Qualitative and quantitative analysis of water‐soluble root exudates in relation to plant species and development

The composition of root‐derived substances is of great importance for the understanding of processes in the rhizosphere. Therefore, methods allowing a comprehensive collection and chemical analysis of the organic root exudates are necessary. In this study, we compare different methods with regard to their suitability to collect and characterize root exudates. Because the percolation or water logging method failed to quantitatively extract root exudates, a dipping method was developed which allowed an almost complete sampling of coldwater‐soluble root exudates. By ¹⁴CO₂ labeling of the shoots the composition of root exudates was found to be influenced by plant species and growth stage. In comparison to pea plants maize plants had a higher share of carboxylic acids and a lower share of sugars. Younger maize plants exuded considerably higher amounts of ¹⁴C labeled organic substances per g root dry matter than older ones. During plant development the relative amount of sugars decreased at the expense of carboxylic acids. The described methods are well suited for the elucidation of the influence of growth factors on root exudation.     

Iron‐reducing root exudates: A modified microanalytical method

Interferences were detected in the PDTS method for the measurement of Fe‐reducing compounds exuded by plant roots into Hoagland nutrient solution. These were caused by a) excess NH₃ which is present in the ammonium acetate buffer used in the assay, which forms precipitates with the KH₂PO₄ especially when nutrient solution aliquots of larger than 2 mL are to be tested, and b) the overriding effect of autoreduction of Fe due to the very high concentration of FeCl₃ present in the assay medium. A modified method is presented whereby up to 20 mL of rooting solution can be tested and the interference due to autoreduction eliminated. Comparative measurements by the proposed and the original method show that the original method overestimates the amount of Fe‐reducing root exudates by Fe‐deficient plants by more than tenfold.     

A novel method for separating root‐derived organic compounds from root respiration in non‐sterilized soils

A novel method of separating exudates from root respiration in non‐sterilized soils has been developed. The method is based on a simultaneous elution of exudates from rhizosphere and the blowout of CO₂ originating from root respiration. The innovation of the method lies in the function of a membrane pump to drive the movement of air and simultaneously the circulation of water according to the Siphon principle. The separation method was tested by means of ¹⁴C pulse labeling of Lolium perenne to track the C dynamics in the production of rhizosphere CO₂ and of exudates, which were eluted. The total ¹⁴C activity of rhizosphere CO₂ and of eluted exudates was found to be 8.5 % and 2.3 % of total assimilated ¹⁴C, respectively. Thus, at least 19 % of root‐derived C can be accounted to root exudation. However, the suggested Siphon method underestimates the amount of exudates and shows only a minimum of organic substances exuded by roots. The diurnal dynamics of exudation was detected, but no significant day‐night changes were measured in root and microbial respiration. Tight coupling of assimilation with exudation, but not with root and microbial respiration, was observed. The advantages, shortcomings, and possible applications of the Siphon method are discussed.     

Release of phytosiderophores from roots of wheat and triticale under nickel‐deficient conditions

Despite numerous studies on phytosiderophores (PS) there is still an open question whether nickel (Ni) deficiency induces release of PS from graminaceous plant roots. Seedlings of two wheat cultivars (Triticum aestivum L. cvs. Rushan and Kavir) and a triticale cultivar (X. triticosecale) were grown in Ni‐free nutrient solution (Ni‐deficient, Ni–) and with 10 µM NiSO₄ (Ni‐sufficient, Ni+, control). Root exudates were collected weekly for 4 weeks and the amount of PS in the root exudates was measured. The response to Ni deficiency on the release of PS differed between species. Roots of Rushan and triticale exuded higher PS in response to Ni‐deficient conditions. Nickel deficiency significantly enhanced shoot Fe and Zn concentrations in wheat, while it decreased shoot Fe and Zn concentrations in triticale. In Kavir, PS exudation was decreased by Ni deficiency at weeks 3 and 4 and the reduced release of PS from roots of Kavir was accompanied by lower concentrations of Fe and Zn in plant roots but higher Fe and Zn concentrations in shoot tissue. The PS release by Kavir was triggered by a Ni‐induced Zn deficiency particularly in the shoots. According to the results, it is suggested that in the studies concerning the phytosiderophore release under Ni deficiency, special attention should be given to different responses among and within cereals and to the plant Zn or Fe nutritional status.     

Copper binding capacity of root exudates of cultivated plants and associated weeds

Cu binding to root exudates of two cultivated plants, wheat (Triticum aestivum) and rape (Brassica napus), and two weeds associated with wheat, dog daisy (Matricaria inodora) and cornflower (Centaurea cyanus), was studied in vitro under hydroponic and sterile conditions. Nutrient solutions were prepared with or without P. A MetPLATE microbiological test was used to assess the metal complexing capacity of root exudates. In the P-deficient solutions, no exudation was observed for any of the four plants; consequently, no Cu binding occurred. When P was present in the nutrient solutions, the plant exudates displayed differing abilities to complex Cu. No difference was detected in the binding capacity of the dog daisy or cornflower, and the blank [heavy metal binding capacity (HMBC)=1.07, 1.40 and 1.00, respectively]; however, the rape and wheat exudates were found to complex Cu in rhizospheric solutions (HMBC=1.73 and 3.00, respectively). The concentrations of exuded organic C were 1.2, 10.8, 15.3 and 15.7 mg l^-1 for the dog daisy, cornflower, wheat and rape, respectively. These results suggest that the nature, as well as the amount, of the organic compounds exuded by plant root, is important in determining the extent of Cu complexation.     

Plant rhizodeposition — an important source for carbon turnover in soils

The soil organic matter plays a key role in ecological soil functions, and has to be considered as an important CO₂ sink on a global scale. Apart from crop residues (shoots and roots), left over on the field after harvest, carbon and nitrogen compounds are also released by plant roots into the soil during vegetation, and undergo several transformation processes. Up to now the knowledge about amount, composition, and turnover of these root‐borne compounds is still very limited. So far it could be demonstrated with different plant species, that up to 20 % of photosynthetically fixed C are released into the soil during vegetation period. These C amounts are ecological relevant. Depending on assimilate sink strength during ontogenesis, the C release varies with plant age. A large percentage of these root‐borne substances were rapidly respired by microorganisms (64—86 %). About 2—5 % of net C assimilation was kept in soil. The root exudates of maize were mainly water‐soluble (79 %), and in this fraction about 64 % carbohydrates, 22 % amino acids/amides and 14 % organic acids could be identified. Plant species and in some cases also plant cultivars varied strongly in their root exudation pattern. Under non‐sterile conditions the exuded compounds were rapidly stabilized in water‐insoluble forms and bound preferably to the soil clay fraction. The binding of root exudates to soil particles also improved soil structure by increasing aggregate stability. Future research should focus on quantification and characterization of root‐borne C compounds during the whole plant ontogenesis. Apart from pot experiments with ¹⁴CO₂ labeling, it is necessary to conduct model field experiments with ¹³CO₂ labeling in order to be able to distinguish between CO₂ originating from the soil C pool and rhizosphere respiration, originating from plant assimilates. Such a separation is necessary to assess if soils are sources or sinks of CO₂. The incorporation of root‐borne C (¹⁴C, ¹³C) into soil organic matter of different stability is also of particular interest.     

Exudation of organic anions by roots of cabbage,carrot, and potato as influenced by environmental factors and plant age

A previous study demonstrated that cabbage was P efficient compared to carrot and potato. However, calculating plant P uptake by a mechanistic simulation model based on P transport by diffusion and mass flow, P uptake of roots according to the Michaelis‐Menten kinetics, and morphological root characteristics including root hairs, revealed that these parameters could explain only 2/5 of the total P uptake of cabbage, but 4/5 of that of carrot and potato (Dechassa et al., 2003). Therefore, it was hypothesized that a higher root exudation of organic anions may enhance P mobilization and hence P uptake of cabbage. The objective of this research was to determine root exudation of organic anions by the three species, and to investigate the influence of plant age and dark/light period on organic‐anion exudation by cabbage. Experiments were conducted in a growth chamber in nutrient solution with or without P. Organic anions were determined in root exudates and in root tissue. With cabbage and potato, P deficiency induced exudation of citrate and succinate, respectively. Citrate‐exudation rate of P‐deficient cabbage plants was correlated with accumulation of citrate in root tissue. In contrast, high succinate‐exudation rates in potato were not correlated with an increased concentration in root tissue. For carrot, no change was observed in the exudation of any of the organic anions in response to P deficiency. The results also showed that succinate‐ and citrate‐exudation rates of cabbage roots increased with increased plant age. There was also a significant increase in exudation rates of organic anions of cabbage roots during the light period of the day. It was concluded that cabbage had the ability to exude large amounts of citrate in response to P deficiency by which it can additionally enhance its P‐uptake efficiency, whereas carrot and potato showed little evidence of possessing such a mechanism.     

The role of root exudates in specific apple (Malus × domestica Borkh.) replant disease (SARD)

In an effort to improve our understanding of the specific apple replant disease (SARD), direct and indirect effects of phytohormones and related compounds (abscisic acid, 6‐benzyladenine, indole‐3‐acetic acid, 1‐naphthaleneacetic acid, and gibberellins GA₃ or GA₄) on root exudates of apple seedlings were evaluated as a potential mediating factor in the infection process. In the first type of experiments, radioactively labeled hormonal substances were applied to the stumps of decapitated apple‐seedlings and the occurrence of radioactivity in root exudates and their qualitative characterization were examined (direct influence). In another set of experiments, the effects of leaf‐applied plant growth regulators on the amount and composition of the predominant organic acids, carbohydrates, and amino acids/amides were studied. Cherry seedlings resistant to SARD and apple seedlings with dormant apical buds and, thus, not susceptible to infection were used for comparison. The results showed no differences in exudation of applied plant growth regulators between growing cherry and apple seedlings. Thus, a direct effect of plant hormones on the infection process is unlikely. However, leaf treatments with growth regulators, in particular with auxin‐type compounds and abscisic acid, increased exudation of alditols. This may indicate that plant hormones are, to some degree, indirectly involved in the infection process.     

Analysis of wild sunflower (Helianthus annuus L.) root exudates using gas chromatography‐mass spectrometry

Plant root systems mediate ecological processes in the rhizosphere through the exudation of organic compounds. Although exudate composition is thought to depend strongly on plant nutrient status, little is known about the influence of multi‐nutrient stresses. In this study, we examined responses to short‐term (3 d) nutrient limitation in Helianthus annuus (common sunflower), and root exudates were collected for 2, 4, or 6 h with the trap‐solution method. Root exudates, analyzed by means of gas chromatography‐mass spectrometry, consisted of over 60 sugars, sugar alcohols, amino acids, organic acids, and phosphates, with sugars and organic acids generally detected in the highest quantities. Twenty‐five of the detected metabolites, including half of the organic acids, sugars, and sugar alcohols, differed in relative abundance among the three sampling intervals, exhibiting higher abundance in sampling intervals greater than 2 h. Similarly, 24 of the detected metabolites, including half of the amino acids, phosphates, and sugar alcohols, were affected by nutrient supply, with 20 exhibiting higher abundance in the high‐nutrient treatment. Fumaric acid, quinic acid, and glucose were detected at significantly higher levels in the low‐nutrient treatment, potentially representing an adaptive response to nutrient limitation in sunflower. However, as sampling interval exerted a strong influence on the apparent effects of nutrient supply, future studies should consider the potential impacts of sampling‐interval length in comparative analyses of genotypes or treatments.     

Release of zinc mobilizing root exudates in different plant species as affected by zinc nutritional status

The effect of zinc nutritional status of the plant on the release of zinc mobilizing root exudates was studied in various dicotyledonous (apple, bean, cotton, sunflower, tomato) and graminaceous (barley, wheat) plant species grown in nutrient solutions. In all species, zinc deficiency increased root exudation of amino acids, sugars and phenolics. However, the root exudates of zinc deficient dicotyledonous species did not enhance zinc mobilization from a synthetic resin (Zn chelite), or a calcareous soil, although mobilization of iron from Fe^III hydroxide was increased. By contrast in the graminaceous species, root exudates from zinc deficient plants greatly increased mobilization of both zinc and iron from the various sources. These differences in capability of mobilization of zinc and iron between the plant species are the result of an enhanced release of phytosiderophores with zinc deficiency in the graminaceous species.     

Influence of microflora and composition of root bathing solution on root exudation of maize plants

Root exudation of carbon (C) plays a major role in processes occurring in the plant rhizosphere. Environmental factors affecting root exudation have been identified but their effects are rarely quantified. The purpose of this work was to evaluate the impact of both the microflora and the chemical composition of the growth medium on root exudation, taking into account soluble exudates and mucilage fraction. Maize plants (Zea mays L.) were grown for 12 days in hydroponic conditions and then transferred in three root bathing solutions (demineralized water, KCl or nutrient solution) during 24 hours. In each case, presence of microflora was tested with a comparison between plants inoculated with maize rhizospheric strain and axenic plants. Exudation was measured in terms of C and biomass production. A strong interaction was noticed between microflora and chemical composition of the root bathing solution. In fact, the presence of rhizospheric microflora induces a stimulation of soluble exudates only in KCl and Nutrient solutions. In demineralized water, a different response was observed with a higher C release for axenic plants, probably due to the osmotic shock induced to the roots. Concerning mucilage fractions, small quantities were recovered on all treatments. This work demonstrates that the chemical composition of the root bathing solution and presence of microorganisms significantly modify the amount of soluble exudates. Attention must therefore be paid to the cultural conditions when exudation is studied because of the sensitivity of this process to root environment.     

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.     

Cadmium Impact on Root Exudates of Sorghum and Maize Plants: A Speciation Study

ABSTRACT

The effect of cadmium (Cd) on root exudates of sorghum and maize was investigated in order to get further insight into the mechanisms of plant tolerance to Cd. Plants were grown hydroponically and supplemented with: 0, 0.5, and 5.0 mg Cd L^{? 1}. Hydroponic solutions containing exudates were analyzed by high performance liquid chromatography (HPLC). The results showed different exudation patterns by sorghum and maize with cadmium supply. While sorghum enhanced malate exudation over the entire range of applied Cd in the uptake solutions, maize increased mainly citrate. Moreover, malate concentration exuded in sorghum rhizosphere presented higher values than citrate (from maize). With the aid of the HYPERQUAD speciation program, a significant decrease in the bioavailable Cd (free Cd plus Cd chloro-complexes) was found due to the increase of Cd organic complexation in the hydroponic solution. Furthermore, similar metal organic complex concentrations were obtained for both plants, which turned the maize and sorghum overall detoxification process equivalent. Exudation of malate and citrate should contribute to tolerance mechanisms of these plants, reducing deleterious effects of free Cd on root growth. These findings support the idea that the metal-binding capabilities of root exudates may be an important mechanism for stabilizing metals in soil.     

Importance of rhizodeposition in the coupling of plant and microbial productivity

Plant roots influence the biological, chemical and physical properties of rhizosphere soil. These effects are a consequence of their growth, their activity and the exudation of organic compounds from them. In natural ecosystems, the linkages between inputs of carbon from plants and microbial activity driven by these inputs are central to our understanding of nutrient cycling in soil and the productivity of these systems. This coupling of plant and microbial productivity is also of increasing importance in agriculture, where the shift towards low‐input systems increases the dependence of plant production on nutrient cycling, as opposed to fertilizers. This review considers the processes by which plants can influence the cycling of nutrients in soil, and in particular the importance of organic inputs from roots in driving microbially mediated transformations of N. This coupling of plant inputs to the functioning of the microbial community is beneficial for acquisition of N by plants, particularly in low‐input systems. This occurs through stimulation of microbes that produce exoenzymes that degrade organic matter, and by promoting cycling of N immobilized in the microbial biomass via predation by protozoa. Also, plants increase the cycling of N by changes in exudation in response to nitrogen supply around roots, and in response to browsing by herbivores. Plants can release compounds in exudates that directly affect the expression of genes in microbes, and this may be an important way of controlling their function to the benefit of the plant.     

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.     

Low molecular weight organic acids released from roots of durum wheat and flax into sterile nutrient solutions

Knowledge of the composition and quantity of organic substances released from roots of different plant species is necessary for understanding the chemical and biological processes in the rhizosphere. The present study was undertaken to quantify low molecular weight organic acids (LMWOAs) released from roots of five cultivars/lines of durum wheat (Triticum turgidum var. durum L.): Kyle, Sceptre, DT618, DT627, and DT637 and four cultivars/ lines of flax (Linum usitatissiumum L.): Somme, Flanders, AC Emerson, and YSED 2. Plants were grown in sterile nutrient solution cultures and amounts of organic acids exuded by roots were analyzed by gas chromatography. The LMWOAs varied significantly among both durum wheat and flax cultivars and oxalic, malonic, fumaric, succinic, acetic, malic, citric and tartaric acids were detected in root exudates of both species. Generally, oxalic and acetic acids were predominant in durum wheat exudates and oxalic, acetic and malic acids were predominant in flax root exudates. High oxalic acid concentrations occurred in root exudates of durum wheat cultivars DT627 and DT637, and flax cultivar YSED 2. Compared with the other durum wheat cultivars, Kyle released the lowest total amount of LMWOAs, whereas among the flax cultivars, YSED 2 had the highest total amount of acids secreted from roots. The data showed that the release of LMWOAs from roots was cultivar dependent. The results provide valuable background information for studying the role of root exudates in soil‐plant relationships.     

大豆根分泌物活化难溶性铝磷的研究

在酸性红壤上，土壤有效磷含量低，大部分磷以难溶性磷形式存在，这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。本文系统研究了大豆根分泌物对难溶性铝磷的活化效果，同时运用阴、阳离子交换树脂将根分泌物分成阴离子组分、中性组分和阳离子组分，活化结果表明．阴离子组分对铝磷的活化量显著高于中性组分和阳离子组分。运用分子膜把根分泌物分成大于8K、8～3．5K、3．5～1K和小于1K组分．发现对铝磷活化量最大的组分为小于1K根分泌物组分。另外．小于1K阴离子组分对铝磷的活化量为缺磷13．2mg／pot，供磷可达9．3mg／pot，分别占总根分泌物活化量的71％和57％。运用离子色谱仪对小于1K阴离子组分根分泌物测定表明，供磷和缺磷处理．根系均能分泌少量的柠檬酸和苹果酸，约占铝处理的10％～20％，但处理之间差异不显著。上述研究表明．除有机酸影响难溶性铝磷活化之外，根分泌物中可能还存在其它物质对铝磷活化有促进作用，相关研究正在深入进行之中。     

Nitrate assay by multiple wavelength spectroscopy and analysis of kinetic parameters of net nitrate uptake by barley seedlings

A method for determination of nitrate concentration and estimation of kinetic parameters of nitrate uptake by spectroscopy based on absorbances at multiple wavelengths has been developed to estimate nitrate uptake by barley (Hordeum vulgare L. cv. Steptoe) seedlings. Nitrate concentration in the nutrient solution was determined from the slope of the linear regression line of the absorbances to nitrate absorption coefficients at 12 wavelengths. Interference by root exudates was only due to an absorption component changing with wavelength in correlation with nitrate absorption. The standard error of the determination decreased in reverse proportion to the square root of the number of the wavelengths. A linear form of the net uptake equation, NUR = I_maxC/K_m+C‐E, could be expressed as NUR = (I_max ‐ E) ‐ K_m NUR/C ‐ K_mE I/C where NUR is net uptake rate, Imax is maximum influx, C is concentration, Km is the Michaelis constant, and E is an efflux constant. The method described here was used to determine the time course of nitrate depletion by barley seedlings from their nutrient solution. The isotherm of net nitrate uptake rates derived from the time course was analyzed after modifications based on the linear form of the net uptake equation. The analysis yielded highly significant results (P<0.0001).     

The effect of mechanical injury on exudation from immature and mature peanut fruits under axenic conditions

Sugars and amino-N compounds exuded from axenically-grown peanut fruits following mechanical injury, were measured by gas-liquid and thin-layer chromatography. Injury of the peanut fruit surface resulted in a 10–20 fold increase in sugar exudation and an increase in the amount of amino N exuded. Analysis of leachates of the weblite (an exploded shale soil substitute) in which axenic fruits developed showed that exudation of sugars had occurred from uninjured fruits in situ. These results support the hypothesis that exudates in the geocarposphere are an important ecological factor affecting microbes in the soil and the colonization of peanut fruits by Aspergillus flarus.     

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.