Showing papers in "Biogeochemistry in 2007"

C:N:P stoichiometry in soil: is there a “Redfield ratio” for the microbial biomass?

Abstract: Well-constrained carbon:nitrogen:phosphorus (C:N:P) ratios in planktonic biomass, and their importance in advancing our understanding of biological processes and nutrient cycling in marine ecosystems, has motivated ecologists to search for similar patterns in terrestrial ecosystems. Recent analyses indicate the existence of “Redfield-like” ratios in plants, and such data may provide insight into the nature of nutrient limitation in terrestrial ecosystems. We searched for analogous patterns in the soil and the soil microbial biomass by conducting a review of the literature. Although soil is characterized by high biological diversity, structural complexity and spatial heterogeneity, we found remarkably consistent C:N:P ratios in both total soil pools and the soil microbial biomass. Our analysis indicates that, similar to marine phytoplankton, element concentrations of individual phylogenetic groups within the soil microbial community may vary, but on average, atomic C:N:P ratios in both the soil (186:13:1) and the soil microbial biomass (60:7:1) are well-constrained at the global scale. We did see significant variation in soil and microbial element ratios between vegetation types (i.e., forest versus grassland), but in most cases, the similarities in soil and microbial element ratios among sites and across large scales were more apparent than the differences. Consistent microbial biomass element ratios, combined with data linking specific patterns of microbial element stoichiometry with direct evidence of microbial nutrient limitation, suggest that measuring the proportions of C, N and P in the microbial biomass may represent another useful tool for assessing nutrient limitation of ecosystem processes in terrestrial ecosystems.

A conceptual model of organo-mineral interactions in soils: self-assembly of organic molecular fragments into zonal structures on mineral surfaces

Abstract: In this paper, we propose a structure for organo-mineral associations in soils based on recent insights concerning the molecular structure of soil organic matter (SOM), and on extensive published evidence from empirical studies of organo-mineral interfaces. Our conceptual model assumes that SOM consists of a heterogeneous mixture of compounds that display a range of amphiphilic or surfactant-like properties, and are capable of self-organization in aqueous solution. An extension of this self-organizational behavior in solution, we suggest that SOM sorbs to mineral surfaces in a discrete zonal sequence. In the contact zone, the formation of particularly strong organo-mineral associations appears to be favored by situations where either (i) polar organic functional groups of amphiphiles interact via ligand exchange with singly coordinated mineral hydroxyls, forming stable inner-sphere complexes, or (ii) proteinaceous materials unfold upon adsorption, thus increasing adhesive strength by adding hydrophobic interactions to electrostatic binding. Entropic considerations dictate that exposed hydrophobic portions of amphiphilic molecules adsorbed directly to mineral surfaces be shielded from the polar aqueous phase through association with hydrophobic moieties of other amphiphilic molecules. This process can create a membrane-like bilayer containing a hydrophobic zone, whose components may exchange more easily with the surrounding soil solution than those in the contact zone, but which are still retained with considerable force. Sorbed to the hydrophilic exterior of hemimicellar coatings, or to adsorbed proteins, are organic molecules forming an outer region, or kinetic zone, that is loosely retained by cation bridging, hydrogen bonding, and other interactions. Organic material in the kinetic zone may experience high exchange rates with the surrounding soil solution, leading to short residence times for individual molecular fragments. The thickness of this outer region would depend more on input than on the availability of binding sites, and would largely be controlled by exchange kinetics. Movement of organics into and out of this outer region can thus be viewed as similar to a phase-partitioning process. The zonal concept of organo-mineral interactions presented here offers a new basis for understanding and predicting the retention of organic compounds, including contaminants, in soils and sediments.

How does fire affect the nature and stability of soil organic nitrogen and carbon? A review

Abstract: After vegetation fires considerable amounts of severely or partly charred necromass (referred to here as char) are incorporated into the soil, with long-term consequences for soil C and N dynamics and thus N availability for primary production and C and N transport within the soil column. Considering results reported in the pyrolysis literature in combination with those obtained from controlled charring of plant material and soil organic matter (SOM), it has become clear that common models claiming char as a graphite-like material composed mainly of highly condensed polyaromatic clusters may be oversimplified. Instead, I suggest a concept in which char is a heterogeneous mixture of heat-altered biopolymers with domains of relatively small polyaromatic clusters, but considerable substitution with N, O and S functional groups. Such a concept allows fast oxidation facilitating both microbial attack and dissolution. Although, char is commonly believed to degrade more slowly than litter, over the long term and under oxic conditions, char may degrade to an extent that it becomes indistinguishable from naturally formed SOM. Oxygen depletion or environments with low microbial activity may be necessary for char to survive without major chemical alteration and in considerable amounts for millennia or longer.

Soil carbon saturation: concept, evidence and evaluation

Abstract: Current estimates of soil C storage potential are based on models or factors that assume linearity between C input levels and C stocks at steady-state, implying that SOC stocks could increase without limit as C input levels increase. However, some soils show little or no increase in steady-state SOC stock with increasing C input levels suggesting that SOC can become saturated with respect to C input. We used long-term field experiment data to assess alternative hypotheses of soil carbon storage by three simple models: a linear model (no saturation), a one-pool whole-soil C saturation model, and a two-pool mixed model with C saturation of a single C pool, but not the whole soil. The one-pool C saturation model best fit the combined data from 14 sites, four individual sites were best-fit with the linear model, and no sites were best fit by the mixed model. These results indicate that existing agricultural field experiments generally have too small a range in C input levels to show saturation behavior, and verify the accepted linear relationship between soil C and C input used to model SOM dynamics. However, all sites combined and the site with the widest range in C input levels were best fit with the C-saturation model. Nevertheless, the same site produced distinct effective stabilization capacity curves rather than an absolute C saturation level. We conclude that the saturation of soil C does occur and therefore the greatest efficiency in soil C sequestration will be in soils further from C saturation.

Environmental constraints on the production and removal of the climatically active gas dimethylsulphide (DMS) and implications for ecosystem modelling

Abstract: Seawater concentrations of the climate-cooling, volatile sulphur compound dimethylsulphide (DMS) are the result of numerous production and consumption processes within the marine ecosystem. Due to this complex nature, it is difficult to predict temporal and geographical distribution patterns of DMS concentrations and the inclusion of DMS into global ocean climate models has only been attempted recently. Comparisons between individual model predictions, and ground-truthing exercises revealed that information on the functional relationships between physical and chemical ecosystem parameters, biological productivity and the production and consumption of DMS and its precursor dimethylsulphoniopropionate (DMSP) is necessary to further refine future climate models. In this review an attempt is made to quantify these functional relationships. The description of processes includes: (1) parameters controlling DMSP production such as species composition and abiotic factors; (2) the conversion of DMSP to DMS by algal and bacterial enzymes; (3) the fate of DMSP-sulphur due to, e.g., grazing, microbial consumption and sedimentation and (4) factors controlling DMS removal from the water column such as microbial consumption, photo-oxidation and emission to the atmosphere. We recommend the differentiation of six phytoplankton groups for inclusion in future models: eukaryotic and prokaryotic picoplankton, diatoms, dinoflagellates, and other phytoflagellates with and without DMSP-lyase activity. These functional groups are characterised by their cell size, DMSP content, DMSP-lyase activity and interactions with herbivorous grazers. In this review, emphasis is given to ecosystems dominated by the globally relevant haptophytes Emiliania huxleyi and Phaeocystis sp., which are important DMS and DMSP producers.

Increases in soil respiration following labile carbon additions linked to rapid shifts in soil microbial community composition

Abstract: Organic matter decomposition and soil CO2 efflux are both mediated by soil microorganisms, but the potential effects of temporal variations in microbial community composition are not considered in most analytical models of these two important processes. However, inconsistent relationships between rates of heterotrophic soil respiration and abiotic factors, including temperature and moisture, suggest that microbial community composition may be an important regulator of soil organic matter (SOM) decomposition and CO2 efflux. We performed a short-term (12-h) laboratory incubation experiment using tropical rain forest soil amended with either water (as a control) or dissolved organic matter (DOM) leached from native plant litter, and analyzed the effects of the treatments on soil respiration and microbial community composition. The latter was determined by constructing clone libraries of small-subunit ribosomal RNA genes (SSU rRNA) extracted from the soil at the end of the incubation experiment. In contrast to the subtle effects of adding water alone, additions of DOM caused a rapid and large increase in soil CO2 flux. DOM-stimulated CO2 fluxes also coincided with profound shifts in the abundance of certain members of the soil microbial community. Our results suggest that natural DOM inputs may drive high rates of soil respiration by stimulating an opportunistic subset of the soil bacterial community, particularly members of the Gammaproteobacteria and Firmicutes groups. Our experiment indicates that variations in microbial community composition may influence SOM decomposition and soil respiration rates, and emphasizes the need for in situ studies of how natural variations in microbial community composition regulate soil biogeochemical processes.

Organic matter stabilization in soil microaggregates: implications from spatial heterogeneity of organic carbon contents and carbon forms

Abstract: This study investigates the spatial distribution of organic carbon (C) in free stable microaggregates (20–250 μm; not encapsulated within macroaggregates) from one Inceptisol and two Oxisols in relation to current theories of the mechanisms of their formation. Two-dimensional micro- and nano-scale observations using synchrotron-based Fourier-transform infrared (FTIR) and near-edge X-ray absorption fine structure (NEXAFS) spectroscopy yielded maps of the distribution of C amounts and chemical forms. Carbon deposits were unevenly distributed within microaggregates and did not show any discernable gradients between interior and exterior of aggregates. Rather, C deposits appeared to be patchy within the microaggregates. In contrast to the random location of C, there were micron-scale patterns in the spatial distribution of aliphatic C–H (2922 cm−1), aromatic C=C and N–H (1589 cm−1) and polysaccharide C–O (1035 cm−1). Aliphatic C forms and the ratio of aliphatic C/aromatic C were positively correlated (r2 of 0.66–0.75 and 0.27–0.59, respectively) to the amount of O–H on kaolinite surfaces (3695 cm−1), pointing at a strong role for organo-mineral interactions in C stabilization within microaggregates and at a possible role for molecules containing aliphatic C-H groups in such interactions. This empirical relationship was supported by nanometer-scale observations using NEXAFS which showed that the organic matter in coatings on mineral surfaces had more aliphatic and carboxylic C with spectral characteristics resembling microbial metabolites than the organic matter of the entire microaggregate. Our observations thus support models of C stabilization in which the initially dominant process is adsorption of organics on mineral surfaces rather than occlusion of organic debris by adhering clay particles.

Quantitative effects of vegetation cover on wind erosion and soil nutrient loss in a desert grassland of southern New Mexico, USA

Abstract: Wind is a key abiotic factor that influences the dynamics of arid and semiarid systems. We investigated two basic relationships on vegetation manipulation (grass cover reduction) plots at the Jornada Experimental Range in southern New Mexico: (1) wind erosion rates (horizontal mass flux and dust emission) versus vegetative cover, and (2) nutrient loss versus vegetative cover. The results indicate that wind erosion rates and nutrient loss by dust emission are strongly affected by plant cover; however, the importance of shrubs and grasses in reducing dust flux may not be equal. The dramatic increase of wind erosion between 75% grass cover reduction and 100% grass cover reduction suggests that sparsely distributed mesquites are relatively ineffective at reducing wind erosion and nutrient loss compared to grasses. Comparisons of nutrients between surface soils and wind blown dust indicate that aeolian transport is a major cause for the loss of soil nutrients in susceptible environments. We found that increased aeolian flux over three windy seasons (March 2004–July 2006) removed up to 25% of total organic carbon (TOC) and total nitrogen (TN) from the top 5 cm of soil, and about 60% of TOC and TN loss occurred in the first windy season (March–July 2004). The balance between net loss of nutrients by aeolian processes and the addition of nutrients by biotic processes changed from negative (net loss) to positive (net accumulation) between 50% grass cover reduction and 25% grass cover reduction. The estimated lifetime of surface soil TOC and TN of about 10 years on the plot with 100% grass cover reduction indicates that impacts of wind erosion on soil resources can occur on very short timescales.

Storage, patterns and environmental controls of soil organic carbon in China

Abstract: Based on the data from China’s second national soil survey and field observations in northwest China, we estimated soil organic carbon (SOC) storage in China and investigated its spatial and vertical distribution. China’s SOC storage in a depth of 1 meter was estimated as 69.1 Pg (1015 g), with an average density of 7.8 kg m−2. About 48% of the storage was concentrated in the top 30 cm. The SOC density decreased from the southeast to the northwest, and increased from arid to semi-humid zone in northern China and from tropical to cold-temperate zone in the eastern part of the country. The vertical distribution of SOC differed in various climatic zones and biomes; SOC distributed deeper in arid climate and water-limited biomes than in humid climate. An analysis of general linear model suggested that climate, vegetation, and soil texture significantly influenced spatial pattern of SOC, explaining 78.2% of the total variance, and that climate and vegetation interpreted 78.9% of the total variance in the vertical SOC distribution.

Role of proteins in soil carbon and nitrogen storage: controls on persistence

Abstract: Mechanisms of soil organic carbon (C) and nitrogen (N) stabilization are of great interest, due to the potential for increased CO2 release from soil organic matter (SOM) to the atmosphere as a result of global warming, and because of the critical role of soil organic N in controlling plant productivity. Soil proteins are recognized increasingly as playing major roles in stabilization and destabilization of soil organic C and N. Two categories of proteins are proposed: detrital proteins that are released upon cell death and functional proteins that are actively released into the soil to fulfill specific functions. The latter include microbial surface-active proteins (e.g., hydrophobins, chaplins, SC15, glomalin), many of which have structures that promote their persistence in the soil, and extracellular enzymes, responsible for many decomposition and nutrient cycling transformations. Here we review information on the nature of soil proteins, particularly those of microbial origin, and on the factors that control protein persistence and turnover in the soil. We discuss first the intrinsic properties of the protein molecule that affect its stability, next possible extrinsic stabilizing influences that arise as the proteins interact with other soil constituents, and lastly controls on accessibility of proteins at coarser spatial scales involving microbial cells, clay particles, and soil aggregates. We conclude that research at the interface between soil science and microbial physiology will yield rapid advances in our understanding of soil proteins. We suggest as research priorities determining the relative abundance and turnover time (age) of microbial versus plant proteins and of functional microbial proteins, including surface-active compounds.

Density fractionation of forest soils: methodological questions and interpretation of incubation results and turnover time in an ecosystem context

Abstract: Soil organic matter (SOM) is often separated by physical means to simplify a com- plex matrix into discrete fractions. A frequent approach to isolating two or more fractions is based on differing particle densities and uses a high density liquid such as sodium polytungstate (SPT). Soil density fractions are often interpreted as organic matter pools with different carbon (C) turnover times, ranging from years to decades or centuries, and with different functional roles for C and nutrient dynamics. In this paper, we discuss the development and mechanistic basis of common density-based methods for dividing soil into distinct organic matter fractions. Further, we directly address the potential effects of dispersing soil in a high density salt solution on the recov- ered fractions and implications for data inter- pretation. Soil collected from forested sites at H. J. Andrews Experimental Forest, Oregon and Bousson Experimental Forest, Pennsylvania was separated into light and heavy fractions by floa- tation in a 1.6 g cm -3 solution of SPT. Mass balance calculations revealed that between 17% and 26% of the original bulk soil C and N content was mobilized and subsequently discarded during density fractionation for both soils. In some cases, the light isotope was preferentially mobilized during density fractionation. During a year-long incubation, mathematically recombined density fractions respired ~40% less than the bulk soil at both sites and light fraction (LF) did not always decompose more than the heavy fraction (HF). Residual amounts of tungsten (W) present even in well-rinsed fractions were enough to reduce microbial respiration by 27% compared to the control in a 90-day incubation of Oa material. However, residual W was nearly eliminated by repeated leaching over the year-long incubation, and is not likely the primary cause of the difference in respiration between summed frac- tions and bulk soil. Light fraction at Bousson, a deciduous site developed on Alfisols, had a radiocarbon-based mean residence time (MRT)

Degradation rates of organic phosphorus in lake sediment

Abstract: Phosphorus (P) binding groups were identified in phytoplankton, settling particles, and sediment profiles by 31P NMR spectroscopy from the Swedish mesotrophic Lake Erken. The 31P NMR analysis revealed that polyphosphates and pyrophosphates were abundant in the water column, but rapidly mineralized in the sediment. Orthophosphate monoesters and teichoic acids degraded more slowly than DNA-P, polyphosphates, and P lipids. Humic acids and organic acids from phytoplankton were precipitated from the NaOH extract by acidification and identified by 31P NMR spectroscopy. The precipitated P was significantly more recalcitrant than the P compound groups remaining in solution, but does not constitute a major sink of P as it did not reach a stable concentration with depth, which indicates that it may eventually be degraded. Since P also precipitated from phytoplankton, the origin of humic-P can not be related solely to allochthonous P.

The saturation of N cycling in Central Plains streams: 15N experiments across a broad gradient of nitrate concentrations

Abstract: We conducted 15NO 3 − stable isotope tracer releases in nine streams with varied intensities and types of human impacts in the upstream watershed to measure nitrate (NO 3 − ) cycling dynamics Mean ambient NO 3 − concentrations of the streams ranged from 09 to 21,000 μg l−1 NO 3 − –N Major N-transforming processes, including uptake, nitrification, and denitrification, all increased approximately two to three orders of magnitude along the same gradient Despite increases in transformation rates, the efficiency with which stream biota utilized available NO 3 − -decreased along the gradient of increasing NO 3 − Observed functional relationships of biological N transformations (uptake and nitrification) with NO 3 − concentration did not support a 1st order model and did not show signs of Michaelis–Menten type saturation The empirical relationship was best described by a Efficiency Loss model, in which log-transformed rates (uptake and nitrification) increase with log-transformed nitrate concentration with a slope less than one Denitrification increased linearly across the gradient of NO 3 − concentrations, but only accounted for ∼1% of total NO 3 − uptake On average, 20% of stream water NO 3 − was lost to denitrification per km, but the percentage removed in most streams was <5% km−1 Although the rate of cycling was greater in streams with larger NO 3 − concentrations, the relative proportion of NO 3 − retained per unit length of stream decreased as NO 3 − concentration increased Due to the rapid rate of NO 3 − turnover, these streams have a great potential for short-term retention of N from the landscape, but the ability to remove N through denitrification is highly variable

The mobilisation of phosphorus, organic carbon and ammonium in the initial stage of fen rewetting (a case study from NE Germany)

Abstract: Currently, more than 10,000 ha of fens have been rewetted to re-establish their function as nutrient sinks in NE Germany. However, field investigations reveal that porewater concentrations of P, dissolved organic carbon (DOC) and ammonium in rewetted fens are orders of magnitude larger than under pristine conditions. Hence, the objective of this study was to investigate the reasons behind enhanced P, organic carbon (OC) and ammonium mobilisation due to rewetting by means of a long-term incubation experiment. Highly, moderately and slightly decomposed peat of a drained fen (polder Zarnekow) was incubated under waterlogged conditions. A time course of concentrations of P, DOC, ammonium, sulphate and other dissolved substances was investigated by means of permanently installed dialysis samplers during 54 weeks of incubation. Simultaneously, the concentrations of these dissolved substances were investigated after rewetting of the field site. Before, and at the end of the incubation study, the amounts of bicarbonate–dithionite (BD) and NaOH soluble P and OC of incubated peat samples were determined by a sequential extraction procedure. The highest mobilisation of P, OC and ammonium occurred in the highly decomposed peat. Final concentrations of P, DOC and ammonium reached about 143 μM, 46 and 1.9 mM, respectively. The initial sulphate concentrations in the rewetting experiment, as well as in the field investigations, were extremely high and ranged between 3 and 13 mM; however, a complete consumption of sulphate was only observed in highly decomposed peat. In conclusion, the reasons for enhanced P, OC and ammonium mobilisation are increased amounts of redox sensitive substances and enhanced availability of decomposable organic matter in the upper highly decomposed peat horizon. These results should be considered in future rewetting management strategies.

Zooplankton grazing on Phaeocystis: a quantitative review and future challenges

Abstract: The worldwide colony-forming hapto- phyte phytoplankton Phaeocystis spp. are key organ- isms in trophic and biogeochemical processes in the ocean. Many organisms from protists to Wsh ingest cells and/or colonies of Phaeocystis. Reports on spe- ciWc mortality of Phaeocystis in natural plankton or mixed prey due to grazing by zooplankton, especially protozooplankton, are still limited. Reported feeding rates vary widely for both crustaceans and protists feeding on even the same Phaeocystis types and sizes. Quantitative analysis of available data showed that: (1) laboratory-derived crustacean grazing rates on monocultures of Phaeocystis may have been overesti- mated compared to feeding in natural plankton communities, and should be treated with caution; (2) formation of colonies by P. globosa appeared to reduce predation by small copepods (e.g., Acartia, Pseudocalanus, Temora and Centropages), whereas large copepods (e.g., Calanus spp.) were able to feed on colonies of Phaeocystis pouchetii; (3) physiologi- cal diVerences between diVerent growth states, spe- cies, strains, cell types, and laboratory culture versus natural assemblages may explain most of the varia- tions in reported feeding rates; (4) chemical signaling between predator and prey may be a major factor con- trolling grazing on Phaeocystis; (5) it is unclear to what extent diVerent zooplankton, especially proto- zooplankton, feed on the diVerent life forms of Phae- ocystis in situ. To better understand the mechanisms controlling zooplankton grazing in situ, future studies

Temperature controls a latitudinal gradient in the proportion of watershed nitrogen exported to coastal ecosystems

Abstract: Increased export of biologically available nitrogen (N) to the coastal zone is strongly linked to eutrophication, which is a major problem in coastal marine ecosystems (NRC (2000) Clean Coastal Waters: Understanding and Reducing the Effects of Nutrient Pollution. National Academy Press, Washington, DC; Bricker et al. (1999) National Estuarine Eutrophication Assessment. Effects of nutrient enrichment in the nation’s estuaries. NOAA-NOS Special Projects Office, Silver Spring, MD). However, not all of the nitrogen input to a watershed is exported to the coast (Howarth et al. (1996) Biogeochemistry 35:75–139; Jordan and Weller (1996) Bioscience 46:655–664). Global estimates of nitrogen export to coasts have been taken to be 25% of watershed input, based largely on northeastern U.S. observations (Galloway et al. (2004) Biogeochemistry 70:153–226; Boyer et al. (2006) Global Biogeochem Cycle 20:Art. No. GB1S91). We applied the N budgeting methodology developed for the International SCOPE Nitrogen project (Howarth et al. (1996) Biogeochemistry 35:75–139; Boyer et al. (2002) Biogeochemistry 57:137–169) to 12 watersheds in the southeastern U.S., and compared them with estimates of N export for 16 watersheds in the northeastern U.S. (Boyer et al. (2002) Biogeochemistry 57:137–169). In southeastern watersheds, average N export was only 9% of input, suggesting the need for downward revision of global estimates. The difference between northern and southern watersheds is not a function of the absolute value of N inputs, which spanned a comparable range and were positively related to export in both cases. Rather, the proportion of N exported was significantly related to average watershed temperature (% N export = 58.41 e−0.11 * temperature; R 2 = 0.76), with lower proportionate nitrogen export in warmer watersheds. In addition, we identified a threshold in proportionate N export at 38°N latitude that corresponds to a reported breakpoint in the rate of denitrification at 10–12°C. We hypothesize that temperature, by regulating denitrification, results in increased proportionate N export at higher latitudes. Regardless of the mechanism, these observations suggest that temperature increases associated with future climate change may well reduce the amount of nitrogen that reaches estuaries, which will have implications for coastal eutrophication.

Soil organic carbon storage in mountain grasslands of the Pyrenees: effects of climate and topography

Abstract: The prediction of soil C stocks across the landscape has been increasingly studied in many areas of the world. Soil organic C storage in mountain areas is highly heterogeneous, mainly as a result of local-scale variability in the soil environment (topography, stoniness, parent material) and microclimate. The aims of the present study are to estimate soil organic C stocks (SOCS) in mineral soils of high-altitude grasslands of the Pyrenees and determine whether climatic and topographic variables can be used as predictors of SOCS and organic C content in the surface soil horizons of these ecosystems. For that purpose we sampled 35 soil profiles in subalpine and alpine grasslands including a range of altitudes, slopes and aspects. We analysed the soils for stoniness, bulk density, total C, texture, and C-to-N ratio and determined topographical variables. We used georeferenced climatic information for climatic descriptions of the sites. SOCS were highly correlated with soil depth. However, we were not able to predict soil depth by using environmental and topographic variables. In spite of this fact, altitude and aspect explained 41.2% of the SOCS variability while summer temperature and precipitation combined with aspect explained 56.9% of the variability of the organic C content of the surface layer (OC). The SOCS were low at high altitudes, probably as a result of an overall temperature limitation of net primary productivity. Under these conditions, the effect of aspect was small. The highest SOCS occurred at the lowest altitudes for ENE or WNW aspects, showing sharper decreases towards the south than to the north. The harsh climatic conditions and low-plant productivity that occur at the northern slopes reduced SOCS at the highest altitudes. In contrast, southern aspects showed similar organic C content along the altitudinal gradient. The OC variability in the surface soils not explained by climatic or topographic variables was partially related to the characteristics of soil organic matter, which may depend on the plant communities.

Soil carbon and nitrogen stores and storage potential as affected by land-use in an agro-pastoral ecotone of northern China

Abstract: Equilibrium carbon stock is the result of a balance between inputs and outflows to the pool. Changes in land-use are likely to alter such balance, resulting in different carbon stores under different land-use types in addition to the impacts of global climate change. In an agro-pastoral ecotone of Inner Mongolia, northern China, we investigated productivity and belowground carbon and nitrogen stores under six different types of land-uses, namely free grazing (FG), grazing exclusion (GE), mowing (MW), corn plantation (CP), fallow (FL), and alfalfa pasture (AP), and their impacts on litter and fine roots in semiarid grassland ecosystems. We found that there were great variations in aboveground net primary production (ANPP) across the six land-use types, with CP having markedly high ANPP; the FG had significantly reduced soil organic carbon (SOC) and nitrogen stores (SON) to 100 cm depth compared with all other types of land uses, while very little litter accumulation was found on sites of the FG and CP. The top 20 cm of soils accounted for about 80% of the root carbon and nitrogen, with very little roots being found below 50 cm. About 60% of SOC and SON were stored in the top 30 cm layer. Land-use change altered the inputs of organic matters, thus affecting SOC and SON stores accordingly; the MW and GE sites had 59 and 56% more SOC and 61% more SON than the FG. Our estimation suggested that restoring severely degraded and overgrazed grasslands could potentially increase SOC and SON stores by more than 55%; conversion from the native grasses to alfalfa could potentially double the aboveground biomass production, and further increase SOC and SON stores by more than 20%. Our study demonstrated significant carbon and nitrogen storage potential of the agro-pastoral ecotone of northern China through land-use changes and improved management in the context of mitigating global climate change.

Fire impact on C and N losses and charcoal production in a scrub oak ecosystem

Abstract: Fire profoundly modifies the terrestrial C cycle of about 40% of the Earth’s land surface. The immediate effect of fire is that of a net loss of C as CO2 gas and soot particles to the atmosphere. Nevertheless, a proportion of the ecosystem biomass is converted into charcoal, which contains highly recalcitrant molecular structures that contribute to long-term C storage. The present study aimed to assess simultaneously losses to the atmosphere and charcoal production rates of C and N compounds as a result of prescription fire in a Florida scrub-oak ecosystem. Pre-fire and post-fire charred and unburned organic matter stocks were determined for vegetation leaves and stems, litter and soil in 20 sub-plots installed in a 30-ha area that was subjected to prescribed fire. Concentrations of C and N were determined, and fluxes among pools and to the atmosphere were derived from these measurements. Soil C and N stocks were unchanged by the fire. Post-fire standing dead biomass contained 30% and 12% of pre-fire vegetation C and N stocks, respectively. In litter, post-fire stocks contained 64% and 83% of pre-fire C and N stocks, respectively. Most of the difference in relative losses between vegetation and litter could be attributed to substantial litter fall of charred and unburned leaves during the fire event. Indeed, an estimated 21% of pre-fire vegetation leaf C was found in the post-fire litter, while the remaining 79% was lost to the atmosphere. About 3/4 of the fire-induced leaf litter fall was in the form of unburned tissue and the remainder was charcoal, which amounted to 5% of pre-fire leaf C stocks. Charcoal production ranged between 4% and 6% of the fire-affected biomass, i.e. the sum of charcoal production and atmospheric losses. This value is below the range of literature values for the transformation of plant tissue into stable soil organic matter through humification processes, which suggests that fire generates a smaller quantity of stable organic C than humification processes over decades and potentially centuries.

The life cycle of Phaeocystis: State of knowledge and presumptive role in ecology

Abstract: Despite numerous investigations, the number and role of morphotypes involved in the life cycle of Phaeocystis species remain under debate. This is partly due to the application of different methodologies such as light, transmission, scanning electron microscopy and flow cytometry on specific samples. This heterogeneity of approaches results in the incomplete morphometric description of the different cell types existing within one species according to relevant criteria and the indetermination of the ploidy level of each observed stage. We review here the different morphotypes observed within each of the six Phaeocystis species recognized up to now. Four different cell types have been observed. In common to all six species is the occurrence of a scaly flagellate producing star-forming filaments (all species except P. jahnii) or not (P. globosa and P. jahnii). In three colony-forming species, P. globosa, P. pouchetii and P. antarctica, three morphotypes are observed: a flagellate with scales and filaments, a colonial cell, and a flagellate devoid of scales and filaments. In the non-colony-forming species, P. scrobiculata and P. cordata, only flagellates with scales and filaments have been observed. While suspected in P. pouchetii and P. antarctica, a haploid–diploid life cycle has only been evidenced for P. globosa. The two main prominent features of this cycle are that sexuality is prevalent in colony bloom formation and termination and that two types of vegetative reproduction exist. The ecological relevance of alternating haploid and diploid stages is not clearly apparent on the basis of existing ecological studies.

Litter decomposition rate is dependent on litter Mn concentrations

Abstract: A statistically significant linear relationship was found between annual mass loss of foliar litter in the late stages of decomposition and Mn concentration in the litter. We used existing decomposition data on needle and leaf decomposition of Scots pine (Pinus sylvestris L.), lodgepole pine (Pinus contorta var. contorta), Norway spruce (Picea abies (L.) Karst.), silver birch (Betula pendula L.), and grey alder (Alnus incana L.) from Sweden and Aleppo pine (Pinus halepensis Mill.) from Libya, to represent boreal, temperate, and Mediterranean climates. The later the decomposition stage as indicated by higher sulfuric-acid lignin concentrations, the better were the linear relationships between litter mass loss and Mn concentrations. We conclude that Mn concentrations in litter have an influence on litter mass-loss rates in very late decomposition stages (up to 5 years), provided that the litter has high enough Mn concentration. The relationship may be dependent on species as the relationship is stronger with species that take up high enough amounts of Mn.

Stabilization and destabilization of soil organic matter—a new focus

Abstract: Interest in soil organic matter (SOM) is ramping up as concern mounts about steadily increasing levels of atmospheric CO2. There are two reasons for this. First, there is hope that improvements in crop, forest, and soil management may allow significant amounts of CO2 to be removed from the atmosphere and sequestered in soil. Second is the possibility that increased soil respiration rates, associated with climate change, will unleash a positive feedback in which temperatures rise even faster than now expected. Other reasons have long existed for understanding SOM dynamics, such as SOM as the source of most of the nonfertilizer N needed for plant growth, but the specter of run-away climate change seems to have now overtaken these other justifications.

Response of dissolved organic matter in the forest floor to long-term manipulation of litter and throughfall inputs

Abstract: Dissolved organic matter (DOM) contributes to organic carbon either stored in mineral soil horizons or exported to the hydrosphere However, the main controls of DOM dynamics are still under debate We studied fresh leaf litter and more decomposed organic material as the main sources of DOM exported from the forest floor of a mixed beech/oak forest in Germany In the field we doubled and excluded aboveground litter input and doubled the input of throughfall From 1999 to 2005 we measured concentrations and fluxes of dissolved organic C and N (DOC, DON) beneath the Oi and Oe/Oa horizon DOM composition was traced by UV and fluorescence spectroscopy In selected DOM samples we analyzed the concentrations of phenols, pentoses and hexoses, and lignin-derived phenols by CuO oxidation DOC and DON concentrations and fluxes almost doubled instantaneously in both horizons of the forest floor by doubling the litter input and DOC concentrations averaged 82 mg C l−1 in the Oe/Oa horizon Properties of DOM did not suggest a change of the main DOM source towards fresh litter In turn, increasing ratios of hexoses to pentoses and a larger content of lignin-derived phenols in the Oe/Oa horizon of the Double litter plots in comparison to the Control plots indicated a priming effect: Addition of fresh litter stimulated microbial activity resulting in increased microbial production of DOM from organic material already stored in Oe/Oa horizons Exclusion of litter input resulted in an immediate decrease in DOC concentrations and fluxes in the thin Oi horizon In the Oe/Oa horizon DOC concentrations started to decline in the third year and were significantly smaller than those in the Control after 5 years Properties of DOM indicated an increased proportion of microbially and throughfall derived compounds after exclusion of litter inputs Dissolved organic N did not decrease upon litter exclusion We assume a microbial transformation of mineral N from throughfall and N mineralization to DON Increased amounts of throughfall resulted in almost equivalently increased DOC fluxes in the Oe/Oa horizon However, long-term additional throughfall inputs resulted in significantly declining DOC concentrations over time We conclude that DOM leaving the forest floor derives mainly from decomposed organic material stored in Oe/Oa horizons Leaching of organic matter from fresh litter is of less importance Observed effects of litter manipulations strongly depend on time and the stocks of organic matter in forest floor horizons Long-term experiments are particularly necessary in soils/horizons with large stocks of organic matter and in studies focusing on effects of declined substrate availability The expected increased primary production upon climate change with subsequently enhanced litter input may result in an increased production of DOM from organic soil horizons

Relations of mineral-soil C and N to climate and texture: regional differences within the conterminous USA

Abstract: Soil is a prominent component of terrestrial C and N budgets. Soil C and N pools are influenced by, and may reciprocally influence, many environmental factors. Our objective was to determine the quantitative relations of surface mineral-soil organic C, N, and C/N ratios to climate and soil texture across seven ecological regions that make up the conterminous USA. Up to 608 soil profiles per region and their corresponding climates were evaluated with regression analysis. The organic C pool (kg C m−2) in the upper 20 cm of mineral soil was positively related to mean annual precipitation, evapotranspiration and clay content in all regions. It was negatively related to a temperature/precipitation index in all regions and negatively related to mean annual temperature, except in the northwest temperate forest region. Soil C/N ratios were negatively related to clay or silt content in all regions. These relations are consistent with concepts of moisture and temperature controls on detrital production, differential effects of temperature on detrital production and decomposition, and stabilization of organic matter by clay and silt. Differences in quantitative relations among regions may be related to vegetation-composition effects on soil organic matter processes, clay mineralogy, and faunal mixing of surface organic horizons with mineral soil. Regional differences also occurred in the importance of climate vs. soil texture in explaining the variability in soil C. The regional differences indicate the importance of using region-specific, rather than generalized, equations for projecting long-term soil responses to climate change and for conducting ecosystem-model calibration or validation.

Are Swedish forest soils sinks or sources for CO2-model analyses based on forest inventory data

Abstract: Forests soils should be neither sinks nor sources of carbon in a long-term perspective. From a Swedish perspective the time since the last glaciation has probably not been long enough to reach a steady state, although changes are currently very slow. In a shorter perspective, climatic and management changes over the past 100 years have probably created imbalances between litter input to soils and organic carbon mineralisation. Using extant data on forest inventories, we applied models to analyse possible changes in the carbon stocks of Swedish forest soils. The models use tree stocks to provide estimates of tree litter production, which are fed to models of litter decomposition and from which carbon stocks are calculated. National soil carbon stocks were estimated to have increased by 3 Tg yr−1 or 12–13 g m−2 yr−1 in the period 1926–2000 and this increase will continue because soil stocks are far from equilibrium with current litter inputs. The figure obtained is likely to be an underestimation because wet sites store more carbon than predicted here and the inhibitory effect of nitrogen deposition on soil carbon mineralisation was neglected. Knowledge about site history prior to the calculation period determines the accuracy of current soil carbon stocks estimates, although changes can be more accurately estimated.

Current understanding of Phaeocystis ecology and biogeochemistry, and perspectives for future research

Abstract: The phytoplankton genus Phaeocystis has well-documented, spatially and temporally extensive blooms of gelatinous colonies; these are associated with release of copious amounts of dimethyl sulphide (an important climate-cooling aerosol) and alterations of material flows among trophic levels and export from the upper ocean. A potentially salient property of the importance of Phaeocystis in the marine ecosystem is its physiological capability to transform between solitary cell and gelatinous colonial life cycle stages, a process that changes organism biovolume by 6–9 orders of magnitude, and which appears to be activated or stimulated under certain circumstances by chemical communication. Both life-cycle stages can exhibit rapid, phased ultradian growth. The colony skin apparently confers protection against, or at least reduces losses to, smaller zooplankton grazers and perhaps viruses. There are indications that Phaeocystis utilizes chemistry and/or changes in size as defenses against predation, and its ability to create refuges from biological attack is known to stabilize predator–prey dynamics in model systems. Thus the life cycle form in which it occurs, and particularly associated interactions with viruses, determines whether Phaeocystis production flows through the traditional “great fisheries” food chain, the more regenerative microbial food web, or is exported from the mixed layer of the ocean.

Atmospheric mercury pollution due to losses of terrestrial carbon pools

Abstract: Plants accumulate significant amounts of atmospheric mercury (Hg) in aboveground biomass, likely sequestering over 1,000 Mg of atmospheric Hg every year This large mercury uptake could be strong enough to affect tropospheric Hg levels and might be partially responsible for seasonal variations in atmospheric Hg observed at Mace Head, Ireland The fluctuations of Hg concentrations coincide temporally with the annual oscillation of carbon dioxide (CO2) in the Northern Hemisphere, which is a result of seasonal growth of vegetation Therefore, declining Hg concentrations in spring and summer may be attributed in part to plant uptake of atmospheric Hg Further, the increase of Hg concentrations during non-active vegetation periods might partially be due to plant-derived Hg emitting back to the atmosphere during carbon mineralization The implications of these propositions are that past and future changes in biomass productivity and organic carbon pools may have had—and may continue to have—significant effects on atmospheric Hg levels Specifically, large losses in soil and biomass carbon pools in the last 150 years could have contributed significantly to observed increases in atmospheric Hg pollution The roles of vegetation and terrestrial carbon pools should receive detailed consideration on how they might attenuate or exacerbate atmospheric Hg pollution

The carbohydrates of Phaeocystis and their degradation in the microbial food web

Abstract: The ubiquity and high productivity associated with blooms of colonial Phaeocystis makes it an important contributor to the global carbon cycle. During blooms organic matter that is rich in carbohydrates is produced. We distinguish five different pools of carbohydrates produced by Phaeocystis. Like all plants and algal cells, both solitary and colonial cells produce (1) structural carbohydrates, (hetero) polysaccharides that are mainly part of the cell wall, (2) mono- and oligosaccharides, which are present as intermediates in the synthesis and catabolism of cell components, and (3) intracellular storage glucan. Colonial cells of Phaeocystis excrete (4) mucopolysaccharides, heteropolysaccharides that are the main constituent of the mucous colony matrix and (5) dissolved organic matter (DOM) rich in carbohydrates, which is mainly excreted by colonial cells. In this review the characteristics of these pools are discussed and quantitative data are summarized. During the exponential growth phase, the ratio of carbohydrate-carbon (C) to particulate organic carbon (POC) is approximately 0.1. When nutrients are limited, Phaeocystis blooms reach a stationary growth phase, during which excess energy is stored as carbohydrates. This so-called overflow metabolism increases the ratio of carbohydrate-C to POC to 0.4–0.6 during the stationary phase, leading to an increase in the C/N and C/P ratios of Phaeocystis organic matter. Overflow metabolism can be channeled towards both glucan and mucopolysaccharides. Summarizing the available data reveals that during the stationary phase of a bloom glucan contributes 0–51% to POC, whereas mucopolysaccharides contribute 5–60%. At the end of a bloom, lysis of Phaeocystis cells and deterioration of colonies leads to a massive release of DOM rich in glucan and mucopolysaccharides. Laboratory studies have revealed that this organic matter is potentially readily degradable by heterotrophic bacteria. However, observations in the field of accumulation of DOM and foam indicate that microbial degradation is hampered. The high C/N and C/P ratios of Phaeocystis organic matter may lead to nutrient limitation of microbial degradation, thereby prolonging degradation times. Over time polysaccharides tend to self-assemble into hydrogels. This may have a profound effect on carbon cycling, since hydrogels provide a vehicle to move DOM up the size spectrum to sizes subject to sedimentation. In addition, it changes the physical nature and microscale structure of the organic matter encountered by bacteria which may affect the degradation potential of the Phaeocystis organic matter.

Sources and fates of dissolved organic carbon in lakes as determined by whole-lake carbon isotope additions

Abstract: Four whole-lake inorganic 13C addition experiments were conducted in lakes of differing trophic status. Inorganic 13C addition enriched algal carbon in 13C and changed the $$\delta^{13}$$ C-DOC by +1.5‰ to +9.5‰, depending on the specific lake. This change in $$\delta^{13}$$ C-DOC represented a significant input of algal DOC that was not completely consumed by bacteria. We modeled the dynamics in $$\delta^{13}$$ C-DOC to estimate the fluxes of algal and terrestrial carbon to and from the DOC pool, and determine the composition of the standing stock. Two experiments in lightly stained, oligotrophic lakes indicated that algal production was the source of about 20% of the DOC pool. In the following year, the experiment was repeated in one of these lakes under conditions of nutrient enrichment, and in a third, more humic lake. Algal contributions to the DOC pool were 40% in the nutrient enriched lake and 5% in the more humic lake. Spectroscopic and elemental analyses corroborated the presence of increased algal DOC in the nutrient enriched lake. Natural abundance measurements of the $$\delta^{13}$$ C of DOC in 32 lakes also revealed the dual contributions of both terrestrial and algal carbon to DOC. From these results, we suggest an approach for inferring the contribution of algal and terrestrial DOC using easily measurable parameters.

Bomb 14C enrichment indicates decadal C pool in deep soil

Abstract: Studies of changes in soil organic carbon (SOC) stocks normally limit their focus to the upper 20–30 cm of soil, yet 0–20 cm SOC stocks are only ∼40% of 0–1 m SOC. Accounting for only the upper 20–30 cm of SOC has been justifiable assuming that deeper SOC is unreactive since it displays 14C-derived mean residence times of hundreds or thousands of years. The dramatic increase in the 14C content of the atmosphere resulting from thermonuclear testing circa 1963 allows the unreactivity of deep SOC to be tested by examining whether deep soils show evidence of ‘bomb-14C’ incorporation. At depths of 40–100 cm, a well-studied New Zealand soil under stable pastoral management displays progressive enrichment of over 200‰ across samplings in 1959, 1974 and 2002, indicating substantial incorporation of bomb 14C. This pattern of deep 14C enrichment—previously observed in 2 well-drained California grassland soils—leads to the hypothesis that roots and/or dissolved organic C transport contribute to a decadally-reactive SOC pool comprising ∼10–40% of SOC below 50 cm. Deep reactive SOC may be important in the global C cycle because it can react to land-use or vegetation change and may respond to different processes than the reactive SOC in the upper 20–30 cm of soil.