Ectomycorrhizal fungi are associated with reduced nitrogen cycling rates in temperate forest soils without corresponding trends in bacterial functional groups.

Microbial processes play a central role in controlling the availability of N in temperate forests. While bacteria, archaea, and fungi account for major inputs, transformations, and exports of N in soil, relationships between microbial community structure and N cycle fluxes have been difficult to detect and characterize. Several studies have reported differences in N cycling based on mycorrhizal type in temperate forests, but associated differences in N cycling genes underlying these fluxes are not well-understood. We explored how rates of soil N cycle fluxes vary across gradients of mycorrhizal abundance (hereafter "mycorrhizal gradients") at four temperate forest sites in Massachusetts and Indiana, USA. We paired measurements of N-fixation, net nitrification, and denitrification rates with gene abundance data for specific bacterial functional groups associated with each process. We find that the availability of NO3 and rates of N-fixation, net nitrification, and denitrification are reduced in stands dominated by trees associated with ECM fungi. On average, rates of N-fixation and denitrification in stands dominated by trees associated with arbuscular mycorrhizal fungi were more than double the corresponding rates in stands dominated by trees associated with ectomycorrhizal fungi. Despite the structuring of flux rates across the mycorrhizal gradients, we did not find concomitant shifts in the abundances of N-cycling bacterial genes, and gene abundances were not correlated with process rates. Given that AM-associating trees are replacing ECM-associating trees throughout much of the eastern US, our results suggest that shifts in mycorrhizal dominance may accelerate N cycling independent of changes in the relative abundance of N cycling bacteria, with consequences for forest productivity and N retention.

Mycorrhiza-mediated competition between plants and decomposers drives soil carbon storage.

Soil contains more carbon than the atmosphere and vegetation combined. Understanding the mechanisms controlling the accumulation and stability of soil carbon is critical to predicting the Earth's future climate. Recent studies suggest that decomposition of soil organic matter is often limited by nitrogen availability to microbes and that plants, via their fungal symbionts, compete directly with free-living decomposers for nitrogen. Ectomycorrhizal and ericoid mycorrhizal (EEM) fungi produce nitrogen-degrading enzymes, allowing them greater access to organic nitrogen sources than arbuscular mycorrhizal (AM) fungi. This leads to the theoretical prediction that soil carbon storage is greater in ecosystems dominated by EEM fungi than in those dominated by AM fungi. Using global data sets, we show that soil in ecosystems dominated by EEM-associated plants contains 70% more carbon per unit nitrogen than soil in ecosystems dominated by AM-associated plants. The effect of mycorrhizal type on soil carbon is independent of, and of far larger consequence than, the effects of net primary production, temperature, precipitation and soil clay content. Hence the effect of mycorrhizal type on soil carbon content holds at the global scale. This finding links the functional traits of mycorrhizal fungi to carbon storage at ecosystem-to-global scales, suggesting that plant-decomposer competition for nutrients exerts a fundamental control over the terrestrial carbon cycle.

Ecosystem responses to elevated CO2 governed by plant-soil interactions and the cost of nitrogen acquisition.

Contents Summary 507 I. Introduction 507 II. The return on investment approach 508 III. CO2 response spectrum 510 IV. Discussion 516 Acknowledgements 518 References 518 SUMMARY: Land ecosystems sequester on average about a quarter of anthropogenic CO2 emissions. It has been proposed that nitrogen (N) availability will exert an increasingly limiting effect on plants' ability to store additional carbon (C) under rising CO2 , but these mechanisms are not well understood. Here, we review findings from elevated CO2 experiments using a plant economics framework, highlighting how ecosystem responses to elevated CO2 may depend on the costs and benefits of plant interactions with mycorrhizal fungi and symbiotic N-fixing microbes. We found that N-acquisition efficiency is positively correlated with leaf-level photosynthetic capacity and plant growth, and negatively with soil C storage. Plants that associate with ectomycorrhizal fungi and N-fixers may acquire N at a lower cost than plants associated with arbuscular mycorrhizal fungi. However, the additional growth in ectomycorrhizal plants is partly offset by decreases in soil C pools via priming. Collectively, our results indicate that predictive models aimed at quantifying C cycle feedbacks to global change may be improved by treating N as a resource that can be acquired by plants in exchange for energy, with different costs depending on plant interactions with microbial symbionts.

Deep peat warming increases surface methane and carbon dioxide emissions in a black spruce-dominated ombrotrophic bog.

Boreal peatlands contain approximately 500 Pg carbon (C) in the soil, emit globally significant quantities of methane (CH4 ), and are highly sensitive to climate change. Warming associated with global climate change is likely to increase the rate of the temperature-sensitive processes that decompose stored organic carbon and release carbon dioxide (CO2 ) and CH4 . Variation in the temperature sensitivity of CO2 and CH4 production and increased peat aerobicity due to enhanced growing-season evapotranspiration may alter the nature of peatland trace gas emission. As CH4 is a powerful greenhouse gas with 34 times the warming potential of CO2 , it is critical to understand how factors associated with global change will influence surface CO2 and CH4 fluxes. Here, we leverage the Spruce and Peatland Responses Under Changing Environments (SPRUCE) climate change manipulation experiment to understand the impact of a 0-9°C gradient in deep belowground warming ("Deep Peat Heat", DPH) on peat surface CO2 and CH4 fluxes. We find that DPH treatments increased both CO2 and CH4 emission. Methane production was more sensitive to warming than CO2 production, decreasing the C-CO2 :C-CH4 of the respired carbon. Methane production is dominated by hydrogenotrophic methanogenesis but deep peat warming increased the δ13 C of CH4 suggesting an increasing contribution of acetoclastic methanogenesis to total CH4 production with warming. Although the total quantity of C emitted from the SPRUCE Bog as CH4 is <2%, CH4 represents >50% of seasonal C emissions in the highest-warming treatments when adjusted for CO2 equivalents on a 100-year timescale. These results suggest that warming in boreal regions may increase CH4 emissions from peatlands and result in a positive feedback to ongoing warming.

Belowground carbon flux links biogeochemical cycles and resource-use efficiency at the global scale.

Nutrient limitation is pervasive in the terrestrial biosphere, although the relationship between global carbon (C) nitrogen (N) and phosphorus (P) cycles remains uncertain. Using meta-analysis we show that gross primary production (GPP) partitioning belowground is inversely related to soil-available N : P, increasing with latitude from tropical to boreal forests. N-use efficiency is highest in boreal forests, and P-use efficiency in tropical forests. High C partitioning belowground in boreal forests reflects a 13-fold greater C cost of N acquisition compared to the tropics. By contrast, the C cost of P acquisition varies only 2-fold among biomes. This analysis suggests a new hypothesis that the primary limitation on productivity in forested ecosystems transitions from belowground resources at high latitudes to aboveground resources at low latitudes as C-intensive root- and mycorrhizal-mediated nutrient capture is progressively replaced by rapidly cycling, enzyme-derived nutrient fluxes when temperatures approach the thermal optimum for biogeochemical transformations.

Reduced snow cover alters root-microbe interactions and decreases nitrification rates in a northern hardwood forest.

Snow cover is projected to decline during the next century in many ecosystems that currently experience a seasonal snowpack. Because snow insulates soils from frigid winter air temperatures, soils are expected to become colder and experience more winter soil freeze-thaw cycles as snow cover continues to decline. Tree roots are adversely affected by snowpack reduction, but whether loss of snow will affect root-microbe interactions remains largely unknown. The objective of this study was to distinguish and attribute direct (e.g., winter snow- and/or soil frost-mediated) vs. indirect (e.g., root-mediated) effects of winter climate change on microbial biomass, the potential activity of microbial exoenzymes, and net N mineralization and nitrification rates. Soil cores were incubated in situ in nylon mesh that either allowed roots to grow into the soil core (2 mm pore size) or excluded root ingrowth (50 µm pore size) for up to 29 months along a natural winter climate gradient at Hubbard Brook Experimental Forest, NH (USA). Microbial biomass did not differ among ingrowth or exclusion cores. Across sampling dates, the potential activities of cellobiohydrolase, phenol oxidase, and peroxidase, and net N mineralization rates were more strongly related to soil volumetric water content (P < 0.05; R2  = 0.25-0.46) than to root biomass, snow or soil frost, or winter soil temperature (R2  < 0.10). Root ingrowth was positively related to soil frost (P < 0.01; R2  = 0.28), suggesting that trees compensate for overwinter root mortality caused by soil freezing by re-allocating resources towards root production. At the sites with the deepest snow cover, root ingrowth reduced nitrification rates by 30% (P < 0.01), showing that tree roots exert significant influence over nitrification, which declines with reduced snow cover. If soil freezing intensifies over time, then greater compensatory root growth may reduce nitrification rates directly via plant-microbe N competition and indirectly through a negative feedback on soil moisture, resulting in lower N availability to trees in northern hardwood forests.

Are above- and below-ground phenology in sync?

Globally, root production accounts for 33-67% of terrestrial net primary productivity and influences decomposition via root production and turnover, carbon (C) allocation to mycorrhizal fungi and root exudation. As recognized above ground, the timing of phenological events affects terrestrial C balance, yet there is no parallel understanding for below-ground phenology. In this paper we examine the phenology of root production and its relationship to temperature, soil moisture, and above-ground phenology. Synthesizing 87 observations of whole-plant phenology from 40 studies, we found that, on average, root growth occurs 25 ± 8 d after shoot growth but that the offset between the peak in root and shoot growth varies > 200 d across biomes (boreal, temperate, Mediterranean, and subtropical). Root and shoot growth are positively correlated with median monthly temperature and mean monthly precipitation in boreal, temperate, and subtropical biomes. However, a temperature hysteresis in these biomes leads to the hypothesis that internal controls over C allocation to roots are an equally, if not more, important driver of phenology. The specific mechanisms are as yet unclear but they are likely mediated by some combination of photoassimilate supply, hormonal signaling, and growth form.

Rhizosphere processes are quantitatively important components of terrestrial carbon and nutrient cycles.

While there is an emerging view that roots and their associated microbes actively alter resource availability and soil organic matter (SOM) decomposition, the ecosystem consequences of such rhizosphere effects have rarely been quantified. Using a meta-analysis, we show that multiple indices of microbially mediated C and nitrogen (N) cycling, including SOM decomposition, are significantly enhanced in the rhizospheres of diverse vegetation types. Then, using a numerical model that combines rhizosphere effect sizes with fine root morphology and depth distributions, we show that root-accelerated mineralization and priming can account for up to one-third of the total C and N mineralized in temperate forest soils. Finally, using a stoichiometrically constrained microbial decomposition model, we show that these effects can be induced by relatively modest fluxes of root-derived C, on the order of 4% and 6% of gross and net primary production, respectively. Collectively, our results indicate that rhizosphere processes are a widespread, quantitatively important driver of SOM decomposition and nutrient release at the ecosystem scale, with potential consequences for global C stocks and vegetation feedbacks to climate.

Fungal functioning in a pine forest: evidence from a ¹5N-labeled global change experiment.

• We used natural and tracer nitrogen (N) isotopes in a Pinus taeda free air CO2 enrichment (FACE) experiment to investigate functioning of ectomycorrhizal and saprotrophic fungi in N cycling. • Fungal sporocarps were sampled in 2004 (natural abundance and (15) N tracer) and 2010 (tracer) and δ(15)N patterns were compared against litter and soil pools. • Ectomycorrhizal fungi with hydrophobic ectomycorrhizas (e.g. Cortinarius and Tricholoma) acquired N from the Oea horizon or deeper. Taxa with hydrophilic ectomycorrhizas acquired N from the Oi horizon (Russula and Lactarius) or deeper (Laccaria, Inocybe, and Amanita). (15)N enrichment patterns for Cortinarius and Amanita in 2010 did not correspond to any measured bulk pool, suggesting that a persistent pool of active organic N supplied these two taxa. Saprotrophic fungi could be separated into those colonizing pine cones (Baeospora), wood, litter (Oi), and soil (Ramariopsis), with δ(15)N of taxa reflecting substrate differences. (15)N enrichment between sources and sporocarps varied across taxa and contributed to δ(15)N patterns. • Natural abundance and (15)N tracers proved useful for tracking N from different depths into fungal taxa, generally corresponded to literature estimates of fungal activity within soil profiles, and provided new insights into interpreting natural abundance δ(15)N patterns.

A big-microsite framework for soil carbon modeling.

Soil carbon cycling processes potentially play a large role in biotic feedbacks to climate change, but little agreement exists at present on what the core of numerical soil C cycling models should look like. In contrast, most canopy models of photosynthesis and leaf gas exchange share a common 'Farquhaur-model' core structure. Here, we explore why a similar core model structure for heterotrophic soil respiration remains elusive and how a pathway to that goal might be envisioned. The spatial and temporal variation in soil microsite conditions greatly complicates modeling efforts, but we believe it is possible to develop a tractable number of parameterizable equations that are organized into a coherent, modular, numerical model structure. First, we show parallels in insights gleaned from linking Arrhenius and Michaelis-Menten kinetics for both photosynthesis and soil respiration. Additional equations and layers of complexity are then added to simulate substrate supply. For soils, model modules that simulate carbon stabilization processes will be key to estimating the fraction of soil C that is accessible to enzymes. Potential modules for dynamic photosynthate input, wetting-event inputs, freeze-thaw impacts on substrate diffusion, aggregate turnover, soluble-C sorption, gas transport, methane respiration, and microbial dynamics are described for conceptually and numerically linking our understanding of fast-response processes of soil gas exchange with longer-term dynamics of soil carbon and nitrogen stocks.

Roots and fungi accelerate carbon and nitrogen cycling in forests exposed to elevated CO2.

A common finding in multiple CO(2) enrichment experiments in forests is the lack of soil carbon (C) accumulation owing to microbial priming of 'old' soil organic matter (SOM). However, soil C losses may also result from the accelerated turnover of 'young' microbial tissues that are rich in nitrogen (N) relative to bulk SOM. We measured root-induced changes in soil C dynamics in a pine forest exposed to elevated CO(2) and N enrichment by combining stable isotope analyses, molecular characterisations of SOM and microbial assays. We find strong evidence that the accelerated turnover of root-derived C under elevated CO(2) is sufficient in magnitude to offset increased belowground inputs. In addition, the C losses were associated with accelerated N cycling, suggesting that trees exposed to elevated CO(2) not only enhance N availability by stimulating microbial decomposition of SOM via priming but also increase the rate at which N cycles through microbial pools.

Inconsistent definitions of "urban" result in different conclusions about the size of urban carbon and nitrogen stocks.

There is conflicting evidence about the importance of urban soils and vegetation in regional C budgets that is caused, in part, by inconsistent definitions of "urban" land use. We quantified urban ecosystem contributions to C stocks in the Boston (Massachusetts, USA) Metropolitan Statistical Area (MSA) using several alternative urban definitions. Development altered aboveground and belowground C and N stocks, and the sign and magnitude of these changes varied by land use and development intensity. Aboveground biomass (live trees, dbh > or = 5 cm) for the MSA was 7.2 +/- 0.4 kg C/m2 (mean +/- SE), reflecting a high proportion of forest cover. Vegetation C was highest in forest (11.6 +/- 0.5 kg C/m2), followed by residential (4.6 +/- 0.5 kg C/m2), and then other developed (2.0 +/- 0.4 kg C/m2) land uses. Soil C (0-10 cm depth) followed the same pattern of decreasing C concentration from forest, to residential, to other developed land uses (4.1 +/- 0.1, 4.0 +/- 0.2, and 3.3 +/- 0.2 kg C/m2, respectively). Within a land use type, urban areas (which we defined as > 25% impervious surface area [ISA] within a 1-km(2) moving window) generally contained less vegetation C, but slightly more soil C, than nonurban areas. Soil N concentrations were higher in urban areas than nonurban areas of the same land use type, except for residential areas, which had similarly high soil N concentrations. When we compared our definition of urban to other commonly used urban extents (U.S. Census Bureau, Global Rural-Urban Mapping Project [GRUMP], and the MSA itself), we found that urban soil (1 m depth) and vegetation C stocks spanned a wide range, from 14.4 +/- 0.8 to 54.5 +/- 3.4 Tg C and from 4.2 +/- 0.4 to 27.3 +/- 3.2 Tg C, respectively. Conclusions about the importance of urban soils and vegetation to regional C and N stocks are very sensitive to the definition of urban used by the investigators. Urban areas, regardless of definition, are rapidly expanding in their extent; a systematic understanding of how our development patterns influence ecosystems is necessary to inform future development choices.

A review of microplastic impacts on seagrasses, epiphytes, and associated sediment communities.

Microplastics have been discovered ubiquitously in marine environments. While their accumulation is noted in seagrass ecosystems, little attention has yet been given to microplastic impacts on seagrass plants and their associated epiphytic and sediment communities. We initiate this discussion by synthesizing the potential impacts microplastics have on relevant seagrass plant, epiphyte, and sediment processes and functions. We suggest that microplastics may harm epiphytes and seagrasses via impalement and light/gas blockage, and increase local concentrations of toxins, causing a disruption in metabolic processes. Further, microplastics may alter nutrient cycling by inhibiting dinitrogen fixation by diazotrophs, preventing microbial processes, and reducing root nutrient uptake. They may also harm seagrass sediment communities via sediment characteristic alteration and organism complications associated with ingestion. All impacts will be exacerbated by the high trapping efficiency of seagrasses. As microplastics become a permanent and increasing member of seagrass ecosystems it will be pertinent to direct future research towards understanding the extent microplastics impact seagrass ecosystems.

Enhanced root exudation induces microbial feedbacks to N cycling in a pine forest under long-term CO2 fumigation.

The degree to which rising atmospheric CO(2) will be offset by carbon (C) sequestration in forests depends in part on the capacity of trees and soil microbes to make physiological adjustments that can alleviate resource limitation. Here, we show for the first time that mature trees exposed to CO(2) enrichment increase the release of soluble C from roots to soil, and that such increases are coupled to the accelerated turnover of nitrogen (N) pools in the rhizosphere. Over the course of 3 years, we measured in situ rates of root exudation from 420 intact loblolly pine (Pinus taeda L.) roots. Trees fumigated with elevated CO(2) (200 p.p.m.v. over background) increased exudation rates (µg C cm(-1) root h(-1) ) by 55% during the primary growing season, leading to a 50% annual increase in dissolved organic inputs to fumigated forest soils. These increases in root-derived C were positively correlated with microbial release of extracellular enzymes involved in breakdown of organic N (R(2) = 0.66; P = 0.006) in the rhizosphere, indicating that exudation stimulated microbial activity and accelerated the rate of soil organic matter (SOM) turnover. In support of this conclusion, trees exposed to both elevated CO(2) and N fertilization did not increase exudation rates and had reduced enzyme activities in the rhizosphere. Collectively, our results provide field-based empirical support suggesting that sustained growth responses of forests to elevated CO(2) in low fertility soils are maintained by enhanced rates of microbial activity and N cycling fuelled by inputs of root-derived C. To the extent that increases in exudation also stimulate SOM decomposition, such changes may prevent soil C accumulation in forest ecosystems.

Increases in the flux of carbon belowground stimulate nitrogen uptake and sustain the long-term enhancement of forest productivity under elevated CO2.

The earth's future climate state is highly dependent upon changes in terrestrial C storage in response to rising concentrations of atmospheric CO2. Here we show that consistently enhanced rates of net primary production (NPP) are sustained by a C-cascade through the root-microbe-soil system; increases in the flux of C belowground under elevated CO2 stimulated microbial activity, accelerated the rate of soil organic matter decomposition and stimulated tree uptake of N bound to this SOM. This process set into motion a positive feedback maintaining greater C gain under elevated CO2 as a result of increases in canopy N content and higher photosynthetic N-use efficiency. The ecosystem-level consequence of the enhanced requirement for N and the exchange of plant C for N belowground is the dominance of C storage in tree biomass but the preclusion of a large C sink in the soil.

Carbon and nitrogen dynamics during forest stand development: a global synthesis.

Our knowledge of carbon (C) and nitrogen (N) dynamics during stand development is not only essential for evaluating the role of secondary forests in the global terrestrial C cycle, but also crucial for understanding long-term C-N interactions in terrestrial ecosystems. However, a comprehensive understanding of forest C and N dynamics over age sequence remains elusive due to the diverse results obtained across individual studies. Here, we synthesized the results of more than 100 studies to examine C and N dynamics during forest stand development. Our results showed that C accumulated in aboveground vegetation, litter and forest floor pools, while the mineral soil C pool did not exhibit significant changes in most studies. The rate of C changes declined with stand age and approached equilibrium during the later stage of stand development. The rate of N changes exhibited linear increases with that of C changes, indicating that N also accrued in various ecosystem components except mineral soil. These results demonstrate that substantial increases in C pools over age sequence are accompanied by N accretion in forest ecosystems. The concurrent C and N dynamics suggest that forest ecosystems may have an intrinsic ability to preclude progressive N limitation during stand development.

Substrate supply, fine roots, and temperature control proteolytic enzyme activity in temperate forest soils.

Temperature and substrate availability constrain the activity of the extracellular enzymes that decompose and release nutrients from soil organic matter (SOM). Proteolytic enzymes are the primary class of enzymes involved in the depolymerization of nitrogen (N) from proteinaceous components of SOM, and their activity affects the rate of N cycling in forest soils. The objectives of this study were to determine whether and how temperature and substrate availability affect the activity of proteolytic enzymes in temperate forest soils, and whether the activity of proteolytic enzymes and other enzymes involved in the acquisition of N (i.e., chitinolytic and ligninolytic enzymes) differs between trees species that form associations with either ectomycorrhizal or arbuscular mycorrhizal fungi. Temperature limitation of proteolytic enzyme activity was observed only early in the growing season when soil temperatures in the field were near 4 degrees C. Substrate limitation to proteolytic activity persisted well into the growing season. Ligninolytic enzyme activity was higher in soils dominated by ectomycorrhizal associated tree species. In contrast, the activity of proteolytic and chitinolytic enzymes did not differ, but there were differences between mycorrhizal association in the control of roots on enzyme activity. Roots of ectomycorrhizal species but not those of arbuscular mycorrhizal species exerted significant control over proteolytic, chitinolytic, and ligninolytic enzyme activity; the absence of ectomycorrhizal fine roots reduced the activity of all three enzymes. These results suggest that climate warming in the absence of increases in substrate availability may have a modest effect on soil-N cycling, and that global changes that alter belowground carbon allocation by trees are likely to have a larger effect on nitrogen cycling in stands dominated by ectomycorrhizal fungi.

Re-assessment of plant carbon dynamics at the Duke free-air CO(2) enrichment site: interactions of atmospheric [CO(2)] with nitrogen and water availability over stand development.

*The potential for elevated [CO(2)]-induced changes to plant carbon (C) storage, through modifications in plant production and allocation of C among plant pools, is an important source of uncertainty when predicting future forest function. Utilizing 10 yr of data from the Duke free-air CO(2) enrichment site, we evaluated the dynamics and distribution of plant C. *Discrepancy between heights measured for this study and previously calculated heights required revision of earlier allometrically based biomass determinations, resulting in higher (up to 50%) estimates of standing biomass and net primary productivity than previous assessments. *Generally, elevated [CO(2)] caused sustained increases in plant biomass production and in standing C, but did not affect the partitioning of C among plant biomass pools. Spatial variation in net primary productivity and its [CO(2)]-induced enhancement was controlled primarily by N availability, with the difference between precipitation and potential evapotranspiration explaining most interannual variability. Consequently, [CO(2)]-induced net primary productivity enhancement ranged from 22 to 30% in different plots and years. *Through quantifying the effects of nutrient and water availability on the forest productivity response to elevated [CO(2)], we show that net primary productivity enhancement by elevated [CO(2)] is not uniform, but rather highly dependent on the availability of other growth resources.

Intact amino acid uptake by northern hardwood and conifer trees.

Empirical and modeling studies of the N cycle in temperate forests of eastern North America have focused on the mechanisms regulating the production of inorganic N, and assumed that only inorganic forms of N are available for plant growth. Recent isotope studies in field conditions suggest that amino acid capture is a widespread ecological phenomenon, although northern temperate forests have yet to be studied. We quantified fine root biomass and applied tracer-level quantities of U-(13)C(2)-(15)N-glycine, (15)NH(4) (+) and (15)NO(3) (-) in two stands, one dominated by sugar maple and white ash, the other dominated by red oak, beech, and hemlock, to assess the importance of amino acids to the N nutrition of northeastern US forests. Significant enrichment of (13)C in fine roots 2 and 5 h following tracer application indicated intact glycine uptake in both stands. Glycine accounted for up to 77% of total N uptake in the oak-beech-hemlock stand, a stand that produces recalcitrant litter, cycles N slowly and has a thick, amino acid-rich organic horizon. By contrast, glycine accounted for only 20% of total N uptake in the sugar maple and white ash stand, a stand characterized by labile litter and rapid rates of amino acid production and turnover resulting in high rates of mineralization and nitrification. This study shows that amino acid uptake is an important process occurring in two widespread, northeastern US temperate forest types with widely differing rates of N cycling.