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

Phase 1 study of sirolimus in combination with oral cyclophosphamide and topotecan in children and young adults with relapsed and refractory solid tumors.

PurposeTo determine the maximum tolerated dose (MTD), toxicities, and pharmacodynamics effects of sirolimus combined with oral metronomic topotecan and cyclophosphamide in a pediatric population.Materials and methodsPatients who were 1 to 30 years of age with relapsed/refractory solid tumors (including CNS) were eligible. Patients received daily oral sirolimus and cyclophosphamide (25-50 mg/m2/dose) on days 1-21 and oral topotecan (0.8 mg/m2/dose) on days 1-14 in 28-day cycles. Sirolimus steady-state plasma trough concentrations of 3-7.9 ng/mL and 8-12.0 ng/mL were evaluated, with dose escalation based on a 3+3 phase 1 design. Biomarkers of angiogenesis were also evaluated.ResultsTwenty-one patients were treated (median age 18 years; range 9-30). Dose-limiting toxicities included myelosuppression, ALT elevation, stomatitis, and hypertriglyceridemia. The MTD was sirolimus with trough goal of 8-12.0 ng/mL; cyclophosphamide 25 mg/m2/dose; and topotecan 0.8 mg/m2/dose. No objective responses were observed. Four patients had prolonged stable disease > 4 cycles (range 4-12). Correlative biomarker analyses demonstrated reductions in thrombospondin-1 (p=0.043) and soluble vascular endothelial growth factor receptor-2 plasma concentrations at 21 days compared to baseline.ConclusionsThe combination of oral sirolimus, topotecan, and cyclophosphamide was well tolerated and biomarker studies demonstrated modulation of angiogenic pathways with this regimen

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

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