Influence of plant roots on C and P metabolism in soil

Summary A technique for studying the modification of soil by plant roots is described. Using it, soil zones differently affected by plant roots can be separated for subsequent analysis. With this method, the transfer of C from roots of¹⁴C-labelled maize plants into soil and the change in soil C and P fractions were investigated.The results show that the C released from roots to soil was 13% of the total assimilated C. The remaining root-derived C in soil was relatively small (15%). Maize roots induced a decrease in organic soil C and in both total and isotopically exchangeable soil P. On the other hand they increased the microbial biomass C, phosphatase activity, bicarbonate extractable organic P and phospholipid P and enhanced the incorporation of³²P into organic P fractions. Both root C and root influences were detectable outside the immediate root zone.These results demonstrate an intensive C turnover and P mobilization in the rhizosphere soil, including some organic P fractions, and suggest that the actual rhizosphere may be greater than is generally assumed.     

Distribution of soil fractions and location of soil bacteria in a vertisol under cultivation and perennial grass

Effects of soil management on soil characteristics were investigated on the rhizosphere (RPP) and the nonrhizosphere (NRPP) soil of a re-grass vertisol underDigitaria decumbens and in the soil under continuous cultivation (CC). A low energy technique allowed to separate eight size and density fractions, including macro- and micro-aggregates while preserving soil bacteria. Organic C and N, microbial biomass C and the number of total bacteria (AODC) and ofAzospirillum brasilense and their distribution were determined in soil fractions isolated from the CC, NRPP and RPP soils. Soil macroaggregates (>2000 m) were similarly predominant in the NRPP and RPP soils when the dispersible clay size fraction (<2 m) respresented more than 25% of the CC soil mass. The main increase of C content in RPP originated from the macroaggregates (> 2000 m) and from the root fraction, not from the finer separates. The proportion of organic C as microbial biomass C revealed the low turnover of microbial C in the PP situations, especially in the clay size fraction of the NRPP soil. A common shift of AODC toward the finer separates from planted soils (CC and RPP) revealed the influence of living plants on the distribution of soil bacteria. The relative abundance ofA. brasilense showed the presence of the active roots ofDigitaria in the macroaggregates and their contact with the dispersible clay size fraction of the rhizosphere soil.     

Modelling the effect of active roots on soil organic matter turnover

The aim of this experiment was to study the effect of living roots on soil carbon metabolism at different decomposition stages during a long-term incubation. Plant material labelled with ¹⁴C and ¹⁵N was incubated in two contrasting soils under controlled laboratory conditions, over two years. Half the samples were cropped with wheat (Triticum aestivum) 11 times in succession. At earing time the wheat was harvested, the roots were extracted from the soil and a new crop was started. Thus the soils were continuously occupied by active root systems. The other half of the samples was maintained bare, without plants under the same conditions. Over the 2 years, pairs of cropped and bare soils were analysed at eight sampling occasions (total-, plant debris-, and microbial biomass-C and -¹⁴C). A five compartment (labile and recalcitrant plant residues, labile microbial metabolites, microbial biomass and stabilised humified compounds) decomposition model was fitted to the labelled and soil native organic matter data of the bare and cropped soils. Two different phases in the decomposition processes showed a different plant effect. (1) During the initial fast decomposition stage, labile ¹⁴C-material stimulated microbial activities and N immobilisation, increasing the ¹⁴C-microbial biomass. In the presence of living roots, competition between micro-organisms and plants for inorganic N weakly lowered the measured and predicted total-¹⁴C mineralisation and resulted in a lower plant productivity compared to subsequent growths. (2) In contrast, beyond 3–6 months, when the labile material was exhausted, during the slow decomposition stage, the presence of living roots stimulated the mineralisation of the recalcitrant plant residue-¹⁴C in the sandy soil and of the humified-¹⁴C in the clay soil. In the sandy soil, the presence of roots also substantially stimulated decomposition of old soil native humus compounds. During this slow decomposition stage, the measured and predicted plant induced decrease in total-¹⁴C and -C was essentially explained by the predicted decrease in humus-¹⁴C and -C. The ¹⁴C-microbial biomass (MB) partly decayed or became inactive in the bare soils, whereas in the rooted soils, the labelled MB turnover was accelerated: the MB-¹⁴C was replaced by unlabelled-C from C derived from living roots. At the end of experiment, the MB-C in the cropped soils was 2.5–3 times higher than in the bare soils. To sustain this biomass and activity, the model predicted a daily root derived C input (rhizodeposition), amounting to 5.4 and 3.2% of the plant biomass-C or estimated at 46 and 41% of the daily net assimilated C (shoot + root + rhizodeposition C) in the clay and sandy soil, respectively. This revised version was published online in June 2006 with corrections to the Cover Date.     

Spatial and temporal distribution of carbon isotopes in soil organic matter at the Dinghushan Biosphere Reserve, South China

The spatial and temporal distribution of carbon isotopes (¹³C, ¹⁴C) in soil organic matter (SOM) were studied based on SOM content, SOM ¹⁴C and SOM ¹³C of thinly layered soil samples for six soil profiles with different elevations at the Dinghushan Biosphere Reserve (DHSBR), South China. The results indicate that variations of SOM ¹³C with depth of the soil profiles at different elevations are controlled by soil development, and correlate well with SOM composition in terms of SOM compartments with different turnover rates, and SOM turnover processes at the DHSBR. The effect of carbon isotope fractionation was obvious during transformation of organic matter (OM) from plant debris to SOM in topsoil and SOM turnover processes after the topsoil was buried, which resulted in great increments of OM ¹³C, respectively. Increments of SOM ¹³C of topsoil from ¹³C of plant debris were controlled by SOM turnover rates. Both topsoil SOM ¹³C and plant debris ¹³C increase with elevation, indicating regular changes in vegetation species and composition with elevation, which is consistent with the vertical distribution of vegetation at the DHSBR. The six soil profiles at different elevations had similar characteristics in variations of SOM ¹³C with depth, alterations of SOM contents with depth and that SOM ¹⁴C apparent ages increasing with depth, respectively. These are presumably attributed to the regular distribution of different SOM compartments with depth because of their regular turnover during soil development. Depth with the maximal SOM ¹³C value is different in mechanism and magnitude with penetrating depth of ¹⁴C produced by nuclear explosion into atmosphere from 1952 to 1962, and both indicate controls of topography and vegetation on the distribution of SOM carbon isotopes with depth. Elevation exerts indirect controls on the spatial and temporal distribution of SOM carbon isotopes of the studied mountainous soil profiles at the DHSBR. This study shows that mountainous soil profiles at different elevations and with distinctive aboveground vegetation are presumably ideal sites for studies on soil carbon dynamics in different climatic-vegetation zones.     

Carbon loss from the roots of tomato and pea seedlings grown in soil

Tomato and pea seedlings were grown for 14d and 28d with shoots in constant specific activity¹⁴CO₂ and the amounts and distribution of carbon within the plants and of that released into the soil from the roots were measured. The estimates of carbon loss were derived from measurements of¹⁴CO₂ respired from both the root and the accompanying microbial population and from the root derived¹⁴C-labelled organic carbon compounds in the soil. The relationship between plant growth and the loss of carbon and distribution of carbon within the plants are discussed.     

Sucrose application,soil microbial respiration and evolved carbon dioxide isotope enrichment under contrasting land uses

Heterotrophic decomposition of organic matter dictates that substrate supply rate, including energy and nutrients, can limit soil microbial activity. In New Zealand, soils are naturally deficient in nitrogen and phosphorus. Fertiliser application is a part of pastoral agriculture, the countrys most widespread land use. We postulated that organic soils under grazed pasture and pristine forest would be at the extremes of substrate quality and supply rate, and thus potential microbial response to food opportunities. Soil microbial responses to the addition of fresh energy (sucrose) were determined by laboratory experiments with root-free samples and intact cores including roots. Responses were quantified by respiration and respired carbon (C) isotope (¹³C) enrichment measurements. A supra-trace sucrose dose (0.002 mol kg^–1 (soil)) caused the forest soils microbial respiration rate to nearly double within 2 h. The peak response took 20 h, and saturation occurred beyond a sucrose dose of 0.05 mol kg^–1 (soil). Intact soil cores from the forest had similar respiration rates and responses. For root-free soil samples from the grazed pasture, respiration response to sucrose was nearly immediate, dose dependent, and there was up to a 9-fold increase in the rate. Intact cores from the pasture had much higher respiration rates, but a similar response to sucrose. For both soils, the similarity of sucrose application effects on respiration and relative ¹³C enrichment of the respired carbon was striking.     

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.     

Environmental manipulation treatment effects on the reactivity of water-soluble phenolics in a subalpine tundra ecosystem

Low soil organic matter content and limited soil water holding are the major natural constraint of dryland cropping on sandy soils in the Quebec boreal regions. We conducted a 3-yr (1994–1996) study in a boreal sandy soil, Ferro-Humic Podzol (Spodosols), to determine the potential of Sphagnum peat for improving soil organic matter (SOM), water holding capacity, bulk density (BD), plant leaf nutrient status, and potato and barley yields. The cropping was a rotation of 2-yr potato (Solanum tuberosum L. Superior) and 1-yr barley (Hordeum vulgare L. Chapais). The treatments consisted of Sphagnum peat at rates of 0, 29, 48, and 68 Mg ha^–1 3-yr^–1 on a dry weight basis, and granular N-P-K fertilizers (12-7.5-7) at rates of 1.4, 1.6, and 1.8 Mg ha^–1 yr^–1, respectively, arranged in a split-block design. The peat-amended soils were higher in water content (SWC), SOM and total porosity but lower in BD and N than neighboring non-peat soils (P < 0.05). Effects of peat and fertilizer treatments and their interaction were significant on potato leaf N, Ca, Mg, and P, tuber yield, dry weight, harvested N and tuber specific gravity (P < 0.05), depending on year. Potato tuber yield and N increased simultaneously up to 30% (compared to the control), and were significantly correlated with SWC, SOM, BD, and NO₃-N (–0.52 r 0.80). In the 3rd year, the linear effect of peat treatments was significant on barley grain yield. In 1995 there was a decline of 4.5–7.3% of SOM of the previous year level. It is suggested that Sphagnum peat at a rate of 48 Mg ha^–1 had the potential for improving sandy soil productivity. A longer-term investigation of soil water, N, SOM pool and crop yield changes is necessary to better understand the physical, chemical and biological processes of peat in cropping systems and to maximize the benefits of peat applications.     

Influence of rhizodeposition under elevated CO2 on plant nutrition and soil organic matter

Atmospheric CO₂ concentrations can influence ecosystem carbon storage through net primary production (NPP), soil carbon storage, or both. In assessing the potential for carbon storage in terrestrial ecosystems under elevated CO₂, both NPP and processing of soil organic matter (SOM), as well as the multiple links between them, must be examined. Within this context, both the quantity and quality of carbon flux from roots to soil are important, since roots produce specialized compounds that enhance nutrient acquisition (affecting NPP), and since the flux of organic compounds from roots to soil fuels soil microbial activity (affecting processing of SOM).From the perspective of root physiology, a technique is described which uses genetically engineered bacteria to detect the distribution and amount of flux of particular compounds from single roots to non-sterile soils. Other experiments from several labs are noted which explore effects of elevated CO₂ on root acid phosphatase, phosphomonoesterase, and citrate production, all associated with phosphorus nutrition. From a soil perspective, effects of elevated CO₂ on the processing of SOM developed under a C4 grassland but planted with C3 California grassland species were examined under low (unamended) and high (amended with 20 g m^–2 NPK) nutrients; measurements of soil atmosphere 13C combined with soil respiration rates show that during vegetative growth in February, elevated CO₂ decreased respiration of carbon from C4 SOM in high nutrient soils but not in unamended soils.This emphasis on the impacts of carbon loss from roots on both NPP and SOM processing will be essential to understanding terrestrial ecosystem carbon storage under changing atmospheric CO₂ concentrations.Abbreviations SOM soil organic matter - NPP net primary productivity - NEP net ecosystem productivity - PNPP p-nitrophenyl phosphate     

Elevated CO2 effects on carbon and nitrogen cycling in grass/clover turves of a Psammaquent soil

Effects of elevated CO₂ (525 and 700 L L^–1), and a control (350 L L^–1 CO₂), on biochemical properties of a Mollic Psammaquent soil in a well-established pasture of C3 and C4 grasses and clover were investigated with continuously moist turves in growth chambers over four consecutive seasonal temperature regimes from spring to winter inclusive. After a further spring period, half of the turves under 350 and 700 L L^–1 were subjected to summer drying and were then re-wetted before a further autumn period; the remaining turves were kept continuously moist throughout these additional three consecutive seasons. The continuously moist turves were then pulse-labelled with ¹⁴C-CO₂ to follow C pathways in the plant/soil system during 35 days.Growth rates of herbage during the first four seasons averaged 4.6 g m^–2 day^–1 under 700 L L^–1 CO₂ and were about 10% higher than under the other two treatments. Below-ground net productivity at the end of these seasons averaged 465, 800 and 824 g m^–2 in the control, 525 and 700 L L^–1 treatments, respectively.in continuously moist soil, elevated CO₂ had no overall effects on total, extractable or microbial C and N, or invertase activity, but resulted in increased CO₂-C production from soil, and from added herbage during the initial stages of decomposition over 21 days; rates of root decomposition were unaffected. CO₂ produced h^–1 mg^–1 microbial C was about 10% higher in the 700 L L^–1 CO₂ treatment than in the other two treatments. Elevated CO₂ had no clearly defined effects on N availability, or on the net N mineralization of added herbage.In the labelling experiment, relatively more ¹⁴C in the plant/soil system occurred below ground under elevated CO₂, with enhanced turnover of ¹⁴C also being suggested.Drying increased levels of extractable C and organic-N, but decreased mineral-N concentrations; it had no effect on microbial C, but resulted in lowered microbial N in the control only. In soil that had been previously summer-dried, CO₂ production was again higher, but net N mineralization was lower, under elevated CO₂ than in the control after autumn pasture growth.Over the trial period of 422 days, elevated CO₂ generally appears to have had a greater effect on soil C turnover than on soil C pools in this pasture ecosystem.     

Moisture retention properties of a mycorrhizal soil

The water relations of arbuscular mycorrhizal plants have been compared often, but virtually nothing is known about the comparative water relations of mycorrhizal and nonmycorrhizal soils. Mycorrhizal symbiosis typically affects soil structure, and soil structure affects water retention properties; therefore, it seems likely that mycorrhizal symbiosis may affect soil water relations. We examined the water retention properties of a Sequatchie fine sandy loam subjected to three treatments: seven months of root growth by (1) nonmycorrhizal Vigna unguiculata given low phosphorus fertilization, (2) nonmycorrhizal Vigna unguiculata given high phosphorus fertilization, (3) Vigna unguiculata colonized by Glomus intraradices and given low phosphorus fertilization. Mycorrhization of soil had a slight but significant effect on the soil moisture characteristic curve. Once soil matric potential (_m) began to decline, changes in _m per unit change in soil water content were smaller in mycorrhizal than in the two nonmycorrhizal soils. Within the range of about –1 to –5 MPa, the mycorrhizal soil had to dry more than the nonmycorrhizal soils to reach the same _m. Soil characteristic curves of nonmycorrhizal soils were similar, whether they contained roots of plants fed high or low phosphorus. The mycorrhizal soil had significantly more water stable aggregates and substantially higher extraradical hyphal densities than the nonmycorrhizal soils. Importantly, we were able to factor out the possibly confounding influence of differential root growth among mycorrhizal and nonmycorrhizal soils. Mycorrhizal symbiosis affected the soil moisture characteristic and soil structure, even though root mass, root length, root surface area and root volume densities were similar in mycorrhizal and nonmycorrhizal soils.     

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.     

Soil organic carbon dynamics in cleared temperate forest spodosols converted to maize cropping

In southwest France, sandy spodosols have developed from Quaternary sandy eolian deposits. On these soils, numerous forest lands have been converted to continuous intensive maize cropping. A chronosequence study is realized by comparing organic C pools and ¹³C natural abundance of one forested and 6 agricultural sites, whose ages of cultivation range from 4 to 32 yr. ¹³C ratio is found to increase with time of cultivation. After 3 decades of intensive maize cropping, about half of the initial organic C content in the forest topsoil layer has disappeared. The fraction of C derived from maize crop increases during the first decades of cultivation, but its level is significantly lower than those observed in other soils, which indicates a high mineralization rate of organic C. In this context, soil characteristics associated to intensive agricultural practices lead to a rapid and large loss of C, whereas inputs from maize seem to have only a very small long-term contribution.     

Analysis of δ13C of CO2 distinguishes between microbial respiration of added C4-sucrose and other soil respiration in a C3-ecosystem

The main aim of this study was to test various hypotheses regarding the changes in ¹³C of emitted CO₂ that follow the addition of C₄-sucrose to the soil of a C₃-ecosystem. It forms part of an experimental series designed to assess whether or not the contributions from C₃-respiration (root and microbial) and C₄-respiration (microbial) to total soil respiration can be calculated from such changes. A series of five experiments, three on sieved (root-free) mor-layer material, and two in the field with intact mor-layer (and consequently with active roots), were performed. Both in the experiments on sieved mor-layer and the field experiments, we found a C₄-sucrose-induced increase in C₃-respiration that accounted for between 30% and 40% of the respiration increase 1 h after sucrose addition. When the course of C₃-, C₄- and total respiration was followed in sieved material over four days following addition of C₄-sucrose, the initially increased respiration of C₃-C was transient, passing within less than 24 h. In a separate pot experiment, neither ectomycorrhizal Pinus sylvestrisL. roots nor non-mycorrhizal roots of this species showed respiratory changes in response to exogenous sucrose. No shift in the ¹³C of the evolved CO₂ after adding C₃-sucrose to sieved mor-layer material was found, confirming that the sucrose-induced increase in respiration of endogenous C was not an artefact of discrimination against ¹³C during respiration. Furthermore, we conclude that the C₄-sucrose induced transient increase in C₃-respiration is most likely the result of accelerated turnover of C in the microbial biomass. Thus, neither respiration of mycorrhizal roots, nor processes discriminating against ¹³C were likely sources of error in the field. The estimated ¹³C of evolved soil CO₂ in three field experiments lay between –25.2 and –23.6. The study shows that we can distinguish between CO₂ evolved from microbial mineralisation of added C₄-sucrose, and CO₂ evolved from endogenous carbon sources (roots and microbial respiration).     

Acclimation of a dominant mangrove plant (<Emphasis Type="Italic">Kandelia candel</Emphasis>) to soil texture and its response to canopy shade

Kandelia candel, a dominant plant species in Hong Kong mangroves, had different growth strategies during its seedling stage for acclimation to various soil types. Leaves of the seedlings grown in sandy soils (coarse texture) were significantly thicker than those in loamy–sandy and silty soils (fine texture). However, leaf weight per unit leaf area of seedlings grown in sandy soils was similar to that in loamy-sandy soils and was 1.60 times that in silty soils. These indicate that K. candel in sandy soils had developed a more loose leaf structure and a xerophilous characteristic during its seedling stage for acclimation to the features of coarse textured soils, which were low water holding capacity and low mineral content. In addition, for young seedlings grown in sandy and loamy–sandy soils, more biomass was allocated to roots than in silty soils, indicating that K. candel seedlings in sandy soils developed stronger roots for anchorage and water absorption; another xerophilous characteristic to acclimate to loose and coarse textured soils. As the plants became more mature, no significant difference in leaf thickness of the saplings was found between loamy-sandy and sandy soils. In order to acclimate to the low water holding capacity in sandy soils, K. candel seedlings had lower physiological activities including lower chlorophyll content; lower activities of root oxidase, nitrate reductase, peroxidase and superoxide dismutase but higher malonaldehyde contents than those in silty soils. Canopy shade is an important factor affecting the growth and physiology of K. candel. The seedling grew worse under the mangrove canopy than that in the open gap, with smaller leaf area, specific leaf area, leaf number and biomass. The seedlings under the canopy eventually died and no saplings were found under canopy shade, implying K. candel is a shade intolerant species and canopy shade might force its newly recruited individuals to expand out of the forest.     

Altered utilization patterns of young and old soil C by microorganisms caused by temperature shifts and N additions

To determine if changes in microbial community composition and metabolic capacity alter decomposition patterns of young and old soil carbon pools, we incubated soils under conditions of varying temperature, N-availability, and water content. We used a soil from a pineapple plantation (CAM; ¹³C litter = –14.1) that had previously been under tropical forest (C3; ¹³C soil carbon = –26.5). Forest derived carbon represented 'old' carbon and plantation inputs represented 'new' carbon. In order to differentiate utilization of young (< 14 years) and old (> 14 years) soil carbon, we measured the ¹³C of respired CO₂ and microbial phospholipid fatty acids (PLFAs) during a 103 day laboratory incubation. We determined community composition (PLFA and bacterial intergenic transcribed spacer (ITS) analysis) in addition to carbon degrading and nutrient releasing enzyme activities. We observed that greater quantities of older carbon were respired at higher temperatures (20 and 35°C) compared to the lower temperature (5°C). This effect could be explained by changes in microbial community composition and accompanying changes in enzyme activities that affect C degradation. Nitrogen addition stimulated the utilization of older soil carbon, possibly due to greater peroxidase activity, but microbial community composition was unaffected by this treatment. Increasing soil moisture had no effect on the utilization of older SOM, but enzyme activity typically declined. Increased oxidative enzyme activities in response to elevated temperature and nitrogen additions point to a plausible mechanism for alterations in C resource utilization patterns.     

Soil organic sulfur dynamics in a coniferous forest

Sulfate microbial immobilization and the mineralization of organic S were measured in vitro in soil horizons (LFH, Ae, Bhf, Bf and C) of the Lake Laflamme watershed (47°17 N, 71°14 O) using ³⁵SO₄. LFH samples immobilized from 23 to 77% of the added ³⁵SO₄ within 2 to 11 days. The ³⁵SO₄ microbial immobilization increased with temperature and reached an asymptote after a few days. The mineral soil generally immobilized less than 20% of the added ³⁵SO₄, and an asymptote was reached after 2 days. An isotopic equilibrium was rapidly reached in mineral horizons. A two-compartment (SO₄ and organic S) model adequately described ³⁵SO₄ microbial immobilization kinetics. The active organic reservoir in the whole soil profile represented less than 1% of the total organic S. The average concentrations of dissolved organic S (DOS) in the soil solutions leaving the LFH, Bhf and Bf horizons were respectively 334, 282 and 143 µgL^–1. Assuming that the DOS decrease with soil depth corresponded to the quantities adsorbed in the B horizons, we estimated that 12 800 kgha^–1 of organic S could have been formed since the last glaciation, which is about 13 times the size of the actual B horizons reservoirs. Our results suggest that the organic S reservoirs present in mineral forest soils are mostly formed by the DOS adsorption resulting from incomplete litter decomposition in the humus layer. The capability of these horizons to immobilize SO₄ from the soil solution would be restricted to a 1% active fraction composed of microorganisms. Despite their refractory nature, these reservoirs can, however, be slowly decomposed by microorganisms and contribute to the S-SO₄ export from the watershed in the long term.     

Microbial activity and soil respiration under nitrogen addition in Alaskan boreal forest

Climate warming could increase rates of soil organic matter turnover and nutrient mineralization, particularly in northern high‐latitude ecosystems. However, the effects of increasing nutrient availability on microbial processes in these ecosystems are poorly understood. To determine how soil microbes respond to nutrient enrichment, we measured microbial biomass, extracellular enzyme activities, soil respiration, and the community composition of active fungi in nitrogen (N) fertilized soils of a boreal forest in central Alaska. We predicted that N addition would suppress fungal activity relative to bacteria, but stimulate carbon (C)‐degrading enzyme activities and soil respiration. Instead, we found no evidence for a suppression of fungal activity, although fungal sporocarp production declined significantly, and the relative abundance of two fungal taxa changed dramatically with N fertilization. Microbial biomass as measured by chloroform fumigation did not respond to fertilization, nor did the ratio of fungi : bacteria as measured by quantitative polymerase chain reaction. However, microbial biomass C : N ratios narrowed significantly from 16.0 ± 1.4 to 5.2 ± 0.3 with fertilization. N fertilization significantly increased the activity of a cellulose‐degrading enzyme and suppressed the activities of protein‐ and chitin‐degrading enzymes but had no effect on soil respiration rates or ¹⁴C signatures. These results indicate that N fertilization alters microbial community composition and allocation to extracellular enzyme production without affecting soil respiration. Thus, our results do not provide evidence for strong microbial feedbacks to the boreal C cycle under climate warming or N addition. However, organic N cycling may decline due to a reduction in the activity of enzymes that target nitrogenous compounds.     

Carbon fluxes to the soil in a mature temperate forest assessed by <Superscript>13</Superscript>C isotope tracing

Photosynthetic carbon uptake and respiratory C release from soil are major components of the global carbon balance. The use of ¹³C depleted CO₂ (¹³C = –30) in a free air CO₂ enrichment experiment in a mature deciduous forest permitted us to trace the carbon transfer from tree crowns to the rhizosphere of 100–120 years old trees. During the first season of CO₂ enrichment the CO₂ released from soil originated substantially from concurrent assimilation. The small contribution of recent carbon in fine roots suggests a much slower fine root turnover than is often assumed.¹³C abundance in soil air correlated best with temperature data taken from 4 to 10 days before air sampling time and is thus rapidly available for root and rhizosphere respiration. The spatial variability of ¹³C in soil air showed relationships to above ground tree types such as conifers versus broad-leaved trees. Considering the complexity and strong overlap of roots from different individuals in a forest, this finding opens an exciting new possibility of associating respiration with different species. What might be seen as signal noise does in fact contain valuable information on the spatial heterogeneity of tree-soil interaction.     

Nitrogen stable isotopic composition of leaves and soil: Tropical versus temperate forests

Several lines of evidence suggest that nitrogen in most tropical forests is relatively more available than N in most temperate forests, and even that it may function as an excess nutrient in many tropical forests. If this is correct, tropical forests should have more open N cycles than temperate forests, with both inputs and outputs of N large relative to N cycling within systems. Consequent differences in both the magnitude and the pathways of N loss imply that tropical forests should in general be more¹⁵N enriched than are most temperate forests. In order to test this hypothesis, we compared the nitrogen stable isotopic composition of tree leaves and soils from a variety of tropical and temperate forests. Foliar ¹⁵N values from tropical forests averaged 6.5 higher than from temperate forests. Within the tropics, ecosystems with relatively low N availability (montane forests, forests on sandy soils) were significantly more depleted in¹⁵N than other tropical forests. The average ¹⁵N values for tropical forest soils, either for surface or for depth samples, were almost 8 higher than temperate forest soils. These results provide another line of evidence that N is relatively abundant in many tropical forest ecosystems.