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.     

Gross fluxes of nitrogen in grassland soil exposed to elevated atmospheric pCO2 for seven years

Plant response to increasing atmospheric CO₂ partial pressure (pCO₂) depends on several factors, one of which is mineral nitrogen availability facilitated by the mineralisation of organic N. Gross rates of N mineralisation were examined in grassland soils exposed to ambient (36 Pa) and elevated (60 Pa) atmospheric pCO₂ for 7 years in the Swiss Free Air Carbon dioxide Enrichment experiment. It was hypothesized that increased below-ground translocation of photoassimilates at elevated pCO₂ would lead to an increase in immobilisation of N due to an excess supply of energy to the roots and rhizosphere. Intact soil cores were sampled from Lolium perenne and Trifolium repens swards in May and September, 2000. The rates of gross N mineralisation (m) and NH₄⁺ consumption (c) were determined using ¹⁵N isotopic dilution during a 51-h period of incubation. The rates of N immobilisation were estimated either as the difference between m and the net N mineralisation rate or as the amount of ¹⁵N released from the microbial biomass after chloroform fumigation. Soil samples from both swards showed that the rates of gross N mineralisation and NH₄⁺ consumption did not change significantly under elevated pCO₂. The lack of a significant effect of elevated pCO₂ on organic N turnover was consistent with the similar size of the microbial biomass and similar immobilisation of applied ¹⁵N in the microbial N pool under ambient and elevated pCO₂. Rates of m and c, and microbial ¹⁵N did not differ significantly between the two sward types although a weak (p<0.1) pCO₂ by sward interaction occurred. A significantly larger amount of NO₃⁻ was recovered at the end of the incubation in soil taken from T. repens swards compared to that from L. perenne swards. Eleven percent of the added ¹⁵N were recovered in the roots in the cores sampled under L. perenne, while only 5% were recovered in roots of T. repens. These results demonstrate that roots remained a considerable sink despite the shoots being cut at ground level prior to incubation and suggest that the calculation of N immobilisation from gross and net rates of mineralisation in soils with a high root biomass does not reflect the actual immobilisation of N in the microbial biomass. The results of this study did not support the initial hypothesis and indicate that below-ground turnover of N, as well as N availability, measured in short-term experiments are not strongly affected by long-term exposure to elevated pCO₂. It is suggested that differences in plant N demand, rather than major changes in soil N mineralisation/immobilisation, are the long-term driving factors for N dynamics in these grassland systems.     

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.     

Changes in forest soil organic matter pools after a decade of elevated CO2 and O3

The impact of rising atmospheric carbon dioxide (CO₂) may be mitigated, in part, by enhanced rates of net primary production and greater C storage in plant biomass and soil organic matter (SOM). However, C sequestration in forest soils may be offset by other environmental changes such as increasing tropospheric ozone (O₃) or vary based on species-specific growth responses to elevated CO₂. To understand how projected increases in atmospheric CO₂ and O₃ alter SOM formation, we used physical fractionation to characterize soil C and N at the Rhinelander Free Air CO₂-O₃ Enrichment (FACE) experiment. Tracer amounts of ¹⁵NH₄⁺ were applied to the forest floor of Populus tremuloides, P. tremuloides-Betula papyrifera and P. tremuloides-Acer saccharum communities exposed to factorial CO₂ and O₃ treatments. The ¹⁵N tracer and strongly depleted ¹³C-CO₂ were traced into SOM fractions over four years. Over time, C and N increased in coarse particulate organic matter (cPOM) and decreased in mineral-associated organic matter (MAOM) under elevated CO₂ relative to ambient CO₂. As main effects, neither CO₂ nor O₃ significantly altered ¹⁵N recovery in SOM. Elevated CO₂ significantly increased new C in all SOM fractions, and significantly decreased old C in fine POM (fPOM) and MAOM over the duration of our study. Overall, our observations indicate that elevated CO₂ has altered SOM cycling at this site to favor C and N accumulation in less stable pools, with more rapid turnover. Elevated O₃ had the opposite effect, significantly reducing cPOM N by 15% and significantly increasing the C:N ratio by 7%. Our results demonstrate that CO₂ can enhance SOM turnover, potentially limiting long-term C sequestration in terrestrial ecosystems; plant community composition is an important determinant of the magnitude of this response.     

Nitrogen Transformation as Influenced by Soil Microbial Nitrogen under Carbon Dioxide Enrichment Induced in Created Riparian Wetlands

Limited research has been conducted on how atmospheric carbon dioxide (CO₂) affects water and soil nitrogen (N) transformation in wetland ecosystems. A stable isotope technique is suitable for conducting a detailed investigation of mechanistic nutrient transformations. Nutrient ammonium sulfate (NH₄)₂SO₄ input in culture water under elevated CO₂ (700 μL L^?1) and ambient CO₂ (380 μL L^?1) was studied to analyze N transformations with N blanks for both water and soil. It was measured by ¹⁵N pool dilution using analytical equations in a riparian wetland during a 3-month period. Soil gross ammonium (NH₄ ⁺) mineralization and consumption rates increased significantly by 22% and 404%, Whereas those of water decreased greatly by??57% and??57% respectively in enriched CO₂. In contrast, gross nitrate (NO₃ ^?) consumption and nitrification rates of soil decreased by??11% and??14% and those of water increased by 29% and 27% respectively in enrichment CO₂. These may be due to the extremely high soil microbial biomass nitrogen (MBN), which increased by 94% in elevated soil. The results can show when CO₂ concentrations are going to rise in the future. Consequently soil microbial activity initiates the decreased N concentration in sediment and increased N concentration in overlying water in riparian wetland ecosystems.     

Atmospheric CO2 enrichment and nutrient additions to planted soil increase mineralisation of soil organic matter, but do not alter microbial utilisation of plant- and soil C-sources

Plants link atmospheric and soil carbon pools through CO₂ fixation, carbon translocation, respiration and rhizodeposition. Within soil, microbial communities both mediate carbon-sequestration and return to the atmosphere through respiration. The balance of microbial use of plant-derived and soil organic matter (SOM) carbon sources and the influence of plant-derived inputs on microbial activity are key determinants of soil carbon-balance, but are difficult to quantify. In this study we applied continuous ¹³C-labelling to soil-grown Lolium perenne, imposing atmospheric CO₂ concentrations and nutrient additions as experimental treatments. The relative use of plant- and SOM-carbon by microbial communities was quantified by compound-specific ¹³C-analysis of phospholipid fatty acids (PLFAs). An isotopic mass-balance approach was applied to partition the substrate sources to soil respiration (i.e. plant- and SOM-derived), allowing direct quantification of SOM-mineralisation. Increased CO₂ concentration and nutrient amendment each increased plant growth and rhizodeposition, but did not greatly alter microbial substrate use in soil. However, the increased root growth and rhizosphere volume with elevated CO₂ and nutrient amendment resulted in increased rates of SOM-mineralisation per experimental unit. As rhizosphere microbial communities utilise both plant- and SOM C-sources, the results demonstrate that plant-induced priming of SOM-mineralisation can be driven by factors increasing plant growth. That the balance of microbial C-use was not affected on a specific basis may suggest that the treatments did not affect soil C-balance in this study.     

Impact of elevated CO2 on mycorrhizal associations and implications for plant growth

The impact of increasing concentrations of atmospheric CO₂ upon plant physiology has been widely investigated. Plant, and in particular root, growth is nearly always enhanced as a direct consequence of CO₂ enrichment, with C₃ species generally more responsive than C₄ species. Such alterations in plant productivity will have consequence for below-ground processes and increased carbon allocation to the roots may favour symbiotic relationships. This paper discusses the current information available for the consequences of these changes upon mycorrhizal relationships. Generally mycorrhizal plants grown under CO₂ enrichment show enhanced phosphorus uptake but nitrogen uptake is unaffected. This increased nutrient uptake is not correlated with increased mycorrhizal colonization of the roots. Similarly root exudation does not increase under CO₂ enrichment but qualitative differences have yet to be assessed. However, it is predicted that total rhizodeposition of materials will increase as will litter inputs, although mineral and biochemical alterations to these plant derived inputs may occur. The consequences of such changes within the rhizosphere are discussed and future research     

Effects of elevated CO2 concentration on rhizodeposition from Lolium perenne grown on soil exposed to 9 years of CO2 enrichment

The effects of enriched CO₂ atmosphere on partitioning of recently assimilated carbon were investigated in a plant-soil-microorganism system in which Lolium perenne seedlings were planted into cores inserted into the resident soil within a sward that had been treated with elevated CO₂ for 9 consecutive years, under two N fertilisation levels (Swiss FACE experiment). The planted cores were excavated from the ambient (35 Pa pCO₂) and enriched (60 Pa pCO₂) rings at two dates, in spring and autumn, during the growing season. The cores were brought back to the laboratory for ¹⁴C labelling of shoots in order to trace the transfer of recently assimilated C both within the plant and to the soil and microbial biomass. At the spring sampling, high N supply stimulated shoot and total dry matter production. Consistently, high N enhanced the allocation of recently fixed C to shoots, and reduced it to belowground compartments. Elevated CO₂ had no consequences for DM or the pattern of C allocation. At the autumn sampling, at high N plot, yield of L. perenne was stimulated by elevated CO₂. Consistently, ¹⁴C was preferentially allocated aboveground and, consequently belowground recent C allocation was depressed and rhizodeposition reduced. At both experimental periods, total soil C content was similar in all treatments, providing no evidence for soil carbon sequestration in the Swiss Free Air CO₂ Enrichment experiment (FACE) after 9 years of enrichment. Recently assimilated C and soil C were mineralised faster in soils from enriched rings, suggesting a CO₂-induced shift in the microbial biomass characteristics (structure, diversity, activity) and/or in the quality of the root-released organic compounds.     

Pflanzen‐mikroben‐interaktionen und c/n‐Stoffumsetzungsprozesse in der rhizosphäre: Aufklärung und quantifizierung

Bis in jüngste Zeit war unklar, ob den in der Rhizosphäre, der Grenzschicht zwischen dem pflanzlichen Wurzelsystem und dem Boden, ablaufenden Vorgängen tatsächlich Relevanz für die C/N‐Umsetzungsprozesse auf ökosystemarer Ebene zukommt. Zielstellung eigener Arbeiten war es daher, auf der Grundlage von Tracern (¹⁴C, ¹⁵N) und immunologischen Techniken (DAS‐ELISA) Verfahren zur entwickeln und zu testen, die zur Klärung dieser Fragestellung beitragen. Wie ^l4C‐Impulsmarkierungsexperimente ergaben, werden bereits im Verlauf der Ontogenese beträchtliche Mengen des neu assimilierten Kohlenstoffs in die organische Bodensubstanz eingebaut (bei Sommerweizen zwischen 51 und 410% und bei Schilf 21% des Wurzel‐C). Mit Hilfe von DAS‐ELISA ausgeführte Studien ließen einen sehr differenzierten Einfluß von Getreidearten und ‐Sorten auf die Besiedlung der Rhizosphäre mit dem phytoeffektiven Bakterienstamm Pantoea agglomérons D5/23 erkennen. In überstautem Niedermoortorf bewirkte die Anwesenheit von Sumpfpflanzen (Schilf, Rohrglanzgras) eine beschleunigte Umsetzung von ¹⁵N‐markiertem Nitrat zu den gasförmigen Denitrifikationsprodukten N₂O und N₂.     

Priming effect of biuret addition on native soil N mineralisation under laboratory conditions

This study was carried out to quantify the priming effect of biuret on native soil nitrogen (N) mineralisation during a 112-day incubation. Addition of biuret (100 mg ¹⁵N-labelled biuret kg⁻¹ soil) increased the turnover rate constant of soil organic matter and had a positive priming effect on native soil N mineralisation in two soils. The additional mineralisation was 0.65% of the total soil N (equivalent to 47.1 kg N ha⁻¹) in a sandy loam soil and 0.62% of the soil N (equivalent to 46.5 kg N ha⁻¹) in a silt loam soil.     

Long term CO2 enrichment stimulates N-mineralisation and enzyme activities in calcareous grassland

Elevated concentration of atmospheric carbon dioxide will affect carbon cycling in terrestrial ecosystems. Possible effects include increased carbon input into the soil through the rhizosphere, altered nutrient concentrations of plant litter and altered soil moisture. Consequently, the ongoing rise in atmospheric carbon dioxide might indirectly influence soil biota, decomposition and nutrient transformations.N-mineralisation and activities of the enzymes invertase, xylanase, urease, protease, arylsulfatase, and alkaline phosphatase were investigated in spring and summer in calcareous grassland, which had been exposed to ambient and elevated CO₂ concentrations (365 and 600 μl l⁻¹) for six growing seasons.In spring, N-mineralisation increased significantly by 30% at elevated CO₂, while there was no significant difference between treatments in summer (+3%). The response of soil enzymes to CO₂ enrichment was also more pronounced in spring, when alkaline phosphatase and urease activities were increased most strongly by 32 and 21%. In summer, differences of activities between CO₂ treatments were greatest in the case of urease and protease (+21 and +17% at elevated CO₂).The stimulation of N-mineralisation and enzyme activities at elevated CO₂ was probably caused by higher soil moisture and/or increased root biomass. We conclude that elevated CO₂ will enhance below-ground C- and N-cycling in grasslands.     

Changes in some enzyme activities and the nicotinamide nucleotide coenzymes in barley seedlings during vernalization

Respiratory metabolism was investigated by determining the following enzyme activities and nicotinamide nucleotide coenzyme contents in barley shoots (Hordeum vulgare L., var. Akashinriki) during vernalization : glucose-6-phosphate dehydrogenase, phosphoglucoisomerase, enolase, isocitrate dehydrogenase, malate dehydrogenase, and glutamic dehydrogenase. The activities of glucose-6-phosphate de hydrogenase and phosphoglucoisomerase in vernalized shoots were slightly lower than those in non-vernalized samples. Malate dehydrogenase activity increased gradually in vernalized shoots and was always higher than in non-vernalized shoots. Activation was found in enolase, isocitrate dehydrogenase and glutamic dehydrogenase after a long vernalization treatment.

The contents of total nicotinamide nucleotide coenzymes increased during the early germination stage in both treatments, then decreased gradually in the succeeding stage.     

Mineralization of carbon and nitrogen from fresh and anaerobically stored sheep manure in soils of different texture

A sandy loam soil was mixed with three different amounts of quartz sand and incubated with (¹⁵NH₄)₂SO₄ (60 g N g^-1 soil) and fresh or anaerobically stored sheep manure (60 g g^-1 soil). The mineralization-immobilization of N and the mineralization of C were studied during 84 days of incubation at 20°C. After 7 days, the amount of unlabelled inorganic N in the manure-treated soils was 6–10 g N g^-1 soil higher than in soils amended with only (¹⁵NH₄)₂SO₄. However, due to immobilization of labelled inorganic N, the resulting net mineralization of N from manure was insignificant or slightly negative in the three soil-sand mixtures (100% soil+0% quartz sand; 50% soil+50% quartz sand; 25% soil+75% quartz sand). After 84 days, the cumulative CO₂ evolution and the net mineralization of N from the fresh manure were highest in the soil-sand mixutre with the lowest clay content (4% clay); 28% fo the manure C and 18% of the manure N were net mineralized. There was no significant difference between the soil-sand mixtures containing 8% and 16% clay, in which 24% of the manure C and -1% to 4% of the manure N were net mineralized. The higher net mineralization of N in the soil-sand mixture with the lowest clay content was probably caused by a higher remineralization of immobilized N in this soil-sand mixture. Anaerobic storage of the manure reduced the CO₂ evolution rates from the manure C in the three soil-sand mixtures during the initial weeks of decomposition. However, there was no effect of storage on net mineralization of N at the end of the incubation period. Hence, there was no apparent relationship between net mineralization of manure N and C.