Isotopic Branchpoints : Linkages and Efficiencies in Carbon and Water Budgets

Forests pass water and carbon through while converting portions to streamflow, soil organic matter, wood production, and other ecosystem services. The efficiencies of these transfers are but poorly quantified. New theory and new instruments have made it possible to use stable isotope composition to provide this quantification of efficiencies wherever there is a measurable difference between the branches of a branchpoint. We present a linked conceptual model that relies on isotopes of hydrogen, carbon, and oxygen to describe these branchpoints along the pathway from precipitation to soil and biomass carbon sequestration and illustrate how it can be tested and generalized. Plain Language Summary The way a forest works can be described in terms of carbon and water budgets, which describe the ways that carbon and water flow through the forest. The paths of such flows are frequently branched and the branches are often different in their stable isotope composition. This means that stable isotopes can be used to describe the branching events. We present isotopic methods of quantifying several such events, then link them in a chain that begins with the evaporation of water and ends with biomass production.Non peer reviewe

Organic nitrogen enhances nitrogen nutrition and early growth of Pinus sylvestris seedlings

Boreal trees are capable of taking up organic nitrogen (N) as effectively as inorganic N. Depending on the abundance of soil N forms, plants may adjust physiological and morphological traits to optimize N uptake. However, the link between these traits and N uptake in response to soil N sources is poorly understood. We examined Pinus sylvestris L. seedlings' biomass growth and allocation, transpiration and N uptake in response to additions of organic N (the amino acid arginine) or inorganic N (ammonium nitrate). We also monitored in situ soil N fluxes in the pots following an addition of N, using a microdialysis system. Supplying organic N resulted in a stable soil N flux, whereas the inorganic N resulted in a sharp increase of nitrate flux followed by a rapid decline, demonstrating a fluctuating N supply and a risk for loss of nitrate from the growth medium. Seedlings supplied with organic N achieved a greater biomass with a higher N content, thus reaching a higher N recovery compared with those supplied inorganic N. In spite of a higher N concentration in organic N seedlings, root-to-shoot ratio and transpiration per unit leaf area were similar to those of inorganic N seedlings. We conclude that enhanced seedlings' nutrition and growth under the organic N source may be attributed to a stable supply of N, owing to a strong retention rate in the soil medium.Peer reviewe

Plant root distributions and nitrogen uptake predicted by a hypothesis of optimal root foraging

CO2-enrichment experiments consistently show that rooting depth increases when trees are grown at elevated CO2 (eCO2), leading in some experiments to increased capture of available soil nitrogen (N) from deeper soil. However, the link between N uptake an

Limited vertical CO2 transport in stems of mature boreal Pinus sylvestris trees

Several studies have suggested that CO2 transport in the transpiration stream can considerably bias estimates of root and stem respiration in ring-porous and diffuse-porous tree species. Whether this also happens in species with tracheid xylem anatomy and lower sap flow rates, such as conifers, is currently unclear. We infused C-13-labelled solution into the xylem near the base of two 90-year-old Pinus sylvestris L. trees. A custom-built gas exchange system and an online isotopic analyser were used to sample the CO2 efflux and its isotopic composition continuously from four positions along the bole and one upper canopy shoot in each tree. Phloem and needle tissue C-13 enrichment was also evaluated at these positions. Most of the C-13 label was lost by diffusion within a few metres of the infusion point indicating rapid CO2 loss during vertical xylem transport. No C-13 enrichment was detected in the upper bole needle tissues. Furthermore, mass balance calculations showed that c. 97% of the locally respired CO2 diffused radially to the atmosphere. Our results support the notion that xylem CO2 transport is of limited magnitude in conifers. This implies that the concerns that stem transport of CO2 derived from root respiration biases chamber-based estimates of forest carbon cycling may be unwarranted for mature conifer stands.Peer reviewe

Effects of externally supplied protein on root morphology and biomass allocation in Arabidopsis

Growth, morphogenesis and function of roots are influenced by the concentration and form of nutrients present in soils, including low molecular mass inorganic N(IN, ammonium, nitrate) and organic N (ON, e.g.amino acids). Proteins, ON of high molecular mass, are prevalent in soils but their possible effects on roots have received little attention. Here, we investigated how externally supplied protein of a size typical of soluble soil proteins influences root development of axenically grown Arabidopsis. Addition of low to intermediate concentrations of protein (bovine serum albumen, BSA) to IN-replete growth medium increased root dry weight, root length and thickness, and root hair length. Supply of higher BSA concentrations inhibited root development. These effects were independent of total N concentrations in the growth medium. The possible involvement of phytohormones was investigated using Arabidopsis with defective auxin (tir1-1 and axr2-1) and ethylene (ein2-1) responses. That no phenotype was observed suggests a signalling pathway is operating independent of auxin and ethylene responses. This study expands the knowledge on N form-explicit responses to demonstrate that ON of high molecular mass elicits specific responses

Long-term nitrogen addition raises the annual carbon sink of a boreal forest to a new steady-state

The boreal forest is an important global carbon (C) sink. Since low soil nitrogen (N) availability is commonly a key constraint on forest productivity, the prevalent view is that increased N input enhances its C sink-strength. This understanding however relies primarily on observations of increased aboveground tree biomass and soil C stock following N fertilization, whereas empirical data evaluating the effects on the whole ecosystem-scale C balance are lacking. Here we use a unique long-term experiment consisting of paired forest stands with eddy covariance measurements to explore the effect of ecosystem-scale N fertilization on the C balance of a managed boreal pine forest. We find that the annual C uptake (i.e. net ecosystem production, NEP) at the fertilized stand was 16 +/- 2% greater relative to the control stand by the end of the first decade of N addition. Subsequently, the ratio of NEP between the fertilized and control stand remained at a stable level during the following five years with an average NEP to N response of 7 & PLUSMN; 1 g C per g N. Our study reveals that this non-linear response of NEP to long-term N fertilization was the result of a cross-seasonal feedback between the N-induced increases in both growing-season C uptake and subsequent winter C emission. We further find that one decade of N addition altered the sensitivity of ecosystem C fluxes to key environmental drivers resulting in divergent responses to weather patterns. Thus, our study highlights the need to account for ecosystem-scale responses to perturbations to improve our understanding of nitrogen-carbon-climate feedbacks in boreal forests.Peer reviewe

Long-term nitrogen addition raises the annual carbon sink of a boreal forest to a new steady-state

The boreal forest is an important global carbon (C) sink. Since low soil nitrogen (N) availability is commonly a key constraint on forest productivity, the prevalent view is that increased N input enhances its C sink-strength. This understanding however relies primarily on observations of increased aboveground tree biomass and soil C stock following N fertilization, whereas empirical data evaluating the effects on the whole ecosystem-scale C balance are lacking. Here we use a unique long-term experiment consisting of paired forest stands with eddy covariance measurements to explore the effect of ecosystem-scale N fertilization on the C balance of a managed boreal pine forest. We find that the annual C uptake (i.e. net ecosystem production, NEP) at the fertilized stand was 16 +/- 2% greater relative to the control stand by the end of the first decade of N addition. Subsequently, the ratio of NEP between the fertilized and control stand remained at a stable level during the following five years with an average NEP to N response of 7 & PLUSMN; 1 g C per g N. Our study reveals that this non-linear response of NEP to long-term N fertilization was the result of a cross-seasonal feedback between the N-induced increases in both growing-season C uptake and subsequent winter C emission. We further find that one decade of N addition altered the sensitivity of ecosystem C fluxes to key environmental drivers resulting in divergent responses to weather patterns. Thus, our study highlights the need to account for ecosystem-scale responses to perturbations to improve our understanding of nitrogen-carbon-climate feedbacks in boreal forests.Peer reviewe

Improving in situ recovery of soil nitrogen using the microdialysis technique

Microdialysis is a technique that can be used to sample fluxes of nitrogen (N) in soils with minimal disturbance. To advance our understanding of the technique and improve N recovery, we compared a common membrane type (10 x 0:5 mm probe length and width, 20 kDa molecular weight cut-off; MWCO) with alternative length and MWCO configurations (30 mm; and 100 kDa MWCO). We hypothesised that the alternative membranes would improve recovery of low molecular weight N via increased surface area and membrane pore size. The test environments, sampled at fixed pump flow rates, were: (i) stirred 100 mu M N standard solution containing organic (amino acid) and inorganic (ammonium, nitrate) N; (ii) soil spiked with 100 mu M standard N solution; and (iii) in situ boreal forest soil. In general, long membranes recovered more N, but the magnitude of improved recovery varied with test environment. Long membranes recovered more inorganic N regardless of flow rate, except ammonium in stirred solution, where length had no effect at slow flow rates. Long membranes also recovered more organic N from stirred solution regardless of flow rate, and recovered most N at slow flow rates in spiked soil. Longer membranes recovered more amino acids in situ in forest soil, with improved resolution of individual amino acids, but were biased towards soluble, mobile forms. MWCO did not affect N recoveries, indicating that in the test conditions, membrane length had greater control than pore size. We discuss the bottlenecks of microdialysis application in soil research and conclude that optimised membrane configurations will advance its use as a tool for quantifying nutrient fluxes in soils. (C) 2017 Elsevier Ltd. All rights reserved

Amino acid transporter mutants of Arabidopsis provides evidence that a non-mycorrhizal plant acquires organic nitrogen from agricultural soil

Although organic nitrogen (N) compounds are ubiquitous in soil solutions, their potential role in plant N nutrition has been questioned. We performed a range of experiments on Arabidopsis thaliana genetically modified to enhance or reduce root uptake of amino acids. Plants lacking expression of the Lysine Histidine Transporter 1 (LHT1) displayed significantly lower contents of C and N label and of U-C,N L-glutamine, as determined by liquid chromatography–mass spectrometry when growing in pots and supplied with dually labelled L-glutamine compared to wild type plants and LHT1-overexpressing plants. Slopes of regressions between accumulation of C-labelled carbon and N-labelled N were higher for LHT1-overexpressing plants than wild type plants, while plants lacking expression of LHT1 did not display a significant regression between the two isotopes. Uptake of labelled organic N from soil tallied with that of labelled ammonium for wild type plants and LHT1-overexpressing plants but was significantly lower for plants lacking expression of LHT1. When grown on agricultural soil plants lacking expression of LHT1 had the lowest, and plants overexpressing LHT1 the highest C/N ratios and natural δN abundance suggesting their dependence on different N pools. Our data show that LHT1 expression is crucial for plant uptake of organic N from soil

Nitrogen fluxes at the root-soil interface show a mismatch of nitrogen fertilizer supply and sugarcane root uptake capacity

Globally only ≈50% of applied nitrogen (N) fertilizer is captured by crops, and the remainder can cause pollution via runoff and gaseous emissions. Synchronizing soil N supply and crop demand will address this problem, however current soil analysis methods provide little insight into delivery and acquisition of N forms by roots. We used microdialysis, a novel technique for in situ quantification of soil nutrient fluxes, to measure N fluxes in sugarcane cropping soils receiving different fertilizer regimes, and compare these with N uptake capacities of sugarcane roots. We show that in fertilized sugarcane soils, fluxes of inorganic N exceed the uptake capacities of sugarcane roots by several orders of magnitude. Contrary, fluxes of organic N closely matched roots’ uptake capacity. These results indicate root uptake capacity constrains plant acquisition of inorganic N. This mismatch between soil N supply and root N uptake capacity is a likely key driver for low N efficiency in the studied crop system. Our results also suggest that (i) the relative contribution of inorganic N for plant nutrition may be overestimated when relying on soil extracts as indicators for root-available N, and (ii) organic N may contribute more to crop N supply than is currently assumed