Rice rhizodeposition and its utilization by microbial groups depends on N fertilization

Rhizodeposits have received considerable attention, as they play an important role in the regulation of soil carbon (C) sequestration and global C cycling and represent an important C and energy source for soil microorganisms. However, the utilization of rhizodeposits by microbial groups, their role in the turnover of soil organic matter (SOM) pools in rice paddies, and the effects of nitrogen (N) fertilization on rhizodeposition are nearly unknown. Rice (Oryza sativa L.) plants were grown in soil at five N fertilization rates (0, 10, 20, 40, or 60 mg N kg^?1 soil) and continuously labeled in a ¹³CO₂ atmosphere for 18 days during tillering. The utilization of root-derived C by microbial groups was assessed by ¹³C incorporation into phospholipid fatty acids. Rice shoot and root biomass strongly increased with N fertilization. Rhizodeposition increased with N fertilization, whereas the total ¹³C incorporation into microorganisms, as indicated by the percentage of ¹³C recovered in microbial biomass, decreased. The contribution of root-derived ¹³C to SOM formation increased with root biomass. The ratio of ¹³C in soil pools (SOM and microbial biomass) to ¹³C in roots decreased with N fertilization showing less incorporation and faster turnover with N. The ¹³C incorporation into fungi (18:2ω6,9c and 18:1ω9c), arbuscular mycorrhizal fungi (16:1ω5c), and actinomycetes (10Me 16:0 and 10Me 18:0) increased with N fertilization, whereas the ¹³C incorporation into gram-positive (i14:0, i15:0, a15:0, i16:0, i17:0, and a17:0) and gram-negative (16:1ω7c, 18:1ω7c, cy17:0, and cy19:0) bacteria decreased with N fertilization. Thus, the uptake and microbial processing of root-derived C was affected by N availability in soil. Compared with the unfertilized soil, the contribution of rhizodeposits to SOM and microorganisms increased at low to intermediate N fertilization rates but decreased at the maximum N input. We conclude that belowground C allocation and rhizodeposition by rice, microbial utilization of rhizodeposited C, and its stabilization within SOM pools are strongly affected by N availability: N fertilization adequate to the plant demand increases C incorporation in all these polls, but excessive N fertilization has negative effects not only on environmental pollution but also on C sequestration in soil.     

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     

Microbial processes and community composition in the rhizosphere of European beech - The influence of plant C exudates

Plant roots strongly influence C and N availability in the rhizosphere via rhizodeposition and uptake of nutrients. This study aimed at investigating the effect of resource availability on microbial processes and community structure in the rhizosphere. We analyzed C and N availability, as well as microbial processes and microbial community composition in rhizosphere soil of European beech and compared it to the bulk soil. Additionally, we performed a girdling experiment in order to disrupt root exudation into the soil. By this novel approach we were able to demonstrate that enhanced resource availability positively affected N mineralization and hydrolytic enzyme activities in the rhizosphere, but negatively affected nitrification rates and oxidative enzyme activities, which are involved in the degradation of soil organic matter. Both rhizosphere effects on N mineralization and oxidative enzyme activities disappeared in the girdling treatment. Microbial community structure in the rhizosphere, assessed by phospholipid fatty acid analysis, differed only slightly from bulk soil but was markedly altered by the girdling treatment, indicating additional effects of the girdling treatment beyond the reduction of root exudation. Differences in oxidative enzyme activities and nitrification rates between rhizosphere soil and bulk soil, however, suggest considerable differences in the (functional) microbial community composition.     

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.     

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.     

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.     

Rhizodeposition and net release of soluble organic compounds by pine and beech seedlings inoculated with rhizobacteria and ectomycorrhizal fungi

Summary A lysimetric experiment was performed in a greenhouse to evalute root deposition and net release of soluble organic compounds after 1 and 2 years from pine and beech seedlings inoculated with an ectomycorrhizal fungus (Laccaria laccata) and/or rhizobacteria (Agrobacterium radiobacter for beech and Agrobacterium sp. for pine). Total C compounds released in the rhizosphere of both plants increased after inoculation with the bacteria or ectomycorrhizal fungus. The rhizobacteria increased root and plant growth and rhizodeposition, but the mycorrhizal fungi appeared to increase only root deposition. Soluble C compounds, collected after 2 years, represented only 0.1–0.3% of the total C compounds released into the rhizosphere, and were modified by inoculation with the microorganisms. After inoculation with the bacteria, levels of sugars and amino acids decreased in pine and beech rhizospheres, whereas organic acids increased, especially in the pine rhizosphere. In the rhizosphere of mycorrhizal beeches, sugar and amino acids increased, and organic acids differed from those released from non-mycorrhizal beeches. In the mycorrhizal pine rhizosphere, however, all compounds decreased. Following dual inoculations, mycorrhizal colonization increased, no effect on plant growth was observed, and virtually no organic acids were detected.     

Microbial biomass and mineralization-immobilization of nitrogen in some agricultural soils

Summary The chloroform fumigation-incubation method (CFIM) was used to measure the microbial biomass of 17 agricultural soils from Punjab Pakistan which represented different agricultural soil series. The biomass C was used to calculate biomass N and the changes occurring in NH₄ ⁺-N and NO₃ ^–-N content of soils were studied during the turnover of microbial biomass or added C source. Mineral N released in fumigated-incubated soils and biomass N calculated from biomass C was correlated with some N availability indexes.The soils contained 427–1240 kg C as biomass which represented 1.2%–6.9% of the total organic C in the soils studied. Calculations based on biomass C showed that the soils contained 64–186 kg N ha^–1 as microbial biomass. Immobilization of NCO₃ ^–-N was observed in different soils during the turnover of microbial biomass and any net increase in mineral N content of fumigated incubated soils was attributed entirely to NH₄ ⁺-N.Biomass N calculated from biomass C showed non-significant correlation with different N availability indexes whereas mineral N accumulated in fumigated-incubated soils showed highly significant correlations with other indexes including N uptake by plants.     

Microbial response to exudates in the rhizosphere of young beech trees (Fagus sylvatica L.) after dormancy

Plants act as an important link between atmosphere and soil: CO2 is transformed into carbohydrates by photosynthesis. These assimilates are distributed within the plant and translocated via roots into the rhizosphere and soil microorganisms. In this study, 3 year old European beech trees (Fagus sylvatica L.) were exposed after the chilling period to an enriched ¹³C–CO₂ atmosphere (δ¹³C = 60‰ – 80‰) at the time point when leaves development started. Temporal dynamics of assimilated carbon distribution in different plant parts, as well as into dissolved organic carbon and microbial communities in the rhizosphere and bulk soil have been investigated for a 20 days period. Photosynthetically fixed carbon could be traced into plant tissue, dissolved organic carbon and total microbial biomass, where it was utilized by different microbial communities. Due to carbon allocation into the rhizosphere, nutrient stress decreased; exudates were preferentially used by Gram-negative bacteria and (mycorrhizal) fungi, resulting in an enhanced growth. Other microorganisms, like Gram-positive bacteria and mainly micro eucaryotes benefited from the exudates via food web development. Overall our results indicate a fast turnover of exudates and the development of initial food web structures. Additionally a transport of assimilated carbon into bulk soil by (mycrorhizal) fungi was observed.     

Mycorrhizal community structure,microbial biomass P and phosphatase activities under Salix polaris as influenced by nutrient availability

Low supply of the nutrients nitrogen (N) and phosphorus (P) limit plant growth and spreading, and increase the plant-microbial nutrient competition in subarctic and arctic regions. We investigated the mycorrhizal community structure of a polar shrub willow (Salix polaris) and the microbial turnover in its rhizosphere to explore the adaptation of a mycorrhizal plant in the subarctic tundra. The ectomycorrhizal colonisation ranged from 35 to 64% of the fine root tips and decreased with an increasing soil C/N ratio. In total, 16 ectomycorrhizal morphotypes were found under S. polaris (eight to 13 morphotypes per site, five morphotypes at all four sites). Cenococcum sp. was the most common EM fungus (32% of the ectomycorrhizal fine roots). The abundance of Cenococcum sp. increased with an increasing organic matter content and N/P ratio in the soil. Arbuscular mycorrhizal colonisation of S. polaris was absent or less than 1% of the fine root length. Microbial biomass P accounted for 21–75% of the organic soil P and 6–49% of the total soil P. Microbial biomass P, alkaline and acid phosphatase activities in the rhizosphere increased with increasing soil N concentration. We conclude that a higher N supply decreases the diversity in the mycorrhizal community on polar willows and increases the role of P turnover from the soil microbial biomass for the nutrient supply.     

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.     

Ecosystem partitioning of N-glycine after long-term climate and nutrient manipulations, plant clipping and addition of labile carbon in a subarctic heath tundra

Low temperatures and high soil moisture restrict cycling of organic matter in arctic soils, but also substrate quality, i.e. labile carbon (C) availability, exerts control on microbial activity. Plant exudation of labile C may facilitate microbial growth and enhance microbial immobilization of nitrogen (N). Here, we studied ¹⁵N label incorporation into microbes, plants and soil N pools after both long-term (12 years) climate manipulation and nutrient addition, plant clipping and a pulse-addition of labile C to the soil, in order to gain information on interactions among soil N and C pools, microorganisms and plants. There were few effects of long-term warming and fertilization on soil and plant pools. However, fertilization increased soil and plant N pools and increased pool dilution of the added ¹⁵N label. In all treatments, microbes immobilized a major part of the added ¹⁵N shortly after label addition. However, plants exerted control on the soil inorganic N concentrations and recovery of total dissolved ¹⁵N (TD¹⁵N), and likewise the microbes reduced these soil pools, but only when fed with labile C. Soil microbes in clipped plots were primarily C limited, and the findings of reduced N availability, both in the presence of plants and with the combined treatment of plant clipping and addition of sugar, suggest that the plant control of soil N pools was not solely due to plant uptake of soil N, but also partially caused by plants feeding labile C to the soil microbes, which enhanced their immobilization power. Hence, the cycling of N in subarctic heath tundra is strongly influenced by alternating release and immobilization by microorganisms, which on the other hand seems to be less affected by long-term warming than by addition or removal of sources of labile C.     

The chemical and functional characterization of soil N and its biotic components

The aim of this review is to describe and discuss the concepts that have been employed to interpret N mineralization-immobilization in soil, and how N turnover is related to the characteristics of organic N and the biota conducting the transformations. A brief survey of the period before the arrival of electronic searches became available provides access to the classical literature that can help interpret today's challenges. Classical (acid hydrolysis) and modern spectrometry and spectroscopy techniques indicate that protein N is the prevalent component of organic N in soil. The presence of heterocyclic N can indicate its abiotic, partial synthesis as in fire-affected soils. Clays and pedogenic oxides can protect organic N against microbial degradation. The evidence for such protection is mostly based on in vitro studies involving pure clays, and proteins and their relevance to field conditions requires further work. The proteomic approach, with extraction and characterization of proteins stabilised by soil colloids (structural proteomics) might give further insights into this area. Functional proteomics can improve our understanding of the degradation of organic pollutants and organic debris as well as identifying the molecular colloquia between microorganisms and between soil biota and plant roots. Subdivision of organic N into sub-pools has helped to interpret mechanistic studies and modelling of N dynamics. Uncomplexed organic matter, obtained by physical fractionation procedures, is considered a labile pool. The interpretation of N mineralization measurements is affected by immobilization during microbial attack especially in high-C environments. Transfer of materials among particle size fractions and changes in microbiological properties of aggregates also can occur during fractionation procedures. Classical mineralization-immobilization turnover (MIT) does not always occur since microorganisms (and plants) can take up amino acid N with intracellular deamination. Protozoa, due to their grazing activities, can influence not only N mineralization but also the composition of rhizosphere-plant growth stimulating communities. Differences between N-poor and N-rich microsites, occurring in the same soil, can markedly affect the competition for N between plants and microorganisms especially the nitrifiers. The use of molecular techniques has allowed the identification of unculturable microorganisms and functional genes in the N cycle. Archeae are probably capable of oxidising NH₄⁺ to NO₃⁻ and anerobic ammonia oxidation (Ammonox) bacteria have been identified in biofilms and probably also occur in soils. The use of nitrate as an electron acceptor is encoded by specific gene clusters but nitrate reduction also occurs in dissimilatory nitrate reduction.     

Earthworm (Aporrectodea trapezoides)–mycorrhiza (Glomus intraradices) interaction and nitrogen and phosphorus uptake by maize

The interactive impacts of arbuscular mycorrhizal fungi (AMF, Glomus intraradices) and earthworms (Aporrectodea trapezoides) on maize (Zea mays L.) growth and nutrient uptake were studied under near natural conditions with pots buried in the soil of a maize field. Treatments included maize plants inoculated vs. not inoculated with AMF, treated or not treated with earthworms, at low (25 mg kg⁻¹) or high (175 mg kg⁻¹) P fertilization rate. Wheat straw was added as feed for earthworms. Root colonization, mycorrhiza structure, plant biomass and N and P contents of shoots and roots, soil available P and NO₃⁻–N concentrations, and soil microbial biomass C and N were measured at harvest. Results indicated that mycorrhizal colonization increased markedly in maize inoculated with AMF especially at low P rate, which was further enhanced by the addition of earthworms. AMF and earthworms interactively increased maize shoot and root biomass as well as N and P uptake but decreased soil NO₃⁻–N and available P concentrations at harvest. Earthworm and AMF interaction also increased soil microbial biomass C, which probably improved root N and P contents and indirectly increased the shoot N and P uptake. At low P rate, soil N mobilization by earthworms might have reduced potential N competition by arbuscular mycorrhizal hyphae, resulting in greater plant shoot and root biomass. Earthworms and AMF interactively enhanced soil N and P availability, leading to greater nutrient uptake and plant growth.     

Comparison of uptake of different N forms by soil microorganisms and two wet-grassland plants: A pot study

Two common plant species of temperate wet grasslands, Carex acuta and Glyceria maxima, were tested for their preferences in the uptake of different nitrogen (N) sources (amino acid, ammonium, nitrate) and their ability to compete for these sources with soil microorganisms. The experiment was a one-day incubation study with plants growing in soil obtained from the field, which was supplied with a solution containing the three N sources, one at a time labeled with ¹⁵N. The bulk of the N demand of both species was covered by nitrate-N, which was the dominant N form in the soil at the time of the experiment. Ammonium-N was taken up less strongly, and organic N formed only a negligible part of their nutrition. The assimilated inorganic N was preferentially transported to apical meristem of the youngest leaf, while organic N remained mostly in the roots. The fast-growing Glyceria took up more N and was a better competitor vis-à-vis soil microbes for rarer N forms than Carex. However, both plants were poor competitors for N vis-à-vis soil microbes, irrespective of the N form. Microbes took up nitrate ca. five times faster and organic N more than a hundred times faster than plants. Correspondingly, the calculated turnover time of microbial N was 17 days, compared to 40 days for N in plant roots. A significant amount of added ¹⁵N was found at non-exchangeable sites in the soil, which points to the importance of microbial N transformation and abiotic N fixation for N retention in soil. In summary, the preferential assimilation of inorganic N by the wetland plants studied here and their poor ability to compete for N with soil microbes over the short term agree with the results of studies carried out with other species from temperate grasslands.     

Rapid soil organic matter loss from forest dieback in a subalpine coniferous ecosystem

Forest dieback caused by climate-change associated stresses and insect outbreaks has emerged as a global concern, and the biogeochemical consequences of this phenomenon need to be elucidated. We measured biological and chemical traits of soil beneath live trees or trees recently killed by a mountain-pine-beetle outbreak in a subalpine coniferous forest in the Front Range of Colorado. We focused on the top 5 cm of mineral soil just beneath the O horizon and measured microbial biomass, soil invertebrate abundance and composition, and soil chemical characteristics. With the termination of inputs from rhizodeposition, mycorrhizal fungal turnover and fine root turnover, soil total carbon (C) and total nitrogen (N) in the mineral soil at three sites decreased by 38–49% and 26–45%, respectively. Tree mortality was associated with reduced soil microbial biomass but soil nematode and microarthropod densities were unchanged. Nematode trophic structure was altered with an increased proportion of bacterial feeders. Soil inorganic N concentrations were inversely correlated to microbial C:N ratios. Tree death was associated with increased soil pH, a possible loss of calcium (Ca²⁺), but an accumulation of soil inorganic N, largely as NH₄⁺. Our results suggest that forest dieback results in rapid C and N loss from surface mineral soils and that the accumulation of soil inorganic N, the reduction in microbial biomass, and the more bacterial-based soil food web increase the potential of enhanced N loss from affected ecosystems.     

Rapid soil organic matter loss from forest dieback in a subalpine coniferous ecosystem

Forest dieback caused by climate-change associated stresses and insect outbreaks has emerged as a global concern, and the biogeochemical consequences of this phenomenon need to be elucidated. We measured biological and chemical traits of soil beneath live trees or trees recently killed by a mountain-pine-beetle outbreak in a subalpine coniferous forest in the Front Range of Colorado. We focused on the top 5 cm of mineral soil just beneath the O horizon and measured microbial biomass, soil invertebrate abundance and composition, and soil chemical characteristics. With the termination of inputs from rhizodeposition, mycorrhizal fungal turnover and fine root turnover, soil total carbon (C) and total nitrogen (N) in the mineral soil at three sites decreased by 38–49% and 26–45%, respectively. Tree mortality was associated with reduced soil microbial biomass but soil nematode and microarthropod densities were unchanged. Nematode trophic structure was altered with an increased proportion of bacterial feeders. Soil inorganic N concentrations were inversely correlated to microbial C:N ratios. Tree death was associated with increased soil pH, a possible loss of calcium (Ca²⁺), but an accumulation of soil inorganic N, largely as NH₄⁺. Our results suggest that forest dieback results in rapid C and N loss from surface mineral soils and that the accumulation of soil inorganic N, the reduction in microbial biomass, and the more bacterial-based soil food web increase the potential of enhanced N loss from affected ecosystems.     

Interactions between tree seedling roots and humus forms in the control of soil C and N cycling

The hypothesis that roots enhance soil-N turnover in humified soil organic matter (SOM) (mull) but not in lignified SOM (mor) was tested in a study involving the growth of eight species of tree seedlings on the two contrasting humus forms. After 12 and 22 weeks of seedling growth, soil-CO₂ efflux was measured with (1) growing seedlings, and after 22 weeks, with (2) roots only, shoots excised, and (3) with roots removed and soils amended with different rates of glucose. Indices of C-flux and of soil available-C were derived and compared to plant-N uptake, extractable soil mineral-N, anaerobically mineralized soil-N, N bioavailability to Agrostis grass following harvest of seedlings, and to seedling fine root C-chemistry. Significant soil x species interactions were found for total soil-CO₂ efflux, root-dependent CO₂, soil available-C and microbial biomass. In all cases, roots were important contributors to C-cycling in the mull soil but not in the mor soil. C was more limiting in the mor than in the mull microbial community. Plant-N uptake and the mineral-N pool was greater in the mor soil, reflecting that soil's higher specific N-supplying capacity (N-mineralized:CO₂). Seedlings decreased the mineral-N pool in both soils, but the presence of roots increased N-mineralization in the mull soil and decreased N-mineralization in the mor soil. Significant positive relationships were observed in the mull soil only between soil respiration and plant N uptake at mid-season, and between soil respiration and N-mineralization at late-season. Birch root activity in the mull soil was greater than that of all other seedlings and this observation is discussed with respect to the autecology of birch. Soil respiration correlated with the non-polar extract content but not the lignin:N ratio of fine roots. Results suggest that root-released C in mull SOM is sufficient to relieve energy limitation to soil microbes and allow them to access appreciable amounts of soil-N, whereas ligninolytic activity, which may ultimately control soil-N turnover in mor SOM, is not increased by rhizodeposition.     

Carbon Turnover in the Rhizosphere

Considerable progress has been made during the last decade towards understanding and quantifying the input and turnover of plant carbon in the rhizosphere. This was made possible by the development (partially by the authors) and combination of appropriate new methods, such as:

• –homogeneous labelling of whole plants with ¹⁴C
• –distinction between root and microbial respiration
• –separation of soil zones of known distances from the roots
• –determination of microbial soil biomass.

These methods were applied to study the following aspects:

• –release of organic plant carbon into the soil by growing roots
• –utilization of this plant carbon by the microbial biomass in the rhizosphere
• –related influence on the turnover of soil organic matter, and
• –spatial range of such root influence in the soil.

About 19% of the total photosynthetic production of the investigated plants was released into the rhizosphere as organic material. Most of this (15%) was transformed by the rhizosphere microorganisms into CO₂, while only a small fraction (4%) remained in the soil, mainly as microbial cells (2.5%). As a result, microbial rhizosphere biomass increased considerably. Relative to the organic C-input, however, the incorporation of root derived carbon by the microbial biomass was remarkably low (13%). Along with the increase in microbial rhizosphere biomass, the presence of plant roots also enhanced the decomposition of soil organic matter and affected soil aggregate stability. Root carbon and root influences were even detected up to 20 mm away from the roots. This may be partially attributed to the contribution of root derived volatiles. Accordingly, both the actual volume of the rhizosphere and its metabolic significance is greater than what has so far been assumed. Possible interactions involving root, soil and microbial carbon are discussed.