Maize varieties can strengthen positive plant-soil feedback through beneficial arbuscular mycorrhizal fungal mutualists.

Plant-soil feedback (PSF) describes the process whereby plant species modify the soil environment, which subsequently impacts the growth of the same or another plant species. Our aim was to explore PSF by two maize varieties (a landrace and a hybrid variety) and three arbuscular mycorrhizal fungi (AMF) species (Funneliformis mosseae, Claroideoglomus etunicatum, Gigaspora margarita, and the mixture). We carried out a pot experiment with a conditioning and a feedback phase to determine PSF with different species of AMF and with a non-mycorrhizal control. Sterilized soil was conditioned separately by each variety, with or without AMF; in the feedback phase, each soil community was used to grow each in its "home" soil and in the "away" soil. Plant performance was assessed as shoot biomass, phosphorus (P) concentration and P content, and fungal performance was assessed as mycorrhizal colonization and hyphal length density. Both maize varieties were differentially influenced by AMF in the conditioning phase. In the feedback phase, PSF was generally negative for non-mycorrhizal plants or when plants were colonized by G. margarita, whereas PSF was positive in the other three AMF treatments. When plants were grown on home soil, hyphal length density was larger than on away soil. We conclude that different maize varieties can strengthen positive plant-soil feedback for themselves through beneficial mutualists for themselves, but not across the maize varieties.

Intercropping enhances soil carbon and nitrogen.

Intercropping, the simultaneous cultivation of multiple crop species in a single field, increases aboveground productivity due to species complementarity. We hypothesized that intercrops may have greater belowground productivity than sole crops, and sequester more soil carbon over time due to greater input of root litter. Here, we demonstrate a divergence in soil organic carbon (C) and nitrogen (N) content over 7 years in a field experiment that compared rotational strip intercrop systems and ordinary crop rotations. Soil organic C content in the top 20 cm was 4% ± 1% greater in intercrops than in sole crops, indicating a difference in C sequestration rate between intercrop and sole crop systems of 184 ± 86 kg C ha(-1) yr(-1). Soil organic N content in the top 20 cm was 11% ± 1% greater in intercrops than in sole crops, indicating a difference in N sequestration rate between intercrop and sole crop systems of 45 ± 10 kg N ha(-1) yr(-1). Total root biomass in intercrops was on average 23% greater than the average root biomass in sole crops, providing a possible mechanism for the observed divergence in soil C sequestration between sole crop and intercrop systems. A lowering of the soil δ(15) N signature suggested that increased biological N fixation and/or reduced gaseous N losses contributed to the increases in soil N in intercrop rotations with faba bean. Increases in soil N in wheat/maize intercrop pointed to contributions from a broader suite of mechanisms for N retention, e.g., complementary N uptake strategies of the intercropped plant species. Our results indicate that soil C sequestration potential of strip intercropping is similar in magnitude to that of currently recommended management practises to conserve organic matter in soil. Intercropping can contribute to multiple agroecosystem services by increased yield, better soil quality and soil C sequestration.

Multi-surface modeling to predict free zinc ion concentrations in low-zinc soils.

Multi-surface models are widely used to assess the potential ecotoxicological risk in metal-contaminated soils. Their accuracy in predicting metal speciation in soils with low metal levels was not yet tested. Now highly sensitive analytical techniques are available to experimentally validate such models at low concentration levels. The objective of this study was to test the accuracy of a multi-surface model to predict the Zn(2+) concentration and to improve our understanding of Zn bioavailability in low-Zn soils. High-Zn soils were included as controls. Model parameters were determined independently on the basis of earlier peer-reviewed publications. Model output was validated against free Zn(2+) concentrations determined with the soil column Donnan membrane technique in a range of soils varying in potentially available Zn, organic matter, clay silicate, and iron (hydr)oxide contents and pH. Deviations between predicted Zn(2+) concentrations and experimentally determined values over the whole Zn concentration range were less or equal to the experimental standard error, except for one low-Zn soil. The Zn(2+) concentration was mainly controlled by adsorption, where organic matter was predicted to be the dominant soil sorbent. The predicted Zn(2+) concentration depends more sensitively upon changes of the reactive Zn pool (application of 0.6, 1.2, 2.4, and 3.6 mg of Zn kg(-1) of soil) and organic matter content (± 0.2 and 0.4%) than pH changes (± 0.5 and 1 pH unit).

Organic inputs to reduce nitrogen export via leaching and runoff: A global meta-analysis.

Organic inputs as a substitution for, or addition to, chemical fertilizers can potentially mitigate N losses. However, it is not well known how their effects on N leaching and runoff depend on application approaches. We conducted a global meta-analysis of 129 studies to compare the effects of organic inputs on N export via leaching and runoff. We compared three application approaches: chemical fertilizer N substituted by organic fertilizer with: 1) equal amounts of total N or, 2) equal amounts of mineral N and 3) additional organic fertilizer N on top of chemical fertilizer. The meta-analysis showed that organic inputs reduced overall N leaching and runoff by 15% and 29%, respectively, without compromising crop yield, and that this effect was significantly influenced by the application approach taken. Organic substitution of chemical fertilizer N with equal amounts of total N decreased both leaching and runoff by more than 30% and hardly affected crop yield. Substitution with equal amounts of mineral N generally increased crop yield by 6% but also increased N leaching by 21%. Organic inputs in addition to chemical fertilizer N did not affect leaching and runoff. The differences between application approaches were reinforced with increased treatment duration. The loss ratios of leaching and runoff were 14% and 4.5%, respectively, from chemical fertilizer, and 9.2% and 2.6%, respectively, from organic fertilizer. The optimal substitution rates differed between leaching (40-60%) and runoff (60-100%) when substitution was based on equal amounts of total N. We conclude that substitution of chemical for organic fertilizer at equal amounts of total N is most effective in reducing N export via leaching and runoff without compromising crop production.

Syndromes of production in intercropping impact yield gains.

Intercropping, the simultaneous production of multiple crops on the same field, provides opportunities for the sustainable intensification of agriculture if it can provide a greater yield per unit land and fertilizer than sole crops. The worldwide absolute yield gain of intercropping as compared with sole crops has not been analysed. We therefore performed a global meta-analysis to quantify the effect of intercropping on the yield gain, exploring the effects of crop species combinations, temporal and spatial arrangements, and fertilizer input. We found that the absolute yield gains, compared with monocultures, were the greatest for mixtures of maize with short-grain cereals or legumes that had substantial temporal niche differentiation from maize, when grown with high nutrient inputs, and using multirow strips of each species. This approach, commonly practised in China, provided yield gains that were (in an absolute sense) about four times as large as those in another, low-input intercropping strategy, commonly practised outside China. The alternative intercropping strategy consisted of growing mixtures of short-stature crop species, often as full mixtures, with the same growing period and with low to moderate nutrient inputs. Both the low- and high-yield intercropping strategies saved 16-29% of the land and 19-36% of the fertilizer compared with monocultures grown under the same management as the intercrop. The two syndromes of production in intercropping uncovered by this meta-analysis show that intercropping offers opportunities for the sustainable intensification of both high- and low-input agriculture.

Malate exudation by six aerobic rice genotypes varying in zinc uptake efficiency.

Zinc (Zn) uptake by plant roots from soils low in plant-available Zn may be increased by Zn-mobilizing rhizosphere processes, including exudation of low-molecular-weight organic anions. A rhizotron experiment with a low Zn clay soil and a nutrient solution experiment were conducted to test if this occurs in six rice (Oryza sativa L.) genotypes varying in tolerance to low Zn supply. In both experiments, low Zn supply resulted in a marked decrease in biomass production of most genotypes compared with adequate Zn supply. The genotypes showed a significant variation in Zn efficiency. Plants responded to low Zn supply with increased root exudation of malate in both experiments. The malate concentration in the rhizosphere of three genotypes ranged from 0.22 to 0.59 mmol L(-1) in rhizotron experiment, and the malate exudation rate of five genotypes ranged from 0.18 to 0.53 nmol g(-1) root dw s(-1) in the nutrient solution experiment. On average, low Zn supply in the rhizotron experiment increased rhizosphere malate concentration by 64% compared with that at adequate Zn supply. The averaged malate exudation rate at low Zn in the nutrient solution experiment was 40% greater than at adequate Zn supply. The malate exudation of rice genotypes at low Zn was not correlated to Zn efficiency or Zn uptake in either experiment. Based on a soil malate extraction experiment, the observed genotypic difference in rhizosphere malate concentration is expected to have a negligible effect on the concentration of Zn in soil solution. These findings suggest that Zn mobilization by rice genotypes cannot be explained by increased malate exudation alone, indicating that other mechanisms contribute to the variation in plant Zn uptake. The results also emphasize that effects of root exudates on soil properties need to be assessed to evaluate the role of root exudation in nutrient mobilization.

Fertilizer response and nitrogen use efficiency in African smallholder maize farms.

Improving fertilizer recommendations for farmers is essential to increase food security in smallholder landscapes. Currently, blanket recommendations are provided across agro-ecological zones, although fertilizer response and nutrient use efficiency by maize crop are spatially variable. We aimed to identify factors that could help to refine fertilizer recommendation by analyzing the variability in fertilizer response (FR) and the agronomic nitrogen use efficiency (N-AE). A literature search for on-farm studies across Kenya and Sub-Sahara Africa (SSA), excluding Kenya, yielded 71 publications. The variability in FR was studied using a meta-analysis whereas key factors that influence FR and N-AE were studied with linear regression models. On average, the FR was 2, but it varied considerably from 1 to 28.5 (excluding outliers). In SSA, 18% of the plots were non-responsive plots with an FR < 1. The main factors affecting N-AE for Kenya were P-Olsen, silt content, soil pH, clay and rainfall, whereas only soil pH, exchangeable K and texture were important for SSA. However, our study indicates that available data on soil, climate and management factors could explain only a small part (< 33%) of the variation in FR and N-AE. Soil pH, P-Olsen, silt content, and rainfall had significant but low levels of power in explaining variation in FR and N-AE. Our findings indicate that strategies to refine fertilizer recommendation should include information on soil types and soil properties.

Phosphate Uptake from Phytate Due to Hyphae-Mediated Phytase Activity by Arbuscular Mycorrhizal Maize.

Phytate is the most abundant form of soil organic phosphorus (P). Increased P nutrition of arbuscular mycorrhizal plants derived from phytate has been repeatedly reported. Earlier studies assessed acid phosphatase rather than phytase as an indication of mycorrhizal fungi-mediated phytate use. We investigated the effect of mycorrhizal hyphae-mediated phytase activity on P uptake by maize. Two maize (Zea mays L.) cultivars, non-inoculated or inoculated with the arbuscular mycorrhizal fungi Funneliformis mosseae or Claroideoglomus etunicatum, were grown for 45 days in two-compartment rhizoboxes, containing a root compartment and a hyphal compartment. The soil in the hyphal compartment was supplemented with 20, 100, and 200 mg P kg-1 soil as calcium phytate. We measured activity of phytase and acid phosphatase in the hyphal compartment, hyphal length density, P uptake, and plant biomass. Our results showed: (1) phytate addition increased phytase and acid phosphatase activity, and resulted in larger P uptake and plant biomass; (2) increases in P uptake and biomass were correlated with phytase activity but not with acid phosphatase activity; (3) lower phytate addition rate increased, but higher addition rate decreased hyphal length density. We conclude that P from phytate can be taken up by arbuscular mycorrhizal plants and that phytase plays a more important role in mineralizing phytate than acid phosphatase.

Ectomycorrhizal weathering of the soil minerals muscovite and hornblende.

Ectomycorrhizal fungi are hypothesized to enhance mineral weathering in forest soils. Several studies have shown an increased uptake of mineral-derived nutrients by trees when in symbiosis with ectomycorrhizal fungi. However, it is difficult to determine from these studies if the improved nutrient uptake is the result of increased weathering or better exploitation of the substrate by the ectomycorrhizal fungi. In a pot experiment, Pinus sylvestris (Scots pine) seedlings were grown with or without ectomycorrhizal fungi, and with or without the mineral muscovite as the only potassium (K) source or the mineral hornblende as the only magnesium (Mg) source. After 27 wk, all pools of non-mineral-bound K or Mg were determined. The ectomycorrhizal fungus Paxillus involutus increased weathering of muscovite but not hornblende. The other ectomycorrhizal fungi tested, Piloderma croceum and Suillus bovinus, did not increase weathering of either muscovite or hornblende compared with the nonmycorrhizal trees. The P. involutus-mediated mobilization of K from muscovite resulted in increased K content of root plus adhering hyphae, but not of shoots. In conclusion, ectomycorrhizal fungi may increase weathering of minerals in response to nutrient deficiencies, but this response is species specific.

Organic anion exudation by ectomycorrhizal fungi and Pinus sylvestris in response to nutrient deficiencies.

Low molecular weight organic anions (LMWOA) can enhance weathering of mineral grains. We tested the hypothesis that ectomycorrhizal (EcM) fungi and tree seedlings increase their exudation of LMWOA when supply of magnesium, potassium and phosphorus is low to enhance the mobilization of Mg, K and P from mineral grains. Ectomycorrhizal fungi and Pinus sylvestris seedlings were cultured in symbiosis and in isolation on glass beads with nutrient solution or with sand as a rooting medium, with a complete nutrient supply or with Mg, K, P or N in low supply. Concentrations of all dicarboxylic LMWOA in the rooting medium were measured. Nonmycorrhizal seedlings released predominantly malonate. Colonization with Hebeloma longicaudum decreased the amount of organic anions exuded, whereas Paxillus involutus and Piloderma croceum increased the concentration of oxalate but not the total amount of LMWOA. Phosphorus deficiency increased the concentration of LMWOA by nonmycorrhizal and EcM seedlings. Magnesium deficiency increased the concentration of oxalate by nonmycorrhizal and EcM seedlings, but not the concentration of total LMWOA. Paxillus involutus grown in pure culture responded differently to low nutrient supply compared with symbiotic growth. Ectomycorrhizal fungi did not increase the total concentration of LMWOA compared with nonmycorrhizal seedlings but, depending on the fungal species, they affected the type of LMWOA found.

Molecular identification of ectomycorrhizal mycelium in soil horizons.

Molecular identification techniques based on total DNA extraction provide a unique tool for identification of mycelium in soil. Using molecular identification techniques, the ectomycorrhizal (EM) fungal community under coniferous vegetation was analyzed. Soil samples were taken at different depths from four horizons of a podzol profile. A basidiomycete-specific primer pair (ITS1F-ITS4B) was used to amplify fungal internal transcribed spacer (ITS) sequences from total DNA extracts of the soil horizons. Amplified basidiomycete DNA was cloned and sequenced, and a selection of the obtained clones was analyzed phylogenetically. Based on sequence similarity, the fungal clone sequences were sorted into 25 different fungal groups, or operational taxonomic units (OTUs). Out of 25 basidiomycete OTUs, 7 OTUs showed high nucleotide homology (> or = 99%) with known EM fungal sequences and 16 were found exclusively in the mineral soil. The taxonomic positions of six OTUs remained unclear. OTU sequences were compared to sequences from morphotyped EM root tips collected from the same sites. Of the 25 OTUs, 10 OTUs had > or = 98% sequence similarity with these EM root tip sequences. The present study demonstrates the use of molecular techniques to identify EM hyphae in various soil types. This approach differs from the conventional method of EM root tip identification and provides a novel approach to examine EM fungal communities in soil.