Mycorrhizal roots in a temperate forest take up organic nitrogen from 13C- and 15N-labeled organic matter

Background and Aims The importance of the uptake of nitrogen in organic form by plants and mycorrhizal fungi has been demonstrated in various ecosystems including temperate forests. However, in previous experiments, isotopically labeled amino acids were often added to soils in concentrations that may be higher than those normally available to roots and mycorrhizal hyphae in situ, and these high concentrations could contribute to exaggerated uptake. Methods We used an experimental approach in which we added 13C-labeled and 15N-labeled whole cells to root-ingrowth cores, allowing proteolytic enzymes to release labeled organic nitrogen at a natural rate, as roots and their associated mycorrhizal fungi grew into the cores. We employed this method in four forest types representing a gradient of soil pH, nitrogen mineralization rate, and mycorrhizal type. Results Intact uptake of organic nitrogen was detected in mycorrhizal roots, and accounted for at least of 1-14% of labeled nitrogen uptake. Forest types did not differ significantly in the importance of organic uptake. Conclusions The estimates of organic N uptake here using 13C-labeled and 15N-labeled whole cells are less than those reported in other temperate forest studies using isotopically labelled amino acids, and likely represent a minimum estimate of organic N-use. The two approaches each have different assumptions, and when used in tandem should complement one another and provide upper and lower bounds of organic N use by plants

Correcting tree-ring δ13C time series for tree-size effects in eight temperate tree species

Stable carbon isotope ratios (δ13C) in tree rings have been widely used to study changes in intrinsic water-use efficiency (iWUE), sometimes with limited consideration of how C-isotope discrimination is affected by tree height and canopy position. Our goals were to quantify the relationships between tree size or tree microenvironment and wood δ13C for eight functionally diverse temperate tree species in northern New England, and to better understand the physical and physiological mechanisms underlying these differences. We collected short increment cores in closed-canopy stands and analyzed δ13C in the most recent 5 years of growth. We also sampled saplings in both shaded and sun-exposed environments. In closed-canopy stands, we found strong tree-size effects on δ13C, with 3.7 – 7.2‰ of difference explained by linear regression vs. height (0.11 – 0.28‰ m-1), which in some cases is substantially stronger than the effect reported in previous studies. However, open-grown saplings were often isotopically more similar to large codominant trees than to shade-grown saplings, indicating that light exposure contributes more to the physiological and isotopic differences between small and large trees than does height. We found that in closed-canopy forests, δ13C correlations with DBH were nonlinear but also strong, allowing a straightforward procedure to correct tree- or stand-scale δ13C-based iWUE chronologies for changing tree size. We demonstrate how to use such data to correct and interpret multi-decadal composite isotope chronologies in both shade-regenerated and open-grown tree cohorts, and we highlight the importance of understanding site history when interpreting δ13C time series

The role of surface roughness, albedo, and Bowen ratio on ecosystem energy balance in the Eastern United States

Land cover and land use influence surface climate through differences in biophysical surface properties, including partitioning of sensible and latent heat (e.g., Bowen ratio), surface roughness, and albedo. Clusters of closely spaced eddy covariance towers (e.g., \u3c10 \u3ekm) over a variety of land cover and land use types provide a unique opportunity to study the local effects of land cover and land use on surface temperature. We assess contributions albedo, energy redistribution due to differences in surface roughness and energy redistribution due to differences in the Bowen ratio using two eddy covariance tower clusters and the coupled (land-atmosphere) Variable-Resolution Community Earth System Model. Results suggest that surface roughness is the dominant biophysical factor contributing to differences in surface temperature between forested and deforested lands. Surface temperature of open land is cooler (−4.8 °C to −0.05 °C) than forest at night and warmer (+0.16 °C to +8.2 °C) during the day at northern and southern tower clusters throughout the year, consistent with modeled calculations. At annual timescales, the biophysical contributions of albedo and Bowen ratio have a negligible impact on surface temperature, however the higher albedo of snow-covered open land compared to forest leads to cooler winter surface temperatures over open lands (−0.4 °C to −0.8 °C). In both the models and observation, the difference in mid-day surface temperature calculated from the sum of the individual biophysical factors is greater than the difference in surface temperature calculated from radiative temperature and potential temperature. Differences in measured and modeled air temperature at the blending height, assumptions about independence of biophysical factors, and model biases in surface energy fluxes may contribute to daytime biases

Partial asynchrony of coniferous forest carbon sources and sinks at the intra-annual time scale.

As major terrestrial carbon sinks, forests play an important role in mitigating climate change. The relationship between the seasonal uptake of carbon and its allocation to woody biomass remains poorly understood, leaving a significant gap in our capacity to predict carbon sequestration by forests. Here, we compare the intra-annual dynamics of carbon fluxes and wood formation across the Northern hemisphere, from carbon assimilation and the formation of non-structural carbon compounds to their incorporation in woody tissues. We show temporally coupled seasonal peaks of carbon assimilation (GPP) and wood cell differentiation, while the two processes are substantially decoupled during off-peak periods. Peaks of cambial activity occur substantially earlier compared to GPP, suggesting the buffer role of non-structural carbohydrates between the processes of carbon assimilation and allocation to wood. Our findings suggest that high-resolution seasonal data of ecosystem carbon fluxes, wood formation and the associated physiological processes may reduce uncertainties in carbon source-sink relationships at different spatial scales, from stand to ecosystem levels

Controls of nitrogen isotope patterns in soil profiles

To determine the dominant processes controlling nitrogen (N) dynamics in soils and increase insights into soil N cycling from nitrogen isotope (δ15N) data, patterns of 15N enrichment in soil profiles were compiled from studies on tropical, temperate, and boreal systems. The maximum 15N enrichment between litter and deeper soil layers varied strongly with mycorrhizal fungal association, averaging 9.6 ± 0.4‰ in ectomycorrhizal systems and 4.6 ± 0.5‰ in arbuscular mycorrhizal systems. The 15N enrichment varied little with mean annual temperature, precipitation, or nitrification rates. One main factor controlling 15N in soil profiles, fractionation against 15N during N transfer by mycorrhizal fungi to host plants, leads to 15N-depleted plant litter at the soil surface and 15N-enriched nitrogen of fungal origin at depth. The preferential preservation of 15N-enriched compounds during decomposition and stabilization is a second important factor. A third mechanism, N loss during nitrification and denitrification, may account for large 15N enrichments with depth in less N-limited forests and may account for soil profiles where maximum δ15N is at intermediate depths. Mixing among soil horizons should also decrease differences among soil horizons. We suggest that dynamic models of isotope distributions within soil profiles that can incorporate multiple processes could provide additional information about the history of nitrogen movements and transformations at a site

Insights into root growth, function, and mycorrhizal abundance from chemical and isotopic data across root orders

Background and aims: Detailed analyses of root chemistry by branching order may provide insights into root function, root lifespan and the abundance of root-associated mycorrhizal fungi in forest ecosystems. Methods: We examined the nitrogen and carbon stable isotopes (δ15N and δ13C) and concentration (%N and %C) in the fine roots of an arbuscular mycorrhizal tree, Fraxinus mandshurica, and an ectomycorrhizal tree, Larix gmelinii, over depth, time, and across five root branching orders. Results and conclusions: Larix δ15N increased by 2.3 ‰ from 4th order to 1st order roots, reflecting the increased presence of 15N-enriched ECM fungi on the lower root orders. In contrast, arbuscular mycorrhizal Fraxinus only increased by 0.7 ‰ from 4th order to 1st order roots, reflecting the smaller 15N enrichment and lower fungal mass on arbuscular mycorrhizal fine roots. Isotopic and anatomical mass balance calculations indicate that first, second, and third order roots in ectomycorrhizal Larix averaged 36 %, 23 %, and 8 % fungal tissue by mass, respectively. Using literature values of root production by root branching order, we estimate that about 25 % of fine root production in ECM species like Larix is actually of fungal sheaths. In contrast to %N, %C, and δ15N, δ13C changed minimally across depth, time, and branching order. The homogeneity of δ13C suggests root tissues are constructed from a large well-mixed reservoir of carbon, although compound specific δ13C data is needed to fully interpret these patterns. The measurements developed here are an important step towards explicitly including mycorrhizal production in forest ecosystem carbon budgets

Tracing metabolic pathways of lipid biosynthesis in ectomycorrhizal fungi from position‐specific 13C‐labelling in glucose

Six position‐specific 13C‐labelled isotopomers of glucose were supplied to the ectomycorrhizal fungi Suillus pungens and Tricholoma flavovirens. From the resulting distribution of 13C among fungal PLFAs, the overall order and contribution of each glucose atom to fatty acid 13C enrichment was: C6 (∼31%) \u3e C5 (∼25%) \u3e C1 (∼18%) \u3e C2 (∼18%) \u3e C3 (∼8%) \u3e C4 (∼1%). These data were used to parameterize a metabolic model of the relative fluxes from glucose degradation to lipid synthesis. Our data revealed that a higher amount of carbon is directed to glycolysis than to the oxidative pentose phosphate pathway (60% and 40% respectively) and that a significant part flows through these pathways more than once (73%) due to the reversibility of some glycolysis reactions. Surprisingly, 95% of carbon cycled through glyoxylate prior to incorporation into lipids, possibly to consume the excess of acetyl‐CoA produced during fatty acid turnover. Our approach provides a rigorous framework for analysing lipid biosynthesis in fungi. In addition, this approach could ultimately improve the interpretation of isotopic patterns at natural abundance in field studies