Estimating N rhizodeposition of grain legumes using a N in situ stem labelling method

Grain legumes in crop rotations cause significant increases in yield for succeeding non-legumes, which cannot be explained simply by the small effect that legumes have on the soil nitrogen balance, as found in the analysis of N in crop residues. Besides known positive non-N-effects, other effects, mainly rhizodeposition and its contribution to the N balance and nitrogen dynamics after harvesting the grain, are poorly understood. In this study, N rhizodeposition, defined as root-derived N in the soil after removal of visible roots, was measured in faba bean (Vicia faba L.), pea (Pisum sativum L.) and white lupin (Lupinus albus L.). In a pot experiment the legumes were pulse labelled in situ with ¹⁵N urea using a cotton wick method. About 84% of the applied ¹⁵N was recovered for the three legume species at maturity. The ¹⁵N was comparatively uniformly distributed among plant parts. The N rhizodeposition constituted 13% of total plant N for faba bean and pea and 16% for white lupin at maturity, about 80% of below ground plant N, respectively. Some 7% (lupin)-31% (pea) of the total N rhizodeposits were recovered as micro-roots by wet sieving (200 μm) the soil after all visible roots had been removed. Only 14-18% of the rhizodeposition N was found in the microbial biomass and a very small amount of 3-7% was found in the mineral N fraction. In pea, 48% and in lupin 72% of N rhizodeposits could not be recovered in the mentioned pools and a major part of the unrecovered N was probably immobilised in microbial residues. The results of this study clearly indicate that N rhizodeposition from grain legumes represent a significant pool for N balance and N dynamics in crop rotations.     

How to calculate nitrogen rhizodeposition: a case study in estimating N rhizodeposition in the pea (Pisum sativum L.) and grasspea (Lathyrus sativus L.) using a continuous N labelling split-root technique

The formulae used in studies with ¹⁵N labelling techniques for estimating the N rhizodeposition (Ndfr) of legumes differ according to the background atom% ¹⁵N values used to determine ¹⁵N excess in the soil and roots grown in soil. Therefore, a continuous ¹⁵N labelling split-root experiment with pea (Pisum sativum L.) and grasspea (Lathyrus sativus L.) was undertaken and the relevant calculations were made to determine a valid method for calculating Ndfr. It is shown that a non-nodulated reference plant or a legume grown on soil without ¹⁵N labelling are required components of experiments which aim to estimate legume-N rhizodeposition, if the ¹⁵N abundance of the total soil N at the start of the experiment and that of the total plant available soil N are different. The standard formula was developed further to calculate Ndfr in a valid way. The impact of using different background atom% ¹⁵N values on the results when estimating Ndfr are demonstrated according to the ¹⁵N abundance of the roots grown in the soil. At physiological maturity, the rhizodeposition of N from roots grown in the soil was 19.8 mg N plant⁻¹ for pea and 14.1 mg N plant⁻¹ for grasspea, which is, respectively, equivalent to 10.5 and 9.2% of their total root and shoot N.     

Rhizodeposition of C and N in peas and oats after C-N double labelling under field conditions

Compounds released by plant roots during growth can make up a high proportion of below-ground plant (BGP) carbon and nitrogen, and therefore influence soil organic matter turnover and plant nutrient availability by stimulating the soil microorganisms. The present study was conducted to examine the amount and fate of C (CdfR) and N rhizodeposits (NdfR), in this study defined as root-derived C or N present in the soil after removal of roots and root fragments, released during reproductive growth. BGP biomass of peas (Pisum sativum L.) and oats (Avena sativa L.) was successfully labelled in situ with a ¹³C-glucose-¹⁵N-urea mixture under field conditions using a stem feeding method. Pea plants were labelled at the beginning of flowering and harvested 36 and 52 days after labelling at pod filling (P_P) and maturity (P_M), respectively. Oat plants were labelled at grain filling and harvested 42 days after labelling at maturity (O_M). CdfR was 24.2% (P_P), 29.6% (P_M) and 30.8% (O_M) of total recovered plant C. NdfR was 32.1% (P_P), 36.4% (P_M) and 30.0% (O_M) of total plant N. Due to higher N assimilation, amounts of NdfR were four times higher in peas in comparison with oats. The results for NdfR in peas were higher than results from other studies. The C-to-N ratio of rhizodeposits was lower under peas (17.3) than under oats (41.9) at maturity. At maturity, microbial CdfR at 0-30 cm soil depth was 37% of the microbial biomass C in peas and 59% in oats. Microbial NdfR was 15% of microbial N in peas and 5% in oats. Furthermore, inorganic NdfR was 34% in peas and 9% in oats at 0-30 cm at maturity. These results show that rhizodeposits of peas provide a more easily available substrate to soil microorganisms, which are incorporated to a greater extent and turned over faster in comparison with oats. Beside the higher amounts of N released from pea roots, this process contributes to the higher N-availability for subsequent crops.     

Does labelling frequency affect N rhizodeposition assessment using the cotton-wick method?

The aim of the present study was to test and improve the reliability of the ¹⁵N cotton-wick method for measuring soil N derived from plant rhizodeposition, a critical value for assessing belowground nitrogen input in field-grown legumes. The effects of the concentration of the ¹⁵N labelling solution and the feeding frequency on assessment of nitrogen rhizodeposition were studied in two greenhouse experiments using the field pea (Pisum sativum L.). Neither the method nor the feeding frequency altered plant biomass and N partitioning, and the method appeared well adapted for assessing the belowground contribution of field-grown legumes to the soil N pool. However, nitrogen rhizodeposition assessment was strongly influenced by the feeding frequency and the concentration of labelling solution. At pod-filling and maturity, despite similar root ¹⁵N enrichment, the fraction of plants' belowground nitrogen allocated to rhizodeposition in both Frisson pea and the non-nodulating isoline P2 was 20 to more than 50% higher when plants were labelled continuously than when they were labelled using fortnightly pulses. Our results suggest that when ¹⁵N root enrichment was high, nitrogen rhizodeposition was overestimated only for plants that were ¹⁵N-fed by fortnightly pulses, and not in plants ¹⁵N-fed continuously. This phenomenon was especially observed for plants that rely on symbiotic N₂ fixation for N acquisition, and it may be linked to the concentration of the labelling solution. In conclusion, the assessment of nitrogen rhizodeposition was more reliable when plants were labelled continuously with a dilute solution of ¹⁵N urea.     

Nitrogen rhizodeposition of young wheat plants under elevated CO<Subscript>2</Subscript> and drought stress

The objective of this study was to determine the effect of drought stress and elevated CO₂ concentrations around the shoots on N rhizodeposition of young wheat plants. In a pot experiment, the plant N pool was labeled through ¹⁵NH₃ application to shoots at nontoxic NH₃ concentrations, and the impact of low water supply (40% field capacity), elevated CO₂ (720 μmol mol⁻¹ CO₂), and the combination of both factors on the ¹⁵N distribution was studied. Total ¹⁵N rhizodeposition ranged from 5 to 11% of the total ¹⁵N recovered in the plant/soil system. Elevated CO₂ concentration as well as drought stress increased the belowground transport of N and increased the relative portion of N rhizodeposition on total ¹⁵N in the plant/soil system. However, while the increased N rhizodeposition with elevated CO₂ was the result of increased total belowground N transport, drought stress additionally increased the portion of ¹⁵N found in rhizodeposition vs roots. Elevated CO₂ intensified the effect of drought stress. The percentage of water soluble ¹⁵N in the ¹⁵N rhizodeposition was very low under all treatments, and it was significantly decreased by the drought-stressed treatments.     

Turnover of grain legume N rhizodeposits and effect of rhizodeposition on the turnover of crop residues

The turnover of N derived from rhizodeposition of faba bean (Vicia faba L.), pea (Pisum sativum L.) and white lupin (Lupinus albus L.) and the effects of the rhizodeposition on the subsequent C and N turnover of its crop residues were investigated in an incubation experiment (168 days, 15 °C). A sandy loam soil for the experiment was either stored at 6 °C or planted with the respective grain legume in pots. Legumes were in situ ¹⁵N stem labelled during growth and visible roots were removed at maturity. The remaining plant-derived N in soil was defined as N rhizodeposition. In the experiment the turnover of C and N was compared in soils with and without previous growth of three legumes and with and without incorporation of crop residues. After 168 days, 21% (lupin), 26% (faba bean) and 27% (pea) of rhizodeposition N was mineralised in the treatments without crop residues. A smaller amount of 15–17% was present as microbial biomass and between 30 and 55% of mineralised rhizodeposition N was present as microbial residue pool, which consists of microbial exoenzymes, mucous substances and dead microbial biomass. The effect of rhizodeposition on the C and N turnover of crop residues was inconsistent. Rhizodeposition increased the crop residue C mineralisation only in the lupin treatment; a similar pattern was found for microbial C, whereas the microbial N was increased by rhizodeposition in all treatments. The recovery of residual ¹⁵N in the microbial and mineral N pool was similar between the treatments containing only labelled crop residues and labelled crop residues + labelled rhizodeposits. This indicates a similar decomposability of both rhizodeposition N and crop residue N and may be attributable to an immobilisation of both N sources (rhizodeposits and crop residues) as microbial residues and a subsequent remineralisation mainly from this pool.Abbreviations C or N_dec C or N decomposed from residues - C or N_mic microbial C or N - C or N_micres microbial residue C or N - C or N_min mineralised C or N - C or N_input added C or N as crop residues and/or rhizodeposits - dfr derived from residues - dfR derived from rhizodeposition - Ndfr N derived from residues - NdfR N derived from rhizodeposition - N_loss losses of N derived from residues - SOM soil organic matter - WHC water holding capacity     

Rhizodeposition-induced decomposition increases N availability to wild and cultivated wheat genotypes under elevated CO2

Elevated CO₂ may increase nutrient availability in the rhizosphere by stimulating N release from recalcitrant soil organic matter (SOM) pools through enhanced rhizodeposition. We aimed to elucidate how CO₂-induced increases in rhizodeposition affect N release from recalcitrant SOM, and how wild versus cultivated genotypes of wheat mediated differential responses in soil N cycling under elevated CO₂. To quantify root-derived soil carbon (C) input and release of N from stable SOM pools, plants were grown for 1 month in microcosms, exposed to ¹³C labeling at ambient (392 μmol mol⁻¹) and elevated (792 μmol mol⁻¹) CO₂ concentrations, in soil containing ¹⁵N predominantly incorporated into recalcitrant SOM pools. Decomposition of stable soil C increased by 43%, root-derived soil C increased by 59%, and microbial-¹³C was enhanced by 50% under elevated compared to ambient CO₂. Concurrently, plant ¹⁵N uptake increased (+7%) under elevated CO₂ while ¹⁵N contents in the microbial biomass and mineral N pool decreased. Wild genotypes allocated more C to their roots, while cultivated genotypes allocated more C to their shoots under ambient and elevated CO₂. This led to increased stable C decomposition, but not to increased N acquisition for the wild genotypes. Data suggest that increased rhizodeposition under elevated CO₂ can stimulate mineralization of N from recalcitrant SOM pools and that contrasting C allocation patterns cannot fully explain plant mediated differential responses in soil N cycling to elevated CO₂.     

Field evaluation of N2 fixation and N partitioning in climbing bean (Phaseolus vulgaris L.) using 15N

Summary The common bean (Phaseolus vulgaris L.) is generally regarded as a poor N₂ fixer. This study assessed the sources of N (fertilizer, soil, and fixed N), N partitioning and mobilization, and soil N balance under field conditions in an indeterminate-type climbing bean (P. vulgaris L. cv. Cipro) at the vegetative, early pod-filling, and physiological maturity stages, using the A-value approach. This involved the application of 10 and 100 kg N ha^-1 of ¹⁵N-labelled ammonium sulphate to the climbing bean and a reference crop, maize (Zea mays L.). At the late pod-filling stage (75 days after planting) the climbing bean had accumulated 119 kg N ha^-1, 84% being derived from fixation, 16% from soil, and only 0.2% from the ¹⁵N fertilizer. N₂ fixation was generally high at all stages of plant growth, but the maximum fixation (74% of the total N₂ fixed) occurred during the interval between early (55 days after planting) and late podfilling. The N₂ fixed between 55 and 75 days after planting bas a major source (88%) of the N demand of the developing pod, and only about 11% was contributed from the soil. There was essentially no mobilization of N from the shoots or roots for pod development. The cultivation of common bean cultivars that maintain a high N₂-fixing capacity especially during pod filling, satisfying almost all the N needs of the developing pod and thus requiring little or no mobilization of N from the shoots for pod development, may lead to a net positive soil N balance.     

Influence of pea root traits modulating soil bioavailable C and N effects upon ammonification activity

Plant functional traits are useful tools for understanding plant impacts on soil nitrogen (N) mineralization. The objective of this study was to examine the root traits that govern the influence of Pisum sativum L. on potential protease and ammonification activities, which are two key microbial activities involved in N mineralization. Ammonification activity was greater during pea reproductive than vegetative stages, whereas potential protease activity did not vary along pea development. Ammonification activity was more strongly affected by root architecture traits (total root length and percentage of fine roots) than by root growth traits (root dry matter content). Pea root traits appear to affect ammonification activity in a complex manner involving variations in rhizodeposition that modulate carbon and N availability for soil microorganisms.     

Tree girdling provides insight on the role of labile carbon in nitrogen partitioning between soil microorganisms and adult European beech

Nitrogen (N) cycling in terrestrial ecosystems is complex since it involves the closely interwoven processes of both N uptake by plants and microbial turnover of a variety of N metabolites. Major interactions between plants and microorganisms involve competition for the same N species, provision of plant nutrients by microorganisms and labile carbon (C) supply to microorganisms by plants via root exudation. Despite these close links between microbial N metabolism and plant N uptake, only a few studies have tried to overcome isolated views of plant N acquisition or microbial N fluxes. In this study we studied competitive patterns of N fluxes in a mountainous beech forest ecosystem between both plants and microorganisms by reducing rhizodeposition by tree girdling. Besides labile C and N pools in soil, we investigated total microbial biomass in soil, microbial N turnover (N mineralization, nitrification, denitrification, microbial immobilization) as well as microbial community structure using denitrifiers and mycorrhizal fungi as model organisms for important functional groups. Furthermore, plant uptake of organic and inorganic N and N metabolite profiles in roots were determined.Surprisingly plants preferred organic N over inorganic N and nitrate (NO₃⁻) over ammonium (NH₄⁺) in all treatments. Microbial N turnover and microbial biomass were in general negatively correlated to plant N acquisition and plant N pools, thus indicating strong competition for N between plants and free living microorganisms. The abundance of the dominant mycorrhizal fungi Cenococcum geophilum was negatively correlated to total soil microbial biomass but positively correlated to glutamine uptake by beech and amino acid concentration in fine roots indicating a significant role of this mycorrhizal fungus in the acquisition of organic N by beech. Tree girdling in general resulted in a decrease of dissolved organic carbon and total microbial biomass in soil while the abundance of C. geophilum remained unaffected, and N uptake by plants was increased. Overall, the girdling-induced decline of rhizodeposition altered the competitive balance of N partitioning in favour of beech and its most abundant mycorrhizal symbiont and at the expense of heterotrophic N turnover by free living microorganisms in soil. Similar to tree girdling, drought periods followed by intensive drying/rewetting events seemed to have favoured N acquisition by plants at the expense of free living microorganisms.     

Mobilization of labelled N in soybean leaves during early podfill

Soybeans accumulate N in vegetative tissues up to pod initiation after which total vegetative N may remain constant during early phases of pod development. Eventually much of the vegetative N is mobilized to the pods. The mobilization of N from vegetative tissue to pods during the first few days of pod development is poorly understood but is important to an overall understanding of soybean N nutrition. The vegetative tissues of field grown soybeans were labelled with ¹⁵N and sampled weekly during the reproductive phase of plant growth. Three foliar applications of (¹⁵NH₄)₂SO₄ were made prior to pod initiation at a combined rate of 3.3 kg N/ha. To immobilize soil N and to increase soybean dependance on N₂‐fixation, sawdust was applied at a rate of 52 t ha^‐1 . Irrigation was required almost weekly because of a shallow soil profile and below normal summer precipitation. Mobilization of vegetative N began immediately upon pod initiation and continued at a linear rate through pod development. It appeared that N₂‐fixation was able to provide approximately half of the N in pods during early podfill. Nitrogen content of vegetative tissue declined as soon as pods began developing.     

Release of C and N from roots of peas and oats and their availability to soil microorganisms

Nutrient mobilisation in the rhizosphere is driven by soil microorganisms and controlled by the release of available C compounds from roots. It is not known how the quality of release influences this process in situ. Therefore, the present study was conducted to investigate the amount and turnover of rhizodeposition, in this study defined as root-derived C or N present in the soil after removal of roots and root fragments, released at different growth stages of peas (Pisum sativum L.) and oats (Avena sativa L.). Plants were grown in soil columns placed in a raised bed under outdoor conditions and simultaneously pulse labelled in situ with a ¹³C-glucose-¹⁵N-urea solution using a stem feeding method. After harvest, ¹³C and ¹⁵N was recovered in plant parts and soil pools, including the microbial biomass. Net rhizodeposition of C and N as a percentage of total plant C and N was higher in peas than in oats. Moreover, the C-to-N ratio of the rhizodeposits was lower in peas, and a higher proportion of the microbial biomass and inorganic N was derived from rhizodeposition. These results suggest a positive plant-soil feedback shaping nutrient mobilisation. This process is driven by the C and N supply of roots, which has a higher availability in peas than in oats.     

Cutting regime determines allocation of fixed nitrogen in white clover

Leguminous leys are important sources of nitrogen (N), especially in forage-based animal production and organic cropping. Models for estimating total N₂ fixation of leys—including below-ground plant-derived N (BGN)—are based on grazed or harvested leys. However, green manure leys can have different proportions of above-ground plant-derived N (AGN) and BGN when subjected to different cutting regimes. To investigate the effects of cutting on N distribution in white clover, a pot experiment was carried out using ¹⁵N techniques to determine N₂ fixation, N rhizodeposition and root C and N content of cut and uncut white clover (Trifolium repens L. cv. Ramona) plants. Percentage N derived from air (%Ndfa) was lower in uncut (63%) than in cut (72%) plants, but total Ndfa was not significantly affected by cutting. The higher reliance on N₂ fixation in cut plants was thus counterbalanced by lower biomass and total N content. With BGN taken into account, total plant-derived N increased by approximately 50% compared with AGN only. Cutting did not affect the proportion of BGN to standing shoot biomass N after regrowth, but decreased the proportion of BGN to total shoot biomass production during the entire growth period. Thus, estimates of N fixation in green manure leys should consider management practices such as cutting regime, as this can result in differences in above- and below-ground proportions of plant-derived N.     

大豆植株苗期至结荚初期对肥料氮的吸收与分配

采用砂培与15N分阶段标记的方法,研究了苗期至结荚初期大豆植株对氮素的同化吸收,尤其是肥料氮在不同器官和节中的积累与分配。结果表明:(1)苗期至结荚初期大豆植株氮素积累量逐渐增加,由标记试验Ⅰ的64.08g/plant逐渐增加到标记试验Ⅴ的307.17mg/plant;(2)不同器官中肥料氮积累存在差异,叶是苗期至结荚初期肥料氮积累的主要器官,所占比例为55.2%,根系次之为25.9%,茎最小为18.9%;(3)苗期至结荚初期肥料氮积累量和所占比例逐渐减小,而根瘤固氮则逐渐增加,其中苗期至盛花期肥料氮是大豆植株氮素主要来源,盛花期以后以根瘤固氮为主;(4)不同成长程度叶对肥料氮的积累存在差异,表现为成长中叶>新生叶>成熟叶;(5)新生茎叶氮素构成随全株氮素构成的变化而变化,苗期至初花期以肥料氮为主,初花期至结荚初期根瘤固氮逐渐转为主体。     

N form and concentration: Effects on reproductive development,growth and N content of southern peas 1

Southern peas [Vigna unguiculata, (L.) Walp.] cultured with 100% NH⁺ ₄ produced no viable flowers, while treatments in which NO^‐ ₃ composed 50% or more of the N form were not significantly different in the number of flowers formed. Flower abortion was least with 100% NO^‐ ₃ at the lower N concentration and with 75% and 100% NO^‐ ₃ at the higher N concentration. Further increments of NH⁺ ₄ resulted in greater flower abortion. The trends in flower survival were reflected in the number of pods and number of seed/plant. At the lower N concentration, the addition of NH⁺ ₄ slowed pod maturity, while at the higher N concentration pod maturity was hastened with the addition of up to 50% NH⁺ ₄. The dry weight and N content of tissues were generally greater with the higher N concentration and with N combinations containing predominantly NO^‐ ₃, but trends varied with the plant part being analyzed. Ammonium appears to adversely influence reproductive development and/or NO^‐ ₃ is essential to complete the reproductive development of southern peas. The observed differences in the response of southern peas to N form may account for previously reported discrepancies concerning the effectiveness of N fertilization on growth and yield parameters. Also, vegetative growth and vegetative N content appear to be poor indicators of final seed yields of southern peas if NH⁺ ₄ supplies a significant portion of the N form utilized by the plant.     

Fate of C- and N-labelled rhizodeposition of Lolium perenne as function of the distance to the root surface

A greenhouse rhizobox experiment was carried out to quantify the incorporation of ¹³C- and ¹⁵N-labelled rhizodeposits into different soil pools, especially into the rhizosphere microbial biomass, with increasing distances to the root surface of Lolium perenne. Five layers were analysed over 0-4.2 mm distance to an artificial root surface. C and N derived from rhizodeposition were 4.2% of total C and 2.8% of total N in soil at 0-1.0 mm distance and decreased rapidly with increasing distance. Microbial biomass C and N increased significantly towards the roots. At 0-1.0 mm distance microbial biomass C and N accounted for 66% and 29% of C and N derived from rhizodeposition, respectively. These percentages declined with increasing distance to the roots, but were still traceable up to 4.2 mm distance. Only small amounts of root released C and N were found in the 0.05 M K₂SO₄-extractable fraction. Extractable C and N derived from rhizodeposition varied around means of 4% of total C and N derived from rhizodeposition and increased only marginally with increasing distance to the roots. C derived from rhizodeposition in the non-extractable soil organic matter increased from 65 to 89% of total C derived from rhizodeposition at 0-3.4 mm distance. Conversely, microbial biomass C derived from rhizodeposition decreased from 33 to 4%. N derived from rhizodeposition in the non-extractable soil organic matter increased from 61 to 79% of total N derived from rhizodeposition at 0-2.6 mm distance, followed by a decline to roughly 55% in the two outer layers. Microbial biomass N decreased from 37 to 16% at 0-2.6 mm distance, followed by an increase to roughly 41% in the two outer layers. The C/N ratio of total C and N derived from rhizodeposition as well as that of extractable C and N derived from rhizodeposition increased with increasing distance to the roots to values above 30. In contrast, the C/N ratio of incorporated rhizodeposition C and N into the microbial biomass decreased to values less than 5 at 2.6-4.2 mm distance. The data indicate differential microbial response to C and N derived from rhizodeposition at a high spatial resolution from the root surface. The turnover of C and N derived from rhizodeposition in the rhizosphere as a function of the distance to the root surface is discussed.     

A model to predict the accuracy of measurements of legume N rhizodeposition using a split-root technique

Legume N rhizodeposition is an important process for understanding the N turnover in legume-based cropping systems. Different ¹⁵N labeling techniques have been developed to estimate the rhizodeposition of legume-derived N into the soil. However, it is not known how the ¹⁵N-based experiments have to be designed to achieve a defined degree of accuracy in measuring the amount of N derived from rhizodeposition (Ndfr). As a consequence therefore, a model for the split-root technique was developed on the basis of experimental data to (i) test the effects of various experimental conditions on soil ¹⁵N enrichment, (ii) evaluate the accuracy of the measurements, and to (iii) deduce instructions for designing efficient experiments using ¹⁵N techniques for quantifying legume N rhizodeposition. It turned out that the coefficient of disproportional ¹⁵N distribution between non-¹⁵N-fed roots grown in soil (Root_Soil) and total plant biomass (D coefficient) is an indispensable component in the development of such a model for ¹⁵N-based split-root experiments. The model showed the coefficients of variation in measuring the Ndfr with regard to the analytical accuracy in determining not only the isotope composition of both the Root_Soil and the soil, but also the N content of the soil itself. Suggestions for designing specific experimental conditions to achieve a high accuracy in quantifying Ndfr were deduced from the model for the split-root technique, particularly in the choice of the amount of soil N at the start of the experiment and the ¹⁵N enrichment of the fertilizer being used. The coefficients of variation in measuring Ndfr are presented regarding the ¹⁵N abundance of Root_Soil and the quotient of the amount of Ndfr and soil N at the start of the experiment.     

Effect of moisture stress on Nitrate Reductase Activity (NRA) in PEA (Pisum sativum L.) [C3] and sorghum (Sorghum vulgare Pers.) [C4] plants

Nitrate reductase activity (NRA) was studied in pea, a C₃ plant, and sorghum, a C₄ plant, at various stages of growth and development. Influence of moisture stress and nitrogen application was also observed since these factors have profound influence on growth and development.

In pea, NRA was maximum at pod maturity stage and minimum at flowering stage. In sorghum plant there was gradual increase in NRA upto grain formation followed by a fall in activity at maturity.

Nitrogen treatment as nitrate and ammonia significantly increased nitrate reductase activity over control in both pea and sorghum. Treatment with potassium nitrate was found to stimulate more NRA in pea than with ammonium sulphate. In sorghum, both forms of nitrogen did not differ much in their influence on NRA.

Influence of moisture stress in reducing NRA was more clear in sorghum, a C₄ plant than in pea, a C₃ plant. In general, control plants recorded low NRA in both the crops when compared to nitrogen treated plants except at pod formation stage in pea.     

CO2 and inorganic N supply modify competition for N between co-occurring grass plants, tree seedlings and soil microorganisms

Plant-plant and plant-soil interactions play a key role in determining plant community structure and ecosystem function. However, the effects of global change on the interplay between co-occurring plants and soil microbes in successional communities are poorly understood. In this study, we investigated competition for nitrogen (N) between soil microorganisms, grass plants and establishing tree seedlings under factorial carbon dioxide (CO₂) and N treatments. Fraxinus excelsior seedlings were germinated in the presence or absence of grass competition (Dactylis glomerata) at low (380 μmol mol⁻¹) or high (645 μmol mol⁻¹) CO₂ and at two levels of N nutrition in a mesocosm experiment. Pulse ¹⁵N labelling was used to examine N partitioning among plant and soil compartments. Dactylis exerted a strong negative effect on Fraxinus biomass, N capture and ¹⁵N recovery irrespective of N and CO₂ treatment. In contrast, the presence of Dactylis had a positive effect on the microbial N pool. Plant and soil responses to N treatment were of a greater magnitude compared with responses to elevated CO₂, but the pattern of Fraxinus- and microbial-N pool response to N and CO₂ varied depending on grass competition treatment. Within the Dactylis competition treatment, decreases in Fraxinus biomass in response to N were not mirrored by decreases in tree seedling N content, suggesting a shift from below- to above-ground competition. In the Dactylis-sown pots, ¹⁵N recovery could be ranked Dactylis > microbial pool > Fraxinus in all N and CO₂ treatment combinations. Inequalities between Fraxinus and soil microorganisms in terms of ¹⁵N recovery were exacerbated by N addition. Contrary to expectations, elevated CO₂ did not increase plant-microbe competition. Nevertheless, microbial ¹⁵N recovery showed a small positive increase in the high CO₂ treatment. Overall, elevated CO₂ and N supply did not interact on plant/soil N partitioning. Our data suggest that the competitive balance between establishing tree seedlings and grass plants in an undisturbed sward is relatively insensitive to CO₂ or N-induced modifications in N competition between plant and soil compartments.     

Pasture type and fertilization effects on N2 fixation, N budgets and external energy inputs in western Canada

A grazing experiment was conducted in Brandon, Manitoba, Canada. The objectives were to examine the effects of including alfalfa and fertilizer management on N₂ fixation by alfalfa and plant N dynamics, and to compare N budgets in the four contrasting pasture systems and external energy inputs between fertilizer-N-based and legume-based pasture systems. Estimates of annual amounts of N₂ fixed, based on shoot herbage production in grazed mixed alfalfa/grass pastures, ranged from 40 to 118 kg N ha⁻¹ y⁻¹. The amounts would be in the range of 52-153 kg N ha⁻¹ y⁻¹, if the amounts of fixed N stored in the roots, were included. Compared to grass-only pastures, total amounts of N₂ fixed in the mixed pastures should be sufficient to improve total external N inputs, replace N fertilizer and sustain plant protein for grazing. The reliance of alfalfa (Medicago sativa L.) on N₂ fixation for growth was high (70-95%), and %N derived from the atmosphere by alfalfa (%Ndfa) was not affected by P fertilizer management. Thus, the amounts of N₂ fixed were predominantly regulated by alfalfa dry matter productivity. The data also indicated that alfalfa fixed 27 kg N t⁻¹ dry matter produced. In mixed alfalfa/grass pastures, high soil mineral N uptake by companion grasses, was essential to effectively utilize N that was fixed by alfalfa and returned to soils through the decomposition of alfalfa litter and roots. Compared to grass-only pastures with or without N fertilizer, alfalfa-based pastures could supply sufficient plant protein for grazing animals through N₂ fixation, and at same time, sustain animal productivity with only 28% of the external energy input of the grass-only pasture with N fertilizer.