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.     

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.     

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.     

Rhizodeposition: Its contribution to microbial growth and carbon and nitrogen turnover within the rhizosphere

A greenhouse rhizobox experiment was carried out to investigate the fate and turnover of ¹³C‐ and ¹⁵N‐labeled rhizodeposits within a rhizosphere gradient from 0–8 mm distance to the roots of wheat. Rhizosphere soil layers from 0–1, 1–2, 2–3, 3–4, 4–6, and 6–8 mm distance to separated roots were investigated in an incubation experiment (42 d, 15°C) for changes in total C and N and that derived from rhizodeposition in total soil, in soil microbial biomass, and in the 0.05 M K₂SO₄–extractable soil fraction. CO₂‐C respiration in total and that derived from rhizodeposition were measured from the incubated rhizosphere soil samples. Rhizodeposition C was detected in rhizosphere soil up to 4–6 mm distance from the separated roots. Rhizodeposition N was only detected in the rhizosphere soils up to 3–4 mm distance from the roots. Microbial biomass C and N was increased with increasing proximity to the separated roots. Beside ¹³C and ¹⁵N derived from rhizodeposits, unlabeled soil C and N (native SOM) were incorporated into the growing microbial biomass towards the roots, indicating a distinct acceleration of soil organic matter (SOM) decomposition and N immobilization into the growing microbial biomass, even under the competition of plant growth. During the soil incubation, microbial biomass C and N decreased in all samples. Any decrease in microbial biomass C and N in the incubated rhizosphere soil layers is attributed mainly to a decrease of unlabeled (native) C and N, whereas the main portion of previously incorporated rhizodeposition C and N during the plant growth period remained immobilized in the microbial biomass during the incubation. Mineralization of native SOM C and N was enhanced within the entire investigated rhizosphere gradient. The results indicate complex interactions between substrate input derived from rhizodeposition, microbial growth, and accelerated C and N turnover, including the decomposition of native SOM (i.e., rhizosphere priming effects) at a high spatial resolution from the roots.     

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.     

Microbial immobilisation and turnover of N labelled substrates in two arable soils under field and laboratory conditions

Microbial biomass N dynamics were studied under field and laboratory conditions in soils of high yield (HY) and low yield (LY) areas in an agricultural field. The objective of the study was to determine the size and activity of soil microbial biomass in the soils of the different yield areas and to compare these data obtained under field and laboratory conditions. Soils were amended with ¹⁵N labelled mustard (Sinapis alba) residues (both experiments) and labelled nitrate (laboratory only) at 30 μg N g⁻¹ dry soil. Soil microbial biomass (SMB) N, mineral N (N_min) and total N content was monitored both in the field and in the laboratory. N₂O efflux was additionally measured in laboratory treatments. Isotope ratios were determined for SMB in both experiments, for all other parameters only in the laboratory treatments. In the laboratory less amounts of added substrate N were immobilised by the SMB in HY soils compared to LY soils, whereas in the field immobilisation of added N by SMB was higher in HY soils initially and slightly lower after 40 days of incubation. Calculated turnover times in the laboratory nitrate, laboratory mustard and field mustard amendments were 0.18, 0.27 and 0.74 years (HY) and 0.22, 0.61 and 1.01 years (LY), respectively. The turnover times of added substrate N always showed the trend to be faster in HY soils compared to LY soils. A faster turnover of nutrients in the HY soils may involve a better nutrient supply of the plants, which coincides with the higher agricultural yield observed in these areas.     

Determination of the fate of C labelled maize and wheat rhizodeposit-C in two agricultural soils in a greenhouse experiment under C-CO2-enriched atmosphere

A deeper understanding of the contribution of carbon (C) released by plant roots (rhizodeposition) to soil organic matter (SOM) can help to increase our knowledge of global C-cycling. These insights can eventually lead to sustainable management of SOM especially in agricultural systems. This study was conducted to determine the fate of ¹³C labelled rhizodeposit-C of maize and wheat plants. They were grown in a greenhouse in permeable nylon bags filled with upper soil material from two agricultural soils of the same location, but with different crop yields. The bags were placed into pots, which were also filled with soil surrounding the bags. Soil inside the bags was considered as rhizosphere soil, wheras the one outside the bags represented bulk soil. The contributions of rhizodeposits to water extractable organic carbon (WEOC), microbial biomass-C (MB-C), CO₂-C evolution, and total organic carbon (C_org) were investigated during a 7-week growing period. The WEOC, MB-C, CO₂-C, C_org contents and the respective δ¹³C values were determined regularly, and a newly developed method for determining δ¹³C values in soil extracts was applied.In both soils, regardless of crop yield potential, significant incorporation of rhizodeposition-derived C was observed in the MB-C, CO₂-C, and C_org pool, but not in the WEOC. The pattern of C incorporation into the different pools was the same for both soils with both plants, and rhizodeposit-derived C was recovered in the order MB-C<C_org<CO₂-C. This showed that rhizodeposits were mainly respired, but since C_org was the second largest pool of the overall balances, they were also stabilized in the soils at least in the short term. It is suggested that the increased SOM mineralization observed in this study (positive priming effects) was probably induced by C exchange processes between the soil matrix and soluble rhizodeposits. Moreover, soluble rhizodeposit-C was detected in MB-C and CO₂-C evolved outside the direct root zone, showing the availability of these C-components in the bulk soil.     

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 availability to grain sorghum from organic and inorganic sources on sandy and clayey soil surfaces in a greenhouse pot study

In a greenhouse pot study, we examined the availability of N to grain sorghum from organic and inorganic N sources. The treatments were¹⁵N-labeled clover residues, wheat residues, and fertilizer placed on a sandy clay loam and loamy sand soil surface for an 8-week period. Soil aggregates formed under each soil texture were measured after 8 weeks for each treatment. Significantly greater ¹⁵N was taken up and recovered by grain sorghum in sandy clay loam pots compared with loamy sand pots. Greater ¹⁵N recovery was consistently observed with the inorganic source than the organic sources regardless of soil texture or time. Microbial biomass C and N were significantly greater for sandy clay loam soil compared with the loamy sand. Microbial biomass ¹⁵N was also significantly greater in the sandy clay loam treatment compared to the loamy sand. The fertilizer treatment initially had the greatest pool of microbial biomass ¹⁵N but decreased with time. The crop residue treatments generally had less microbial biomass ¹⁵N with time. The crop residues and soil texture had a significant effect on the water-stable aggregates formed after 8 weeks of treatments. Significantly greater water-stable aggregates were formed in the sandy clay loam than the loamy sand. Approximately 20% greater water-stable aggregates were formed under the crop residue treatments compared to the fertilizer only treatment. Soil texture seemed to be one of the most important factors affecting the availability of N from organic or inorganic N sources in these soils.Contribution from the MissouriAgricultural Experiment Station, Journal Series No.12131     

Characterization of the N benefit of a grain legume (Lupinus angustifolius L.) to a cereal (Hordeum vulgare L.) by an in situ 15N isotope dilution technique

Summary A crop of barley was grown on plots which had previously supported pure stands of lupins, canola, ryegrass, and wheat. The plots were labelled with ¹⁵N-enriched fertilizers at the time of sowing of the antecedent crops. The crop of lupins, which derived 79% of its N from symbiotic N₂ fixation at physiological maturity, conferred an N benefit to barley of 3.4 g N m^-2 when compared to barley following wheat. Lupins used less fertilizer N and less unlabelled soil N compared to the other crops, but the ratios of these sources of N in the plant tops were similar. The apparent sparing of soil+fertilizer N under lupins compared with wheat was 13.6 g N m^-2, which was much larger than the measured N benefit. Barley following lupins was less enriched in ¹⁵N compared to barley following wheat, and the measured isotope dilution was used to estimate the proportion of barley N derived from biologically fixed N in the lupin residues. This in turn enabled the N benefit to be partitioned between the uptake of spared N and the uptake of fixed N derived from the mineralization of legume residues. Spared N and fixed N contributed in approximately equal proportions to the N benefit measured in barley following lupins compared to barley following wheat.     

Frequent addition of wheat straw residues to soil enhances carbon mineralization rate

In many ecosystems, residues are added frequently to soil, in the form of root turnover and litter fall. However, in most studies on residue decomposition, residues are added once and there are few studies that have investigated the effect of frequent residue addition on C mineralization and N dynamics. To close this knowledge gap, we mixed mature wheat residue (C/N 122) into soil at a total rate of 2% w/w once at the start (R1×), every 16 days (R4×), every 8 days (R8×) or every 4 days (R16×). Un-amended soil served as control. All treatments were mixed every 4 days. Soil respiration was measured continuously over the 80-day incubation. Inorganic N, K₂SO₄-extractable C and N, chloroform-labile C and N (as an estimate of microbial biomass C and N), soil pH and microbial community composition were assessed every 16 days. Increasing frequency of residue addition increased C mineralization per g residue. Compared to R1×, cumulative respiration per g residue at the end of the incubation (day 80) was increased by 57, 82 and 92% in R4×, R8× and R16×, respectively. The largest differences in soil respiration per g residue occurred in the first 30 days. Despite large increases in cumulative respiration, frequent residue addition did not affect inorganic N or K₂SO₄-extractable N concentrations, chloroform-labile C and N or soil pH. Compared to the control, all residue treatments resulted in increases in chloroform-labile C and N and soil pH but decreased inorganic and K₂SO₄-extractable N. Microbial community composition was affected by residue addition, however there were no consistent differences among residue treatments. It is concluded that experiments with single residue additions may underestimate residue decomposition rates in the field. The increased C mineralization caused by frequent residue additions does not appear to be due to an increased microbial biomass or changes in microbial community composition, but rather to increased C mineralization per unit biomass.     

Microbial use of maize cellulose and sugarcane sucrose monitored by changes in the C/C ratio

An arable soil with organic matter formed from C₃-vegetation was amended initially with maize cellulose (C₄-cellulose) and sugarcane sucrose (C₄-sucrose) in a 67-day laboratory incubation experiment with microcosms at 25 °C. The amount and isotopic composition (¹³C/¹²C) of soil organic C, CO₂ evolved, microbial biomass C, and microbial residue C were determined to prove whether the formation of microbial residues depends on the quality of the added C source adjusted with NH₄NO₃ to the same C/N ratio of 15. In a subsequent step, C₃-cellulose (3 mg C g⁻¹ soil) was added without N to soil to determine whether the microbial residues formed initially from C₄-substrate are preferentially decomposed to maintain the N-demand of the soil microbial community. At the end of the experiment, 23% of the two C₄-substrates added was left in the soil, while 3% and 4% of the added C₄-cellulose and C₄-sucrose, respectively, were found in the microbial biomass. The addition of the two C₄-substrates caused a significant 100% increase in C₃-derived CO₂ evolution during the 5-33 day incubation period. The addition of C₃-cellulose caused a significant 50% increase in C₄-derived CO₂ evolution during the 38-67 day incubation period. The decrease in microbial biomass C₄-C accounted for roughly 60% of this increase. Cellulose addition promoted microorganisms strongly able to recycle N immediately from their own tissue by “cryptic growth” instead of incorporating NO₃⁻ from the soil solution. The differences in quality of the microbial residues produced by C₄-cellulose and C₄-sucrose decomposing microorganisms are also reflected by the difference in the rates of CO₂ evolution, but not in the rates of net N mineralization.     

Priming effects: Interactions between living and dead organic matter

In this re-evaluation of our 10-year old paper on priming effects, I have considered the latest studies and tried to identify the most important needs for future research. Recent publications have shown that the increase or decrease in soil organic matter mineralization (measured as changes of CO₂ efflux and N mineralization) actually results from interactions between living (microbial biomass) and dead organic matter. The priming effect (PE) is not an artifact of incubation studies, as sometimes supposed, but is a natural process sequence in the rhizosphere and detritusphere that is induced by pulses or continuous inputs of fresh organics. The intensity of turnover processes in such hotspots is at least one order of magnitude higher than in the bulk soil. Various prerequisites for high-quality, informative PE studies are outlined: calculating the budget of labeled and total C; investigating the dynamics of released CO₂ and its sources; linking C and N dynamics with microbial biomass changes and enzyme activities; evaluating apparent and real PEs; and assessing PE sources as related to soil organic matter stabilization mechanisms. Different approaches for identifying priming, based on the assessment of more than two C sources in CO₂ and microbial biomass, are proposed and methodological and statistical uncertainties in PE estimation and approaches to eliminating them are discussed. Future studies should evaluate directions and magnitude of PEs according to expected climate and land-use changes and the increased rhizodeposition under elevated CO₂ as well as clarifying the ecological significance of PEs in natural and agricultural ecosystems. The conclusion is that PEs - the interactions between living and dead organic matter - should be incorporated in models of C and N dynamics, and that microbial biomass should regarded not only as a C pool but also as an active driver of C and N turnover.     

Nitrous oxide emissions after incorporation of winter oilseed rape (Brassica napus L.) residues under two different tillage treatments

The aim of this study was to investigate the effect of crop residues from winter oilseed rape on N₂O emissions from a loamy soil and to determine the effect of different tillage practices on N₂O fluxes. We therefore conducted a field experiment in which crop residues of winter oilseed rape (Brassica napus L., OSR) were replaced with ¹⁵N labelled OSR residues. Nitrous oxide (N₂O) emissions and ¹⁵N abundance in the N₂O were determined for a period of 11 months after harvest of OSR and in the succeeding crop winter wheat (Triticum aestivum L.) cultivated on a Haplic Luvisol in South Germany. Measurements were carried out with the closed chamber method in a treatment with conventional tillage (CT) and in a treatment with reduced soil tillage (RT). In both tillage treatments we also determined N₂O fluxes in control plots where we completely removed the crop residues. High N₂O fluxes occurred in a short period just after OSR residue replacement in fall and after N‐fertilization to winter wheat in the following spring. Although N₂O emissions differed for distinct treatments and sub‐periods, cumulative N₂O emissions over the whole investigation period (299 days) ranged between 1.7 kg and 2.4 kg N₂O‐N ha^?1 with no significant treatment effects. More than half of the cumulative emissions occurred during the first eight weeks after OSR replacement, highlighting the importance of this post‐harvest period for annual N₂O budgets of OSR. The contribution of residue N to the N₂O emission was low and explained by the high C/N‐ratio fostering immobilization of mineral N. In total only 0.03% of the N₂O‐N emitted in the conventional tillage treatment and 0.06% in the reduced tillage treatment stemmed directly from the crop residues. The ¹⁵N recovery in the treatments with crop residues was 62.8% (CT) and 75.1% (RT) with more than 97% of the recovered ¹⁵N in the top soil. Despite our measurements did not cover an entire year, the low contribution of the OSR residues to the direct N₂O emissions shows, that the current IPCC tier 1 approach, which assumes an EF of 1%, strongly overestimated direct emissions from OSR crop residues. Furthermore, we could not observe any relationship between tillage and crop residues on N₂O emission, only during the winter period were N₂O emissions from reduced tillage significantly higher compared to conventional tillage. Annual N₂O emission from RT and CT did not differ.     

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.     

Nitrogen immobilization and mineralization during initial decomposition of15N-labelled pea and barley residues

The immobilization and mineralization of N following plant residue incorporation were studied in a sandy loam soil using¹⁵N-labelled field pea (Pisum sativum L.) and spring barley (Hordeum vulgare L.) straw. Both crop residues caused a net immobilization of soil-derived inorganic N during the complete incubation period of 84 days. The maximum rate of N immobilization was found to 12 and 18 mg soil-derived N g^–1 added C after incorporation of pea and barley residues, respectively. After 7 days of incubation, 21% of the pea and 17% of the barley residue N were assimilated by the soil microbial biomass. A comparison of the¹⁵N enrichments of the soil organic N and the newly formed biomass N pools indicated that either residue N may have been assimilated directly by the microbial biomass without entering the soil inorganic N pool or the biomass had a higher preference for mineralized ammonium than for soil-derived nitrate already present in the soil. In the barley residue treatment, the microbial biomass N was apparently stabilized to a higher degree than the biomass N in the pea residue treatment, which declined during the incubation period. This was probably due to N-deficiency delaying the decomposition of the barley residue. The net mineralization of residue-derived N was 2% in the barley and 22% in the pea residue treatment after 84 days of incubation. The results demonstrated that even if crop residues have a relative low C/N ratio (15), transient immobilization of soil N in the microbial biomass may contribute to improved conservation of soil N sources.     

Plant-N incorporation into microbial amino sugars as affected by inorganic N addition: A microcosm study of N-labeled maize residue decomposition

Carbon (C) and/or nitrogen (N) in plant residues can be assimilated into microbial biomass during the plant residue decomposition before incorporation into SOM in the form of microbial residues. Yet, microbial transformation of plant residue-N into microbial residues and the effects of inorganic N inputs on this process have not been well documented. Here, we undertook a 38-week incubation with a silt loam soil amended with a ¹⁵N-labeled maize (Zea mays L.) residue to determine how the transformation of maize residue-N into soil amino sugars was affected by rates of inorganic N addition. The newly metabolized amino sugars derived from maize residue-N were differentiated and quantified by using an isotope-based gas chromatography-mass spectrometry technique. We found that greater amounts of maize residue-N were transformed into amino sugars with lower inorganic N addition at the early stages of the plant residue degradation. However, the trend was reversed during later stages of decay as greater percentage of maize residue-N (8.6-9.4%) were enriched in amino sugars in the N_med and N_high soils, as compared with N₀ and N_low (7.5-8.2%). This indicated that higher availability of inorganic N could delay the transformation process of plant-N into microbial residues during the mineralization of plant residues. The dynamic transformations of the plant residue-N into individual amino sugars were compound-specific, with very fast incorporation into bacterial MurA_M-new found during the initial weeks, while the dynamics of maize residue-derived GluN exhibited a delayed response to assimilate plant-N into fungal products. The findings indicated differential contributions of maize residue decomposing microorganisms over time. Moreover, we found no preferential utilization of inorganic N over plant residue-N into amino sugars during the incubation course, but inorganic N inputs altered the rate of plant-N accumulation in microbial-derived organic matters. Our results indicated that higher N availability had a positive impact on the accumulation or stabilization of newly-produced microbial residues in the long term.     

Dynamics of maize (Zea mays L.) leaf straw mineralization as affected by the presence of soil and the availability of nitrogen

An incubation experiment was carried out with maize (Zea mays L.) leaf straw to analyze the effects of mixing the residues with soil and N amendment on the decomposition process. In order to distinguish between soil effects and nitrogen effects for both the phyllospheric microorganisms already present on the surface of maize straw and soil microorganisms the N amendment was applied in two different placements: directly to the straw or to the soil. The experiment was performed in dynamic, automated microcosms for 22 days at 15 °C with 7 treatments: (1) untreated soil, (2) non-amended maize leaf straw without soil, (3) N amended maize leaf straw without soil, (4) soil mixed with maize leaf straw, (5) N amended soil, (6) N amended soil mixed with maize leaf straw, and (7) soil mixed with N amended maize leaf straw. ¹⁵NH₄¹⁵NO₃ (5 at%) was added. Gas emissions (CO₂, ¹³CO₂ and N₂O) were continuously recorded throughout the experiment. Microbial biomass C, biomass N, ergosterol, δ¹³C of soil organic C and of microbial biomass C as well as ¹⁵N in soil total N, mineral N and microbial biomass N were determined in soil samples at the end of the incubation. The CO₂ evolution rate showed a lag-phase of two days in the non-amended maize leaf straw treatment without soil, which was completely eliminated when mineral N was added. The addition of N generally increased the CO₂ evolution rate during the initial stages of maize leaf straw decomposition, but not the cumulative CO₂ production. The presence of soil caused roughly a 50% increase in cumulative CO₂ production within 22 days in the maize straw treatments due to a slower decrease of CO₂ evolution after the initial activity peak. Since there are no limitations of water or N, we suggest that soil provides a microbial community ensuring an effective succession of straw decomposing microorganisms. In the treatments where maize and soil was mixed, 75% of microbial biomass C was derived from maize. We concluded that this high contribution of maize using microbiota indicates a strong influence of organisms of phyllospheric origin to the microbial community in the soil after plant residues enter the soil.     

Tracing the source and fate of dissolved organic matter in soil after incorporation of a C labelled residue: A batch incubation study

While dissolved organic matter (DOM) in soil solution is a small but reactive fraction of soil organic matter, its source and dynamics are unclear. A laboratory incubation experiment was set up with an agricultural topsoil amended with ¹³C labelled maize straw. The dissolved organic carbon (DOC) concentration in soil solution increased sharply from 25 to 186 mg C L⁻¹ 4 h after maize amendment, but rapidly decreased to 42 mg C L⁻¹ and reached control values at and beyond 2 months. About 65% of DOM was straw derived after 4 h, decreasing to 29% after one day and only 1.3% after 240 days. A significant priming effect of the straw on the release of autochthonous DOM was found. The DOM fractionation with DAX-8 resin revealed that 98% of the straw derived DOM was hydrophilic in the initial pulse while this hydrophilic fraction was 20-30% in control samples. This was in line with the specific UV absorbance of the DOM which was significantly lower in the samples amended with maize residues than in the control samples. The δ¹³C of the respired CO₂ matched that of DOC in the first day after amendment but exceeded it in following days. The straw derived C fractions in respired CO₂ and in microbial biomass were similar between 57 and 240 days after amendment but were 3-10 fold above those in the DOM. This suggests that the solubilisation of C from the straw is in steady state with the DOM degradation or that part of the straw is directly mineralised without going into solution. This study shows that residue application releases a pulse of hydrophilic DOM that temporarily (<3 days) dominates the soil DOM pool and the degradable C. However, beyond that pulse the majority of DOM is derived from soil organic matter and its isotope signature differs from microbial biomass and respired C, casting doubt that the DOM pool in the soil solution is the major bioaccessible C pool in soil.     

Organic fertilizers derived from plant materials Part I: Turnover in soil at low and moderate temperatures

The aim was to investigate different organic fertilizers derived from plant materials with respect to their nitrogen and carbon turnover in soil in comparison with organic fertilizers derived from animal‐waste products. In a 64‐day incubation study at 5°C and 15°C, the following fertilizers were used: coarse faba bean–seed meal (Vicia faba L.), coarse meals of yellow and white lupin seeds (Lupinus albus L. and Lupinus luteus L.), Phytoperls^® (waste products of maize [Zea mays L.] processing), coarse meal of castor cake (Ricinus communis L.) as a widely used organic fertilizer, and horn meal as a reference fertilizer‐derived from animal waste products. At 15°C, horn meal showed the highest apparent net N mineralization of fertilizer‐derived N, followed by castor cake and the two lupin meals. At 5°C, apparent net N mineralization of fertilizer‐derived N from horn meal and coarse meal of yellow lupin seeds was nearly identical, followed by castor‐cake meal. Net N mineralization from legume‐seed meals showed no or even a negative temperature response, at least temporarily. In contrast, the other fertilizers showed a positive temperature response of net N mineralization. The content in recalcitrant structural components and the decoupling of decomposition of N‐rich and C‐rich tissue components in time are discussed as controlling factors of fertilizer‐N turnover at low temperature. Microbial residues seem to be an important temporary sink of fertilizer‐derived C and N. Legume‐seed meals induced considerable N‐priming effects. Temperature induced differences in the decomposition of total fertilizer C, indicated by changes in the sum of cumulative CO₂‐C evolution, total K₂SO₄‐soluble organic C and microbial‐biomass C were much smaller than indicated by cumulative CO₂‐C evolution alone. Our results indicate that legume‐seed meals have the potential to replace horn meal and castor‐cake meal in organic vegetable production, especially when soil temperatures in early spring are still low.