Carbon input by plants into the soil. Review

The methods used for estimating below‐ground carbon (C) translocation by plants, and the results obtained for different plant species are reviewed. Three tracer techniques using C isotopes to quantify root‐derived C are discussed: pulse labeling, continuous labeling, and a method based on the difference in ¹³C natural abundance in C3 and C4 plants. It is shown, that only the tracer methods provided adequate results for the whole below‐ground C translocation. This included roots, exudates and other organic substances, quickly decomposable by soil microorganisms, and CO₂ produced by root respiration. Advantages due to coupling of two different tracer techniques are shown. The differences in the below‐ground C translocation pattern between plant species (cereals, grasses, and trees) are discussed. Cereals (wheat and barley) transfer 20%—30% of total assimilated C into the soil. Half of this amount is subsequently found in the roots and about one‐third in CO₂ evolved from the soil by root respiration and microbial utilization of rootborne organic substances. The remaining part of below‐ground translocated C is incorporated into the soil microorganisms and soil organic matter. The portion of assimilated C allocated below the ground by cereals decreases during growth and by increasing N fertilization. Pasture plants translocated about 30%—50% of assimilates below‐ground, and their translocation patterns were similar to those of crop plants. On average, the total C amounts translocated into the soil by cereals and pasture plants are approximately the same (1500 kg C ha^—1), when the same growth period is considered. However, during one vegetation period the cereals and grasses allocated beneath the ground about 1500 and 2200 kg C ha^—1, respectively. Finally, a simple approach is suggested for a rough calculation of C input into the soil and for root‐derived CO₂ efflux from the soil.     

Dissolved and colloidal phosphorus fluxes in forest ecosystems—an almost blind spot in ecosystem research

Understanding and quantification of phosphorus (P) fluxes are key requirements for predictions of future forest ecosystems changes as well as for transferring lessons learned from natural ecosystems to croplands and plantations. This review summarizes and evaluates the recent knowledge on mechanisms, magnitude, and relevance by which dissolved and colloidal inorganic and organic P forms can be translocated within or exported from forest ecosystems. Attention is paid to hydrological pathways of P losses at the soil profile and landscape scales, and the subsequent influence of P on aquatic ecosystems. New (unpublished) data from the German Priority Program 1685 “Ecosystem Nutrition: Forest Strategies for limited Phosphorus Resources” were added to provide up‐to‐date flux‐based information. Nitrogen (N) additions increase the release of water‐transportable P forms. Most P found in percolates and pore waters belongs to the so‐called dissolved organic P (DOP) fractions, rich in orthophosphate‐monoesters and also containing some orthophosphate‐diesters. Total solution P concentrations range from ca. 1 to 400 µg P L^?1, with large variations among forest stands. Recent sophisticated analyses revealed that large portions of the DOP in forest stream water can comprise natural nanoparticles and fine colloids which under extreme conditions may account for 40–100% of the P losses. Their translocation within preferential flow passes may be rapid, mediated by storm events. The potential total P loss through leaching into subsoils and with streams was found to be less than 50 mg P m^?2 a^?1, suggesting effects on ecosystems at centennial to millennium scale. All current data are based on selected snapshots only. Quantitative measurements of P fluxes in temperate forest systems are nearly absent in the literature, probably due to main research focus on the C and N cycles. Therefore, we lack complete ecosystem‐based assessments of dissolved and colloidal P fluxes within and from temperate forest systems.     

Effect of land use types on decomposition of 14C-labelled maize residue (Zea mays L.)

The effect of three land use types on decomposition of ¹⁴C-labelled maize (Zea mays L.) residues and soil organic matter were investigated under laboratory conditions. Samples of three Dystric Cambisols under plow tillage (PT), reduced tillage (RT) and grassland (GL) collected from the upper 5 cm of the soil profile were incubated for 159 days at 20 °C with or without ¹⁴C-labelled maize residue. After 7 days cumulative CO₂ production was highest in GL and lowest in PT, reflecting differences in soil organic C (SOC) concentration among the three land use types and indicating that mineralized C is a sensitive indicator of the effects of land use regime on SOC. ¹⁴CO₂ efflux from maize residue decomposition was higher in GL than in PT, possibly due to higher SOC and microbial biomass C (MBC) in GL than in PT. ¹⁴CO₂ efflux dynamics from RT soil were different from those of PT and GL. RT had the lowest ¹⁴CO₂ efflux from days 2 to 14 and the highest from days 28 to 159. The lowest MBC in RT explained the delayed decomposition of residues at the beginning. A double exponential model gave a good fit to the mineralization of SOC and residue-¹⁴C (R² > 0.99) and allowed estimation of decomposition rates as dependent on land use. Land use affected the decomposition of labile fractions of SOC and of maize residue, but had no effect on the decomposition of recalcitrant fractions. We conclude that land use affected the decomposition dynamics within the first 1.5 months mainly because of differences in soil microbial biomass but had low effect on cumulative decomposition of maize residues within 5 months.     

Three‐source partitioning of CO2 efflux from maize field soil by 13C natural abundance

A natural‐¹³C‐labeling approach—formerly observed under controlled conditions—was tested in the field to partition total soil CO₂ efflux into root respiration, rhizomicrobial respiration, and soil organic matter (SOM) decomposition. Different results were expected in the field due to different climate, site, and microbial properties in contrast to the laboratory. Within this isotopic method, maize was planted on soil with C₃‐vegetation history and the total CO₂ efflux from soil was subdivided by isotopic mass balance. The C₄‐derived C in soil microbial biomass was also determined. Additionally, in a root‐exclusion approach, root‐ and SOM‐derived CO₂ were determined by the total CO₂ effluxes from maize (Zea mays L.) and bare‐fallow plots. In both approaches, maize‐derived CO₂ contributed 22% to 35% to the total CO₂ efflux during the growth period, which was comparable to other field studies. In our laboratory study, this CO₂ fraction was tripled due to different climate, soil, and sampling conditions. In the natural‐¹³C‐labeling approach, rhizomicrobial respiration was low compared to other studies, which was related to a low amount of C₄‐derived microbial biomass. At the end of the growth period, however, 64% root respiration and 36% rhizomicrobial respiration in relation to total root‐derived CO₂ were calculated when considering high isotopic fractionations between SOM, microbial biomass, and CO₂. This relationship was closer to the 50% : 50% partitioning described in the literature than without fractionation (23% root respiration, 77% rhizomicrobial respiration). Fractionation processes of ¹³C must be taken into account when calculating CO₂ partitioning in soil. Both methods—natural ¹³C labeling and root exclusion—showed the same partitioning results when ¹³C isotopic fractionation during microbial respiration was considered and may therefore be used to separate plant‐ and SOM‐derived CO₂ sources.     

Soil microbial biomass and its activity estimated by kinetic respiration analysis – Statistical guidelines

The use of kinetic respiration analysis to determine soil microbial biomass its active part and maximum specific growth rate has recently increased. With this method, the increase in soil respiration rate initiated by application of carbon growth substrate, e.g. glucose, and mineral nutrients is used to estimate parameters describing microbial growth in soil. This study refines the method by developing statistical guidelines for the data analysis and processing. The kinetic respiration analysis assumes that microbial growth is not limited by substrate and energy. That is why it is critically important to identify the time period corresponding to the unlimited growth. In this work, we studied how the unlimited growth phase can be identified in less subjective ways by examining 121 datasets of respiration time series of 44 different soil samples taken from field plots. Deflection of the respiratory curve from the exponential pattern indicates growth limitation. Subjective selection of the part of respiratory curve which fits to the exponential pattern resulted in a 30% bias in specific microbial growth rates. We propose rules that are based on inspecting the patterns in a series of plots of residuals of fitted respiration rate. By comparing those rules with a set of statistical criteria we find that the weighted-coefficient of determination (r²) can be used to objectively constrain the unlimited growth phase in those cases where double-limitation does not occur. Furthermore, we discuss how the uncertainty of estimated microbial parameters is influenced by a) measurement uncertainty, b) biased measurement at the beginning of the experiment, and c) the number and timing of respiration measurements. We recommend checking plots of fits and residuals as well as reporting uncertainty bounds together with the estimated microbial parameters. A free statistical package is provided to easily deal with those aspects.     

Nutrient limitation of alpine plants: Implications from leaf N : P stoichiometry and leaf δ15N

Nitrogen (N) deposition can affect grassland ecosystems by altering biomass production, plant species composition and abundance. Therefore, a better understanding of the response of dominant plant species to N input is a prerequisite for accurate prediction of future changes and interactions within plant communities. We evaluated the response of seven dominant plant species on the Tibetan Plateau to N input at two levels: individual species and plant functional group. This was achieved by assessing leaf N : P stoichiometry, leaf δ¹⁵N and biomass production for the plant functional groups. Seven dominant plant species—three legumes, two forbs, one grass, one sedge—were analyzed for N, P, and δ¹⁵N 2 years after fertilization with one of the three N forms: NO$ _3^- $ , NH$ _4^+ $ , or NH₄NO₃ at four application rates (0, 7.5, 30, and 150 kg N ha^–1 y^–1). On the basis of biomass production and leaf N : P ratios, we concluded that grasses were limited by available N or co‐limited by available P. Unlike for grasses, leaf N : P and biomass production were not suitable indicators of N limitation for legumes and forbs in alpine meadows. The poor performance of legumes under high N fertilization was mainly due to strong competition with grasses. The total above‐ground biomass was not increased by N fertilization. However, species composition shifted to more productive grasses. A significant negative correlation between leaf N : P and leaf δ¹⁵N indicated that the two forbs Gentiana straminea and Saussurea superba switched from N deficiency to P limitation (e.g., N excess) due to N fertilization. These findings imply that alpine meadows will be more dominated by grasses under increased atmospheric N deposition.