants and soil microbial communities in other ecosystems and agricultural systems. The hierarchical filtering and enriching effect observed in desert leguminous plants could be applicable to other arid and semi-arid regions, where soil nutrient enhancement and microbial diversity are critical for plant growth and soil health. Furthermore, understanding the role of soil physicochemical factors in shaping microbial communities can inform sustainable agricultural practices, such as the use of leguminous plants to improve soil fertility and microbial diversity in degraded soils (Schlatter et al., 2015; Zhang et al., 2019).

In summary, this case study highlights the complex interactions between leguminous plants and soil microbial communities in the Mu Us Desert. The findings underscore the importance of rhizocompartments and soil physicochemical factors in influencing microbial diversity and community structure, with broader implications for ecosystem restoration and sustainable agriculture.

7 Factors Affecting Plant-Microbe Interactions

7.1 Soil properties

Soil properties such as pH, texture, and organic matter content play a crucial role in shaping the interactions between leguminous plants and soil microbial communities. For instance, soil pH significantly influences the composition of rhizomicrobiomes associated with different leguminous plants. In a study examining soybean and alfalfa, it was found that soil pH, along with phosphorus (P) and potassium (K), significantly affected the rhizomicrobiome composition of soybean, while soil pH and nitrogen (N) were more influential for alfalfa (Xiao et al., 2017).

Additionally, soil physicochemical factors, including pH and ammonium nitrogen (NH4+-N), were found to significantly impact the structure and composition of microbial communities in various rhizocompartments of desert leguminous plants (Zhou et al., 2020a).

7.2 Environmental conditions

Environmental conditions such as temperature and moisture also affect plant-microbe interactions. Soil moisture, for example, can influence the selection pressures on plant traits mediated by microbial community structure. In an experiment with Brassica rapa, it was observed that plants grown in soils with simplified microbial communities exhibited reduced growth and fecundity, and these effects were consistent across different soil moisture treatments (Lau and Lennon, 2011). This indicates that environmental stressors like soil moisture can modulate the impact of microbial communities on plant traits and interactions.

7.3 Plant species and their specific traits

The specific traits of leguminous plant species can differentially influence soil microbial communities. Different legume species have been shown to affect soil microbial community structure and function in species-specific ways. For example, the introduction of Medicago sativa and Astragalus adsurgens into abandoned fields significantly restored soil carbon and nitrogen content and microbial biomass within 3~5 years, whereas Melilotus suaveolens had a retarding effect on these parameters (Li et al., 2012). Additionally, legume species were found to enrich soil fungal communities more than grass species, and the variation in soil microbial community structure was greater among different legume species compared to grass species (Zhou et al., 2017).

7.4 Agricultural practices

Agricultural practices such as crop rotation, use of fertilizers, and pesticides can significantly influence plant-microbe interactions. These practices can impose selective pressures on microbial communities, thereby affecting their composition and function. For instance, the use of fertilizers can alter soil nutrient levels, which in turn can influence the microbial communities associated with leguminous plants. Studies have shown that soil nutrients, including carbon, nitrogen, phosphorus, and potassium, are negatively correlated with bacterial diversity, while the proportion of antagonistic bacteria is positively correlated with soil bacterial diversity (Schlatter et al., 2015). This suggests that agricultural practices that modify soil nutrient levels can have profound effects on the microbial communities and their interactions with leguminous plants.

In summary, the interactions between leguminous plants and soil microbial communities are influenced by a complex interplay of soil properties, environmental conditions, plant species traits, and agricultural practices. Understanding these factors is essential for optimizing plant-microbe interactions to enhance soil health and plant productivity.

8 Applications and Implications for Agriculture

8.1 Use of Leguminous plants in sustainable agriculture

Leguminous plants play a crucial role in sustainable agriculture due to their ability to form symbiotic relationships with soil microbes, such as rhizobia and arbuscular mycorrhizal fungi (AMF). These interactions enhance nitrogen fixation and phosphorus solubilization, reducing the need for chemical fertilizers and improving soil health (Shtark et al., 2012; Swarnalakshmi et al., 2020). The introduction of leguminous plants into agricultural systems can increase soil nutrient content and microbial diversity, which are essential for maintaining soil fertility and promoting plant growth. Additionally, leguminous plants can improve the resilience of soil microbial communities, making them more robust against environmental stresses (Zhang et al., 2019).

8.2 Strategies for enhancing beneficial plant-microbe interactions

To maximize the benefits of leguminous plants in agriculture, several strategies can be employed to enhance beneficial plant-microbe interactions. The application of plant growth-promoting rhizobacteria (PGPR) and AMF can significantly improve nutrient uptake and plant growth. Co-inoculation of PGPR with rhizobia has been shown to enhance nodulation and symbiotic interactions, leading to better crop yields (Shtark et al., 2012; Swarnalakshmi et al., 2020). Incorporating organic fertilizers such as compost and humic substances can promote the growth of beneficial soil microorganisms. These amendments act as prebiotics, enhancing the establishment and activity of inoculated microbes (Cozzolino et al., 2021). Advances in microbiome research and genetic tools allow for the manipulation of microbial communities to favor beneficial interactions. This can involve selecting and breeding legume varieties that are more effective at recruiting and maintaining beneficial microbes (Shtark et al., 2012; Finkel et al., 2017). Combining microbial inoculants with other sustainable practices, such as crop rotation and reduced tillage, can create a more favorable environment for beneficial microbes, further enhancing soil health and crop productivity (George et al., 2016; Cozzolino et al., 2021).

8.3 Potential challenges and limitations

While the benefits of using leguminous plants and enhancing plant-microbe interactions are well-documented, several challenges and limitations need to be addressed for practical application. The effectiveness of microbial inoculants can vary depending on environmental conditions, soil types, and crop species. This variability can make it challenging to achieve consistent results across different agricultural settings (Timmusk et al., 2017). Inoculated microbes may face competition from native soil microbial communities, which can limit their establishment and effectiveness. Strategies to mitigate this competition, such as using high-quality inoculants and optimizing application methods, are essential (Shtark et al., 2012). The development and commercialization of microbial inoculants require rigorous testing and regulatory approval. Ensuring the safety, efficacy, and stability of these products is crucial for their widespread adoption (Timmusk et al., 2017). Farmers may lack the knowledge or resources to implement these strategies effectively. Extension services and educational programs are needed to promote the benefits of leguminous plants and microbial inoculants and provide guidance on their use (Dubey et al., 2015).

9 Conclusion and Future Research Directions

The interactions between leguminous plants and soil microbial communities are complex and multifaceted, involving various rhizocompartments and microbial taxa. Studies have shown that leguminous plants significantly influence the diversity and structure of soil microbial communities through hierarchical filtering and enrichment processes. The presence of leguminous plants enhances soil nutrient content and microbial diversity, particularly in younger and less mature ecosystems. Additionally, plant community richness and microbial interactions play crucial roles in shaping soil bacterial communities, with plant-derived resources and antagonistic bacteria being key mediators. The role of microbial symbionts, such as arbuscular mycorrhizal fungi and rhizobia, in plant growth and nutrient acquisition has been well-documented, highlighting the importance of multi-component symbiosis in sustainable agriculture.

Despite significant advancements, several gaps remain in our understanding of legume-microbe interactions. One major gap is the limited knowledge of the specific mechanisms underlying the hierarchical filtering and enrichment of microbial communities by leguminous plants. Additionally, the interactions between multiple microbial symbionts and their collective impact on plant health and soil ecology are not fully understood. The influence of environmental variables, such as soil pH and nutrient availability, on these interactions also requires further investigation. Moreover, the role of microbial antagonists in shaping soil bacterial communities and their potential applications in biocontrol and soil health management need to be explored in greater detail.

Emerging technologies and methods are poised to revolutionize the study of plant-microbe interactions. High-throughput sequencing techniques, such as 16S rRNA gene sequencing and metagenomics, have already provided valuable insights into microbial diversity and community structure. Advances in bioinformatics and data analysis tools are enabling more comprehensive and accurate interpretations of complex microbial datasets. Additionally, molecular and genetic tools are being developed to unravel the intricate mechanisms of symbiotic interactions at the molecular level. The integration of these technologies with ecological and evolutionary frameworks will enhance our understanding of plant-microbe-soil interactions and their applications in sustainable agriculture and environmental management.

Future research should focus on elucidating the specific mechanisms of microbial community filtering and enrichment by leguminous plants. This includes investigating the roles of different rhizocompartments and soil physicochemical factors in shaping microbial communities. Studies should also explore the interactions between multiple microbial symbionts and their collective impact on plant health and soil ecology. Additionally, research on the influence of environmental variables, such as soil pH and nutrient availability, on plant-microbe interactions will provide valuable insights for optimizing agricultural practices. The development and application of advanced molecular and genetic tools will be crucial for unraveling the complex mechanisms of symbiotic interactions and their evolutionary and ecological dynamics. Finally, interdisciplinary approaches that integrate microbial ecology, plant biology, and soil science will be essential for advancing our understanding of plant-microbe interactions and their applications in sustainable agriculture and environmental management.

Conflict of Interest Disclosure

The authors affirm 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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