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Understanding the diversity and community structure of arbuscular mycorrhizal fungi (AMF) is important for potentially optimizing their role in mining phosphorus (P) in agricultural ecosystems. Here, we conduct a comprehensive study to investigate the vertical distribution of AMF in a calcareous field and their temporal structure in maize-roots with fertilizer P application over a three-year period. The results showed that soil available-P response to P fertilization but maize yields did not. Phosphorus fertilization had no-significant effect on richness of AMF except at greater soil-depths. High P-supply reduced root colonization while optimum-P tended to increase colonization and fungal richness on all sampling occasions. Crop phenology might override P-supply in determining the community composition of active root inhabiting fungi. Significant differences in the community structure of soil AMF were observed between the controls and P treatments in surface soil and the community shift was attributable mainly to available-P, N/P and pH. Vertical distribution was related mainly to soil electrical conductivity and Na content. Our results indicate that the structure of AMF community assemblages is correlated with P fertilization, soil depth and crop phenology. Importantly, phosphorus management must be integrated with other agricultural-practices to ensure the sustainability of agricultural production in salinized soils.

Background: Phosphorus (P), iron (Fe) and zinc (Zn) are essential elements for plant growth and development, but their availability in soil is often limited. Intercropping contributes to increased P, Fe and Zn uptake and thereby increases yield and improves grain nutritional quality and ultimately human health. A better understanding of how intercropping leads to increased plant P, Fe and Zn availability will help to improve P-fertilizer-use efficiency and agronomic Fe and Zn biofortification., Scope: This review synthesizes the literature on how intercropping of legumes with cereals increases acquisition of P, Fe and Zn from soil and recapitulates what is known about root-to-shoot nutrient translocation, plant-internal nutrient remobilization and allocation to grains., Conclusions: Direct interspecific facilitation in intercropping involves below-ground processes in which cereals increase Fe and Zn bioavailability while companion legumes benefit. This has been demonstrated and verified using isotopic nutrient tracing and molecular analysis. The same methodological approaches and field studies should be used to explore direct interspecific P facilitation. Both niche complementarity and interspecific facilitation contribute to increased P acquisition in intercropping. Niche complementarity may also contribute to increased Fe and Zn acquisition, an aspect poorly understood. Interspecific mobilization and uptake facilitation of sparingly soluble P, Fe and Zn from soil, however, are not the only determinants of the concentrations of P, Fe and Zn in grains. Grain yield and nutrient translocation from roots to shoots further influence the concentrations of these nutrients in grains., (© The Author 2016. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

The mechanistic understanding of the dynamic processes linking nutrient acquisition and biomass production of competing individuals can be instructive in optimizing intercropping systems. Here, we examine the effect of inoculation with Funneliformis mosseae on competitive dynamics between wheat and faba bean. Wheat is less responsive to mycorrhizal inoculation. Both inoculated and uninoculated wheat attained the maximum instantaneous N and P capture approximately five days before it attained the maximum instantaneous biomass production, indicating that wheat detected the competitor and responded physiologically to resource limitation prior to the biomass response. By contrast, the instantaneous N and P capture by uninoculated faba bean remained low throughout the growth period, and plant growth was not significantly affected by competing wheat. However, inoculation substantially enhanced biomass production and N and P acquisition of faba bean. The exudation of citrate and malate acids and acid phosphatase activity were greater in mycorrhizal than in uninoculated faba bean, and rhizosphere pH tended to decrease. We conclude that under N and P limiting conditions, temporal separation of N and P acquisition by competing plant species and enhancement of complementary resource use in the presence of AMF might be attributable to the competitive co-existence of faba bean and wheat.

The effects of arbuscular mycorrhizal (AM) fungus (Glomus mosseae) and phosphorus (P) addition (100 mg/kg soil) on arsenic (As) uptake by maize plants (Zea mays L.) from an As-contaminated soil were examined in a glasshouse experiment. Non-mycorrhizal and zero-P addition controls were included. Plant biomass and concentrations and uptake of As, P, and other nutrients, AM colonization, root lengths, and hyphal length densities were determined. The results indicated that addition of P significantly inhibited root colonization and development of extraradical mycelium. Root length and dry weight both increased markedly with mycorrhizal colonization under the zero-P treatments, but shoot and root biomass of AM plants was depressed by P application. AM fungal inoculation decreased shoot As concentrations when no P was added, and shoot and root As concentrations of AM plants increased 2.6 and 1.4 times with P addition, respectively. Shoot and root uptake of P, Mn, Cu, and Zn increased, but shoot Fe uptake decreased by 44.6%, with inoculation, when P was added. P addition reduced shoot P, Fe, Mn, Cu, and Zn uptake of AM plants, but increased root Fe and Mn uptake of the nonmycorrhizal ones. AM colonization therefore appeared to enhance plant tolerance to As in low P soil, and have some potential for the phytostabilization of As-contaminated soil, however, P application may introduce additional environmental risk by increasing soil As mobility.

In a multifactorial pot experiment, maize (Zea mays L.) with or without inoculation with the arbuscular mycorrhizal (AM) fungus Glomus mosseae BEG167 was grown in a sterilized soil spiked with three levels of zinc (0, 300 and 900 mg Zn kg(-1) soil) and three levels of cadmium (0, 25 and 100 mg Cd kg(-1) soil). At harvest after 8 weeks of growth, the proportion of root length of inoculated plants colonized decreased with increasing Zn or Cd addition, and was 56% in the absence of both metals and was reduced significantly to 27% in the presence of the higher levels of both metals. Mycorrhizal plants had higher biomass than non-mycorrhizal controls except at the highest soil level of Cd. Cadmium had more pronounced effects on plant biomass than did Zn at the levels studied and the two metals showed a significant interaction. The data suggest that mycorrhizal inoculation increased plant growth with enhancement of P nutrition, perhaps increasing plant tolerance to Zn and Cd by a dilution effect. AM inoculation also led to higher soil solution pH after harvest, possibly reducing the availability of the metals for plant uptake, and lowered the concentrations of soluble Zn and Cd in the soil solution, perhaps by adsorption onto the extrametrical mycelium.

Foraging strategies in arbuscular mycorrhizal fungi (AMF) for heterogeneously distributed resources in the soil remain to be explored. We used nylon-mesh bags of 30 μm to simulate patches of different phosphorus (P) supply levels (Expt 1) and P forms (organic υ. inorganic, Expts 1 and 2). In Expt 1, host maize (Zea mays) was unfertilised; in each pot, five P-enriched bags were supplied with either Na-phytate or KH2PO4 at P rates of 0 (P0), 50 (P50), 100 (P100), 150 (P150) and 200 (P200) mg P kg-1. In Expt 2, maize plants were supplied with 20 (P20) or 50 (P50) mg P kg-1, and five P-enriched bags were supplied with different P forms (Na-phytate, lecithin, RNA, KH2PO4) and a nil-P control. Three fungal species (Funneliformis mosseae, Rhizophagus irregularis, and Glomus etunicatum) were compared in Expt 1, and the first two species in Expt 2. In Expt 1, the hyphal-length density (HLD) of G. etunicatum was not significantly different among different P levels when supplied with KH2PO4, whereas theHLDof R. irregularis tended to increase at higherPsupply (above P50) in the Na-phytate treatment. TheHLDof F. mosseae increased at P150 when supplied with KH2PO4, and increased at P100 and P150 in the Naphytate treatment relative to P0. APase activity levels were more related to P supply level, in particular with F. mosseae inoculation and uninoculated control, showing that P200 significantly reduced APase relative to P0. In Expt 2, greater hyphal growth of both fungal species tended to occur with KH2PO4. At P20, the HLD of R. irregularis in treatments with KH2PO4 and lecithin, and of F. mosseae with KH2PO4, were higher than in P0. At P50, the HLD of F. mosseae was higher than of R. irregularis; but P form had no significant influence on HLD of F. mosseae, whereas the HLD of R. irregularis in the P-amended treatment (except with Na-phytate) was higher than in P0. APase activity did not differ significantly between the two fungal species. Highest APase activity generally occurred with lecithin, with no significant difference among the other P forms. Our results indicate that the response of AMF to P-enriched patches is complex, and both the form and amount of P supplied should be considered. Variations between AMF in the proliferation of hyphae to heterogeneous nutrient patches might be a mechanism by which these species can maintain diversity in intensive agricultural ecosystems. [ABSTRACT FROM AUTHOR]

Background Phosphorus (P), iron (Fe) and zinc (Zn) are essential elements for plant growth and development, but their availability in soil is often limited. Intercropping contributes to increased P, Fe and Zn uptake and thereby increases yield and improves grain nutritional quality and ultimately human health. A better understanding of how intercropping leads to increased plant P, Fe and Zn availability will help to improve P-fertilizer-use efficiency and agronomic Fe and Zn biofortification. Scope This review synthesizes the literature on how intercropping of legumes with cereals increases acquisition of P, Fe and Zn from soil and recapitulates what is known about root-to-shoot nutrient translocation, plant-internal nutrient remobilization and allocation to grains. Conclusions Direct interspecific facilitation in intercropping involves below-ground processes in which cereals increase Fe and Zn bioavailability while companion legumes benefit. This has been demonstrated and verified using isotopic nutrient tracing and molecular analysis. The same methodological approaches and field studies should be used to explore direct interspecific P facilitation. Both niche complementarity and interspecific facilitation contribute to increased P acquisition in intercropping. Niche complementarity may also contribute to increased Fe and Zn acquisition, an aspect poorly understood. Interspecific mobilization and uptake facilitation of sparingly soluble P, Fe and Zn from soil, however, are not the only determinants of the concentrations of P, Fe and Zn in grains. Grain yield and nutrient translocation from roots to shoots further influence the concentrations of these nutrients in grains. [ABSTRACT FROM AUTHOR]

A pot experiment was carried out to investigate the tolerance of cucumber plants (Cucumis sativus L.) to root-knot nematode after inoculation with Glomus intraradices. Plants were inoculated with G. intraradices for four weeks and then transplanted in soil treated with Meloidogyne incognita for a further five weeks. The low phosphorus (P) loamy soil was amended with 50 and 100 mg P kg-1 soil. Mycorrhizal colonization increased shoot dry weight, shoot length, leaf numbers, root fresh weight and shoot P concentration, whereas nematode penetration and reproduction were significantly decreased. Similarly, P fertilization usually increased shoot growth and significantly decreased the number of galls and the number of egg masses and eggs per g root. Our results indicate that inoculation with G. intraradices and P fertilizer confer tolerance of cucumber plants to M. incognita by enhancing plant growth and by suppressing reproduction and/or galling of nematodes during the early stages of plant growth. [ABSTRACT FROM AUTHOR]

A pot experiment investigated the response of two maize inbred lines with contrasting root morphology and phosphorus (P) efficiency to inoculation with Glomus mosseae or Glomus etunicatum compared with non-mycorrhizal controls. Soil phosphorus was supplied at rates of 10, 50, and 100 mg P kg -1 soil. Root length, specific root length, and specific phosphorus uptake of maize line 178 (P-efficient) were significantly higher than of line Hc (P-inefficient). Percentage of root length colonized showed the opposite trend regardless of soil P supply level. The two maize lines did not differ significantly in growth response to mycorrhizal colonization. Root colonization rate decreased with increasing soil phosphorus supply. The beneficial effect of the two AM fungi on plant growth and P uptake was greatest at low soil P level and the responses were negative at high P supply. Mycorrhizal responsiveness also decreased with increasing P supply and differed between the two mycorrhizal fungal isolates. [ABSTRACT FROM AUTHOR]

Three ectomycorrhizal fungal isolates, Cenococcum geophilum SIV, a putative isolate of Pisolithus microcarpus and a Pisolithus sp., were cultured in vitro without potassium (K) and with three levels of K supply (as K2SO4). Some growth stimulation by K was observed among the fungi. Potassium in the liquid medium led to increased concentrations of nitrogen (N) and K in the fungi compared to K-free controls, and to higher phosphorus (P) concentrations in Pisolithus sp., but lower P in C. geophilum SIV. In a second experiment, the K influx rates of the fungi from external solutions followed Michaelis-Menten kinetics, indicating that K influx was an active process. Mycorrhizal fungi showed quite small Cmin values for K influx, ranging from 2.12 to 5.39 µmol L-1, in relation to K concentrations in the external solution. Such low Cmin values might allow the external hyphae to extract K efficiently from K-deficient soils. Furthermore, the fungi had low Km values for K uptake (<14.85 µmol L-1), indicating high affinities and efficiencies of K carriers. Both Pisolithus sp. and P. microcarpus exhibited similar Cmin and Km values but were different from C. geophilum SIV. Furthermore, Imax values varied significantly among the fungi, ranging from 9.60 to 21.63 µmol g-1 DM h-1, perhaps due to variation in the intracellular K concentrations in the ectomycorrhizal fungi. [ABSTRACT FROM AUTHOR]

Reports on a study of the anaerobic phosphate flux and mineralization of lake sediment organic carbon over a period of 182 days. Importance of lake sediment porewater composition and redox status that control material exchange at the sediment-water interface; Indication that under anaerobic conditions, organic-rich lake sediment can be an important source of phosphorus and methane.

Three wheat (Triticum aestivum L.) genotypes with high, intermediate and low phosphorus (P) efficiency were grown in a pot experiment with low P supply and adequate P supply either inoculated with the arbuscular mycorrhizal fungus Glomus versiforme or uninoculated. The mycorrhizal dependency of the genotype with relatively high P efficiency was lower than that of the genotypes with lower P efficiencies. Linear correlation analysis revealed that mycorrhizal dependency was primarily controlled by P uptake efficiency. More carbohydrate was translocated to the roots of genotypes with low P efficiency than of those with high P efficiency. In a second pot experiment the same three genotypes were grown in low P soils. Higher hyphal length density arising from carbohydrate translocation led to more P uptake, and this may account for higher mycorrhizal dependency. [ABSTRACT FROM AUTHOR]

Red clover (Trifolium pratense L.) and Glomus versiforme (Karsten) Berch growing in rhizoboxes were employed in two glasshouse experiments to study the mobilization of sparingly soluble phosphates by arbuscular mycorrhizal fungal (AMF) mycelium. In one experiment, four inorganic sources of phosphate, CaHPO4.2H2O (Ca2-P), Ca8H2(PO4)6.5H2O (Ca8-P), Ca10 (PO4)6.F2 (Ca10-P) and AlPO4.nH2O (Al-P), were chemically synthesized, labelled with 32P in an atomic pile and applied to the hyphal compartments of the rhizoboxes. Shoot yield, 32P and total P uptake weremeasured in clover growing in the root compartments. A similar experiment was conducted simultaneously using the same phosphate sources unlabelled and clover mycorrhizal infection and soil pH were determined. Although AMF inoculation increased the P uptake and biomass of clover shoots, the contribution of AMF to shoot P uptake and biomass varied with phosphate source, and was greatest with Ca2-P andleast with Ca10-P. 32P measurements indicated that external hyphae could mobilize Ca2-P, Ca8-P and Al-P, but not Ca10-P. This indicates that AMF not only mobilize the same types of phosphates that plants mobilize under stress conditions of low P, but give increased contact with phosphatesin the soil compared with non-mycorrhizal root systems. [ABSTRACT FROM AUTHOR]

Abstract: Rational soil/rhizosphere-plant phosphorus (P) management strategies in intercropping systems are poorly understood. Three years of field experiments with different rates of P fertilizer (0, 40 and 80kgha−1) as main effects and maize (Zea mays L.) intercropping with oilseed rape (Brassica napus L.), turnip (Brassica campestris L.), faba bean (Vicia faba L.), chickpea (Cicer arietinum L.) and soybean (Glycine max L.) with the respective monocultures as subplot effects were carried out to study the influence of fertilizer P application on productivity, shoot P content, apparent recovery of fertilizer P and soil Olsen-P in intercrops and monocultures. Average total grain yields and shoot P contents of maize/turnip, maize/faba bean, maize/chickpea and maize/soybean intercropping increased by 30.7%, 24.8%, 24.4%, and 25.3% and by 44.6%, 30.7%, 39.1%, and 28.6%, respectively, compared with the weighted means of the corresponding monocultures, and were highest at 40kgPha−1. Moreover, the average apparent recovery of fertilizer P of the intercropping systems increased from 6.1% to 30.6% at 40kgPha−1 and from 4.8% to 14.5% at 80kgPha−1 compared with overall monoculture systems on average over three years. The results indicate that intercropping and a rational P application rate (e.g. 40kgPha−1) maintained maximum total grain production and shoot P content, P balance of inputs/outputs and soil Olsen-P at an appropriate level (21.3mgkg−1), and maximum apparent recovery of fertilizer P (30.6%) through exploitation of the biological potential for efficient acquisition of P and other resources by interspecific interactions toward a sustainable and productive agricultural system. [Copyright &y& Elsevier]

Highlights: [•] A ‘competition–recovery–overyielding’ process was found in dynamics of root length growth and distribution of intercropped maize. [•] The stronger the associated crop roots, the more depressed the intercropped maize root development and shoot biomass. [•] Root lengths in 0–1.0m at jointing and in 0–0.3m of soil profiles at grain-filling were response to P fertilization. [•] Dynamics of root length growth and distribution correlated well with corresponding shoot biomass accumulation of maize. [ABSTRACT FROM AUTHOR]

Arbuscular mycorrhizal (AM) fungi are very abundant in many agricultural soils and are important in increasing and maintaining the sustainability of agricultural systems. However, high soil fertility, especially high available soil phosphorus (P) status, often limits AM fungal development in intensively managed agricultural systems and results in the C-costs to the host plant outweighing any benefits from AM colonization. Some promising approaches can minimize negative effects on soil biota while providing the desired agricultural benefits (e.g. reduced soil tillage and organic amendments). However, the influence of these agricultural practices on the benefits of root mutualistic associations remains poorly understood. Here, we collected indigenous AM fungi from a long-term field experiment with different application rates of cow manure and conducted a pot experiment with different soil P levels. Using high throughput sequencing we demonstrate that increasing cow manure application changed the community composition of AM fungi and reduced community diversity in the field. Phosphorus addition in a bioassay experiment produced a mycorrhizal growth response (MGR) and the mycorrhizal P response (MPR) shifted from positive to negative, regardless of the origin of the inoculum. An increase in intraradical and extraradical AM fungal structures enhanced maize shoot biomass and P uptake under low (no P added) and moderate (20 mg P added kg−1 soil) P levels but the opposite occurred at a high P level (250 mg P added kg−1 soil). The MGR was higher in the treatment with higher community diversity at the moderate P level but there was no significant correlation at the other two P levels examined. We conclude that available soil P may regulate the abundance and diversity of AM fungi and their symbiotic function in intensive cropping systems. Our results indicate that managing the soil phosphorus level to favor a rich and diverse AM fungal community can enhance crop production and result in a more sustainable agricultural system. • Indigenous AM fungi showed great potential for promoting plant growth and P uptake.. • Increased fertility in the field soil changed the community composition of AM fungi and reduced community diversity. • A higher indigenous AM fungal diversity was more beneficial to plants at a moderate soil P level. [ABSTRACT FROM AUTHOR]