Changes in soil aggregation and glomalin-related soil protein content as affected by the arbuscular mycorrhizal fungal species Glomus mosseae and Glomus intraradices

Arbuscular mycorrhizal (AM) fungi are key organisms of the soil/plant system, influencing soil fertility and plant nutrition, and contributing to soil aggregation and soil structure stability by the combined action of extraradical hyphae and of an insoluble, hydrophobic proteinaceous substance named glomalin-related soil protein (GRSP). Since the GRSP extraction procedures have recently revealed problems related to co-extracting substances, the relationship between GRSP and AM fungi still remains to be verified. In this work the hypothesis that GRSP concentration is positively correlated with the occurrence of AM fungi was tested by using Medicago sativa plants inoculated with different isolates of Glomus mosseae and Glomus intraradices in a microcosm experiment. Our results show that (i) mycorrhizal establishment produced an increase in GRSP concentration - compared to initial values - in contrast with non-mycorrhizal plants, which did not produce any change; (ii) aggregate stability, evaluated as mean weight diameter (MWD) of macroaggregates of 1-2 mm diameter, was significantly higher in mycorrhizal soils compared to non-mycorrhizal soil; (iii) GRSP concentration and soil aggregate stability were positively correlated with mycorrhizal root volume and weakly correlated with total root volume; (iv) MWD values of soil aggregates were positively correlated with values of total hyphal length and hyphal density of the AM fungi utilized.The different ability of AM fungal isolates to affect GRSP concentration and to form extensive and dense mycelial networks, which may directly affect soil aggregates stability by hyphal enmeshment of soil particles, suggests the possibility of selecting the most efficient isolates to be utilized for soil quality improvement and land restoration programs.     

短期放牧对高寒草甸土壤水稳性团聚体构成及稳定性的影响

土壤水稳性团聚体是土壤肥力的基础。以青藏高原东北边缘地区高寒草甸为对象,采用放牧控制试验研究不同放牧强度下各粒级水稳性团聚体含量、根系和微生物数量的变化特征,以期阐明短期放牧对水稳性团聚体及其稳定性的影响。结果表明,中牧能够增加土壤内水稳性大团聚体的含量。土壤水稳性团聚体有机碳含量随土层深度增加而减少,中牧时在0—10cm土层,粒径1mm的土壤水稳性团聚体有机碳含量显著低于对照,但在10—20cm土层,粒径1mm土壤水稳性团聚体有机碳含量大于对照和其他放牧处理;随着放牧梯度的增大,地下生物量也递增,在0—10cm土层表现最为明显;中牧处理下土壤中真菌数量最大,而放线菌的数量随着放牧强度的增大而增大。     

Soil aggregate stabilization by a saprophytic lignin-decomposing basidiomycete fungus I. Microbiological aspects

 We studied the effects of a saprophytic lignin-decomposing basidiomycete isolated from plant litter on soil aggregation and stabilization. The basidiomycete produced large quantities of extracellular materials that bind soil particles into aggregates. These binding agents are water-insoluble and heat-resistant. Water stability of aggregates amended with the fungus and the degrees of biodegradation of the binding agents by native soil microorganisms were determined by the wet-sieving method. The data demonstrated that aggregates supplemented with a source of C (millet or lentil straw) were much more water-stable and resisted microbial decomposition longer than when they were prepared with fungal homogenates alone. Moreover, retrieval of fungal-amended aggregates supplemented with millet during the first 4 weeks of incubation in natural soil exhibited more large aggregate fractions (>2 mm) than the ones supplemented with lentil straw. The possible relationship of the role of basidiomycetes in litter decomposition and soil aggregation is discussed. Received: 27 September 1999     

Fraxinus-Glomus-Pisolithus symbiosis: Plant growth and soil aggregation effects

It has been established that arbuscular mycorrhizal (AM) fungi are involved in the conservation of soil structure. However, the effect of ectomycorrhizal (EM) fungi alone or in interaction with AM fungi in soil structure has been much less studied. This experiment evaluated EM and AM fungi effects on soil aggregation and plant growth. Ash plants (Fraxinus uhdei) were grown in pots, and were inoculated with Glomus intraradices and Pisolithus tinctorius separately but also in combination. Our results showed that F. uhdei established a symbiotic association with EM and AM fungi, and that these organisms, when interacting, showed synergistic and additive effects on plant growth compared to singly inoculated treatments. EM and AM fungi prompted changes in root morphology and increased water-stable aggregates. AM fungi affect mainly small-sized macroaggregates, while EM and EM-AM fungi interaction mainly affected aggregates bigger than 0.5 mm diameter. These results suggest that ectomyccorrhizal as well as arbuscular mycorrhizal fungi should be considered in restoration programs with Fraxinus plants.     

Stabilisation of soil against wind erosion by six saprotrophic fungi

Soils with biological crusts that consist of fungal hyphae, bacteria and other small organisms usually resist erosion. However, the processes by which soil organisms stabilise air-dry aggregates against wind erosion are not well understood. We used saprotrophic fungi to examine some of these processes in a sandy clay loam (Hypercalcic Calcarosol). Soil aggregates, wetted with distilled water or glucose solution, were inoculated with one of six fungi, and incubated in darkness at 24 °C for 7 d in petri dishes under sterile conditions. Abrasion resistance (taken as resistance to wind erosion), tensile strength of soil, hot-water extractable carbohydrates (HWEC), dispersion index, pH, and hyphal length density (HLD) were each measured across all treatments. In all treatments, stability (abrasion resistance) and tensile strength, were positively related to HLD. Such relationships have not been reported elsewhere. All fungi enlarged the aggregates of the soil by cross-linkage and entanglement of particles, but with different processes, or different intensity of the processes, between species (for the same amount of substrate). The skins seen in scanning electron micrographs of stabilised soil were probably extracellular polysaccharides also produced by the fungi. We propose that the ductile failure of disks of soil, particularly those inoculated with Mucor sp., under tensile stress was due to movement of enmeshed particles, whereas the brittle failure of disks of soil inoculated with the other fungal species was due to metabolites or dispersed clay on the surface of the hyphae which limited deformation.     

麦茬复种饲料油菜对耕层土壤团聚体的影响

通过田间小区试验，研究了不同油菜种植密度与施氮肥对麦茬复种饲料油菜耕层（0～20cm）土壤颗粒组成、水稳性团聚体、微团聚体、粘粒分散率和团聚度的影响。试验结果表明，麦茬复种油菜能提高耕层土壤水稳性团聚体，改善土壤颗粒组成状况；与对照（CK）相比，耕层土壤粘粒分散率呈显著或极显著性降低，〈0．001mm的土壤团聚体破坏率和特征微团聚体比例（〈0．01mm／〉0．01mm）显著降低，而土壤团聚度在油菜苗期为降低趋势，收获期呈显著或极显著性增加，〉0．25mm的土壤团聚体百分含量除收获期的水稳性团聚体外均呈不显著性降低。复种油菜对土壤水稳性团聚体改善贡献高于土壤微团聚体。种植密度和施肥处理均能不同程度地影响和改变耕层土壤水稳性团聚体、微团聚体和粘粒分散率的含量，处理间规律性不强，差异性并不十分显著。说明麦茬复种饲料油菜对农田土壤肥力的改善和提高具有积极作用。     

Soil biota effects on soil structure: Interactions between arbuscular mycorrhizal fungal mycelium and collembola

Soil aggregation is an important ecosystem process mediated by soil organisms. Collembola and arbuscular mycorrhizal (AM) fungi are major soil biota representing different functional groups, and are known as two key promoters of soil aggregation. Although several studies have experimentally demonstrated that AM fungi and, more recently, collembola affect soil structure, there is no study investigating how both soil organisms affect soil aggregation excluding the influence of plant roots, another important driver of soil aggregation. Considering the importance of AM fungi and collembola in terrestrial ecosystems, here we asked if both organisms have any influence on soil aggregation when roots are not present.In order to examine this question we conducted a completely factorial greenhouse study manipulating the presence of both collembola and AM fungi and excluded the roots of Plantago lanceolata using a 38 μm nylon screen compartment. We quantified soil aggregation as water stable soil aggregates in four size classes in the hyphal compartment and monitored a number of other explanatory variables, including AM (and non-AM) fungal soil hyphal length.The soil in the hyphal compartment showed greater soil aggregation with larger mean weight diameter when collembola were present, and a similar result was found in the presence of AM fungi, compared to control treatments. Moreover, combined presence of both AM fungi and collembola resulted in a non-additive increase of soil aggregation.Our study clearly indicated that collembola can enhance soil aggregation, that they can partially complement effects of AM fungi, and that these effects are independent of roots.     

The capacity of soil particles for spontaneous formation of macroaggregates after a wetting-drying cycle

The capacity of soil particles for spontaneous formation of aggregates >0.25 mm was studied in a laboratory experiment. The particles from soil aggregates (3-1 mm) (initially aggregated particles, APs) and initially free particles (FPs) of <0.25 mm in size were isolated from the soddy-podzolic and chernozemic soils under fallow and from the arable soddy-podzolic soil. The aggregates of 3-1 mm were ground and passed through a 0.25-mm sieve. Then, the aggregates and free particles were poured with water and dried, and the content of the formed aggregates and their water stability were determined; in the samples from the arable soddy-podzolic soil, the organic carbon content was also determined in the newly formed aggregates. The FPs from the untilled soils formed almost no aggregates. At the same time, the APs from these soils manifested the ability for the spontaneous formation of aggregates, including water-stable aggregates. In the arable soddy-podzolic soil, on the contrary, both FPs and APs demonstrated the capacity for spontaneous self-organization into aggregates. The water stability of the self-organized aggregates from the arable soil was similar regardless of their source (APs or FPs). It was supposed that the ability of the FPs from the arable soil to form macroaggregates reflects the mechanical degradation of the aggregates in the soil: tillage results in the degradation of the aggregates, and the particles capable of spontaneously aggregation temporarily fall in the fraction of <0.25 mm. The water-stable aggregates produced from the APs or FPs of the arable soil contained more organic carbon (1.89%) in comparison with the water-stable aggregates separated from the initial 3- to 1-mm aggregates of this soil (1.31%).     

护坡人工混凝土壤不同粒级团聚体分布特征及其抗蚀性研究

为了定量评价人工混凝土壤生态护坡基材的抗蚀性,室内测定了其团聚体分布特征和各抗蚀性指标,并与立地条件相似的自然土壤进行了对照分析。结果表明:与自然土壤相比,人工混凝土壤中 > 0.25 mm机械稳定性团聚体含量、> 0.25 mm水稳性团聚体含量R_0.25、平均重量直径MWD、几何平均直径GWD与有机质含量均显著提升,可蚀性因子K、结构破坏率PAD、分形维数D与分散率均显著降低;同时,抗蚀性主成分值提高了2.53倍。这均证实人工混凝土壤团聚状况和团聚度有所改善,抗蚀性较自然土壤有较大的提升。相关性分析表明 > 0.25 mm水稳性大团聚体含量R_0.25和有机质含量与其他抗蚀性指标之间均存在极显著的相关性。综合分析可知,由于添加了水泥和天然有机物料,人工混凝土壤的水稳性大团聚体含量和有机质含量明显提升,因此导致其稳定性与抗蚀性显著优于原材料之一的自然土壤。     

The formation of water-stable coprolite aggregates in soddy-podzolic soils and the participation of fungi in this process

The water-stability of soil and coprolite aggregates in soddy-podzolic soils and the participation of fungi in the formation of water-stable aggregates from earthworm (Aporrectodea caliginosa) coprolites were assessed. The water stability of the soil and coprolite aggregates in the soils increased in the following sequence: potato field—mown meadow—mixed forest. The fungal mycelium reserves increased in the same sequence. The water stability of the coprolite aggregates of Aporrectodea caliginosa inhabiting these soils is 2–2.5 times higher than that of the soil aggregates of the same size (3–5 mm). The inhibition of the growth of fungi by cycloheximide decreased the water stability of the coprolite aggregates, on the average, by 15–20%.     

石灰岩区土壤分形特征及其与土壤性质的关系

研究了岩溶坡地不同生态系统土壤颗粒组成和团粒结构的分形特征.结果表明,土壤颗粒组成分形维数与黏粒及物理性黏粒含量显著正相关,与砂砾含量显著负相关.团粒结构分形维数与水稳性团聚体含量显著负相关,与团聚体湿筛后的破坏率显著正相关,即分形维数愈高,>0.25 mm水稳性团聚体和水稳性大团聚体含量愈低;团粒结构的分形维数与土壤有机质有负相关趋势,与土壤阳离子交换量显著负相关,与土壤体积质量(容重)呈正相关趋势.次生灌丛岩溶生态系统退化后,土壤黏粒减少,体积质量上升,土壤水稳性团聚体含量及其稳定性下降,土壤颗粒组成分形维数降低,土壤团粒结构分形维数则呈上升趋势.颗粒组成分形维数与团粒结构分形维数对土壤质量和岩溶生态环境状况的反映是一致的.     

Using lipid analysis and hyphal length to quantify AM and saprotrophic fungal abundance along a soil chronosequence

We evaluate the use of signature fatty acids and direct hyphal counts as tools to detect and quantify arbuscular mycorrhizal (AM) and saprotrophic fungal (SF) biomass in three Hawaiian soils along a natural soil fertility gradient. Phospholipids16:1ω5c and 18:2ω6,9c were used as an index of AM and saprotrophic fungal biomass, respectively. Both phospholipid analysis and hyphal length indicated that the biomass of AMF was greatest at the highest fertility site, and lowest where phosphorus limits plant growth. Saprotrophic fungal biomass did not vary. Hyphal length counts appeared to under-estimate SF abundance, while the phospholipid AMF:SF ratio was in line with expectations. This study indicates that phospholipids may be a valuable and reliable tool for studying the abundance, distribution, and interactions between AM and saprotrophic fungi in soil.     

Relationships between soil aggregation and mycorrhizae as influenced by soil biota and nitrogen nutrition

 The effect of the form of N nutrition on soil stability is an important consideration for the management of sustainable agricultural systems. We grew soybean [Glycine max (L.) Merr.] plants in pot cultures in unsterilized soil, and treated them by (1) inoculating them with Bradyrhizobium japonicum, fertilizing with (2) nitrate or (3) ammonia, or (4) by providing only minimum N amendment for the controls. The soils were sampled at 3-week intervals to determine changes in water-stable soil aggregates (WSA), soil pH, the development of roots, arbuscular mycorrhizal (AM) soil and root colonization, and selected functional groups of soil bacteria. The soil fauna was assayed at the end of the experiment (9 weeks). WSA was correlated positively with root and AM soil mycelium development, but negatively with total bacterial counts. Soil arthropod (Collembola) numbers were negatively correlated with AM hyphal length. Soils of nodulated and ammonia-fertilized plants had the highest levels of WSA and the lowest pH at week 9. Sparse root development in the soils of the N-deficient, control plants indicated that WSA formation was primarily influenced by AM hyphae. The ratio of bacterial counts in the water-stable versus water-unstable soil fractions increased for the first 6 weeks and then declined, while counts of anaerobic bacteria increased with increasing WSA. The numbers of soil invertebrates (nematodes) and protozoans did not correlate with bacterial counts or AM soil-hyphal lengths. Soil pH did not affect mycorrhiza development, but actinomycete counts declined with decreasing soil pH. AM fungi and roots interacted as the factors that affect soil aggregation, regardless of N nutrition. Received: 20 December 1997     

Mineralogy of the clay fraction of water-stable aggregates from dark gray forest soil

Regular structural changes of mixed-layered illite-smectite phases depending on aggregate sizes were revealed by x-ray powder diffraction analysis in the clay fraction of water-stable aggregates of dark gray forest soil. They indicate the differences in the aggregation mechanisms at different depths of the soil profile, which are mutually related and restrict the growth of the water-stable aggregates in the lower part of the A1 horizon to a certain optimal size, because only the smectite surface (unlike the mica one) of the process-determining organomineral composites is capable of interacting. This ensures the kinetic stabilization of the soil as a system and its self-regulation ability.     

Improved soil structure and citrus growth after inoculation with three arbuscular mycorrhizal fungi under drought stress

In a controlled potted experiment, citrus (Poncirus trifoliata) seedlings were inoculated with three species of arbuscular mycorrhizal (AM) fungi, Glomus mosseae, G. versiforme or G. diaphanum. Two soil-water levels (ample water, −0.10 MPa; drought stress, −0.44 MPa) were applied to the pots 4 months after transplantation. Eighty days after water treatments, the soils and the citrus seedlings were well colonized by the three AM fungi. Mycorrhizal fungus inoculation improved plant biomass regardless of soil-water status but decreased the concentrations of hot water-extractable and hydrolyzable carbohydrates of soils. Mycorrhizal soils exhibited higher Bradford-reactive soil protein concentrations than non-mycorrhizal soils. Mycorrhizas enhanced >2 mm, 1–2 mm and >0.25 mm water-stable aggregate fractions but reduced 0.25–0.5 mm water-stable aggregates. Peroxidase activity was higher in AM than in non-AM soils whether drought stressed or not, whereas catalase activity was lower in AM than non-AM soils. Drought stress and AM fungus inoculation did not affect polyphenol oxidase activity of soils. A positive correlation between the Bradford-reactive soil protein concentrations, soil hyphal length densities, and water-stable aggregates (only >2 mm, 1–2 mm and >0.25 mm) suggests beneficial effects of the AM symbiosis on soil structure. It concluded that AM fungus colonization enhanced plant growth under drought stress indirectly through affecting the soil moisture retention via glomalin's effect on soil water-stable aggregates, although direct mineral nutritional effects could not be excluded.     

种植果树对土壤物理性状的双重效应

在渭北果区选择不同园龄(<10 年、10~20 年、>20 年)果园, 分层测定0~60 cm 土层土壤容重、土壤坚实度、土壤含水量以及表层土壤团聚体组成等物理性状, 进一步分析了果园土壤物理性状随园龄的变化特征。结果表明: 土壤容重在0~30 cm 土层随园龄增长而降低; 在30 cm 以下土层随园龄增长而增加, 超过了健康园艺土壤的质量标准1.30 g·cm^-3; 与休闲农田相比, 种植果树可降低10~30 cm 土层土壤容重; 但30 cm 以下土层土壤坚实度急剧增大, 接近或达到了限制根系延伸的土壤质量标准1 000 kPa; 与休闲农田相比, 种植果树对于降低17.5~27.5 cm 土层的坚实度具有明显作用。果园表层土壤团聚体状况整体较差, 水稳性优势团聚体直径为0.5~0.25 mm, >0.25 mm 水稳性团聚体含量随园龄增加而增大, >20 年果园比<10 年果园高1 倍。种植果树对表层土壤具有明显的保护和改善作用, 却在深层发生着紧实化和坚硬化过程。果树对土壤物理状况的双重效应体现在对0~30 cm 土层土壤结构具有改善作用, 对30 cm 以下土层土壤结构有破坏作用。果园土壤“深层的隐蔽性退化过程”影响着果树根系健康生长, 应当给予极大关注。     

Role of active aluminum in the formation of water-stable macroaggregates

Soil structure is determined by the arrangement of particles in soil and the particles of sand, silt, and clay bind together into aggregates of various sizes by organic and inorganic materials. Structural stability which is the ability of the aggregates and pores to remain intact when subjected to stress, markedly affects crop production and soil erosion (Tisdall 1996). Since water, either directly as rainfall or as surface runoff is the main agent of aggregate breakdown, in the analyzes of stable soil aggregation, the term water-stable aggregation is generally used (Lynch and Bragg 1985). Water-stable aggregates have been divided into micro aggregates < 0.25 mm dia.) and macro aggregates (> 0.25 mm dia.) (Edwards and Bremner 1967; Tisdall and Oades 1982). Microaggregates show a relatively high stability against physical disruption (Edwards and Bremner 1967). On the other hand, macro aggregates are sensitive to soil management (Tisdall and Oades 1982).

There are many reports on the relationships between the aggregate stability and the soil physicochemical properties. For example, significant correlations were found between the aggregate stability and the amounts of organic C (Tisdall and Oades 1982), total N, and carbohydrates or the CEC (Chaney and Swift 1984). However, most of these studies were conducted in non-volcanic ash soils. Volcanic ash soils are widely distributed in Japan and are very important soils for crop production. The objective of this study was, therefore, to obtain more information on the relationship between the degree of macro aggregation and the soil physicochemical properties in non-volcanic and volcanic ash soils.     

Arbuscular mycorrhizal fungi and collembola non-additively increase soil aggregation

Soil aggregation is a principal ecosystem process mediated by soil biota. Collembola and arbuscular mycorrhizal (AM) fungi are important groups in the soil, and can interact in various ways. Few studies have examined collembola effects on soil aggregation, while many have quantified AM effects. Here, we asked if collembola have any effect on soil aggregation, and if they alter AM fungi-mediated effects on soil aggregation.We carried out a factorial greenhouse study, manipulating the presence of both collembola and AM fungi, using two different plant species, Sorghum vulgare and Daucus carota. We measured root length and biomass, AMF (and non-AMF) soil hyphal length, root colonization, and collembolan populations, and quantified water stable soil aggregates (WSA) in four size classes.Soil exposed to growth of AMF hyphae and collembola individually had higher WSA than control treatments. Moreover, the interaction effects between AMF and collembola were significant, with non-additive increases in the combined application compared to the single treatments.Our findings show that collembola can play a crucial role in maintaining ecological sustainability through promoting soil aggregation, and point to the importance of considering organism interactions in understanding formation of soil structure.     

Contributions of biotic and abiotic factors to soil aggregation across a land use gradient

Soil aggregation is a major ecosystem process that can be impacted by intensified land use directly through soil disturbances, or indirectly through impacts on biotic and abiotic factors that affect soil aggregation. We collected soils from 27 grassland sites across a range of land use intensities including varying levels of mowing, grazing, and fertilization in order to test the importance of selected direct and indirect effects on soil aggregation. We measured root length and mass, root colonization by arbuscular mycorrhizal fungi (AMF), extraradical AMF hyphal length, soil aggregation, and soil hydrophobicity. We also quantified levels of phosphorus, nitrogen, organic carbon, carbonate carbon, and sand in the soil.As land use intensity (defined as a multivariate index combining mowing, grazing, and fertilization intensities) increased, root mass decreased and length of extraradical hyphae increased. Total colonization by AMF was unaffected by land use intensity, but vesicular colonization tended to increase while arbuscular colonization declined. Soil aggregation increased with increasing land use intensity. We used structural equation models to explore mechanisms of soil aggregation and found that extraradical AMF hyphal length contributed to soil aggregation in models containing only biotic explanatory factors. When we also included abiotic factors in the model, no biotic factor was significant, and soil aggregation decreased as levels of sand and carbonate increased, likely due to concurrent decreases in levels of clay in the soil.In summary, we have shown that agricultural measures such as mowing, grazing, and fertilization can increase soil aggregation in managed grasslands. Furthermore, abiotic factors can be more important for determining soil aggregation than biotic factors, especially in highly aggregated soils. Aggregate turnover may be reduced in such highly aggregated soils past the point required to ensure efficient integration of new labile C into stable aggregates.     

铵、钾同时存在时, 土壤对铵的优先吸附

The water stability of aggregates in various size classes separated from 18 samples of red soils under different managements, and the mechanisms responsible for the formation of water-stable soil aggregates were studied. The results showed that the water stability of soil aggregates declined with increasing size, especially for the low organic matter soils. Organic matter plays a key role in the formation of water-stable soil aggregates. The larger the soil aggregate size, the greater the impact of organic matter on the water stability of soil aggregates. Removal of organic matter markedly disintegrated the large water-stable aggregates (> 2.0 mm) and increased the small ones (< 0.25-0.5mm) to some extent, whereas removal of free iron(aluminium) oxides considerably destroyed aggregates of all sizes, especially the < 0.25-0.5 mm classes. The contents of organic matter in water-stable aggregates increased with aggregate sizes. It is concluded from this study that small water-stable aggregates (< 0.25-0.5 mm) were chiefly cemented by Fe and Al oxides whilst the large ones (> 2.0 mm) were mainly glued up by organic matter. Both free oxides and organic matter contribute to the formation and water stability of aggregates in red soils.