Carbon flow in an upland grassland: effect of liming on the flux of recently photosynthesized carbon to rhizosphere soil

The effect of liming on the flow of recently photosynthesized carbon to rhizosphere soil was studied using (CO2)-C-13 pulse labelling, in an upland grassland ecosystem in Scotland. The use of C-13 enabled detection, in the field, of the effect of a 4-year liming period of selected soil plots on C allocation from plant biomass to soil, in comparison with unlimed plots. Photosynthetic rates and carbon turnover were higher in plants grown in limed soils than in those from unlimed plots. Higher delta(13)C% values were detected in shoots from limed plants than in those from unlimed plants in samples clipped within 15 days of the end of pulse labelling. Analysis of the aboveground plant production corresponding to the 4-year period of liming indicated that the standing biomass was higher in plots that received lime. Lower delta(13)C% values in limed roots compared with unlimed roots were found, whereas no significant difference was detected between soil samples. Extrapolation of our results indicated that more C has been lost through the soil than has been gained via photosynthetic assimilation because of pasture liming in Scotland during the period 1990-1998. However, the uncertainty associated with such extrapolation based on this single study is high and these estimates are provided only to set our findings in the broader context of national soil carbon emissions.

A review of rhizosphere carbon flow modelling

Rhizosphere processes play a key role in nutrient cycling in terrestrial ecosystems. Plant rhizodeposits supply low-molecular weight carbon substrates to the soil microbial community, resulting in elevated levels of activity surrounding the root. Mechanistic compartmental models that aim to model carbon flux through the rhizosphere have been reviewed and areas of future research necessary to better calibrate model parameters have been identified. Incorporating the effect of variation in bacterial biomass physiology on carbon flux presents a considerable challenge to experimentalists and modellers alike due to the difficulties associated with differentiating dead from dormant cells. A number of molecular techniques that may help to distinguish between metabolic states of bacterial cells are presented. The calibration of growth, death and maintenance parameters in rhizosphere models is also discussed. A simple model of rhizosphere carbon flow has been constructed and a sensitivity analysis was carried out on the model to highlight which parameters were most influential when simulating carbon flux. It was observed that the parameters that most heavily influenced long-term carbon compartmentalisation in the rhizosphere were exudation rate and biomass yield. It was concluded that future efforts to simulate carbon flow in the rhizosphere should aim to increase ecological realism in model structure.

A critical review of labelling techniques used to quantify rhizosphere carbon-flow

The rhizosphere is a major sink for photo-assimilated carbon and quantifying inputs into this sink is one of the main goals of rhizosphere biology as organic carbon lost from plant roots supports a higher microbial population in the rhizosphere compared to bulk soil. Two fundamentally different¹⁴CO₂ labelling strategies have been developed to estimate carbon fluxes through the rhizosphere - continuous feeding of shoots with labelled carbon dioxide and pulse-chase experiments. The biological interpretation that can be placed on the results of labelling experiments is greatly biased by the technique used. It is the purpose of this paper to assess the advantages, disadvantages and the biological interpretation of both continuous and pulse labelling and to consider how to partition carbon fluxes within the rhizosphere. © 1994 Kluwer Academic Publishers.

The effect of soil pH on rhizosphere carbon flow of Lolium perenne

Perennial rye-grass plants were grown at 15°C in microcosms containing soil sampled from field plots that had been maintained at constant pH for the last 30 years. Six soil pH values were tested in the experiment, with pH ranging from 4.3-6.5. After 3 weeks growth in the microcosms, plant shoots were exposed to a pulse of ¹⁴C-CO₂. The fate of this label was determined by monitoring ¹⁴C-CO₂ respired by the plant roots/soil and by the shoots. The ¹⁴C remaining in plant roots and shoots was determined when the plants were harvested 7 days after receiving the pulse label. The amount of ¹⁴C (expressed as a percentage of the total ¹⁴C fixed by the plant) lost from the plant roots increased from 12.3 to 30.6% with increasing soil pH from 4.3 to 6. Although a greater percentage of the fixed ¹⁴C was respired by the root/soil as soil pH increased, plant biomass was greater with increasing soil pH. Possible reasons for observed changes in the pattern of ¹⁴C distribution are discussed and, it is suggested that changes in the soil microbial biomass and in plant nitrogen nutrition may, in particular be key factors which led to increased loss of carbon from plant roots with increasing soil pH. © 1990 Kluwer Academic Publishers.

A comparison of carbon flow from pre-labelled and pulse-labelled plants

Perennial rye-grass was subjected to two different¹⁴C labelling regimes to enable a partitioning of the carbon sources contributing to rhizosphere carbon-flow. Plant/soil microcosms were designed which enabled rye-grass plants to either receive a single pulse of¹⁴C-CO₂ or to be pre-labelled using a series of¹⁴C-CO₂ pulses, allowing the fate of newly photoassimilated carbon and carbon lost by root decomposition to be followed into the soil. For young rye-grass plants grown over a short period, rhizosphere carbon flow was found to be dominated by newly photoassimilated carbon. Evidence for this came from the observed percentage of the total¹⁴C budget (i.e. total¹⁴C-CO₂ fixed by the plants) lost from the root/soil system, which was 30 times greater for the pulse labelled compared to pre-labelled plants. Root decomposition was found to be less at 10°C compared to 20-25°C, though input of¹⁴C into the soil was the same at both temperatures. © 1988 Kluwer Academic Publishers.

Carbon fluxes and pelagic ecosystem dynamics near two western Antarctic Peninsula Adélie penguin colonies: an inverse model approach

An inverse food-web model for the western Antarctic Peninsula (WAP) pelagic food web was constrained with data from Palmer Long Term Ecological Research (PAL-LTER) project annual austral summer sampling cruises. Model solutions were generated for 2 regions with Adelie penguin Pygoscelis adeliae colonies presenting different population trends (a northern and a southern colony) for a 12 yr period (1995-2006). Counter to the standard paradigm, comparisons of carbon flow through bacteria, microzooplankton, and krill showed that the diatom-krill-top predator food chain is not the dominant pathway for organic carbon exchanges. The food web is more complex, including significant contributions by microzooplankton and the microbial loop. Using both inverse model results and network indices, it appears that in the northern WAP the food web is dominated by the microbial food web, with a temporal trend toward its increasing importance. The dominant pathway for the southern WAP food web varies from year to year, with no detectable temporal trend toward dominance of microzooplankton versus krill. In addition, sensitivity analyses indicated that the northern colony of Adelie penguins, whose population size has been declining over the past 35 yr, appears to have sufficient krill during summer to sustain its basic metabolic needs and rear chicks, suggesting the importance of other processes in regulating the Adelie population decline.

Assessment of lux-marked Pseudomonas fluorescens for reporting on organic carbon compounds

Carbon-flow from plant roots to the rhizosphere provides a major source of nutrients for the soil microbial population. However, quantification of carbon-flow is problematic due to its complex composition. This study investigated the potential of lux-marked Pseudomonas fluorescens to discriminate between forms of carbon present in the rhizosphere by measuring the light response to a range of carbon compounds. Results indicate that bioluminescence of short-term carbon-starved P. fluorescens is dependent upon the source and concentration of carbon. This system, therefore, has the potential to both quantify and qualify organic acids present in rhizodeposits.

Balancing the (carbon) budget: Using linear inverse models to estimate carbon flows and mass-balance 13C:15N labelling experiments in low oxygen sediments.

Over 1 million km² of seafloor experience permanent low-oxygen conditions within oxygen minimum zones (OMZs). OMZs are predicted to grow as a consequence of climate change, potentially affecting oceanic biogeochemical cycles. The Arabian Sea OMZ impinges upon the western Indian continental margin at bathyal depths (150 - 1500 m) producing a strong depth dependent oxygen gradient at the sea floor. The influence of the OMZ upon the short term processing of organic matter by sediment ecosystems was investigated using in situ stable isotope pulse chase experiments. These deployed doses of ¹³C:¹⁵N labeled organic matter onto the sediment surface at four stations from across the OMZ (water depth 540 - 1100 m; [O₂] = 0.35 - 15 μM). In order to prevent experimentally anoxia, the mesocosms were not sealed. ¹³C and ¹⁵N labels were traced into sediment, bacteria, fauna and ¹³C into sediment porewater DIC and DOC. However, the DIC and DOC flux to the water column could not be measured, limiting our capacity to obtain mass-balance for C in each experimental mesocosm. Linear Inverse Modeling (LIM) provides a method to obtain a mass-balanced model of carbon flow that integrates stable-isotope tracer data with community biomass and biogeochemical flux data from a range of sources. Here we present an adaptation of the LIM methodology used to investigate how ecosystem structure influenced carbon flow across the Indian margin OMZ. We demonstrate how oxygen conditions affect food-web complexity, affecting the linkages between the bacteria, foraminifera and metazoan fauna, and their contributions to benthic respiration. The food-web models demonstrate how changes in ecosystem complexity are associated with oxygen availability across the OMZ and allow us to obtain a complete carbon budget for the stationa where stable-isotope labelling experiments were conducted.

Concentrations and carbon isotopic compositions of organic and inorganic compounds in sediment porewaters at IODP Site U1329

Ocean drilling has revealed the existence of vast microbial populations in the deep subseafloor, but to date little is known about their metabolic activities. To better understand the biogeochemical processes in the deep biosphere, we investigate the stable carbon isotope chemistry of acetate and other carbon-bearing metabolites in sediment pore-waters. Acetate is a key metabolite in the cycling of carbon in anoxic sediments. Its stable carbon isotopic composition provides information on the metabolic processes dominating acetate turnover in situ. This study reports our findings for a methane-rich site at the northern Cascadia Margin (NE Pacific) where Expedition 311 of the Integrated Ocean Drilling Program (IODP) sampled the upper 190 m of sediment. At Site U1329, d13C values of acetate span a wide range from -46.0 per mill to -11.0 per mill vs. VPDB and change systematically with sediment depth. In contrast, d13C values of both the bulk dissolved organic carbon (DOC) (-21.6 ± 1.3 per mill vs. VPDB) and the low-molecular-weight compound lactate (-20.9 ± 1.8 per mill vs. VPDB) show little variability. These species are interpreted to represent the carbon isotopic composition of fermentation products. Relative to DOC, acetate is up to 23.1 per mill depleted and up to 9.1 per mill enriched in 13C. Broadly, 13C-depletions of acetate relative to DOC indicate flux of carbon from acetogenesis into the acetate pool while 13C-enrichments of pore-water acetate relative to DOC suggest consumption of acetate by acetoclastic methanogenesis. Isotopic relationships between acetate and lactate or DOC provide new information on the carbon flow and the presence and activity of specific functional microbial communities in distinct biogeochemical horizons of the sediment. In particular, they suggest that acetogenic CO2-reduction can coexist with methanogenic CO2-reduction, a notion contrary to the hypothesis that hydrogen levels are controlled by the thermodynamically most favorable electron-accepting process. Further, the isotopic relationship suggests a relative increase in acetate flow to acetoclastic methanogenesis with depth although its contribution to total methanogenesis is probably small. Our study demonstrates how the stable carbon isotope biogeochemistry of acetate can be used to identify pathways of microbial carbon turnover in subsurface environments. Our observations also raise new questions regarding the factors controlling acetate turnover in marine sediments.

Carbon cycling in the northern Humboldt Current off Chile

The structure of the zooplankton foodweb and their dominant carbon fluxes were studied in the upwelling system off northern Chile (Mejillones Bay; 23°S) between October 2000 and December 2002. High primary production (PP) rates (18 gC/m**2 d) were mostly due to the net-phytoplankton size fraction (>23 µm). High PP has been traditionally associated with the wind-driven upwelling fertilizing effect of equatorial subsurface waters, which favour development of a short food chain dominated by a few small clupeiform fish species. The objective of the present work was to study the trophic carbon flow through the first step of this 'classical chain' (from phytoplankton to primary consumers such as copepods and euphausiids) and the carbon flow towards the gelatinous web composed of both filter-feeding and carnivorous zooplankton. To accomplish this objective, feeding experiments with copepods, appendicularians, ctenophores, and chaetognaths were conducted using naturally occurring plankton prey assemblages. Throughout the study, the total carbon ingestion rates showed that the dominant appendicularian species and small copepods consumed an average of 7 and 5 µgC/ind d, respectively. In addition, copepods ingested particles mainly in the size range of nano- and microplankton, whereas appendicularians ingested in the range of pico- and nanoplankton. Small copepods and appendicularians removed a small fraction of total daily PP (range 6-11%). However, when the pico- + nanoplankton fractions were the major contributors to total PP (oligotrophic conditions), grazing by small copepods increased markedly to 86% of total PP. Under these more oligotrophic conditions, the euphausiids grazing increased as well, but only reached values lower than 5% of total PP. During this study, chaetognaths and ctenophores ingested an average of 1 and 14 copepods/ind d, respectively. In terms of biomass consumed, the potential impact of carnivorous gelatinous zooplankton on the small-size copepod community (preferred prey) was important (2-12% of biomass removed daily). However, their impact produced more significant results on copepod abundance (up to 33%), which suggests that carnivorous gelatinous zooplankton may even modulate (control) the abundance of some species as well as the size structure of the copepod community.

黄土区人工与天然植被对凋落物量和土壤有机碳变化的影响

在黄土丘陵区选择从耕地、草地、灌木林到乔木林样地,不同样地内设立1m×1m(乔木10m×10m)的样方,分析样方内凋落物积累量、碳氮含量、土壤有机碳(SOC)和可溶性碳(DOC)含量变化。结果表明:天然草地、灌木林、乔木林凋落物积累量依次为5.3,12.1和32.4t.hm-2;但人工灌木林和乔木林的凋落物积累量分别为6.7和11.4t.hm-2,分别是天然灌木林和乔木林的1/2和1/3。随着植被的恢复,天然植被凋落物的C/N高于人工植被(刺槐林除外)。与耕地SOC(4.67g·kg-1)相比,天然灌木林地SOC提高5.9倍,人工灌木林地提高1.8倍;天然乔木林地提高8.0倍,而人工乔木林地仅提高4.0倍。凋落物积累量与0～20cm土层土壤有机碳存在显著线性相关关系(R2>0.83),但20cm以下线性相关关系不显著。凋落物积累量与0～10cm土壤可溶性碳含量存在显著线性相关关系(R2>0.893),与10～60cm土层线性相关关系不显著,与80～100cm土壤可溶性碳存在显著线性负相关关系。

Characterization of stable isotope fractionation during methane production in the sediment of a eutrophic lake,lake dagow,germany

We measured delta C-13 of CO2, CH4, and acetate-methyl in profundal sediment of eutrophic Lake Dagow by incubation experiments in the presence and absence of methanogenic inhibitors chloroform, bromoethane sulfonate (BES), and methyl fluoride, which have different specificities. Methyl fluoride predominantly inhibits acetoclastic methanogenesis and affects hydrogenotrophic methanogenesis relatively little. Optimization of methyl fluoride concentrations resulted in complete inhibition of acetoclastic methanogenesis. Methane was then exclusively produced by hydrogenotrophic methanogenesis and thus allowed determination of the fractionation factors specific for this methanogenic pathway. Acetate, which was then no longer consumed, accumulated and allowed determination of the isotopic signatures of the fermentatively produced acetate. BES and chloroform also inhibited CH4 production and resulted in accumulation of acetate. The fractionation factor for hydrogenotrophic methanogenesis exhibited variability, e. g., it changed with sediment depth. The delta C-13 of the methyl group of the accumulated acetate was similar to the delta C-13 of sedimentary organic carbon, while that of the carboxyl group was by about 12 parts per thousand higher. However, the delta C-13 of the acetate was by about 5 parts per thousand lower in samples with uninhibited compared with inhibited acetoclastic methanogenesis, indicating unusual isotopic fractionation. The isotope data were used for calculation of the relative contribution of hydrogenotrophic vs. acetoclastic methanogenesis to total CH4 production. Contribution of hydrogenotrophic methanogenesis increased with sediment depth from about 35% to 60%, indicating that organic matter was only partially oxidized in deeper sediment layers.

Intra-habitat heterogeneity of microbial food web structure under the regime of eutrophication and sediment resuspension in the large subtropical shallow Lake Taihu, China

Planktonic microbial community structure and classical food web were investigated in the large shallow eutrophic Lake Taihu (2338 km(2), mean depth 1.9 m) located in subtropical Southeast China. The water column of the lake was sampled biweekly at two sites located 22 km apart over a period of twelve month. Site 1 is under the regime of heavy eutrophication while Site 2 is governed by wind-driven sediment resuspension. Within-lake comparison indicates that phosphorus enrichment resulted in increased abundance of microbial components. However, the coupling between total phosphorus and abundance of microbial components was different between the two sites. Much stronger coupling was observed at Site 1 than at Site 2. The weak coupling at Site 2 was mainly caused by strong sediment resuspension, which limited growth of phytoplankton and, consequently, growth of bacterioplankton and other microbial components. High percentages of attached bacteria, which were strongly correlated with the biomass of phytoplankton, especially Microcystis spp., were found at Site 1 during summer and early autumn, but no such correlation was observed at Site 2. This potentially leads to differences in carbon flow through microbial food web at different locations. Overall, significant heterogeneity of microbial food web structure between the two sites was observed. Site-specific differences in nutrient enrichment (i.e. nitrogen and phosphorus) and sediment resuspension were identified as driving forces of the observed intra-habitat differences in food web structure.

Spatial Variability of Durum Wheat Yield as Related to Soil Parameters in an Organic Field

The yield in organic farming is generally much lower than its potential, which is due to its specificity. The objective of the present study was to quantify the yield spatial variation of wheat and relate it to soil parameters in an organic farm located in the north of the Negev Desert. Soil samples were gathered in a triangular grid at three time intervals. Yields were measured at 73 georeferenced points before the actual harvest. Several thematic maps of soil and yield parameters were produced using geographic information system and geostatistical methods. The strongest spatial correlation was found in the weight of 1000 grains and the weakest was in carbon flow. Temporal relationships were found between soil nitrate concentration, soil water content, and leaf area index. Wheat yield varied from 1.11 to 2.84 Mg ha(-1) and this remarkable variation indicates that the spatial analysis of soil and yield parameters is significant in organic agriculture.