Distribution of assimilated carbon in plants and rhizosphere soil of basket willow (Salix viminalis L.)

Willow is often used in bio-energy plantations for its potential to function as a renewable energy source, but knowledge about its effect on soil carbon dynamics is limited. Therefore, we investigated the temporal variation in carbon dynamics in willow, focusing on below-ground allocation and sequestration to soil carbon pools. Basket willow plants (Salix viminalis L.) in their second year of growth were grown in pots in a greenhouse. At five times during the plants growth, namely 0, 1, 2, 3 and 4 months after breaking winter dormancy, a subset of the plants were continuously labelled with ¹⁴CO₂ in an ESPAS growth chamber for 28 days. After the labelling, the plants were harvested and separated into leaves, first and second year stems and roots. The soil was analysed for total C and ¹⁴C content as well as soil microbial biomass. Immediately after breaking dormancy, carbon stored in the first year stems was relocated to developing roots and leaves. Almost half the newly assimilated C was used for leaf development the first month of growth, dropping to below 15% in the older plants. Within the second month of growth, secondary growth of the stem became the largest carbon sink in the system, and remained so for the older age classes. Between 31 and 41% of the recovered ¹⁴C was allocated to below-ground pools. While the fraction of assimilated ¹⁴C in roots and root+soil respiration did not vary with plant age, the amount allocated to soil and soil microbial biomass increased in the older plants, indicating an increasing rhizodeposition. The total amount of soil microbial biomass was 30% larger in the oldest age class than in an unplanted control soil. The results demonstrate a close linkage between photosynthesis and below-ground carbon dynamics. Up to 13% of the microbial biomass consisted of carbon assimilated by the willows within the past 4 weeks, up to 11% of the recovered ¹⁴C was found as soil organic matter.     

Contribution by two arbuscular mycorrhizal fungi to P uptake by cucumber (Cucumis sativus L.) from32P-labelled organic matter during mineralization in soil

An experiment was set up to investigate the role of arbuscular mycorrhiza (AM) in utilization of P from organic matter during mineralization in soil. Cucumber (Cucumis sativus L.) inoculated with one of two AM fungi or left uninoculated were grown for 30 days in cross-shaped PVC pots. One of two horizontal compartments contained 100 g soil (quartz sand: clay loam, 1:1) with 0.5 g ground clover leaves labelled with³²P. The labelled soil received microbial inoculum without AM fungi to ensure mineralization of the added organic matter. The labelling compartment was separated from a central root compartment by either 37 m or 700 m nylon mesh giving only hyphae or both roots and hyphae, respectively, access to the labelled soil. The recovery of³²P from the hyphal compartment was 5.5 and 8.6% for plants colonized withGlomus sp. andG. caledonium, respectively, but only 0.6 % for the non-mycorrhizal controls. Interfungal differences were not related to root colonization or hyphal length densities, which were lowest forG. caledonium. Both fungi depleted the labelled soil of NaHCO₃-extractable P and³²P compared to controls. A 15–25% recovery of³²P by roots was not enhanced in the presence of mycorrhizas, probably due to high root densities in the labelled soil. The experiment confirms that AM fungi differ in P uptake characteristics, and that mycorrhizal hyphae can intercept some P immobilization by other microorganisms and P-sorbing clay minerals.     

Seed source variation in 32P uptake in inus taeda L. seedlings: A possible regulation by efflux

Short-term ³²P uptake experiments were conducted with intact seedlings of loblolly pine (Pinus taeda L.) to examine possible seed source variation in net accumulation of ³²P in roots and shoots, and in rates of unidirectional influx. Seed source had a highly significant effect on biomass and P concentrations of shoots and roots. Seedlings from two seed sources representing fast-growing populations (a broadly-adapted and wet-site seed source) accumulated over 60% more total seedling P than smaller seedlings from a drought-hardy seed source, reflecting higher biomass and root P concentrations. Rates of unidirectional ³²P influx in seedlings from the drought-hardy seed source were more than twice the rates of the seedlings from the broadly-adapted seed source. However, after 24 h in labeled uptake solution, net accumulation of ³²P was similar, suggesting that rates of unidirectional efflux from roots of the drought-hardy seed source were also high. Although there were no significant differences in biomass and tissue P concentrations between the two fast-growing seed sources, rates of unidirectional influx in seedlings from the broadly-adapted seed source were 42% lower than rates in seedlings from the wet-site source. Yet, after 24 h in labeled uptake solution, net accumulation of ³²P in seedlings from the broadly-adapted seed source was 50% higher. Unidirectional efflux out of the root may regulate net uptake of P as much, if not more, than influx in loblolly pine seedlings-at least under high-P growth conditions. The results in this study do not support previous studies with herbaceous plants suggesting that fast-growing species typically exhibit higher rates of nutrient uptake than slow-growing species.     

Nutrient acquisition from different soil depths by pedunculate oak

Eight oak trees (Quercus robur L.) received ³²P at a soil depth of 50 cm and ³³P at a soil depth of 15 cm at the end of June 2002 through plastic tubes inserted into the mineral soil. The phosphorus uptake from different soil depths was estimated by analysing the concentration of ³²P and ³³P in the foliage of oak growing in a mixed stand in southern Sweden. ³²P and ³³P were recovered in the leaves/needles after 21 and 39 days. The recovery of labelled P in oak was higher from 15 cm soil depth than from 50 cm, however, more than 4% of the total amount of labelled P was taken up from 50 cm. This indicates that oak can utilize deep soil layers for nutrient uptake. A study on the uptake of Cs (as an analogue to K) and ¹⁵N into the leaves was performed on the same trees and detectable amounts of ¹⁵N and Cs were recovered in leaves and buds. This indicates that ¹⁵N and Cs can be used to study nutrient uptake of mature trees from the mineral soil.     

Carbon distribution and nitrogen partitioning in a soil-plant system with barley (Hordeum vulgare L.), ryegrass (Lolium perenne) and rape (Brassica napus L.) grown in a14CO2-atmosphere

To examine the influence of plant-microorganism interactions on soil-N transformations (e.g. net mineralization, net immobilization) a pot experiment was conducted in a¹⁴C-labelled atmosphere by using different (two annuals, one perennial) plants species. It was assumed that variation in below-ground, microorganism-available C would influence N transformations in soil. Plant species were fertilized (low rate) with¹⁵N-labelled nitrogen and grown, during days 13 and 62 after germination, in a growth chamber with a¹⁴C-labelled atmosphere. Nitrification was inhibited by using nitrapyrin (N-Serve). During the chamber period, shoots were harvested, and associated roots and soil were collected on two sampling occasionm, e.g. after 4 and 7 weeks in the growth chamber.The distribution of net (%) assimilated¹⁴C was significantly affected by both plant and time factors, and there was a significant plant × time interaction. There were significant differences between plants in all plant-soil compartments examined as well as in the degree of the plant × time interaction.Differences in the¹⁴C distribution between plants were due to both interspecific and developmental variation. In general, when comparing¹⁵N and¹⁴C quantities between species, many of the differences found between plants can be explained by the differences determined in the weight of shoot or root parts. Despite the fact that amounts of C released were greater in ryegrass than in the other plant-treatments no unequivocal evidence was found to show that the effects of plant-microorganism interactions on soil-N mineralization were greater under ryegrass. Possible mechanisms accounting for the partitioning of N found among plant biomass, soil biomass and soil residues are discussed.     

Temporal and spatial distribution of roots and competition for nitrogen in pea-barley intercrops – a field study employing 32P technique

Root system dynamics, productivity and N use were studied in inter- and sole crops of field pea (Pisum sativum L.) and spring barley (Hordeum vulgare L.) on a temperate sandy loam. A ³²P tracer placed at a depth of 12.5, 37.5, 62.5 or 87.5 cm was employed to determine root system dynamics by sampling crop leaves at 0, 15, 30 and 45 cm lateral distance. ¹⁵N addition was used to estimate N₂ fixation by pea, using sole cropped barley as reference crop. The Land Equivalent Ratio (LER), which is defined as the relative land area under sole crops that is required to produce the yields achieved in intercropping, were used to compare the crop growth in intercrops relative to the respective sole crops.The ³²P appearance in leaves revealed that the barley root system grows faster than that of pea. P uptake by the barley root system during early growth stages was approximately 10 days ahead of that of the pea root system in root depth and lateral root distribution. More than 90% of the P uptake by the pea root system was confined to the top 12.5 cm of soil, whereas barley had about 25–30% of tracer P uptake in the 12.5 – 62.5 cm soil layer. Judging from this P uptake, intercropping caused the barley root system to grow deeper and faster lateral root development of both species was observed. Barley accumulated similar amounts of aboveground N when grown as inter- and sole crop, whereas the total aboveground N acquired by pea in the intercrop was only 16% of that acquired in the pea sole crop. The percentage of total aboveground N derived from N₂ fixation in sole cropped pea increased from 40% to 80% during the growth period, whereas it was almost constant at 85% in intercropped pea. The total amounts of N₂ fixed were 95 and 15 kg N ha^–1 in sole cropped and intercropped pea, respectively. Barley was the dominant component of the pea-barley intercrop, obtaining 90% of its sole crop yield, while pea produced only 15% of the grains of a sole crop pea. Intercropping of pea and barley improved the utilization of plant growth resources (LER > 1) as compared to sole crops. Root system distribution in time and space can partly explain interspecific competition. The ³²P methodology proved to be a valuable tool for determining root dynamics in intercropping systems.     

Effect of soil compaction on root growth and uptake of phosphorus

Summary Zea mays L. andLolium rigidum Gaud. were grown for 18 and 33 days respectively in pots containing three layers of soil each weighing 1 kg. The top and bottom layers were 100 mm deep and they had a bulk density of 1200 kg m^–3, while the central layer of soil was compacted to one of 12 bulk densities between 1200 and 1750 kg m^–3. The soil was labelled with³²P and³³P so that the contribution of the different layers of soil to the phosphorus content of the plant tops could be determined. Soil water potential was maintained between –20 and –100 kPa.Total dry weight of the plant tops and total root length were slightly affected by compaction of the soil, but root distribution was greatly altered. Compaction decreased root length in the compacted soil but increased root length in the overlying soil. Where bulk density was 1550 kg m^–3, root length in the compacted soil was about 0.5 of the maximum. At that density, the penetrometer resistance of the soil was 1.25 and 5.0 MPa and air porosity was 0.05 and 0.14 at water potentials of –20 and –100 kPa respectively, and daytime oxygen concentrations in the soil atmosphere at time of harvest were about 0.1 m³m^–3. Roots failed to grow completely through the compacted layer of soil at bulk densities 1550 kg m^–3. No differences were detected in the abilities of the two species to penetrate compacted soil.Ryegrass absorbed about twice as much phosphorus from uncompacted soil per unit length of root as did maize. Uptake of phosphorus from each layer of soil was related to the length of root in that layer, but differences in uptake between layers existed. Phosphorus uptake per unit length of root was higher from compacted than from uncompacted soil, particularly in the case of ryegrass at bulk densities of 1300–1500 kg m^–3.     

Measurement of microbial biomass phosphorus in rhizosphere soil

³²P-labelled monocalcium phosphate solution was supplied by point injection to the root system of wheat plants grown in soil cores in a controlled environment. There was no detectable incorporation of³²P into organic P fractions in the soil remaining after roots were removed, confirming field observations. The techniques used to measure organic P (including biomass P) could detect an incorporation of³²P into soil microbial biomass equivalent to 0.3 μgP.g^?1 soil, compared to a total soil biomass P content estimated to be ca. 6.5 μgP.g^?1 soil. The limited incorporation of the added P into microbial biomass in the root-free soil may be due partly to a limited diffusion of³²P into the non-rhizosphere soil and partly to the removal of³²P-labelled microbial biomass adhering to or in very close association with the root surface. it is proposed that in studies of soil nutrient status, total soil biomass P (roots + soil flora + microfauna) should be measured, rather than attempting an estimate of microbial P. A sequential extraction procedure using a single soil sample, where a biocide is added to the extracting solution, is proposed as an alternative to the conventional procedure for measuring soil biomass P where two soil samples, one treated with a biocide, are extracted simultaneously.     

Phosphorus efficiency and phosphorus remobilization in two sorghum (Sorghum bicolor (L.) Moench) cultivars

With two sorghum cultivars differing in P efficiency a P uptake experiment (³²P/³³P labelling) was carried out followed by a period of P deficiency. The tendency of the total P distribution and redistribution pattern was rather similar in both sorghum cultivars. Although in the cultivar with a greater P absorbing capacity per unit root weight a higher proportion of the P was found in the inorganic P soluble fraction this is not necessarily an indication of a higher vacuolar affinity for P. Under P deficiency in both cultivars a rapid decrease of the TCA soluble P fraction in the leaves was observed. Before complete exhaustion of this fraction the TCA insoluble P fraction was also markedly reduced. In the roots the total P content was maintained fairly constant with a distinct shift in favour of the insoluble fraction occurring during the period of P deficiency. It is assumed that in the P efficient sorghum cultivar producing more dry matter per increment of P absorbed, rather inherent growth promoting factors contribute to the intraspecific P efficiency by a stimulation of the intensity of P redistribution and thus compensate for the lower P absorbing capacity of its roots.     

How does nitrogen availability alter rhizodeposition in Lolium multiflorum Lam. during vegetative growth?

The objective of this work was to determine if the impact of nitrogen (N) on the release of organic carbon (C) into the soil by roots (rhizodeposition) correlated with the effect of this nutrient on some variables of plant growth. Lolium multiflorum Lam. was grown at two levels of N supply, either in sterile sand percolated with nutrient solution or in non-sterile soil. The axenic sand systems allowed continuous quantification of rhizodeposition and accurate analysis of root morphology whilst the soil microcosms allowed the study of ¹⁴C labelled C flows in physico-chemical and biological conditions relevant to natural soils. In the axenic sand cultures, enhanced N supply strongly increased the plant biomass, the plant N content and the shoot to root ratio. N supply altered the root morphology by increasing the root surface area and the density of apices, both being significantly positively correlated with the rate of organic C release by plant roots before sampling. This observation is consistent with the production of mucilage by root tips and with mechanisms of root exudation reported previously in the literature, i.e. the passive diffusion of roots solutes along the root with increased rate behind the root apex. We proposed a model of root net exudation, based on the number of root apices and on root soluble C that explained 60% of the variability in the rate of C release from roots at harvest. The effects of N on plant growth were less marked in soil, probably related to the relatively high supply of N from non-fertiliser soil-sources. N fertilization increased the shoot N concentration of the plants and the shoot to root ratio. Increased N supply decreased the partitioning of ¹⁴C to roots. In parallel, N fertilisation increased the root soluble ¹⁴C and the ¹⁴C recovered in the soil per unit of root biomass, suggesting a stimulation of root exudation by N supply. However, due to the high concentration of N in our unfertilised plants, this stimulation was assumed to be very weak because no significant effect of N was observed on the microbial C and on the bacterial abundance in the rhizosphere. Considering the difficulties in evaluating rhizodeposition in non sterile soil, it is suggested that the root soluble C, the root surface area and the root apex density are additional relevant variables that should be useful to measure along with the variables that are commonly determined when investigating how plant functioning impacts on the release of C by roots (i.e soil C, C of the microbial biomass, rhizosphere respiration).     

Role of root hairs in phosphorus depletion from a macrostructured soil

One rape (Brassica napus cv. Wesroona) plant and four cotton (Gossypium hirsutum cv. Sicot 3) plants were grown in plastic cells containing soil labelled with 407 kBq of³³P g⁻¹ soil. After 5–8 days of growth, the³³P depletion zones of all plants were autoradiographed and³³P uptake by plants was measured. The autoradiographs were scanned with a microdensitometer and the optical densities at several places within the³³P depletion zones of roots were obtained. The volume of soil explored by root hairs was estimated from measurements of root diameters and lengths of roots and root hairs. About half of the total³³P depleted by cotion roots came from outside the root hair cylinder whereas most of³³P taken up by rape was from within the root hair cylinder. Plants grown in a macrostructured soil may have roots growing in voids, within aggregates or on the surfaces of aggregates. The results of this study demonstrate that root hairs have a strong influence on the accessibility of phosphorus to roots in such a soil, and thus on the phosphorus nutrition of plants.     

Influence of light intensity on the distribution of carbon and consequent effects on mineralization of soil nitrogen in a barley (Hordeum vulgare L.)-soil system

A pot experiment was conducted in a ¹⁴C-labelled atmosphere to study the influence of living plants on organic-N mineralization. The soil organic matter had been labelled, by means of a 200-days incubation, with ¹⁵N. The influence of the carbon input from the roots on the formation of microbial biomass was evaluated by using two different light intensities (I). Mineralization of ¹⁵N-labelled soil N was examined by following its fate in both the soil biomass and the plants. Less dry matter accumulated in shoots and roots at the lower light intensity. Furthermore, in all the plant-soil compartments examined, with the exception of rhizosphere respiration, the proportion of net assimilated ¹⁴C was lower in the low-I treatment than in the high-I treatment. The lower rates of ¹⁴C and ¹⁵N incorporation into the soil biomass were associated with less root-derived ¹⁴C. During the chamber period (¹⁴CO₂-atmosphere), mineralized amounts of ¹⁵N (measured as plant uptake of ¹⁵N) were small and represented about 6.8 to 7.8% of the initial amount of organic ¹⁵N in the soil. Amounts of unlabelled N found in the plants, as a percentage of total soil N, were 2.5 to 3.3%. The low availability of labelled N to microorganisms was the result of its stabilization during the 210 days of soil incubation. Differences in carbon supply resulted in different rates of N mineralization which is consistent with the hypothesis that roots induce N mineralization. N mineralization was higher in the high-I treatment. On the other hand, the rate of mineralization of unlabelled stable soil N was lower than labelled soil ¹⁵N which was stabilized. The amounts of ¹⁵N mineralized in planted soil during the chamber period (43 days) which were comparable with those mineralized in unplanted soil incubated for 210 days, also suggested that living plants increased the turnover rate of soil organic matter.     

Carbon distribution in grass (Festuca pratensis L.) during regrowth after cutting—utilization of stored and newly assimilated carbon

Distribution of net assimilated C in meadow fescue (Fectuca pratensi L.) was followed before and after cutting of the shoots. Plants were continuously labelled in a growth chamber with ¹⁴C-labelled CO₂ in the atmosphere from seedling to cutting and with ¹³C-labelled CO₂ in the atmosphere during regrowth after the cutting. Labelled C, both ¹⁴C and ¹³C, was determined at the end of the two growth periods in shoots, crowns, roots, soil and rhizosphere respiration. Distribution of net assimilated C followed almost the same pattern at the end of the two growth periods, i.e. at the end of the ¹⁴C- and the ¹³C-labelling periods. Shoots retained 71–73% of net assimilated C while 9% was detected in the roots and 11–14% was released from the roots, determined as labelled C in soil and as rhizosphere respiration. At the end of the 2nd growth period, after cutting and regrowth, 21% of the residual plant ¹⁴C at cutting (¹⁴C in crowns and roots) was found in the new shoot biomass. A minor part of the residual plant ¹⁴C, 12%, was lost from the plants. The decreases in ¹⁴C in crowns and roots during the regrowth period suggest that ¹⁴C in both crowns and roots was translocated to new shoot tissue. Approximately half of the total root C at the end of the regrowth period after cutting was ¹³C-labelled C and thus represents new root growth. Root death after cutting could not be determined in this experiment, since the decline in root ¹⁴C during the regrowth period may also be assigned to root respiration, root exudation and translocation to the shoots. ei]{gnH}{fnLambers} ei]{gnA C}{fnBorstlap}     

Phosphorus retention and movement across an ombrotrophic-minerotrophic peatland gradient

An understanding of the mechanisms controlling nutrient availability and retention in and across ecosystems allows for a greater understanding of the role of nutrients in maintaining ecosystem structure and function. To examine the underlying mechanisms of phosphorus (P) cycling in northern peatlands, we compared the retention and movement of P across a natural hydrologic/pH gradient in nine peatlands by applying as a light rain an in situ tracer amount of ³²PO₄ ^–3 to track changes in P pools (vegetation, soil, microbial) over 30 days. The ³¹P concentrations of available P, microbial P, and the root P at 10–20 cm did not differ across the gradient, although total soil P and aboveground vegetation P content (g m^–2) increased from bog to rich fen. Total retention of ³²P in the first 24 hours of application was greatest in the bogs and intermediate fens (90–100%) and was very low (20–50%) in the rich fens. Retention of P in the different pools was dependent on the type of peatland and changed with time. In the first 24 hours in the bogs and intermediate fens, the microbial pool contained the largest amount of ³²P, but by the seventh day, the aboveground vegetation contained the largest amount. In the rich fen, the recovered ³²P was almost equally divided between the aboveground vegetation and the litter layer with very little in other pools. Therefore, although bogs and intermediate fens have a small total P pool, they have similar P availability to rich fens because of rapid cycling and efficient retention of P.     

Effect of rice plants on methane production and rhizospheric metabolism in paddy soil

In order to elucidate the effects of rice plants on CH₄ production, we conducted experiments with soil slurries and planted rice microcosms. Methane production in anoxic paddy soil slurries was stimulated by the addition of rice straw, of unsterile or autoclaved rice roots, and of the culture fluid in which rice plants had axenically been cultivated. The addition of these compounds also increased the concentrations of acetate and H₂, precursors of CH₄ production, in the soil. Planted compared to unplanted paddy soil microcosms exhibited lower porewater CH₄ concentrations but higher CH₄ emission rates. They also exhibited higher sulfate concentrations but similar nitrate concentrations. Concentrations of acetate, lactate and H₂ were not much different between planted and unplanted microcosms. Pulse labeling of rice plants with¹⁴CO₂ resulted during the next 5 days in transient accumulation of radioactive lactate, propionate and acetate, and after the second day of incubation in the emission of¹⁴CH₄. Most of the radioactivity (40–70%) was incorporated into the above-ground biomass of rice plants. However, during a total incubation of 16 days about 3–6% of the applied radioactivity was emitted as¹⁴CH₄, demonstrating that plant-derived carbon was metabolized and significantly contributed to CH₄ production. The sequence of the appearance of radioactive products and their specific radioactivities indicate that CH₄ was produced from root exudates by a microbial community consisting of fermenting and methanogenic bacteria.     

Rhizosphere Properties of Poaceae Genotypes Under P-limiting Conditions

Plant genotypes differ in P efficiency, i.e. their capacity to grow in soil with low P availability. Plant properties such as root and root hair length, release of P mineralising and mobilising compounds by the roots and P requirement for optimal growth are known to influence P efficiency. In order to improve the understanding of the role of rhizosphere properties in plant P uptake, we grew three Poaceae genotypes [two wheat (Triticum aestivum L.) genotypes (the P-efficient Goldmark and the P-inefficient Janz), and the Australian native grass Austrostipa densiflora L.] to maturity in an acidic loamy sand with low P availability. Addition of 120 mg P as FePO₄ kg⁻¹ (P120) improved the growth of all three genotypes. In both P0 and P120, growth and P uptake were smaller in Janz than in Goldmark. During the vegetative phase, growth and P uptake of Austrostipa were smaller than in Goldmark in P0 but greater in P120. These differences can be explained by plant properties such as root growth, specific P uptake, mobilisation of inorganic and organic P by root exudates and P utilisation efficiency. In P120, P availability in the rhizosphere was least in Janz and greatest in Austrostipa. Microbial biomass P in the rhizosphere was least in Janz. Acid phosphatase activity was greatest in the rhizosphere of Austrostipa and least in Janz. Plant growth and P uptake were positively correlated with microbial P, acid phosphatase activity and resin P in the rhizosphere, suggesting that microorganisms contribute to uptake of P by plants in this soil. Microbial community composition in the rhizosphere [analysed by fatty acid methylester (FAME) analysis and denaturing gradient gel electrophoresis (DGGE)] differed among genotypes, changed during plant development and was affected by P addition to the soil. Genotype-specific microbial community composition in the rhizosphere may have contributed to the observed differential capacity of plants to grow at low P availability.     

Turnover of residual 15N-labelled fertilizer N in soil following harvest of oilseed rape (t Brassica napus L.)

We studied the fate of ¹⁵N-labelled fertilizer nitrogen in a sandy loam soil after harvest of winter oilseed rape (Brassica napus L. cv. Ceres) given 100 or 200 kg N ha^-1 in spring, with or without irrigation. Our main objective was to quantify the temporal variations of the soil mineral N, the extractable soil organic N and soil microbial biomass N, and fertilizer derived N in these pools during autumn and winter. Nitrogen use efficiency of the oilseed rape crop varied from 47% of applied N in the 100N, irrigated treatment to 34% in the 200N, non-irrigated treatment. However, only in the latter treatment did we find significantly higher fertilizer derived soil mineral N than in the three other treatments which all had low soil mineral N contents at the first sampling after harvest (8 days after stubble tillage). Between 31% and 42% of the applied N could not be accounted for in the harvested plants or 0-15 cm soil layer at this first sampling. Over the following autumn and winter none of the remaining fertilizer derived soil N was lost from the 0–5 cm depth, but from the 5–15 cm depth a marked proportion of N derived from fertilizer was lost, probably by leaching. Negligible amounts of fertilizer derived extractable soil organic and mineral N (<1 kg N ha^-1, 0-15 cm) were found in all treatments after the first sampling.Soil microbial biomass N was not significantly affected by treatments and showed only small temporal variability (±11% of the mean 76 kg N ha^-1, 0- 15 cm depth). Surprisingly, the average amount of soil microbial biomass N derived from fertilizer was significantly affected by the treatments, with the extremes being 5.5 and 3.1 kg N ha^-1 in the 200N, non-irrigated and 100N, irrigated treatments, respectively. Also, the estimated exponential decay rate of microbial biomass N derived from fertilizer, differed greatly (2 fold) between these two treatments, indicating highly different microbial turnover rates in spite of the similar total microbial biomass N values. In studies utilising ¹⁵N labelling to estimate turnover rates of different soil organic matter pools this finding is of great importance, because it may question the assumption that turnover rates are not affected by the insertion of the label.     

Climate Change Affects Carbon Allocation to the Soil in Shrublands

Climate change may affect ecosystem functioning through increased temperatures or changes in precipitation patterns. Temperature and water availability are important drivers for ecosystem processes such as photosynthesis, carbon translocation, and organic matter decomposition. These climate changes may affect the supply of carbon and energy to the soil microbial population and subsequently alter decomposition and mineralization, important ecosystem processes in carbon and nutrient cycling. In this study, carried out within the cross-European research project CLIMOOR, the effect of climate change, resulting from imposed manipulations, on carbon dynamics in shrubland ecosystems was examined. We performed a ¹⁴C-labeling experiment to probe changes in net carbon uptake and allocation to the roots and soil compartments as affected by a higher temperature during the year and a drought period in the growing season. Differences in climate, soil, and plant characteristics resulted in a gradient in the severity of the drought effects on net carbon uptake by plants with the impact being most severe in Spain, followed by Denmark, with the UK showing few negative effects at significance levels of p 0.10. Drought clearly reduced carbon flow from the roots to the soil compartments. The fraction of the ¹⁴C fixed by the plants and allocated into the soluble carbon fraction in the soil and to soil microbial biomass in Denmark and the UK decreased by more than 60%. The effects of warming were not significant, but, as with the drought treatment, a negative effect on carbon allocation to soil microbial biomass was found. The changes in carbon allocation to soil microbial biomass at the northern sites in this study indicate that soil microbial biomass is a sensitive, early indicator of drought- or temperature-initiated changes in these shrubland ecosystems. The reduced supply of substrate to the soil and the response of the soil microbial biomass may help to explain the observed acclimation of CO₂ exchange in other ecosystems.     

Utilization of organic phosphorus in calcium chloride extracts of soil by barley plants and hydrolysis by acid and alkaline phosphatases

Organic phosphorus is often a major part of total phosphorus in soil solution. The role of this fraction as a P source for plants and the mechanism involved in its transfer from soil to plant is still unclear. We studied the utilization of organic phospharus in 0.01 M calcium chloride extracts by barley and its hydrolysis by isolated acid and alkaline phosphatases. Calcium chloride extracts were used as a nutrient solution in 24 hrs assays. Concentration of organic and inorganic P in equilibrium calcium chloride extracts was 7.8 and 1.8 µmol P L^-1, respectively, which was similar to the soil solution P concentration. When soil microbial biomass was destroyed by autoclaving, organic P concentration increased to 64.8 µmol P L^-1 whereas the inorganic P was hardly changed. Inoculation of the autoclaved soil with non-sterile soil and incubation for 5 days decreased the organic P concentration to 27.9 µmol P L^-1 but did not change inorganic P. In this study barley plants utilized organic P from all extracts. The greatest reduction of organic P concentration occurred in fresh extracts of the autoclaved soil. Inorganic P was depleted to traces in all extracts. Organic P was hydrolyzed by isolated acid and alkaline phosphatases. We conclude that organic P in soil solution is a heterogeneous pool of organic P compounds originating from microbial biomass. Its initial availability to plants was nigh but its susceptibility to phosphatase hydrolysis was quickly reduced but not completely lost.