Abstract
Winter planting is promising for improving the utilization rate of fallow paddy fields in southern China by establishing arbuscular mycorrhizal fungi (AMF) communities. However, the effects of different winter forage crops on AMF community construction remain unknown. The AMF community establishment of different winter planting forage crops were conducted in oat, rye, Chinese milk vetch, and ryegrass, with winter fallow as a control. The AMF colonization rate, soil AMF spore density, community structure and diversity, and soil physicochemical properties were determined. The results showed that the total nitrogen and available nitrogen in winter Chinese milk vetch were 11.11% and 16.92% higher than those in winter fallow (P < 0.05). After planting winter forage crops, the AMF spore density in winter oat, rye, Chinese milk vetch, and ryegrass soil were 127.90%, 64.37%, 59.91%, and 73.62% higher than that before planting, respectively (P < 0.05). Claroideoglomus was the dominant AMF genus in the soil of winter planting oat, rye, and ryegrass. The average membership function value of winter Chinese milk vetch was the highest, indicating that it had the best comprehensive effect on soil physicochemical properties, AMF community structure and diversity, and fresh forage yield. Winter forage crops could increase the spore pool of soil AMF and improve the soil AMF community structure and diversity. Winter Chinese milk vetch in paddy field had the best comprehensive effect on soil physicochemical properties and soil AMF community according to the comprehensive evaluation. These findings provide a theoretical basis for sustainable development and utilization of the southern rice paddy ecosystem.
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1 Introduction
Arbuscular mycorrhizal fungi (AMF) are endosymbiotic fungi capable of forming stable symbiotic structures in most terrestrial plants (Smith and Read 2008). The mycorrhizal structure formed by AMF and plant roots can promote the growth, development, nutrient uptake, and stress resistance of host plants (Neumann and George 2010; Orlowska et al. 2012; Estrada et al. 2013). The ecological functions of the soil AMF community structure and diversity include the improvement of plant community stability and plant system productivity (Yang et al. 2016). When the AMF community diversity is low, the plant community structure and composition are unstable (Maherali et al. 2007). An increase in AMF community diversity can reduce competition between legumes and herbaceous plants and promote community stability (Wagg et al. 2011). With the increase of AMF species diversity, the corresponding functional diversity also increases, and the productivity of plant community increases through "functional complementarity" (Vogelsang et al. 2006; Wagg et al. 2011). Because AMF are symbiotic microorganisms in plant roots, host plant species determine AMF diversity to a certain extent (Aidee et al. 2021). In flooded paddy fields, the AMF colonization rate is low owing to a lack of oxygen, but AMF spores can be preserved in the soil (Lumini et al. 2011), which would contribute to the improvement of rice production (Cao et al. 2024). Few studies have investigated the effects of planting different crops on AMF communities in paddy soils during the winter fallow period.
There are abundant fallow paddy fields in southern China during winter that are suitable for crop growth with sufficient light, water, and heat resources (Wu et al. 2018). After long-term continuous double cropping rice in southern China, the soil is prone to problems such as compaction and deterioration of its physical and chemical properties (Gao et al. 2013). In the absence of vegetation cover, the soil is more susceptible to erosion, which accelerates the leaching of various elements (Xin et al. 2011). The introduction of crop rotation systems in winter fallow paddy fields can effectively improve soil conditions and avoid continuous cropping obstacles. Winter planting of green manure crops in paddy fields not only make effective use of winter fallow rice fields but also improve soil physicochemical properties and microbial communities, and produce a large amount of high-quality green feed to alleviate the shortage of forage crops during winter in southern China (Huang et al. 2022).
Common paddy rotation systems in southern China are generally constructed by the forage crops such as "rice-rice-Chinese milk vetch" and "rice-rice-ryegrass". Italian ryegrass (Lolium multiflorum L.) is widely cultivated in winter paddy fields to provide sufficient fodder (Yang et al. 2008; Tao et al. 2017), which can alleviate soil acidification and increase soil organic matter content (He et al. 2019), soil porosity, and soil water retention capacity (Madari et al. 2005; Qiao et al. 2017). Chinese milk vetch (Astragalus sinicus L.) is a legume green manure with high nutritional content and good quality that serves as a forage crop (Liu et al. 2020a), which can be symbiotic with rhizobia for biological nitrogen fixation to supplement soil nitrogen reservoirs (Li et al. 2019b). Similarly, other forage crops, such as oat (Avena sativa L.) and rye (Secale cereale L.), have become the dominant crops to be popularized and planted in southern winter fallow fields because of their cold resistance, strong adaptability, high yield, and high feeding value characteristics (Wang et al. 2016; Ma et al. 2020). Compared with cash crops, these forage crops not only provide a large amount of green fodder during winter, but can also be returned to the field as green manure, increasing the soil nutrient content and thus reducing fertilizer application (Zhang et al. 2017; Xu et al. 2021). The composition and diversity of the AMF community in the soil were affected by the different forage treatments. The relative abundances of Acaulospora and Diversispora increased significantly in soil planted with alfalfa (Medicago sativa L.) for a long time (Pellegrino et al. 2019). Legume nitrogen fixation can affect the soil AMF community composition (Xiao et al. 2019). Acaulospora and Septoglomus have high colonization rates and adaptability to leguminous forage-growing areas (Xiao et al. 2020). Planting ryegrass increased the AMF colonization rate and AMF community richness and increased the relative abundance of soil Glomerales and Archaeosporales (Garcia-González et al. 2023). However, few studies have focused on AMF communities from the winter planting of different forage crops in paddy fields.
The influences of different forage crops on AMF communities in winter paddy fields in southern China were explored using winter fallow, winter ryegrass, winter Chinese milk vetch, winter rye, and winter oat. Our hypotheses were as follows: (1) Winter planting of different forage crops has distinct effects on soil physicochemical properties. (2) Winter planting of forage crops can increase the AMF community diversity in paddy fields. (3) Different forage crops domesticate different AMF communities in the paddy fields.
2 Materials and methods
2.1 Experimental site description and design
The field experiment was conducted in Ai-rice Daoxiang Town (23°55’N, 113°56’E), Conghua District, Guangzhou, Guangdong Province, from November 2021 to March 2022. The experimental site was located approximately 35 m above sea level and had a subtropical monsoon climate. The average monthly precipitation during the experiment was only 42.4 mm. The highest temperature during the experiment period was 23 °C and the lowest temperature was 9 °C. The soil type was red soil, the paddy fields were planted with rice for two seasons per year before the experiment began, and field fallow in winter. The soil fertility at the experimental site was uniform. The soil background pH was 5.71 and soil organic matter (SOM), total nitrogen (TN), total phosphorus (TP), total potassium (TK), available nitrogen (AN), available phosphorus (AP), and available potassium (AK) were 23.14 g/kg, 1.32 g/kg, 0.67 g/kg, 18.52 g/kg, 102.15 mg/kg, 55.67 mg/kg, and 110.01 mg/kg, respectively.
Four forage crops suitable for winter planting in southern China were selected: Astragalus sinicus L. (MV), Avena sativa L. (O), Secale cereale L. (R), and Lolium multiflorum L. (RG). The sowing amount was 25 kg/ha. A random block design was adopted; each plot had an area of 20 m2 and three replicates of each crop were planted. The winter fallow treatment was performed using the same number of repeats. The winter forage crops were sown on November 15, 2021, and harvested on March 30, 2022. No fertilizer was applied during planting. The AMF colonization rate was determined by collecting root samples from plants in the crop harvest period. The soil samples were collected using a five-point sampling method. Each repeated soil sample was divided into three parts. A portion of the soil was air-dried and screened to 0.149 mm for the determination of the soil physical and chemical properties. A part of samples was stored in -80 ℃ refrigerator for soil AMF community structure and diversity analysis. The other part of the soil samples was temporarily stored in -20 ℃ refrigerator to count the AMF spore density in the soil.
2.2 Determination of soil chemical properties and forage yield
Soil pH was determined using a pH meter (PE-10; Sartorious, Germany) with a fresh soil to water ratio of 1:2.5 (w/v). Soil organic matter (SOM) was measured by combustion using a CNS elemental analyzer (CNS-2000; LECO, St. Joseph, MI, USA). Soil total nitrogen (TN) was measured using the Kjeldahl digestion method, soil total phosphorus (TP) was measured using molybdenum-antimony resistance colorimetry, and flame photometry was used to determine soil total potassium (TK) (Bao 2000). Available nitrogen (AN) was measured using the alkaline hydrolysis diffusion method with 1 mol/L NaOH, soil available phosphorus (AP) was determined by HCl-H2SO4 digestion, and soil available potassium (AK) was determined by NH4OAc solution extraction and flame photometry (Lu 2000).
During forage harvesting, five 20 cm × 20 cm quadrats were selected in each plot, all forage ground parts in the quadrat were harvested, fresh weight was measured, and yield was calculated. The collected fresh grass was dried for 1 h at 105℃, and then at 75℃ to constant weight.
2.3 Determination of AMF colonization rate and spore density
Fresh plant roots were selected and cut into 1 cm long root segments. After clearing with KOH, the roots were stained with trypan blue (Philips and Hayman 1970). Stained root segments were mounted on slides and examined under a microscope. The mycorrhizal colonization rate was calculated as the number of fields with mycorrhiza/total root segments × 100% (Cheng et al. 2019). 10 g of dried soil were weighed for each replicate. The AMF spores were separated from the 10 g soil by wet sieve decanting and sucrose density gradient centrifugation through 0.8 mm, 0.25 mm, and 0.045 mm screens from top to bottom, and then counted under a dissecting microscope (Trejo-Aguilar and Banuelos 2020).
2.4 DNA extraction and sequencing of soil AMF community
Soil DNA was extracted using an E.Z.N.A.® soil Kit (Omega Bio-tek, Norcross, GA, U.S.). DNA concentration and purity were measured by NanoDrop2000 UV–vis spectrophotometer (Thermo Scientific, Wilmington, DE, USA), and the quality of the extracted DNA was measured using 1% agarose gel electrophoresis. The fungal 18S rDNA gene was amplified by nested PCR, and AML1(5'-ATCAACTTTCGATGGTAGGATAGA-3') and AML2(5'-GAACCCAAACACTTTGGTTTCC-3') were selected as the first pair of primers. The amplified fragment was approximately 800 bp. AMV4-5NF (5'-AAGCTCGTAGTTGAATTTCG-3') and AMDGR (5'-CCCAACTATCCCTATTAATCAT-3') served as the second pair of primers-amplified fragments of about 300 bp (Tian et al. 2021). In the second round of PCR, Barcode was added to the primers AMV4-5NF and AMDGR for sample differentiation. The PCR products were extracted using a 2% agarose gel and purified using an AxyPrep DNA gel extraction kit (Axygen Biosciences, Union City, CA, USA). The purified PCR products were QuantiFluor™-ST (Promega, USA). These amplifications were subsequently combined and sequenced on the Illumina MiSeq platform in equal molar quantities (Zhang et al. 2020).
2.5 Applying fuzzy membership function to comprehensive evaluation
The fuzzy membership function was used to evaluate the effects of different treatments on the soil physicochemical properties and AMF communities (Wang et al. 2023). The membership function value of a certain index for each treatment was calculated, and then the average membership function value for each treatment was calculated. A larger value indicates a higher comprehensive evaluation of the treatment. The calculation method is as follows:
Uij is the membership function value of j index in i treatment, Xij is the average value of j index in i treatment, Ximax is the maximum value of this index in all treatments. Ximin is the minimum value of this index in all treatments. Ui is the average membership function value for each index in i treatment. If a certain index is negatively correlated with the variety selection goal, the membership function value is calculated using the inverse membership function method, as follows:
2.6 Data analysis
The obtained sequencing data were assigned to different samples according to the barcode sequence, which was then removed. Quality control was performed before analyzing the raw data to obtain high-quality sequences. Cluster analysis was performed on the processed sequences. Operational taxonomic units (OTU) were divided by 97% similarity using UPARSE software. BLAST software was used to compare the OTU representative sequences (the most abundant sequences in each OTU as representative sequences) with the NCBI microbial species classification database (ftp://ftp.ncbi.nih.gov/pub/taxonomy/), and the species information with the highest identity value was selected as the species classification information of the OTU. The α diversity analysis (Chao, Shannon) and β diversity analysis of OTU were performed by QIIME software. R Studio V3.6.1 software to perform principal component analysis (principal co-ordinates analysis, PCoA; based on the Bray–Curtis distance algorithm), distance-based redundancy analysis (db-RDA), and correlation heat maps. R Studio V3.6.1 and Gephi V0.9.2 were used to map the correlation networks between the soil AMF communities. One-way analysis of variance (ANOVA) was performed using SPSS (version 23.0; IBM, Armonk, NY, USA). The least significant difference (LSD) was used to compare means at the 5% confidence level. Origin 2021 was used to draw the bar and box charts. Numerical values are the means ± standard errors.
3 Results
3.1 Effects of different winter forage crops on soil physicochemical properties
The physical and chemical properties of the soil varied among the forage crops planted in winter (Table 1). The soil pH in winter fallow was the highest (5.81) and was the lowest after winter planting rye (5.57). With the exception of winter oat, planting other winter forage crops reduced soil pH to different degrees without significant differences compared with the background soil (P > 0.05). The SOM content in winter Chinese milk vetch was the highest, 15.13% higher than that in the background soil (P < 0.05); however, the differences among the five treatments were not significant (P > 0.05). The soil TN content of winter Chinese milk vetch was 26.64 g/kg, which was significantly higher than that of the background, winter fallow, winter oat, and winter ryegrass (P < 0.05). The highest AN content was found in winter rye (111.55 mg/kg), which was significantly higher than that in winter fallow, winter oat, and winter ryegrass (P < 0.05). The soil TP content of winter Chinese milk vetch was the highest (0.72 g/kg), and that of winter oat was the lowest (0.62 g/kg). There were no significant differences among the TP contents of the other treatments (P > 0.05). The AP content in the background soil was the highest and decreased to different degrees after the treatments. The AP content in the winter oat soil was significantly lower than those in the other treatments (P < 0.05). The TK content in winter forage crops decreased and the TK content in winter oat soil was the lowest (16.43 g/kg). The soil AK content of winter ryegrass was 69.78 mg/kg, which was significantly lower than that of other treatments (P < 0.05).
3.2 Yield of different forage crops
The results for different forage yields are shown in Table 2. The fresh forage yield and dry forage yield of ryegrass were the highest at 54,533.79 kg/ha and 7918.33 kg/ha, respectively. The fresh forage yield of ryegrass was 29.55%, 71.42%, and 43.96% higher than those of oat, rye, and Chinese milk vetch, respectively (P < 0.05). From high to low, the fresh and dry forage yields of different forage types were ryegrass, oat, Chinese milk vetch, and rye, and the difference among the different forage types was significant (P < 0.05).
3.3 AMF colonization rate and soil AMF spore density of different winter forage crops
The AMF colonization rates of different winter forage crops are shown in Fig. 1(a). The AMF colonization rate of oat was the highest (66.15%), and was significantly higher than those of rye, Chinese milk vetch, and ryegrass (P < 0.05). After winter forage crops, the soil AMF spore density increased to a certain extent (Fig. 1b). The AMF spore densities in winter oat, rye, Chinese milk vetch, and ryegrass soil were 127.90%, 64.37%, 59.91%, and 73.62% higher than that before planting, respectively (P < 0.05). The number of AMF spore density in the winter oat treatment reached 364.13, which was the highest among all the treatments.
Characteristics of the arbuscular mycorrhizal fungi (AMF) communities in different winter forage crops. a AMF colonization rate. b Soil AMF spore density. Different lowercase letters indicate significant differences between the same indicators under different treatments (P < 0.05). B: background soil; F: winter fallow; O: winter oat; R: winter rye; MV: winter Chinese milk vetch; RG: winter ryegrass
3.4 AMF community structure and diversity under different treatments
The AMF community composition in the soil under different treatments is shown in Fig. 2. Glomus was the dominant genus in both background and winter fallow soils. After planting forage crops in winter, the soil AMF community composition changed with an increased relative abundance of Claroideoglomus, Paraglomus and Acaulospora. In the soil of winter oat, rye, and ryegrass, Claroideoglomus was the dominant genus, Glomus was the secondary dominant genera. The relative abundance of Glomus (37.05%) was the highest in the soil of winter Chinese milk vetch, but was not significantly different from that of Claroideoglomus (33.74%).
Composition of AMF community in different treatments. B: background soil; F: winter fallow; O: winter oat; R: winter rye; MV: winter Chinese milk vetch; RG: winter ryegrass. uG1: unclassified_Gigasporaceae; uG2: unclassified_Glomeromycetes; uG3: unclassified_Glomeromycota; uG4: unclassified_Glomerales.
The α diversity indices (Chao1 and Shannon indices) of AMF community under different treatments were shown in Fig. 3. The Chao1 index of the winter oat was the highest, and the Chao1 index of the background soil was the lowest. The Shannon index of winter rye was the highest; however, there was no significant difference among winter rye, winter oat, and winter Chinese milk vetch (P > 0.05). The Chao1 and Shannon indices of the winter ryegrass treatments were not significantly different from those of the background soil and winter fallow treatments (P > 0.05). The principal coordinate analysis profiled by Bray–Curtis distances illustrated a significant difference in the AMF community structure among the different treatments (Fig. 4). The AMF community structure of the background soil was similar to that of the winter fallow. The AMF community structure in winter Chinese milk vetch was different from that of the other treatments.
α-diversity index of AMF communities in different winter forage crops a Chao1 index of AMF community. b Shannon index of AMF community. Different lowercase letters indicate significant differences between the same indicators under different treatments (P < 0.05). B: background soil; F: winter fallow; O: winter oat; R: winter rye; MV: winter Chinese milk vetch; RG: winter ryegrass
3.5 Correlations between the AMF genera and environmental factors
Redundancy analysis (RDA) revealed the correlation between AMF genera and environmental factors in the different treatments (Fig. 5). The first two redundancy analysis components explained 43.4% of the total variation, with RDA1 and RDA2 accounting for 33.3% and 10.1%, respectively. TK had the greatest effect on AMF genera, followed by TP and SOM. The RDA results indicated that differences in certain environmental factors regulated AMF genera. Spearman correlation analysis showed that soil TK content was significantly positively correlated with Glomus (P < 0.01) and significantly negatively correlated with Archaeospora (P < 0.01), Claroideoglomus (P < 0.01), Paraglomus (P < 0.01), and Acaulospora (P < 0.05) (Fig. 6). Soil pH was significantly and negatively correlated with Archaeospora and Acaulospora (P < 0.05). Soil AK was significantly negatively correlated with Archaeospora (P < 0.01) and significantly positively correlated with Glomus (P < 0.01). Acaulospora was also significantly positively correlated with the soil SOM and TN contents (P < 0.05) and significantly negatively correlated with the soil AP content (P < 0.05). Claroideoglomus was significantly and positively correlated with soil SOM content (P < 0.05).
Redundancy analysis (RDA) based on the relationship between the environmental factors and AMF genera. The environmental factors are denoted by red lines with arrows, and the AMF genera are denoted by blue lines with arrows. Dots of different colors represent the different soil samples. B: background soil; F: winter fallow; O: winter oat; R: winter rye; MV: winter Chinese milk vetch; RG: winter ryegrass. uG1: unclassified_Gigasporaceae; uG2: unclassified_Glomeromycetes; uG3: unclassified_Glomeromycota; uG4: unclassified_Glomerales.
Heatmap of Spearman correlation analysis between the soil physicochemical properties and AMF genera. Significance codes: *P < 0.05; **P < 0.01. B: background soil; F: winter fallow; O: winter oat; R: winter rye; MV: winter Chinese milk vetch; RG: winter ryegrass. uG1: unclassified_Gigasporaceae; uG2: >unclassified_Glomeromycetes; uG3: unclassified_Glomeromycota; uG4: unclassified_Glomerales.
3.6 Comprehensive evaluation of effects of different winter forage crops on soil physicochemical properties and AMF community
The comprehensive evaluation of the effects of the different treatments on soil physicochemical properties, AMF communities, and forage yield are shown in Table 3. The average membership function values were ranking from high to low as follows: winter Chinese milk vetch > winter rye > winter oat = winter ryegrass > winter fallow. The average membership function value of winter Chinese milk vetch was 0.55, indicating that it had the best comprehensive effect on soil physicochemical properties, AMF community structure, diversity, and forage yield.
4 Discussion
4.1 Winter forage crops improve soil nutrient cycling in paddy fields
Crop rotation is considered a sustainable agricultural strategy for improving soil ecological functions and crop yields (Zhao et al. 2020; Liu et al. 2023). Crop rotation also increases soil carbon and nitrogen contents and reduces fertilizer inputs for agricultural production (Cernay et al. 2018; Li et al. 2019a). Winter forage crops in paddy fields affect the physical and chemical properties of the soil. Our results showed that winter forage crops decreased soil pH, which may be due to the roots secrete organic acids in the process of forage planting, thus reducing the pH of rhizosphere soil (Adeleke et al. 2017). Legume forage crops, such as Chinese milk vetch, can use nitrogenase in root nodules to fix N2 in the air to form NH4+, resulting in the release of H+ from plant roots and a reduction in rhizosphere soil pH (Jarvis and Hatch 1985).
The increase in TN and AN content in winter Chinese milk vetch may be due to symbiosis with rhizobia for biological nitrogen fixation in Chinese milk vetch, which is a leguminous plant (Li et al. 2019b). The increase in TN and AN content in winter rye soil might be due to the low nitrogen absorption and utilization of rye as a barren-tolerant crop, and planting rye could reduce soil nitrogen loss (Krueger et al. 2011; Bauer et al. 2017). The decrease in TN and AN in the winter fallow treatment might be due to the loss of gaseous nitrogen caused by nitrification and denitrification of the soil or the loss of nitrogen in farmland runoff caused by rainfall (Fu et al. 2004; Wang et al. 2019). Winter forage crops decrease the contents of AP and AK in the soil, which are consumed to sustain forage crop growth. The different degrees of reduction in AP and AK indicate differences in the absorption and utilization of nutrients by different winter forage crops. The decrease in soil AP content during winter fallow may be duec (Liu et al. 2020b). The increase in SOM content under different treatments may have been due to the decomposition of the previous rice. The increase in SOM content under the treatment with winter rye and winter Chinese milk vetch might also be due to sufficient nitrogen nutrients in the soil, which could promote the growth of soil microorganisms and the decomposition of plant residues, thus leading to the accumulation of SOM (Xie et al. 2013; Bian et al. 2024).
4.2 Winter forage crops increased AMF spore pool in paddy soil and improved AMF community structure and diversity
Previous studies have generally considered that AMF, as aerobic fungi, rarely exists in the flooded environment of paddy fields; therefore, the importance of AMF in paddy fields has often been overlooked (Ilag et al. 1987). However, recent studies have shown that AMF spores exist in flooded paddy environments, and because rice roots secrete oxygen, they could give AMF the oxygen needed for symbiosis (Lumini et al. 2011; Li et al. 2022). Soil spore density was related to the AMF colonization rate of the plants. AMF spores are thick-walled asexual reproductive organs formed on the mycelium outside the root when the AMF develops to a certain stage (Smith and Read 2008). In natural environments, AMF can only produce spores by forming symbiotic structures with plants (Kokkoris et al. 2020). Our results showed that winter forage crops increased AMF spore density in paddy soils. Forage crops planted in paddy fields without flooding in winter can form a symbiotic structure with native AMF in the soil. After winter planting, the mycelia develop and mature to produce spores, thereby increasing the AMF spore pool in paddy fields. The AMF spore pool was the highest in the winter oat soil. A possible reason for this is that the AMF colonization rate of winter oat was the highest, and the mycelial structure was higher; therefore, the sporulation ability was stronger. The high AMF colonization rate of oat might be due to their well-developed root systems, which provide more AMF colonization sites (Ma et al. 2023). Therefore, winter forage crops in paddy fields were conducive to the enhancement of the soil AMF spore pool, and thus the AMF colonization rate of later rice increased. In conclusion, improvement of the AMF spore pool in paddy soils using winter forage crops will be beneficial for the production of post-cropping rice.
The diversity of AMF communities in the soils of different winter forage crops differed. Higher AMF diversity improves host plant resilience to the environment and improves plant community productivity (Vogelsang et al. 2006; da Silva et al. 2014). Winter oat, rye, and Chinese milk vetch improved the soil AMF community diversity. This might be due to the differences in root morphology and root exudates of different plants, which affect the recognition and infection of AMF, and thus affect the community composition and diversity of AMF (Bao et al. 2023). There are also differences in plant growth, physiological metabolism, and nutrient absorption that affect AMF diversity (Parvin et al. 2019). The growth and development stages of winter forage crops at the time of harvest would affect nutrient requirements, and thus affect the AMF species infected. Therefore, AMF community composition and diversity were affected. For example, soil AMF diversity is higher during reproductive growth than during vegetative growth (Wang et al. 2015). Winter forage crops increased the relative abundance of Claroideoglomus in soil. The dominant AMF genus in the soil of winter ryegrass, oat, and rye was Claroideoglomus, whereas the dominant AMF genera in the soil of winter Chinese milk vetch were Glomus and Claroideoglomus. Studies have shown that Claroideoglomus is helpful for improving the drought resistance of host plants. In the case of low rainfall in winter in southern China, the increase in Claroideoglomus infecting plants increases the drought resistance of plants, resulting in large spore production in the soil and an increase in relative abundance (Deng et al. 2020). Glomus and rhizobium co-exist more easily to promote the growth of legumes; therefore, Glomus is also the dominant genus in the soil of winter Chinese milk vetch (Franzini et al. 2010). Some studies have found that Claroideoglomus can improve the absorption of Ca and K, root development, and plant biomass, which are conducive to better plant growth (Jamiolkowska et al. 2020; Aguilera et al. 2022). Claroideoglomus has also been found to increase the tolerance of host plants to drought as well as to some heavy metals, such as cadmium and lanthanum (Deng et al. 2020; Hao et al. 2022). Therefore, an increase in the relative abundance of Claroideoglomus in paddy soils may be beneficial for rice growth and development. In conclusion, winter forage crops increase AMF diversity and alter the AMF community composition, which may improve the stability and productivity of paddy ecosystems.
4.3 Winter forage crops affected AMF community diversity by influencing soil physicochemical properties
Soil physicochemical properties are important factors that affect the AMF community structure and diversity (Mbodj et al. 2018). Soil pH, moisture, SOM content, and AP content have important effects on AMF community composition and diversity (Parvin et al. 2019). Soil pH plays a key role in the AMF community, and the most suitable pH for AMF infection is around pH 6; an increase or decrease in pH inhibits AMF infection (Qin et al. 2015; Liu et al. 2021). This finding is consistent with the results of our study. Many studies have shown that AMF community composition is significantly correlated with soil AP content (Sarkodee-Addo et al. 2020; Ibne et al. 2021). Our results showed no significant correlation between soil AP content and AMF. This may be because the AP content of the soil meets the plant growth and development needs. Under these conditions, available phosphorus was not the main factor limiting AMF colonization. Because oat roots are well-developed and could provide more infection sites for AMF (Ma et al. 2023), the colonization rate of AMF was higher, which promoted its absorption of nitrogen and phosphorus nutrients from the soil; therefore, the content of nitrogen and phosphorus in the soil was low. Our results showed that most AMF genera in paddy fields were significantly correlated with total and available K contents. The extraneous mycelia of AMF can activate soil potassium by secreting organic acids and oxalic acids, so that the potassium in soil that cannot be directly absorbed by plants can be converted into AK (Rouphael et al. 2015). Recent studies have shown that AMF can promote the absorption of potassium ions and reduce the absorption of sodium ions in plant tissues, thereby improving plant tolerance to salt stress (Wu et al. 2013). Therefore, the decrease in soil TK content might be due to the activation of potassium in the soil by mycelia after AMF infection, which is then absorbed by plants. The lowest soil AK content in winter ryegrass might be due to the differences between plant species, and the absorption and utilization of soil potassium by ryegrass was more obvious (Li et al. 2016).
It has been reported that the addition of soil nitrogen and phosphorus nutrients could change the composition of AMF community and promote the formation of dominant species (Camenzind et al. 2014). Glomerales are less sensitive to soil nitrogen content, and the relative abundance of most other AMF species decreases with increasing soil nitrogen content; however, there is no significant effect on Glomerales (Egerton-Warburton et al. 2007; Camenzind et al. 2014). Therefore, the increase in soil nitrogen content due to nitrogen fixation in winter Chinese milk vetch makes Glomus the dominant genus in the soil.
4.4 Comprehensive effects of winter forage crops on soil physicochemical properties and AMF community
Winter forage crops can produce green feed, increase soil nutrient content, and improve AMF communities without fertilization, making them ideal sustainable agricultural production models. The fuzzy membership function method has been widely used in the introduction and comprehensive evaluation of agronomic traits (Jin et al. 2021). Our results showed that Chinese milk vetch had the best comprehensive effects on soil physicochemical properties, AMF communities, and yield compared with other forage crops. Although winter oat had the greatest effect on improving the soil AMF community structure, they consumed a large amount of soil nutrients. The yield of winter rye was low, and although winter ryegrass had obvious advantages in terms of yield, it consumed more soil nutrients and reduced AMF community diversity. Winter Chinese milk vetch consumes less soil nutrients and could increase the soil nitrogen content owing to nitrogen fixation. In addition, the improvement in the soil AMF community structure was good. Although the forage crop yield was low, winter milk vetch had the greatest effect.
In practical production, Chinese milk vetch is widely applied to feed livestock because of its abundance as a legume crop. In addition, Chinese milk vetch is used as green manure to increase soil nutrient content and reduce fertilizer application through nitrogen fixation (Zhou et al. 2012; Yang et al. 2024). The aboveground parts of ryegrass are typically used as feed, whereas the stubble or the rest of the plant is returned to the field to decompose and increase the nutrient content of the soil (He et al. 2019b; He et al. 2020). Oat and rye are commonly used as fodder. However, little research has been conducted on their use as green manure. Therefore, Chinese milk vetch and ryegrass are considered more suitable for production owing to their diverse functions.
In post-cropping rice, increasing the AMF spore pool in the soil of winter forage crops can increase the AMF colonization rate, thus directly promoting nutrient absorption (Cao et al. 2024). AMF infection can promote the absorption and transfer of ammonium nitrogen and nitrate nitrogen, which can improve nitrogen nutrition in rice (Koegel et al. 2017). Therefore, nitrogen-fixing plants such as Chinese milk vetch can increase soil nitrogen levels, which may be more helpful for the growth of post-cropping rice. Changes in the soil AMF community can also increase post-cropping rice production. The dominant AMF genus Claroideoglomus in the soil of winter forage crops promotes the absorption of Ca and K by plants and improves the resistance of host plants to drought and heavy metals (Aguilera et al. 2022; Hao et al. 2022). The dominant AMF genus, Glomus, in winter Chinese milk vetch soil can improve rice yield and quality (Cao et al. 2024). Nutrient consumption by planting winter forage crops is gradually released by the root stubble decomposition of winter forage crops, thus reducing the impact on post-cropping rice (He et al. 2019b).
5 Conclusion
In this study, the effects of forage crops on soil physicochemical properties and AMF communities in winter paddy fields in southern China were compared. The results showed that winter forage crops could improve soil physicochemical properties and AMF community structure and diversity. The fuzzy membership function was used for a comprehensive evaluation, and the results showed that winter Chinese milk vetch had the most obvious improvement in soil physicochemical properties and the AMF community. These results provide a theoretical basis for the sustainable utilization of rice paddy ecosystems and the promotion of winter agriculture in southern China.
Availability of data and materials
The datasets during and/or analysed during the current study available from the corresponding author on reasonable request.
References
Adeleke R, Nwangburuka C, Oboirien B. Origins, roles and fate of organic acids in soils: A review. S Afr J Bot. 2017;108:393–406.
Aguilera P, Becerra N, Alvear M, Ortiz N, Turrini A, Azcón-Aguilar C, López-Gómez M, Romero JK, Massri M, Seguel A, Mora MdLL, Borie F. Arbuscular mycorrhizal fungi from acidic soils favors production of tomatoes and lycopene concentration. J Sci Food Agric. 2022;102:2352–8.
Aidee G, Marisol M, Leslie H, Gisel D, Paula Y, Anne K, Ruth MDW, Mary KF, Timothy N, Claire K. Crop diversity enriches arbuscular mycorrhizal fungal communities in an intensive agricultural landscape. New Phytol. 2021;231:447–59.
Bao SD. Analytical Methods of Soil Agro-chemistry. Beijing: China agriculture press; 2000. p. 87–162.
Bao XZ, Ma YY, Zou JX, Wu LM, Yang TT, Huang Q, Zhang B, Zhu HY. Research progress of arbuscular mycorrhizal fungi in paddy fields. Microbiol China. 2023;50:392–412.
Bauer E, Schmutzer T, Barilar I, Mascher M, Gundlach H, Martis MM, Twardziok SO, Hackauf B, Gordilo A, Wilde P, Schmidt M, Korzun V, Mayer KFX, Schmid K, Schön CC, Scholz U. Towards a whole-genome sequence for rye (Secale cereale L.). Plant J. 2017;89:853–69.
Bian Q, Zhao LX, Cheng K, Jiang YJ, Li DM, Xie ZB, Sun B, Wang XY. Divergent accumulation of microbe- and plant-derived carbon in different soil organic matter fractions in paddy soils under long-term organic amendments. Agric Ecosyst Environ. 2024;366: 108934.
Camenzind T, Hempel S, Homeier J, Horn S, Velescu A, Wilcke W, Rillig MC. Nitrogen and phosphorus additions impact arbuscular mycorrhizal abundance and molecular diversity in a tropical montane forest. Global Change Biol. 2014;20:3646–59.
Cao MY, Ye SP, Jin C, Cheng JK, Xiang Y, Song Y, Xin GR, He CT. Arbuscular mycorrhizal fungi communities established by different winter green manures in paddy field promotes the post-cropping rice production. J Integr Agric. 2024; https://doi.org/10.1016/j.jia.2024.07.035.
Cernay C, Makowski D, Pelzer E. Preceding cultivation of grain legumes increases cereal yields under low nitrogen input conditions. Environ Chem Lett. 2018;16:631–6.
Cheng JK, Yue MF, Yang HR, Chen BM, Xin GR. Do arbuscular mycorrhizal fungi help the native species Bidens biternata resist the invasion of Bidens alba? Plant Soil. 2019;444:443–55.
da Silva IR, de Mello CMA, Neto RAF, da Silva DKA, de Melo AL, Oehl F, Maia LC. Diversity of arbuscular mycorrhizal fungi along an environmental gradient in the Brazilian semiarid. Appl Soil Ecol. 2014;84:166–75.
Deng J, Li F, Duan TY. Claroideoglomus etunicatum reduces leaf spot incidence and improves drought stress resistance in perennial ryegrass. Australas Plant Pathol. 2020;49:147–57.
Egerton-Warburton LM, Johnson NC, Allen EB. Mycorrhizal community dynamics following nitrogen fertilization: a cross-site test in five grasslands. Ecol Monogr. 2007;77:527–44.
Estrada B, Aroca R, Maathuis FJM, Barea JM, Ruiz-Lozano JM. Arbuscular mycorrhizal fungi native from a Mediterranean saline area enhance maize tolerance to salinity through improved ion homeostasis. Plant Cell Environ. 2013;36:1771–82.
Franzini VI, Azcón R, Mendes FL, Aroca R. Interactions between Glomus species and Rhizobium strains affect the nutritional physiology of drought-stressed legume hosts. J Plant Physiol. 2010;167:614–9.
Fu BJ, Meng QH, Qiu Y, Zhao WW, Zhang QJ, Davidson DA. Effects of land use on soil erosion and nitrogen loss in the hilly area of the Loess Plateau. China Land Degrad Dev. 2004;15:87–96.
Gao JS, Xu MG, Dong CH, Huang J, Cao WD, Zeng XB, Wen SL, Nie J. Effects of long term rice-rice-green manure cropping rotation on rice yield and soil fertility. Acta Agron Sinica. 2013;39:343–9.
Garcia-González I, Martínez-García LB, Barel JM, Martens H, Snoek LB, Hontoria C, De Deyn GB. Cover crop identity determines root fungal community and arbuscular mycorrhiza colonization in following main crops. Eur J Soil Sci. 2023;74: e13427.
Hao BH, Zhang ZC, Bao ZH, Hao LJ, Diao FW, Li FYH, Guo W. Claroideoglomus etunicatum affects the structural and functional genes of the rhizosphere microbial community to help maize resist Cd and La stresses. Environ Pollut. 2022;307: 119559.
He HB, Zhang YW, Cheng JK, Jia XY, Xin GR, Chen WG. Effects of planting Italian ryegrass (Lolium multiflorum L.) during winter leisure on soil properties in paddy fields. Ecol Sci. 2019a;38:155–61.
He HB, Zhang YW, Cheng JK, Zhang Y, Li JN, Xin GR. Nutrient release regulation of stubble in the Italian ryegrass–rice rotation (IRR) system. Pratac Sci. 2019b;36:1852–60.
He HB, Li WX, Zhang YW, Cheng JK, Jia XY, Li S, Yang HR, Chen BM, Xin GR. Effects of Italian ryegrass residues as green manure on soil properties and bacterial communities under an Italian ryegrass (Lolium multiflorum L.)-rice (Oryza sativa L.) rotation. Soil Tillage Res. 2020;196:104487.
Huang GQ. Achievements, problems and strategic countermeasures of paddy farming system development in south China. J Huazhong Agric Univ. 2022;41:1–20.
Ibne Baki MZ, Suzuki K, Takahashi K, Chowdhury SA, Asiloglu R, Harada N. Molecular genetic characterization of arbuscular mycorrhizal fungi associated with upland rice in Bangladesh. Rhizosphere. 2021;18: 100357.
Ilag LL, Rosales AM, Elazegui FA, Mew TW. Changes in the population of infective endomycorrhizal fungi in a rice-based cropping system. Plant Soil. 1987;103:67–73.
Jamiolkowska A, Thanoon AH, Skwarylo-Bednarz B, Patkowska E, Mielniczuk E. Mycorrhizal inoculation as an alternative in the ecological production of tomato (Lycopersicon esculentum Mill.). Int Agrophys. 2020;34:253–64.
Jarvis SC, Hatch DJ. The effects of aluminium on the growth of white clover dependent upon fixation of atmospheric nitrogen. J Exp Bot. 1985;36:1075–86.
Jin HX, Guo DD, Yang QH, Yu XM, Fu XJ, Yuan FJ. Comprehensive evaluation of salt tolerance in soybean germination period by fuzzy subordinate function method. Mol Plant Breed. 2021;19:8265–71.
Koegel S, Mieulet D, Baday S, Chatagnier O, Lehmann M, Wiemken A, Boller T, Wipf D, Bernèche S, Guiderdoni E, Courty PE. Phylogenetic, structural, and functional characterization of AMT3;1, an ammonium transporter induced by mycorrhization among model grasses. Mycorrhiza. 2017;27:695–708.
Kokkoris V, Stefani F, Dalpé Y, Dettman J, Corradi N. Nuclear dynamics in the arbuscular mycorrhizal fungi. Trends Plant Sci. 2020;25:765–78.
Krueger ES, Ochsner TE, Porter PM, Baker JM. Winter rye cover crop management influences on soil water, soil nitrate, and corn development. Agron J. 2011;103:316–23.
Li T, Wang HY, Zhou ZJ, Chen XQ, Zhou JM. A new grading system for plant-available potassium using exhaustive cropping techniques combined with chemical analyses of soils. Sci Rep. 2016;6:37327.
Li XG, Jousset A, Boer WD, Carrión VJ, Zhang TL, Wang XX, Kuramae EE. Legacy of land use history determines reprogramming of plant physiology by soil microbiome. ISME J. 2019a;13:738–51.
Li YY, Pan FX, Yao HY. Response of symbiotic and asymbiotic nitrogen-fixing microorganisms to nitrogen fertilizer application. J Soil Sediment. 2019b;19:1948–58.
Li SY, Chen Y, Yu F, Zhang YJ, Liu K, Zhuo XX, Qiu YY, Zhang H, Gu JF, Wang WL, Yang JC, Liu LJ. Reducing methane emission by promoting its oxidation in rhizosphere through nitrogen-induced root growth in paddy fields. Plant Soil. 2022;474:541–60.
Liu CL, He CM, Wang LM, Zhang H, Wang F, Huang YB, Cao WD, Lin GT. The Study on Comprehensive Value of Chinese Chinese milk vetch as Forage and Green manure. Acta Agrestia Sin. 2020a;28:857–62.
Liu WF, Ciais P, Liu XC, Yang H, Hoekstra AY, Tang QH, Wang XH, Li XD, Cheng L. Global phosphorus losses from croplands under future precipitation scenarios. Envion Sci Technol. 2020b;54:14761–71.
Liu MH, Shen YK, Li Q, Xiao WF, Song XZ. Arbuscular mycorrhizal fungal colonization and soil pH induced by nitrogen and phosphorus additions affects leaf C:N: P stoichiometry in Chinese fir (Cunninghamia lanceolata) forests. Plant Soil. 2021;461:421–40.
Lu RK. Agriculture chemical analysis methods of soil. Beijing: China Agricultural Science and Technology Press; 2000. p. 106–95.
Lumini E, Vallino M, Alguacil MM, Romani M, Bianciotto V. Different farming and water regimes in Italian rice fields affect arbuscular mycorrhizal fungal soil communities. Ecol Appl. 2011;21:1696–707.
Ma XD, Sun JJ, Wang PB, Zhao YS, Song MJ, Yu XJ. Screening of the best mixed ratio of Avena sativa and Triticale Wittmack in the high-cold area of Guoluo. Qinghai Province Pratac Sci. 2020;37:753–61.
Ma QJ, Luan F, Jia B, Zhang Q, Wang L, Cui ZT, Li XG. Agricultural soil aggregation is affected by the crop root biomass rather than morphological characteristics. J Plant Nutr Soil Sci. 2023;186:339–50.
Madari B, Machado PL, Torres E, de Andrade AG, Valencia LI. No tillage and crop rotation effects on soil aggregation and organic carbon in a Rhodic Ferralsol from southern Brazil. Soil Tillage Res. 2005;80:185–200.
Liu Q, Zhao YX, Li T, Chen L, Chen YQ, Sui P. Changes in soil microbial biomass, diversity, and activity with crop rotation in cropping systems: A global synthesis. Appl Soil Ecol. 2023;186: 104815.
Maherali H, Klironomos JN. Influence of Phylogeny on fungal community assembly and ecosystem functioning. Sci. 2007;316:1746–8.
Mbodj D, Effa-Effa B, Kane A, Manneh B, Gantet P, Laplaze L, Diedhiou AG, Grondin A. Arbuscular mycorrhizal symbiosis in rice: establishment, environmental control and impact on plant growth and resistance to abiotic stresses. Rhizosphere. 2018;8:12–26.
Neumann E, George E. Nutrient uptake: the arbuscular mycorrhiza fungal symbiosis as a plant nutrient acquisition strategy. In: Koltai H, Kapulnik Y, editors. Arbuscular mycorrhizas: physiology and function. New York: Springer; 2010. p. 137–67.
Orlowska E, Godzik B, Turnau K. Effect of different arbuscular mycorrhizal fungal isolates on growth and arsenic accumulation in Plantago lanceolata L. Environ Pollut. 2012;168:121–30.
Parvin S, van Geel M, Yeasmin T, Lievens B, Honnay O. Variation in arbuscular mycorrhizal fungal communities associated with lowland rice (Oryza sativa) along a gradient of soil salinity and arsenic contamination in Bangladesh. Sci Total Environ. 2019;686:546–54.
Pellegrino E, Gamper HA, Ciccolini V, Ercolo L. Forage Rotations Conserve Diversity of Arbuscular Mycorrhizal Fungi and Soil Fertility. Front Microbiol. 2019;10:2969.
Phillips JM, Hayman DS. Improved procedures for clearing roots and staining parasitic and vesicular-arbuscular mycorrhizal fungi for rapid assessment of infection. Trans Br Mycol Soc. 1970;55:158-61.
Qiao WY, Gu HR, Shen YH. Effects of planting Italian ryegrass in winter fallow fields on soil fertility and microorganisms. Pratac Sci. 2017;34:240–5.
Qin H, Lu K, Strong PJ, Xu Q, Wu Q, Xu Z, Xu J, Wang H. Long-term fertilizer application effects on the soil, root arbuscular mycorrhizal fungi and community composition in rotation agriculture. Appl Soil Ecol. 2015;89:35–43.
Rouphael Y, Franken P, Schneider C, Schwarz D, Giovannetti M, Agnolucci M, De Pascale S, Bonini P, Colla G. Arbuscular mycorrhizal fungi act as biostimulants in horticultural crops. Sci Hortic. 2015;196:91–108.
Sarkodee-Addo E, Yasuda M, Lee CG, Kanasugi M, Fujii Y, Omari RA, Abebrese SO, Bam R, Asuming-Brempong S, Dastogeer KMG, Okazaki S. Arbuscular mycorrhizal fungi associated with rice (Oryza sativa L.) in Ghana: effect of regional locations and soil factors on diversity and community assembly. Agron J. 2020;10:559.
Smith SE, Read DJ. Mycorrhizal symbiosis. 3rd ed. New York: Elsevier; 2008. p. 13–41.
Tao JM, Liu XD, Liang YL, Niu JJ, Xiao YH, Gu YB, Ma LY, Meng DL, Zhang YG, Huang WK, Peng DL, Yin HQ. Maize growth responses to soil microbes and soil properties after fertilization with different green manures. Appl Microbiol Biot. 2017;101:1289–99.
Tian BL, Pei YC, Huang W, Ding J, Siemann E. Increasing flavonoid concentrations in root exudates enhance associations between arbuscular mycorrhizal fungi and an invasive plant. ISME J. 2021;15:1919–30.
Trejo-Aguilar D, Banuelos J. Isolation and culture of arbuscular mycorrhizal fungi from field samples. In: Ferrol N, Lanfranco L, editors. Arbuscular Mycorrhizal Fungi: Methods and Protocols. New York, NY, USA: Springer; 2020. p. 1–18.
Vogelsang KM, Reynolds HL, Bever JD. Mycorrhizal fungal identity and richness determine the diversity and productivity of a tallgrass prairie system. New Phytol. 2006;172:554–62.
Wagg C, Jansa J, Stadler M, Schmid B, van der Heijden MGA. Mycorrhizal fungal identity and diversity relaxes plant-plant competition. Ecology. 2011;92:1303–13.
Wang YT, Li T, Li YW, Björn LO, Rosendahl S, Olsson PA, Li SS, Fu XL. Community dynamics of arbuscular mycorrhizal fungi in high-input and intensively irrigated rice cultivation systems. Appl Environ Microb. 2015;81:2958–65.
Wang XX, Zhu DJ, Li Y, Xu ZM, Li Y. Screening of oat varieties in winter fallow field in South. Seed. 2016;35:112–4.
Wang HL, Han JQ, Deng N, An SS. Effects of the natural restoration time of abandoned farmland in a semiarid region on the soil denitrification rates and abundance and community structure of denitrifying bacteria. Appl Microbiol Biot. 2019;103:1939–51.
Wang L, Gao R, Zhuo XL, Bao JZ, Zhan WY, Geng JL, Yu Z. Membership function analysis for concentrate-to-forage ratio and ensiling time in total mixed ration fermentation. Pratac Sci. 2023;40:2711–20.
Wu QS, Zou YN, He XH. Mycorrhizal symbiosis enhances tolerance to NaCl stress through selective absorption but not selective transport of K+ over Na+ in trifoliate orange. Sci Hortic. 2013;160:366–74.
Wu MX, Zhao XL, Liu W, Zhang L, He L. Characteristics of hydrothermal conditions and their impacts on agricultural production in winter of 2017/2018. Chin J Agrometeorol. 2018;39:354–6.
Xiao D, Tan YJ, Liu X, Yang R, Zhang W, He XY, Wang KL. Effects of different legume species and densities on arbuscular mycorrhizal fungal communities in a karst grassland ecosystem. Sci Total Environ. 2019;678:551–8.
Xiao D, Liu X, Yang R, Tan YJ, Zhang W, Deng CX, He XY, Wang KL. Nitrogen input through chemical and biological pathway affects arbuscular mycorrhizal fungal communities in forage grass cultivation in southwest China. Appl Soil Ecol. 2020;150: 103454.
Xie ZB, Xu YP, Liu G, Liu Q, Zhu JG, Tu C, Amonette JE, Cadisch G, Yong JWH, Hu SJ. Impact of biochar application on nitrogen nutrition of rice, greenhouse-gas emissions and soil organic carbon dynamics in two paddy soils of China. Plant Soil. 2013;370:527–40.
Xin GR, Li J, Yang ZY. Effect of winter cropping Italian ryegrass on N and P leakage from soils. Acta Pratac Sinica. 2011;20:118–25.
Xu LX, Tang GJ, Tian K, Wang XY, Zhang JG. Effects of no-tillage sowing on soil properties and forage wheat and Italian ryegrass yields in winter fallow paddy fields. PeerJ. 2021;9: e10573.
Yang ZY, Xin GR, Yuan JG, Fang W, Li GX. Ecological fertilization: an example for paddy rice performed as a crop rotation system in southern China. In: Elsworth LR, Paley WO, editors. Fertilizers: properties, applications and effects. New York: Nova Science Publishers Inc; 2008. p. 1–28.
Yang HS, Xiong YQ, Wang Q, Guo Y, Dai YJ, Xu MM. Arbuscular mycorrhizal fungi species diversity: ecological functioning, determinants and assembling mechanisms. Acta Ecol Sin. 2016;36:2826–32.
Yang ZP, Rao ZX, Li HL, Zeng XJ, Xie J. Effects of Returning Green Manure-Chinese Milk Vetch on the Availability and Transformation of Zinc in Purple Tidal Mud Soil under Rice Cultivation. Agron J. 2024;14:1126.
Zhang XX, Zhang RJ, Gao JS, Wang XC, Fan FL, Ma XT, Yin HQ, Zhang CW, Feng K, Deng Y. Thirty-one years of rice-rice-green manure rotations shape the rhizosphere microbial community and enrich beneficial bacteria. Soil Biol Biochem. 2017;104:208–17.
Zhang RJ, Mu Y, Li XR, Li SM, Sang P, Wang XR, Wu HL, Xu N. Response of the arbuscular mycorrhizal fungi diversity and community in maize and soybean rhizosphere soil and roots to intercropping systems with different nitrogen application rates. Sci Total Environ. 2020;740: 139810.
Zhao J, Yang YD, Zhang K, Jeong JH, Zeng ZH, Zang HD. Does crop rotation yield more in China? A meta-analysis. Field Crops Res. 2020;245:107659.
Zhou XQ, Wu HW, Koetz E, Xu ZH, Chen CR. Soil labile carbon and nitrogen pools and microbial metabolic diversity under winter crops in an arid environment. Appl Soil Ecol. 2012;53:49–55.
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Cao, M., Xiang, Y., Huang, L. et al. Winter forage crops influence soil properties through establishing different arbuscular mycorrhizal fungi communities in paddy field. Adv. Biotechnol. 2, 30 (2024). https://doi.org/10.1007/s44307-024-00037-5
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DOI: https://doi.org/10.1007/s44307-024-00037-5