Root exudation of sugars,amino acids,and organic acids by maize as affected by nitrogen,phosphorus, potassium,and iron deficiency

Root exudates play a major role in the mobilization of sparingly soluble nutrients in the rhizosphere. Since the amount and composition of major metabolites in root exudates from one plant species have not yet been systematically compared under different nutrient deficiencies, relations between exudation patterns and the type of nutrient being deficient remain poorly understood. Comparing root exudates from axenically grown maize plants exposed to N, K, P, or Fe deficiency showed a higher release of glutamate, glucose, ribitol, and citrate from Fe‐deficient plants, while P deficiency stimulated the release of γ‐aminobutyric acid and carbohydrates. Potassium‐starved plants released less sugars, in particular glycerol, ribitol, fructose, and maltose, while under N deficiency lower amounts of amino acids were found in root exudates. Principal‐component analysis revealed a clear separation in the variation of the root‐exudate composition between Fe or P deficiency versus N or K deficiency in the first principal component, which explained 46% of the variation in the data. In addition, a negative correlation was found between the amounts of sugars, organic and amino acids released under deficiency of a certain nutrient and the diffusion coefficient of the respective nutrient in soils. We thus hypothesize that the release of dominant root exudates such as sugars, amino acids, and organic acids by roots may reflect an ancient strategy to cope with limiting nutrient supply.     

Influence of enhanced malate dehydrogenase expression by alfalfa on diversity of rhizobacteria and soil nutrient availability

Transgenic alfalfa over-expressing a nodule-enhanced malate dehydrogenase (neMDH) cDNA and untransformed alfalfa plants were grown at the same field site and rhizosphere soils collected after 53 weeks of plant growth. These alfalfa lines differ in the amount and composition of root organic acids produced and exuded into the rhizosphere. Nucleotide sequencing of PCR-based 16S ribosomal DNA (rDNA) clone libraries and Biolog™ GN microtiter plates were employed to assess the activity of naturally occurring rhizobacteria in the two alfalfa rhizospheres. Selected macro- and micro-elements in the two alfalfa rhizosphere soils were also measured. Analysis of 240 16S rDNA clone sequences indicated the existence of about 11 bacterial phyla and their major subdivisions in the two alfalfa rhizosphere samples. There were qualitative changes in the abundance of bacterial phylogenetic groups between rhizosphere soils of transgenic and untransformed alfalfa. Carbon substrate utilization profiles suggested that rhizosphere samples from transgenic alfalfa had significantly greater microbial functional diversity compared with rhizosphere samples from untransformed alfalfa. The concentrations of nitric acid extractable P, K, Mn, Zn and Cu increased significantly in the transgenic alfalfa rhizosphere compared with the untransformed alfalfa rhizosphere. These observations indicate that organic acids produced by plant roots significantly influence rhizosphere microbial diversity and availability of macro- and micro-nutrients and demonstrate the utility of such trangenic plants as tools for studying the potential impact of plant root exudates on soil microbial ecosystems.     

Rhizosphere interactions between microorganisms and plants govern iron and phosphorus acquisition along the root axis - model and research methods

Iron and phosphorus availability is low in many soils; hence, microorganisms and plants have evolved mechanisms to acquire these nutrients by altering the chemical conditions that affect their solubility. In plants, this includes exudation of organic acid anions and acidification of the rhizosphere by release of protons in response to iron and phosphorus deficiency. Grasses (family Poaceae) and microorganisms further respond to Fe deficiency by production and release of specific chelators (phytosiderophores and siderophores, respectively) that complex Fe to enhance its diffusion to the cell surface. In the rhizosphere, the mutual demand for Fe and P results in competition between plants and microorganisms with the latter being more competitive due to their ability to decompose plant-derived chelators and their proximity to the root surface; however microbial competitiveness is strongly affected by carbon availability. On the other hand, plants are able to avoid direct competition with microorganisms due to the spatial and temporal variability in the amount and composition of exudates they release into the rhizosphere. In this review, we present a model of the interactions that occur between microorganisms and roots along the root axis, and discuss advantages and limitations of methods that can be used to study these interactions at nanometre to centimetre scales. Our analysis suggests mechanisms such as increasing turnover of microbial biomass or enhanced nutrient uptake capacity of mature root zones that may enhance plant competitiveness could be used to develop plant genotypes with enhanced efficiency in nutrient acquisition. Our model of interactions between plants and microorganisms in the rhizosphere will be useful for understanding the biogeochemistry of P and Fe and for enhancing the effectiveness of fertilization.     

Metabolite profiling of shoot extract,root extract,and root exudate of rice under nitrogen and phosphorus deficiency

ABSTRACT

Root exudate is derived from plant metabolites and its composition is affected by plant nutrient status. A deficiency of mineral nutrients, such as nitrogen (N) and phosphorus (P), strongly affects the type and amount of plant metabolites. We applied a metabolite profiling technique to investigate root exudates of rice plants under N and P deficiency. Oryza sativa was grown in culture solution containing two N levels (0 and 60 mg N L^?1) or two P levels (0 and 8 mg P L^?1). Shoot extracts, root extracts, and root exudates were obtained from the rice plants 5 and 15 days after transplanting and their metabolites were determined by capillary electrophoresis/time-of-flight mass spectrometry. Shoot N concentration and dry weight of rice plants grown at ?N level were lower than those of plants grown at +N level. Shoot P concentration and dry weight of rice plants grown at ?P level were lower than those of plants grown at +P level. One hundred and thirty-two, 127, and 98 metabolites were identified in shoot extracts, root extracts, and root exudates, respectively, at the two N levels. One hundred and thirty-two, 128, and 99 metabolites were identified in shoot extracts, root extracts, and root exudates, respectively, at the two P levels. Seventy-seven percent of the metabolites were exuded to the rhizosphere. The concentrations of betaine, gamma-aminobutyric acid, and glutarate in root exudates were higher at both ?N and ?P levels than at their respective high levels. The concentration of spermidine in root exudates was lower at both ?N and ?P levels than at their respective high levels. The concentrations of the other metabolites in root exudates were affected differently by plant N or P status. These results suggest that rice roots actively release many metabolites in response to N and P deficiency.     

植物高效利用土壤磷营养的化学机理

本文闸述了植物根系分泌物与土壤磷素有效性的相互关系。根据根系分泌物的化学特性,从七个方而讨论了植物活化土壤磷素的机理。     

Organic phosphorus compounds as a phosphorus source for higher plants through the activity of phosphatases produced by plant roots and microorganisms

Summary The efficiency of phosphatases produced by clover, barley, oats and wheat was investigated in soils treated with sodium glycerophosphate, lecithin and phytin. Root exudates of aseptically grown clover were also examined for the breakdown of different organic P compounds in order to test the efficiency of plant-produced phosphatases. In general, the plants were able to use P from all the organic sources used in the study almost as efficiently as inorganic sources. Dry-matter yield, P uptake, acid and alkaline phosphatase activity and microbial population were increased in all the P treatments. Organic P enhanced alkaline phosphatase activity. Lecithin increased fungal, and phytin bacterial growth. There was no alkaline phosphatase activity in the asepticallly grown clover root exudates. Phosphatase released in aseptic culture after 4 weeks of clover growth was able to efficiently hydrolyse sodium glycerophosphate, lecithin and phytin. The amount of organic P hydrolysed in this and in the soil experiment surpassed plant uptake by a factor of 20. This suggests that the limiting factor on plant utilization of organic P is the availability of hydrolysable organic P sources.     

The bacterial community of tomato rhizosphere is modified by inoculation with arbuscular mycorrhizal fungi but unaffected by soil enrichment with mycorrhizal root exudates or inoculation with Phytophthora nicotianae

Arbuscular mycorrhizal (AM) fungi have been shown to induce the biocontrol of soilborne diseases, to change the composition of root exudates and to modify the bacterial community structure of the rhizosphere, leading to the formation of the mycorrhizosphere. Tomato plants were grown in a compartmentalized soil system and were either submitted to direct mycorrhizal colonization or to enrichment of the soil with exudates collected from mycorrhizal tomato plants, with the corresponding negative controls. Three weeks after planting, the plants were inoculated or not with the soilborne pathogen Phytophthora nicotianae growing through a membrane from an adjacent infected compartment. At harvest, a PCR-Denaturing gradient gel electrophoresis analysis of 16S rRNA gene fragments amplified from the total DNA extracted from each plant rhizosphere was performed. Root colonization with the AM fungi Glomus intraradices or Glomus mosseae induced significant changes in the bacterial community structure of tomato rhizosphere, compared to non-mycorrhizal plants, while enrichment with root exudates collected from mycorrhizal or non-mycorrhizal plants had no effect. Our results support that the effect of AM fungi on rhizosphere bacteria would not be mediated by compounds present in root exudates of mycorrhizal plants but rather by physical or chemical factors associated with the mycelium, volatiles and/or root surface bound substrates. Moreover, infection of mycorrhizal or non-mycorrhizal plants with P. nicotianae did not significantly affect the bacterial community structure suggesting that rhizosphere bacteria would be less sensitive to the pathogen invasion than to mycorrhizal colonization. Of 96 unique sequences detected in the tomato rhizosphere, eight were specific to mycorrhizal fungi, including two Pseudomonas, a Bacillus simplex, an Herbaspirilium and an Acidobacterium. One Verrucomicrobium was common to rhizospheres of mycorrhizal plants and of plants watered with mycorrhizal root exudates.     

Sour Orange (Citrus Aurantium L.) Growth is Strongly Mycorrhizal Dependent in Terms of Phosphorus (P) Nutrition Rather than Zinc (Zn)

The partial sterilization of soil eliminates useful microorganisms, resulting in the reduced growth of mycorrhizae-dependent citrus plants, which are often unresponsive to the application of fertilizer. Research was conducted to test the hypothesis that indigenous mycorrhizae (IM) inoculation is as efficient as selected mycorrhizal inoculation under sterile and non-sterile soil conditions. Rhizophagus clarus and indigenous mycorrhiza spores, isolated from citrus orchards, were used as arbuscular mycorrhizae fungi under greenhouse conditions with sterile and non-sterile Çanakçi series (Typic xerofluvent) soils with low phosphorus (P) fertility. Different P (0 and 100 mg kg^?1) and zinc (Zn) (0, 5 and 10 mg kg^?1) concentrations were used at the start of the experiments. The shoot, root dry weight (RDW), root colonization, and P, Zn, iron (Fe), copper (Cu) and manganese (Mn) concentrations of the shoot were determined; mycorrhizae dependency (MD) was also calculated.

The results indicate that R. clarus and indigenous mycorrhiza in sterile and non-sterile soil conditions considerably increased the growth of citrus plants. Owing to existing beneficial indigenous rhizosphere microorganisms, citrus plant growth without inoculation was better in non-sterile soils than in the sterile soils. In non-sterilized soil, the plant growth parameters of R. clarus-inoculated soils were higher than those of indigenous mycorrhiza-inoculated soils. Mycorrhizae infection increased certain citrus plant growth parameters, such as root infection, biomass and nutrient uptake (P, Zn, Fe, Mn and Cu). In sterile soil, the addition of up to 5 mg kg^?1 soil Zn and the inoculation of R. clarus significantly increased plant growth; inoculation with indigenous mycorrhiza produced more dry weight upon the addition of up to 100 mg kg^?1 phosphorus pentoxide (P₂O₅). Under sterile soil conditions, without considering fertilizer addition, MD was found to be higher than that of non-sterile soils. In general, the contribution of the indigenous soil spores is significant. However, indigenous soil mycorrhizae may need to be managed for better efficiency in increasing plant growth and nutrient uptake. The major finding was that the inoculation of citrus seedlings with mycorrhiza is necessary under both sterilized and non-sterilized soil conditions.     

石灰性土壤难溶态磷的微生物转化和利用

目前农业生产中大多通过施用可溶性磷肥为植物提供有效磷。磷酸根化学性质活泼,施入土壤后能很快与土壤中的其它成分发生反应,使植物对其利用的有效度随时间延长而降低,最终以难溶性磷酸盐或吸附态形式滞留于土壤中,难以被植物直接吸收。据估计,在石灰性土壤中约有80% 的磷肥以难溶性磷酸盐存在。为此人们采用了许多方法提高磷肥的利用率,其中利用植物根际与磷循环相关的生物学系统来调节植物根际磷的有效性是重要的途径之一。这个生物学系统包括植物本身对土壤难溶态磷的吸收与利用以及土壤中某些微生物参与的难溶态磷的释放与利用。本文论述了微生物( 细菌和真菌) 转化和利用石灰性土壤中难溶态磷的研究进展。     

Rhizospheric organic compounds in the soil–microorganism–plant system: their role in iron availability

Poor iron (Fe) availability in soil represents one of the most important limiting factors of agricultural production and is closely linked to physical, chemical and biological processes within the rhizosphere as a result of soil–microorganism–plant interactions. Iron shortage induces several mechanisms in soil organisms, resulting in an enhanced release of inorganic (such as protons) and organic (organic acids, carbohydrates, amino acids, phytosiderophores, siderophores, phenolics and enzymes) compounds to increase the solubility of poorly available Fe pools. However, rhizospheric organic compounds (ROCs) have short half‐lives because of the large microbial activity at the soil–root interface, which might limit their effects on Fe mobility and acquisition. In addition, ROCs also have a selective effect on the microbial community present in the rhizosphere. This review aims therefore to unravel these complex dynamics with the objective of providing an overview of the rhizosphere processes involved in Fe acquisition by soil organisms (plants and microorganisms). In particular, the review provides information on (i) Fe availability in soils, including mineral weathering and Fe mobilization from soil minerals, ligand and element competition and plant‐microbe competition; (ii) microbe–plant interactions, focusing on beneficial microbial communities and their association with plants, which in turn influences plant mineral nutrition; (iii) plant–soil interactions involving the metabolic changes triggered by Fe deficiency and the processes involved in exudate release from roots; and (iv) the influence of agrochemicals commonly used in agricultural production systems on rhizosphere processes related to Fe availability and acquisition by crops.     

Phosphorus Efficiency of Buckwheat (Fagopyrum esculentum)

The ability of buckwheat (Fagopyrum esculentum) roots to acquire phosphorus (P) was characterized by investigating P uptake, morphological features, and chemical changes in the rhizosphere. Over a range of nutrient solution P concentrations (5–500 μmol · L^?1), maximum shoot growth was achieved with a P supply between 5 and 100 μmol · L^?1. Root weight and root length, as well as length and frequency of root hairs, were higher at low P levels. Root surface and the root surface/shoot dry weight ratio reached high values. Though P uptake rates were only moderate (0.15 pmol · cm^?1 root · sec^?1), shoot P concentrations were high (1.8% of dry weight with 100 μM P) predominantly being inorganic (80%). Phosphorus efficiency was characterized by a high specific absorption rate (810 mmol P · kg^?1 root dry wt · d^?1) rather than by an efficient utilization for dry weight production. Root exudates of low-P plants had lower pH values than exudates of high-P plants and increased the solubility of FePO₄ and MnO₂ to a greater extent. Amounts of exuded organic acids and phenolics were low and could not account for the observed solubilization of FePO₄ and MnO₂. Enhanced hydrolysis of glucose-6-phosphate by exudates from low-P plants was due to an increased “soluble” acid phosphatase activity, and root surface phosphatase activity was also slightly enhanced with P deficiency. In the rhizosphere soil of buckwheat, some depletion of organic P forms was observed, and in pot trials with quartz sand, buckwheat utilized glucose- 6-phosphate as a P source at the same rate as inorganic P.     

Intercropping promotes the ability of durum wheat and chickpea to increase rhizosphere phosphorus availability in a low P soil

Cereal–legume intercropping can promote plant growth (i.e. facilitation) through an increase in the amount of phosphorus (P) taken up, especially in low P soils. The aim of this study was to test the hypothesis that these positive interactions are supported by rhizosphere processes that increase P availability, such as root-induced pH changes. In neutral and alkaline soils legumes are assumed to increase inorganic P availability by rhizosphere acidification due to N₂ fixation which benefit to the intercropped cereal. Growth, P uptake, changes in inorganic P availability and pH in the rhizosphere of intercropped species were thus investigated in a greenhouse pot experiment with durum wheat and chickpea either grown alone or intercropped. We used a neutral soil from a P fertilizer long-term field trial exhibiting either low (−P) or high (+P) P availability. Phosphorus availability was increased in the rhizosphere of both species, especially when intercropped in −P. Such increase was associated with alkalization. Rhizosphere pH changes could not fully explain the observed changes of P availability though. Low rates of N₂ fixation may explain why no rhizosphere acidification was observed. Increases in P availability did not lead to enhanced P uptake but growth promotion was observed for durum wheat intercropped with chickpea in −P soil. Our hypothesis of an increase in inorganic P availability in intercropping as a consequence of root-induced acidification by the legume was not validated, and we suggested that root-induced alkalization was involved instead, as well as other root-induced processes. Thus, the cereal through rhizosphere alkalization may also enhance P uptake and growth of the intercropped legume. Facilitation can thus occur in both ways.     

土壤－水分－植物体系中铅和镉的交互作用及其机制: Ⅱ. 根际中Pb-Cd的交互作用

The interaction of Pb-Cd can be observed not only in the uptake process of elements by plants and in their influence on the growth,but also in rhizosphere.The changes in extractable Cd and Pb concentrations in the rhizosphere soil of rice plants ,root exudates from wheat and wheat plant and their complexing capacity,with Pa and Cd were investigated under different Pb and Cd treatments.Results showed that the concentration of extractable Cd in the rhizosphere of rice in red soil was markedly increased by Pb-Cd interaction,It increased by 56% in the treatment with Pb and Cd added against that in the treatment with only Cd added in soil . The considerable differences in both composition and amount of root exudate from wheat and rice were found among different treatments.Pb and Cd might be complexed by root exudates ,The concentrations of free Pb and Cd in the solution were increased markedly by adding root exudate from wheat and decreased by that from rice due to Pd-Cd interaction.The distribution patterns of Pb and Cd in roots were affected by Pb-Cd interaction,which accelerated transport of Pb into internal tissue and retarded accumulation of Cd in external tissue.     

Growth and Phosphorus Uptake of Oat (Arena nuda L.) as Affected by Mineral Nitrogen Forms Supplied in Hydroponics and Soil Culture

Plants show different growth responses to N sources supplied with either NH4+ or NO3-.The uptake of different N sources also affects the rhizosphere pH and therefore the bioavailability of soil phosphorus,particularly in alkaline soils.The plant growth,P uptake,and P availability in the rhizosphere of oat (Arena nuda L.) grown in hydroponics and in soil culture were investigated under supply with sole NH4+-N,sole NO3--N,or a combination.Sole NO3-fed oat plants accumulated more biomass than sole NH4+-fed ones.The highest biomass accumulation was observed when N was supplied with both NH4+-N and NO3--N.Growth of the plant root increased with the proportion of NO3-in the cultural medium.Better root growth and higher root/shoot ratio were consistently observed in NO3--fed plants.However,root vigor was the highest when N was supplied with NO3-+NH4+.NH4+ supply reduced the rhizosphere pH but did not affect P uptake by plants grown in soils with CaHPO4 added as P source.No P deficiency was observed,and plant P concentrations were generally above 2 g kg-1.P uptake was increased when N was supplied partly or solely as NO3--N,similarly as biomass accumulation.The results suggested that oat was an NO3-prcferring plant,and NO3--N was essential for plant growth and the maintenance of root absorption capacity.N supply with NH4+-N did not improve P nutrition,which was most likely due to the absence of P deficiency.     

水培和土培条件下氮素供应对燕麦生长和磷素吸收的影响

Plants show different growth responses to N sources supplied with either NH₄⁺ or NO₃^-. The uptake of different N sources also affects the rhizosphere pH and therefore the bioavailability of soil phosphorus, particularly in alkaline soils. The plant growth, P uptake, and P availability in the rhizosphere of oat (Avena nuda L.) grown in hydroponics and in soil culture were investigated under supply with sole NH₄⁺-N, sole NO₃^--N, or a combination. Sole NO₃^--fed oat plants accumulated more biomass than sole NH₄⁺-fed ones. The highest biomass accumulation was observed when N was suppliedw ith both NH₄⁺-N and NO₃^--N. Growth of the plant root increased with the proportion of NO₃^- in the cultural medium. Better root growth and higher root/shoot ratio were consistently observed in NO₃^--fed plants. However, root vigor was the highest when N was supplied with NO₃^-+NH₄⁺. NH₄⁺ supply reduced the rhizosphere pH but did not affect P uptake by plants grown in soils with CaHPO₄ added as P source. No P deficiency was observed, and plant P concentrations were generally above 2 g kg^-1. P uptake was increased when N was supplied partly or solely as NO₃^--N, similarly as biomass accumulation. The results suggested that oat was an NO₃^--preferring plant, and NO₃^--N was essential for plant growth and the maintenance of root absorption capacity. N supply with NH₄⁺-N did not improve P nutrition, which was most likely due to the absence of P deficiency.     

根分泌物对活化土壤中难溶性磷的作用

土壤大部分磷以难溶性磷形式存在,是影响作物生产的重要限制因素之一。作物根分泌物活化难溶性磷的能力对改善其磷素营养具有重要意义。采用室内培养方法,研究3种土壤不同磷(KH2PO4)处理后有效磷动态变化情况;同时将不同玉米基因型分别置于全磷和低磷的营养液中生长15 d后,收集根系分泌物,然后加入预先采用磷酸钙和磷酸铝处理并已培养95 d的土壤样品中进行试验。不同磷处理研究结果表明,所有施磷处理短期内速效磷含量急剧上升,并随培养时间推移不断下降,60 d后趋于稳定。酸性土对磷的吸附和固定作用较强,磷处理后有效磷增加量显著小于中性土、碱性土。土壤加根系分泌物的试验表明,磷酸钙和磷酸铝有效磷的含量较对照有所提高,其中加入耐低磷基因型M02低磷根系分泌物浸提的土壤有效磷含量显著高于去离子水的浸提量,且根系分泌物对磷酸铝的活化能力大于磷酸钙。     

Soil sickness of peanuts is attributable to modifications in soil microbes induced by peanut root exudates rather than to direct allelopathy

The quantity and quality of peanut yields are seriously compromised by consecutive monoculture in the subtropical regions of China. Root exudates, which represent a growth regulator in peanut–soil feedback processes, play a principal role in soil sickness. The growth inhibition of a species in an in vitro bioassay enriched with root exudates and allelochemicals is commonly viewed as evidence of an allelopathic interaction. However, for some of these putative examples of allelopathy, the results have not been verified in more natural settings with plants continuously growing in soil. In this study, the phenolic acids in peanut root exudates, their retention characteristics in an Udic Ferrosol, and their effects on rhizosphere soil microbial communities and peanut seedling growth were studied. Phenolic acids from peanut root exudates were quickly metabolized by soil microorganisms and did not accumulate to high levels. The peanut root exudates selectively inhibited or stimulated certain communal bacterial and fungal species, with decreases in the relative abundance of the bacterial taxa Gelria glutamica, Mitsuaria chitosanitabida, and Burkholderia soli and the fungal taxa Mortierella sp. and Geminibasidium hirsutum and increases in the relative abundance of the bacterial taxon Desulfotomaculum ruminis and the fungal taxa Fusarium oxysporum, Bionectria ochroleuca and Phoma macrostoma. The experimental application of phenolic acids to non-sterile and sterile soil revealed that the poor performance of the peanut plants was attributed to changes in the soil microbial communities promoted by phenolic acids. These results suggest that pathogenic fungal accumulation at the expense of such beneficial microorganisms as plant growth promoting rhizobacteria, mycorrhizal fungi induced by root exudates, rather than direct autotoxicity induced by root exudates, might represent the principal cause underlying the soil sickness associated with peanut plants. We hope that our study will motivate researchers to integrate the role of soil microbial communities in allelopathic research, such that their observed significance in soil sickness during continuous monocropping of fields can be further explored.     

Contribution of exudates,arbuscular mycorrhizal fungi and litter depositions to the rhizosphere priming effect induced by grassland species

The presence of plants induces strong accelerations in soil organic matter (SOM) mineralization by stimulating soil microbial activity – a phenomenon known as the rhizosphere priming effect (RPE). The RPE could be induced by several mechanisms including root exudates, arbuscular mycorrhizal fungi (AMF) and root litter. However the contribution of each of these to rhizosphere priming is unknown due to the complexity involved in studying rhizospheric processes. In order to determine the role of each of these mechanisms, we incubated soils enclosed in nylon meshes that were permeable to exudates, or exudates & AMF or exudates, AMF and roots under three grassland plant species grown on sand. Plants were continuously labeled with ¹³C depleted CO₂ that allowed distinguishing plant-derived CO₂ from soil-derived CO₂. We show that root exudation was the main way by which plants induced RPE (58–96% of total RPE) followed by root litter. AMF did not contribute to rhizosphere priming under the two species that were significantly colonized by them i.e. Poa trivialis and Trifolium repens. Root exudates and root litter differed with respect to their mechanism of inducing RPE. Exudates induced RPE without increasing microbial biomass whereas root litter increased microbial biomass and raised the RPE mediating saprophytic fungi. The RPE efficiency (RPE/unit plant-C assimilated into microbes) was 3–7 times higher for exudates than for root litter. This efficiency of exudates is explained by a microbial allocation of fresh carbon to mineralization activity rather than to growth. These results suggest that root exudation is the main way by which plants stimulated mineralization of soil organic matter. Moreover, the plants through their exudates not only provide energy to soil microorganisms but also seem to control the way the energy is used in order to maximize soil organic matter mineralization and drive their own nutrient supply.     

A conceptual framework and an empirical test of complementarity and facilitation with respect to phosphorous uptake by plant species mixtures

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.     

Manipulating the soil microbiome to increase soil health and plant fertility

A variety of soil factors are known to increase nutrient availability and plant productivity. The most influential might be the organisms comprising the soil microbial community of the rhizosphere, which is the soil surrounding the roots of plants where complex interactions occur between the roots, soil, and microorganisms. Root exudates act as substrates and signaling molecules for microbes creating a complex and interwoven relationship between plants and the microbiome. While individual microorganisms such as endophytes, symbionts, pathogens, and plant growth promoting rhizobacteria are increasingly featured in the literature, the larger community of soil microorganisms, or soil microbiome, may have more far-reaching effects. Each microorganism functions in coordination with the overall soil microbiome to influence plant health and crop productivity. Increasing evidence indicates that plants can shape the soil microbiome through the secretion of root exudates. The molecular communication fluctuates according to the plant development stage, proximity to neighboring species, management techniques, and many other factors. This review seeks to summarize the current knowledge on this topic.