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Aim: Cereal/legume intercropping often increases yield, partly because of increased nitrogen (N) and phosphorus (P) acquisition. The aim of this paper was to investigate the role of arbuscular mycorrhizal (AM) fungal common mycorrhizal networks (CMNs) in overyielding by the millet (Setaria italica L.) and chickpea (Cicer arietinum L.) mixture and to find out if the effect of a CMN depends on which of the two species was first colonized by AM fungi (AMF). Methods: Microcosms with two compartments were used, separated by a 30-μm nylon mesh. Both compartments contained either chickpea or millet, in monoculture or mixed. One or none of the two compartments was inoculated with the AMF species Funneliformis mosseae. The plant in the inoculated compartment was referred to as the donor, and the plant in the neighboring, non-inoculated compartment as the receiver. Results: Inoculation in one compartment resulted in mycorrhiza formation in the other compartment, providing evidence for the formation of CMNs. Inoculation of chickpea in the mixture increased N and P acquisition and biomass of both chickpea (donor) and millet (receiver) leading to overyielding of the mixture, whereas inoculation of millet increased biomass of chickpea (receiver) only, but did not increase N or P acquisition by any of the two species, and there was no overyielding. Chickpea as donor had higher numbers of phosphate-solubilizing bacteria in its rhizosphere compared to chickpea as receiver. The shoot N:P ratio of chickpea as donor was lower than as receiver. Conclusion: Our study demonstrated asymmetry in nutrient gains by a mixture of a cereal and a legume, dependent on which plant species was the donor or receiver. This suggests that initiating mycorrhizal networks by legumes in intercropping could be an important factor contributing to the magnitude of the intercropping effect.

Plant species have different traits for mobilizing sparingly soluble phosphorus (P) resources, which could potentially lead to overyielding in P uptake by plant species mixtures compared to monocultures due to higher P uptake as a result of resource (P) partitioning and facilitation. However, there is circumstantial evidence at best for overyielding as a result of these mechanisms. Overyielding (the outcome) is easily confused with underlying mechanisms because of unclear definitions. We aimed to define a conceptual framework to separate outcome from underlying mechanisms and test it for facilitation and complementarity with respect to P acquisition by three plant species combinations grown on four soils. Our conceptual framework describes both mechanisms of complementarity and facilitation and outcomes (overyielding of mixtures or no overyielding) depending on the competitive ability of the species to uptake the mobilized P. Millet/chickpea mixtures were grown in pots on two calcareous soils mixed with calcium-bound P (CaP) and phytate P (PhyP). Cabbage/faba bean mixtures were grown on both acid and neutral soils mixed with P-coated iron (hydr)oxide (FeP) and PhyP. Wheat/maize mixtures were grown on all four soils. Rhizosphere carboxylate concentration and acid phosphatase activity (mechanisms) as well as plant P uptake and biomass (outcome) were determined for monocultures rhizosphere and species mixtures. Facilitation of P uptake occurred in millet/chickpea mixtures on one calcareous soil. We found no indications for P acquisition from different P sources, neither in millet/chickpea, nor in cabbage/faba bean mixtures. Cabbage and faba bean on the neutral soil differed in rhizosphere acid phosphatase activity and carboxylate concentration, but showed no overyielding. Wheat and maize, with similar root exudates, showed overyielding (the observed P uptake being 22% higher than the expected P uptake) on one calcareous soil. We concluded that although differences in plant physiological traits (root exudates) provide necessary conditions for complementarity and facilitation with respect to P uptake from different P sources, they do not necessarily result in increased P uptake by species mixtures, because of the relative competitive ability of the mixed species.

Aims: The benefits of arbuscular mycorrhizal fungi (AMF) on yield and phosphorus (P) uptake of crops have commonly been studied by inoculating a single mycorrhizal fungal species in pot experiments. Yet, how the native AMF community affects the performance of different maize varieties under field conditions remains obscure. Methods: In-growth cores with and without rotation were used in three soils that differed in P application to assess shoot biomass, P uptake, and mycorrhizal colonization of three maize varietal groups, encompassing four inbred lines, 12 hybrids, and four landraces. Results: Rotating cores drastically reduced mycorrhizal colonization, biomass and P uptake for each varietal group at every P level. Performance of plants at natural mycorrhizal colonization at 30 kg P ha−1 was equal to that of reduced-mycorrhizal plants at 60 kg P ha−1, suggesting the potential for adequate mycorrhizal management to save P fertilizer. Conclusion: There were no significant differences between varietal groups for mycorrhizal responsiveness, confirming that the ability to associate with and benefit from AMF has been maintained in modern breeding. Mycorrhizal plants both exhibited higher P acquisition efficiency and higher P use efficiency than reduced-mycorrhizal plants. Disadvantages of in-growth cores should be duly considered.

Ecological intensification seeks to achieve crop yield increases by intensifying complementary or facilitative interactions between plant species or varieties. Different species of arbuscular mycorrhizal fungi (AMF) exhibit niche differentiation and show selectivity towards certain plants, which can further enhance complementarity. It is not clear whether in the presence of one AMF species, where mycelial networks connect crop species, opportunities for complementarity effects may be reduced. We grew monocultures and mixtures of maize varieties in a greenhouse with one species of AMF, Funneliformis mosseae, during two consecutive years to investigate whether under such conditions the mycorrhizal symbiosis would affect complementarity and overyielding compared to non-mycorrhizal plants. Variety mixtures showed increased phosphatase activity and mycorrhizal colonization, enhanced phosphorus uptake and overyielding when plants were mycorrhizal. There was no overyielding when plants were non-mycorrhizal. The increase in relative yield total was due to complementarity effects. Synthesis and applications. Our study implies that appropriate agricultural management that enhances mycorrhizal fungal contribution to ecosystem services may result in overyielding in terms of yield or phosphorus uptake through mixing varieties within one crop species.

Aim Cereal/legume intercropping is known to increase yield, partly because of increased nitrogen (N) and phosphorus (P) acquisition. The aim of this paper was to investigate the role of common mycorrhizal networks (CMNs) in overyielding by the crop species mixture and to find out if the effect of a CMN depends on which of the two species was colonized by AM fungi.Methods Microcosms with two compartments were used, separated by a 30-μm nylon mesh. Both compartments contained either chickpea or millet, in monoculture or mixed. One or none of the two compartments was inoculated with the AMF species Funneliformis mosseae. The plant in the inoculated compartment was referred to as the AMF donor, and the plant in the neighboring, non-inoculated compartment as the AMF receiver. Results Inoculation in one compartment resulted in mycorrhiza formation in the other compartment, providing evidence for the formation of CMNs. Inoculation of chickpea in the mixture increased N and P acquisition and biomass of both chickpea (AMF donor) and millet (AMF receiver), whereas inoculation of millet increased biomass of chickpea (AMF receiver) only, but did not increase N or P acquisition by any of the two species. Chickpea as AMF donor had higher numbers of phosphate-solubilizing bacteria in its rhizosphere compared to chickpea as receiver. The shoot N:P ratio of chickpea as AMF donor was lower than as receiver. Conclusion Our study demonstrated asymmetry in nutrient gains by a mixture of cereal and a legume, dependent on which plant species was the AMF donor or receiver. This suggests that initiating mycorrhizal networks by legumes in intercropping could be an important factor contributing to the magnitude of the intercropping effect.

Aims: The phosphorus (P) resource partitioning hypothesis assumes that dissimilarity in P acquisition traits among plant species leads to enhanced P uptake by crop combinations compared with their sole crops. We developed and implemented a test for this hypothesis. Methods: Two pot experiments were conducted with quartz sands. In Experiment 1, the ability of the crop species to acquire P from sparingly soluble sources (Ca phosphate (CaP), phytate (PhyP) and P-coated Fe (hydr)oxide (FeP)) was tested. In accordance with the species performances in Experiment 1, combinations of millet/chickpea and cabbage/faba bean (which have dissimilar P acquisition traits) and wheat/maize (which have similar traits) were selected for Experiment 2. The biomass production and P uptake were compared between the sole crops and species combinations as well as between the single and mixed P sources. Results: A dissimilarity in P acquisition traits enhanced P uptake by millet/chickpea on CaP/PhyP (as expected) but not by cabbage/faba bean on FeP/PhyP. Despite their similar P acquisition traits, we found enhanced P uptake by wheat/maize on CaP/PhyP. Conclusions: Because of complicating factors such as unstable P acquisition traits and competitive inequality between species, the conditions under which the P resource partitioning hypothesis can be tested are limited. This challenge complicates designing for complementarity in soil P pools by intercrops.

There is increasing evidence that the soil microbial community produces a suite of volatile organic compounds that suppress plant pathogens. However, it remains unknown which soil properties and management practices influence volatile-mediated pathogen suppression. The aim of this study was to relate soil properties to growth suppression of three plant pathogens by soil volatiles. We measured the effect of volatiles emitted from a broad range of agricultural soils on the in vitro growth of the plant pathogenic fungi Rhizoctonia solani and Fusarium oxysporum, and the oomycete Pythium intermedium. Growth suppression of pathogens by soil volatiles could be linked to various soil properties, and some aspects of microbial community composition and field history, using multiple linear regression. Volatile-mediated suppression of mycelial development occurred for each pathogen type, but the magnitude of inhibition differed among soils as well as pathogens. On average R. solani and P. ultimum appeared more sensitive to volatile suppression than F. oxysporum. Suppression of R. solani by volatiles was positively correlated with organic matter content, microbial biomass and proportion of litter saprotrophs in the microbial community, but negatively correlated with pH, microbial diversity (Shannon), and the proportion of Acidobacteria in the community. R. solani, F. oxysporum, and P. intermedium suppression by volatiles was affected by various management practices occurring in the soil's field history, such as reduced tillage, the presence of certain crops in the crop rotation, and the application of solid manure. P. intermedium suppression was also negatively correlated with soil sulphur content. This study identifies pathogen-specific drivers of growth-suppressive volatiles, a critical step in integrating soil volatiles into prediction and management of soil-borne plant diseases.

Plant roots have a profound effect on soil microbial activity, particularly in the rhizosphere. Hence, it is important to understand the potential effects of genetically modified (GM) crops on soil microbial activity and related processes such as litter decomposition. In this study, we compared the effects of GM potato Modena on soil microbial activity and carbon (C) and nitrogen (N) mineralization to effects induced by Modena’s parental isoline (Karnico) and a conventional potato cultivar (Aventra). A field experiment was conducted at two sites to assess microbial catabolic diversity (using MicroRespTM) in the rhizosphere and in bulk soil, during flowering and senescence of the potato plants. In a laboratory experiment with soil and potato litter from the field experiment, we investigated whether the cultivars had modified the activity of soil microbial communities to such an extent that this affected C and N mineralization. Results of the field experiment showed no GM-induced effects on microbial catabolic diversity, while effects of field site location and sampling date were significant. Multivariate analysis including plant traits and soil characteristics revealed that microbial catabolic activities in rhizosphere soil were strongly correlated with soil organic matter and tuber sucrose content, whereas in bulk soil, they were primarily correlated with soil moisture. In the laboratory experiment, we found that Modena induced a "home-field advantage" in N mineralization, yet this effect was inconsistent across locations and was also observed for other cultivars. Based on our data and results from previous studies, we conclude that the effects of GM cultivar Modena on soil microbial activity and litter decomposition fall within the normal range of effects found for conventional potato cultivars.

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.

The aim of this study was to assess the feasibility of controlling the potato cyst nematode Globodera pallida through biofumigation with glucosinolate-rich Brassica juncea genotypes. The main glucosinolate of B. juncea is 2-propenyl glucosinolate which is the precursor of 2-propenyl isothiocyanate. Toxicity of 2-propenyl isothiocyanate to encysted G. pallida was tested in vitro . Fifty percent reduction in hatching was found within 2 h of exposure to 0.002% 2-propenyl isothiocyanate. Based on the in vitro results, we hypothesized that biofumigation with B. juncea would reduce hatching of G. pallida in vivo and higher 2-propenyl glucosinolate levels would have a stronger effect. Plant genotype, sulfur fertilization and insect herbivory affected 2-propenyl glucosinolate concentration of B. juncea . However, increasing 2-propenyl glucosinolate concentration of B. juncea did not affect G. pallida hatching after biofumigation. The absence of a biofumigation effect was most likely due to lower concentrations of 2-propenyl isothiocyanate in vivo compared to in vitro conditions. These results show that it is difficult to reach sufficiently high levels of toxicity to reduce hatching of G. pallida under realistic conditions.

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.

Zinc (Zn) deficiency is a widely occurring constraint for rice production and for human nutrition. Scarcity of water is leading to a shift from flooded to aerobic rice production, which can have an impact on Zn deficiency in rice. Zinc bioavailability is a function of both soil and plant factors that can be altered by water management, particularly in relation to conditions in the rhizosphere. Biogeochemical modeling based on bulk soil conditions failed to predict the effect of water management on Zn bioavailability, but revealed that dissolved organic anions, pH, and redox conditions were major determinants. Rhizosphere sampling is needed to understand the difference in Zn mobilization and uptake between flooded and aerobic cultivation systems. Zinc bioavailability is not only affected by changes in the chemical properties of the soils, but also by biological processes such as mycorrhizal inoculation and root release of organic compounds into rhizosphere. Phytosiderophores and organic acids are two classes of Zn chelators secreted from roots that have been linked to the release of Zn from soil-bound forms and its subsequent uptake by plants. A shift to aerobic condition provides a favorable environment for activity of mycorrhizal fungi and enhanced mycorrhizal inoculation under aerobic conditions has been shown to increase plant Zn uptake. Aerobic rice genotypes with varying tolerance to Zn deficiency display a trade-off between mycorrhizal Zn responsiveness and root exudation of Zn chelator in the rhizosphere, which is probably due to a competition for carbon. Potential agronomic management practices in aerobic rice production systems are discussed, with an emphasis on their roles in improving bioavailability of Zn. Addition of Zn fertilizers by soil or foliar application have been shown to increase Zn concentration in cereal grains but the extent of the increase differs among crop species. The shift from flooded to aerobic condition can cause significant N transformations, which may consequently affect Zn mobilization and uptake. An appropriate N management strategy, including an effective combination of source, rate, application method, and timing, should consider the effects on soil pH. Application of P fertilizer should be done with careful consideration to the effect on Zn uptake. A reasonable cropping system (intercropping and crop rotation) could prevent Zn deficiency and offer an effective and sustainable pathway to Zn biofortification. Keeping these points in mind, this review describes our current knowledge of Zn bioavailability as affected by changes in soil–plant interactions caused by the transition from flooded to aerobic rice cultivation.

The transition from anaerobic to aerobic rice (Oryza sativa L.) cultivation has been reported to decrease Zn bioavailability. To determine and understand the differences in plant Zn uptake between anaerobic and aerobic rice cultivation systems, a field plot experiment was conducted with direct-seeded rice grown on a low-Zn soil under both anaerobic and aerobic conditions. Results showed that the mass fraction of Zn in shoots and Zn uptake of plants in the aerobic field was significantly lower than in the anaerobic field. Visual Zn deficiency symptoms as whitish-brown necrotic patches on leaves were more severe on plants grown in aerobic soils than plants in anaerobic soils, indicating reduced Zn bioavailability. To determine whether the underlying causes could be of a geochemical nature, a controlled soil incubation experiment was done. Using a fully parameterized model, we found that the difference in plant Zn uptake between the two cultivation systems was small compared with that predicted by geochemical modeling, assuming Zn uptake is controlled by the soluble fraction in the soil. Whereas individual effects of soil factors (e.g., pH) were demonstrated to cause large variations in Zn concentration, only about a 15% difference in plant Zn uptake was observed. This large discrepancy reveals that the consideration of separate soil chemical processes, instead of their integrated assessment, may be inappropriate. We speculated that the discrepancy could also be associated with the chemical conditions in the rhizosphere, where the local acidity, dissolved organic C, and redox conditions may deviate significantly from the bulk soil on which the geochemical analysis was based

Previous studies have shown that soil fungal biomass increases towards more natural, mature systems. Shifts to a fungal-based soil food web have previously been observed with abandonment of agricultural fields and extensification of agriculture. In a previous field experiment we found increased fungal biomass with reduced N fertilisation. Here, we explore relationships between fungi, bacteria, N input and grassland age on real dairy farms in the Netherlands. We hypothesised that also in pastures that are still in production there is a negative relationship between fungal biomass and fertilisation, and that fungal biomass increases with grassland age in pastures that are still in production. We expected the fungal/bacterial biomass ratio to show the same responses, as this ratio has often been used as an indicator for management changes. We sampled 48 pastures from eight organic dairy farms. Sites differed in age and fertilisation rate. We determined fungal and bacterial biomass, as well as ergosterol (a fungal biomarker). Fungal and bacterial biomass and ergosterol, showed a negative relationship with N application rate, and correlated positively with organic matter percentage. In old pastures, fungal biomass and ergosterol were higher than in younger pastures. Because bacterial biomass responded in the same way as fungal biomass, the F/B ratio remained constant, and can therefore¿in our data set¿not be used as an indicator for changing management. We conclude that the changes in fungal and bacterial biomass were driven by changes in organic matter quality and quantity. The negative relationship we found between N application rate and fungal biomass adds to earlier work and confirms the presence of this relationship in pastures with relatively small differences in management intensities. Earlier studies on shifts in fungal biomass focused on ex-agricultural fields or restoration projects. Here we show that fungal biomass is also higher in older agricultural pastures.

Increasing plant species richness generally enhances plant biomass production, which may enhance accumulation of carbon (C) in soil. However, the net change in soil C also depends on the effect of plant diversity on C loss through decomposition of organic matter. Plant diversity can affect organic matter decomposition via changes in litter species diversity and composition, and via alteration of abiotic and/or biotic attributes of the soil (soil legacy effect). Previous studies examined the two effects on decomposition rates separately, and do therefore not elucidate the relative importance of the two effects, and their potential interaction. Here we separated the effects of litter mixing and litter identity from the soil legacy effect by conducting a factorial laboratory experiment where two fresh single root litters and their mixture were mixed with soils previously cultivated with single plant species or mixtures of two or four species. We found no evidence for litter-mixing effects. In contrast, root litter-induced CO2 production was greater in soils from high diversity plots than in soils from monocultures, regardless of the type of root litter added. Soil microbial PLFA biomass and composition at the onset of the experiment was unaffected by plant species richness, whereas soil potential nitrogen (N) mineralization rate increased with plant species richness. Our results indicate that the soil legacy effect may be explained by changes in soil N availability. There was no effect of plant species richness on decomposition of a recalcitrant substrate (compost). This suggests that the soil legacy effect predominantly acted on the decomposition of labile organic matter. We thus demonstrated that plant species richness enhances root litter-induced soil respiration via a soil legacy effect but not via a litter-mixing effect. This implies that the positive impacts of species richness on soil C sequestration may be weakened by accelerated organic matter decomposition.

The aim of this study was to investigate the effect of tissue nitrogen concentration, as a consequence of nitrogen supply rate, on the susceptibility of tomato plants to three pathogens. We varied tissue N concentration by supplying N at different rates by adding nitrate in different, exponentially increasing amounts to the nutrient solution on which the tomato plants were grown. Separate experiments were carried out to test susceptibility of tomato plants to the bacterial speck-causing Pseudomonas syringae pv tomato, to the wilt agent Fusarium oxysporum f.sp. lycopersici and to tomato powdery mildew caused by Oidium lycopersicum. The effect of tissue N concentration appeared to be highly pathogen-dependent: there was no effect on susceptibility to F. oxysporum, but susceptibility to P. syringae and O. lycopersicum increased significantly with increasing N concentration. We have previously demonstrated the opposite for susceptibility to Botrytis cinerea: decreasing susceptibility with increasing N concentration. The apparent contradictory effects are discussed in relation to the effect of N supply on both the nutritional value of the plant tissue to the pathogen and on the concentration of resistance-related compounds. We conclude that the effect of changing both characteristics on disease susceptibility is highly pathogen-specific and is probably dependent on differences in resource requirements of the pathogen or the sensitivity of the pathogen to plant resistance reactions or on both these factors.

The aim of this work was to study how nitrogen availability affects within-plant allocation to growth and secondary metabolites and how this allocation affects host selection by herbivores. Tomato plants (Lycopersicon esculentum) were grown at six levels of nitrogen availability. When nitrogen availability increased, plant relative growth rate increased, but tissue carbon/nitrogen ratio in the second oldest true leaf and allocation to large glandular trichomes (type VI) as well as to the defense compounds rutin and chlorogenic acid decreased. Leaf protein concentration increased. Two-spotted spider mite (Tetranychus urticae) females responded significantly to these changes: in dual choice tests they preferred leaf disks from plants grown at high nitrogen availability, with a low C/N ratio. This preference persisted in an olfactometer in which the mites were offered only the odors released by leaves with damaged trichomes. We conclude that in a tomato leaf increased nitrogen availability leads to decreased allocation to defenses, and that repellent volatiles released by trichomes play a key role in affecting leaf selection by two-spotted spider mite females.

The aim of this study was to investigate the effect of nitrogen availability on susceptibility of tomato leaves to the fungal pathogen Botrytis cinerea. Plants with varying nitrogen availability were grown by adding N daily in exponentially increasing amounts to a nutrient solution at different rates. Leaves of plants grown at low nitrogen availability had a high leaf C/N ratio (21 g g-1) and were about 2.5 times more susceptible to primary lesion formation by B. cinerea compared to plant grown at high nitrogen availability, which had a low leaf C/N ratio (11 g g-1). Leaf C/N ratio accounted for 95% of variation in susceptibility. This relationship between C/N ratio and susceptibility persisted when plants were grown with exponential P addition and optimal N supply, and was thus independent of plant growth rate or related factors. We could not explain the effect of nitrogen availability by variation in the most obvious N-based resistance compound α-tomatine because more susceptible leaves with a high C/N ratio contained more α-tomatine. These leaves also contained more soluble carbohydrates. The level of soluble carbohydrates correlated positively with susceptibility, independent of the growth method. We therefore suggest that the effect of N availability on susceptibility must be explained by variation in levels of soluble carbohydrates and speculate about the role of these carbohydrates in the infection process.

Aim Our aim was to improve the prediction of Zn bioavailability to wheat grown on low-Zn soils. The classical approach that directly relates Zn in a certain soil extract to Zn uptake has been shown to be inadequate in many cases. We tested a stepwise approach where the steps of the uptake process are characterized with, respectively, Zn solid-solution distribution, adsorption of Zn to root surface, Zn uptake into root and Zn translocation to shoot. Methods Two pot experiments were done with wheat grown on nine low-Zn soils varying widely in pH, clay and organic matter content. Soluble Zn concentrations in two soil extracts (DTPA and CaCl2) were measured. Free Zn ion concentrations in CaCl2 soil extracts were determined with the Donnan Membrane Technique. These Zn concentrations were then related to plant Zn uptake following both the direct and the stepwise approach. Results In the direct approach, Zn in the DTPA extract was a better predictor for shoot Zn uptake than Zn in the CaCl2 extract. In the stepwise approach, the relationship between Zn in CaCl2 extracts and the root surface adsorbed Zn was pH-dependent and nonlinear. Root surface adsorbed Zn was linearly related to root Zn uptake, and the latter was linearly related to the shoot Zn uptake. The stepwise approach improved the Zn uptake prediction compared to the direct approach and was also validated for different wheat cultivars. Conclusions The adsorption of Zn on the root surface is pH dependent and nonlinear with respect to the soil Zn concentration, and a useful proxy for bioavailable Zn over a wide range of soils.

To test the hypothesis that rhizosphere acidification would enhance the hydrolyzation of organic phosphates by increasing phosphatase activity. A Petri dish experiment with sterile agar and a pot experiment with a low P soil were used. In the Petri dish experiment, roots of each plant were cultured in two compartments, each of which contained agar with one of three nitrogen combinations: NH4+/N0 (N0 = nitrogen free), NH4+/NO3-, and NO3-/N0. Phytin was supplied as the sole phosphorus (P) source to all compartments. In the pot experiment, the soil in each pot was treated with N0, KNO3, or (NH4)2SO4) together with 0 or 75 mg kg−1 phytin-P. Dry weight, P concentration, and P content of roots were highest in the NH4+ compartments in the Petri dish experiment. In the pot experiment, dry weight, P concentration, and P content of both shoots and roots were higher with NH4+ than with NO3-. NH4+ treatments reduced rhizosphere pH, promoted the hydrolization of phytin, enhanced acid phosphatase activity in the rhizosphere, and increased phytin-P utilization relative to N0 and NO3- treatments. Phosphatase activity was negatively correlated with rhizosphere pH but was positively correlated with plant P content in both experiments. Rhizosphere acidification optimized the activity of acid phosphatase excreted by maize roots and promoted phytin mineralization. NH4+-induced acidification in the maize rhizosphere improved the growth of maize roots by improving P uptake from phytin; the improved growth, in turn, increased NH4+ uptake and acidification.

summary Rape (Brassica napus L.) plants are efficient users of rock phosphates. This has been ascribed to the excretion of malic and citric acids from the roots during phosphorus deficiency, resulting in a decrease of the pH in the rhizosphere and a consequent solubilization of the rock phosphate. In this paper the biosynthesis and excretion of organic acids were investigated, partly by comparing physiological reactions to P deficiency of rape with those of an inefficient user of rock phosphate from the same family, hedge mustard (Sisymbrium officinale Scop.). In the leaves of both species, both citrate levels and phosphoenolypyruvate carboxylase were nearly doubled in P–deficient plants, and the citrate/sugar ratio in the phloem exudate into EDTA solution was doubled as well. After exposure of the shoots of rape plants to 14CO2 in the light, the specific activities of citrate in the root and in the nutrient solution were ten times higher than those of malate, and also much higher than the respective activities of citrate in P–sufficient controls. It is suggested that citrate produced in the leaf is exported to the roots, where it is accumulated in the excretion region. Malic acid which is prevailing in the exudate is probably newly–synthesized within the excreting root segment. When no phosphate is supplied, this excretion zone is located 1–2 cm behind the root tips. However, when rock phosphate is applied locally this zone shifts along the root to that part which is in direct contact with the rock phosphate particles. No accumulation and excretion of organic acids could be observed with the roots of P–deficient Sisymbrium.

Plant Zn uptake from low Zn soils can be increased by Zn-mobilizing chemical rhizosphere processes. We studied whether inoculation with arbuscular mycorrhizal fungi (AMF) can be an additional or an alternative strategy. We determined the effect of AMF inoculation on growth performance and Zn uptake by rice genotypes varying in Zn uptake when nonmycorrhizal. A pot experiment was conducted with six aerobic rice genotypes inoculated with Glomus mosseae or G. etunicatum or without AMF on a low Zn soil. Plant growth, Zn uptake and mycorrhizal responsiveness were determined. AMF-inoculated plants produced more biomass and took up more Zn than nonmycorrhizal controls. Mycorrhizal inoculation, however, significantly increased Zn uptake only in genotypes that had a low Zn uptake in the nonmycorrhizal condition. We conclude that genotypes that are less efficient in Zn uptake when nonmycorrhizal are more responsive to AMF inoculation. We provide examples from literature allowing generalization of this conclusion on a trade off between mycorrhizal responsiveness and nutrient uptake efficiency.

The objectives of this paper were to determine (1) if lowland rice (Oryza sativa L.) plants respond similarly to low zinc (Zn) and phosphorus (P) availability by increased root exudation of low-molecular weight organic anions (LMWOAs) and (2) if genotypic variation in tolerance to low soil supply of either Zn or P is related to LMWOA exudation rates. Exudation of LMWOAs can increase bioavailability of both Zn and P to the plant, through partly similar chemical mechanisms. We used seven lowland rice genotypes and showed in two experiments that genotypes that grow relatively well on a soil with low Zn availability also grow well on a sparingly soluble Ca-phosphate (r = 0.80, P = 0.03). We measured exudation rates of LMWOAs on nutrient solution and found that both Zn and P deficiency induced significant increases. Among the LMWOAs detected oxalate was quantitatively the most important, but citrate is considered more effective in mobilizing Zn. Citrate exudation rates correlated with tolerance to low soil levels of Zn (P=0.05) and P (P = 0.07). In a low-Zn-field we found an increased biomass production at higher plant density, which is supportive for a concentration-dependent rhizosphere effect on Zn bioavailability such as LMWOA exudation. We, for the first time, showed that tolerance to low Zn availability is related to the capacity of a plant to exude LMWOAs and confirmed that exudation of LMWOAs must be regarded a multiple stress response.

Scarcity of water causes a shift from flooded to aerobic conditions for rice production in zinc deficient areas in Northern China. This shift alters soil conditions that affect zinc availability to the crop. This paper concerns the effect of aerobic compared to flooded conditions on crop biomass production, grain yield and zinc content. A field experiment was done with six rice genotypes (Oryza sativa L.) grown on a calcareous soil, both with (23 kg Zn ha¿1) and without Zn fertilization. Sampling was conducted at tillering and physiological mature stage. Zn concentration in the shoots was significantly lower at both stages in plants grown in the aerobic field. At maturity, Zn uptake, biomass production, grain yield and Zn-harvest index [grain Zn/(shoot + grain Zn)] were lower under aerobic cultivation. Rice genotypes including aerobic rice and lowland rice differ in degree of response to low Zn supply. A twofold difference was found among aerobic genotypes in grain yield and Zn uptake. Also Zn-harvest index varied significantly. Zn application affected neither grain yield nor grain Zn content, although it significantly improved biomass production in both systems in most genotypes. These results demonstrate that introduction of aerobic rice systems on calcareous soils may increase Zn deficiency problems.

To study variation in zinc efficiency (ZE) among current Chinese rice genotypes, a pot experiment was conducted with 15 aerobic and 8 lowland rice genotypes. Aerobic rice is currently bred by crossing lowland with upland rice genotypes, for growth in an aerobic cultivation system, which is saving water and producing high yields. A Zn deficient clay soil was used in our screening. Zn deficiency resulted in a marked decrease in shoot dry matter production of most genotypes after 28 days of growth. Genotypes were ranked according to their tolerance to Zn deficiency based on ZE, expressed as the ratio of shoot dry weight at Zn deficiency over that at adequate Zn supply. Substantial genotypic variation in ZE (50¿98%) was found among both lowland and aerobic genotypes. ZE correlated significantly (P

Plant nutrients, with the exception of nitrogen, are ultimately derived from weathering of primary minerals. Traditional theories about the role of ectomycorrhizal fungi in plant nutrition have emphasized quantitative effects on uptake and transport of dissolved nutrients. Qualitative effects of the symbiosis on the ability of plants to access organic nitrogen and phosphorus sources have also become increasingly apparent. Recent research suggests that ectomycorrhizal fungi mobilize other essential plant nutrients directly from minerals through excretion of organic acids. This enables ectomycorrhizal plants to utilize essential nutrients from insoluble mineral sources and affects nutrient cycling in forest systems.

To determine the origin of organic acids exuded by the roots of P deficient rape plants, phosphoenolpyruvate carboxylase (PEPC) activity was measured in plants after deprivation of P in the nutrient solution. PEPC activity in the shoot increased as a reaction to P stress. This increase coincided with accumulation of citrate in the shoot and with a higher citrate/sugar ratio in the phloem. Application of 14CO2 to the shoots resulted in a ninefold increase in specific activity of organic acids exuded by —P roots as compared with +P roots. These results indicate that exuded organic acids resulted from increased PEPC activity in the shoot of P stressed rape plants.

This thesis focuses on the interaction between fungi and the nitrogen cycle in grassland soils, and attempts to unravel causes and consequences of changing fungal biomass.

Keywords:Arbuscular mycorrhiza, Exudation, Oryza sativa , Rhizosphere, Rice, Soil, ZincZinc deficiency is a wide-spread constraint for crop production and human health. This thesis should contribute to alleviation of Zn deficiency problems and aimed at identifying soil and plant factors affecting Zn bioavailability in rice ( Oryza sativa L.). Two main research questions were: 1) what is the consequence of a cultivation shift from flooded to aerobic rice on Zn bioavailability and 2) which mechanisms control Zn mobilization by aerobic rice? Field experiments demonstrated that the cultivation shift from flooded to aerobic may increase Zn deficiency problems. Lower Zn uptake and mass fraction in shoot and brown rice were observed in aerobic fields. Results of a soil incubation experiment and modeling showed that the difference found in plant Zn uptake between the two cultivation systems was orders of magnitude lower compared to what can be expected based on chemical equilibria in the bulk soil. This discrepancy suggests that soil chemical properties such as pH, DOC level and redox conditions in rhizosphere need consideration. We observed considerable variation among aerobic rice genotypes in tolerance to Zn deficiency under both field and pot conditions. In a pot experiment we found that variation in tolerance was mainly associated with Zn uptake. Plant Zn uptake from low Zn soils can be increased by both Zn-mobilizing rhizosphere processes and inoculation with arbuscular mycorrhizal fungi. We demonstrated that mycorrhizal inoculation significantly increased Zn uptake, but only in genotypes with a low inherent Zn uptake. High Zn mobilization by a combination of chemical rhizosphere and mycorrhizal effects seems impossible and may not be feasible as a target for breeders. In a rhizotron experiment and a nutrient solution experiment, aerobic rice genotypes responded to Zn deficiency with increased root exudation of malate.Genotypes with a higher Zn uptake showed a stronger increase in malate exudation in response to Zn deficiency.These results confirm our hypothesis that genotypic variation in Zn uptake of aerobic rice can partly be explained by root exudation of malate.