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In this Brief Communications Arising Reply, the affiliation for author P. H. Templer was incorrectly listed as 'Department of Ecology & Evolutionary Biology, University of California Irvine, Irvine, California 92697, USA' instead of 'Department of Biology, Boston University, Boston, Massachusetts 02215, USA'. This has been corrected online.

Air temperatures are rising and the winter snowpack is getting thinner in many high-latitude and high-elevation ecosystems around the globe. Past studies show that soil warming accelerates microbial metabolism and stimulates soil carbon (C) and nitrogen (N) cycling. Conversely, winter snow removal to simulate loss of snow cover leads to increased soil freezing and reductions in soil microbial biomass, exoenzyme activity, and N cycling. The Climate Change Across Seasons Experiment (CCASE), located at Hubbard Brook Experimental Forest, NH (USA) is designed to evaluate the combined effects of growing season soil warming and an increased frequency of winter soil freeze-thaw cycles on a northern forest ecosystem. Soils were collected from CCASE over two years (2014 and 2015) and extractable C and N pool sizes, as well as microbial biomass, exoenzymes, and potential net N mineralization and microbial respiration were measured. Soil warming alone did not stimulate microbial activity at any sampling time. Extractable amino acid N and organic C, proteolytic and acid phosphatase activity, and microbial respiration were reduced by the combination of warming in the growing season and winter soil freeze-thaw cycles during the period following snowmelt through tree leaf out in spring. The declines in microbial activity also coincided with an 85% decline in microbial biomass N at that time. Growing season warming and winter soil freeze-thaw cycles also resulted in a two-fold reduction in phenol oxidase activity and a 20% reduction in peroxidase activity and these declines persisted throughout the snow-free time of the year. The results from this study suggest that positive feedbacks between warming and rates of soil C and N cycling over the next 100 years will be partially mitigated by an increased frequency of winter soil freeze-thaw cycles, which decrease microbial biomass and rates of soil microbial activity.

The majority of the Earth's terrestrial carbon is stored in the soil. If anthropogenic warming stimulates the loss of this carbon to the atmosphere, it could drive further planetary warming. Despite evidence that warming enhances carbon fluxes to and from the soil, the net global balance between these responses remains uncertain. Here we present a comprehensive analysis of warming-induced changes in soil carbon stocks by assembling data from 49 field experiments located across North America, Europe and Asia. We find that the effects of warming are contingent on the size of the initial soil carbon stock, with considerable losses occurring in high-latitude areas. By extrapolating this empirical relationship to the global scale, we provide estimates of soil carbon sensitivity to warming that may help to constrain Earth system model projections. Our empirical relationship suggests that global soil carbon stocks in the upper soil horizons will fall by 30 ± 30 petagrams of carbon to 203 ± 161 petagrams of carbon under one degree of warming, depending on the rate at which the effects of warming are realized. Under the conservative assumption that the response of soil carbon to warming occurs within a year, a business-as-usual climate scenario would drive the loss of 55 ± 50 petagrams of carbon from the upper soil horizons by 2050. This value is around 12-17 per cent of the expected anthropogenic emissions over this period. Despite the considerable uncertainty in our estimates, the direction of the global soil carbon response is consistent across all scenarios. This provides strong empirical support for the idea that rising temperatures will stimulate the net loss of soil carbon to the atmosphere, driving a positive land carbon-climate feedback that could accelerate climate change.

A defining feature of the redwood forest in coastal California is the presence of fog in the summer months, a time when there is typically little rainfall. Our goal was to determine the role of summer fog in canopy transformation of nitrogen, nitrogen uptake by trees and photosynthesis within a coastal redwood forest ecosystem. We measured horizontal and vertical inputs of nitrogen, the isotopic composition of nitrogen in a variety of atmospheric sources (summer fog, winter rain and throughfall throughout the year), nitrogen pools (soil solution) and plant tissue (roots and foliage), as well as rates of photosynthesis and nitrogen uptake by trees. Throughfall nitrogen fluxes were greater at the forest edge compared to the interior both within the canopy (sampled 10m above-ground) and onto the forest floor (sampled 1m above-ground; P0.05). Nitrification in the forest floor, rather than the canopy, is the primary source of NO3- in these soils throughout the year. Synthesis. Summer fog provides nitrogen directly and indirectly to redwood trees, especially those at the forest edge, and affects the physiologic function of redwood trees. Summer fog provides nitrogen directly and indirectly to redwood trees, especially those at the forest edge, and affects the physiologic function of redwood trees.

Anthropogenic nitrogen (N) deposition is unequally distributed across space and time, with inputs to terrestrial ecosystems impacted by industry regulations and variations in human activity. Soil carbon (C) content normally controls the fraction of mineralized N that is nitrified (ƒ nitrified ), affecting N bioavailability for plants and microbes. However, it is unknown whether N deposition has modified the relationships among soil C, net N mineralization, and net nitrification. To test whether N deposition alters the relationship between soil C and net N transformations, we collected soils from coniferous and deciduous forests, grasslands, and residential yards in 14 regions across the contiguous United States that vary in N deposition rates. We quantified rates of net nitrification and N mineralization, soil chemistry (soil C, N, and pH), and microbial biomass and function (as beta-glucosidase (BG) and N-acetylglucosaminidase (NAG) activity) across these regions. Following expectations, soil C was a driver of ƒ nitrified across regions, whereby increasing soil C resulted in a decline in net nitrification and ƒ nitrified . The ƒ nitrified value increased with lower microbial enzymatic investment in N acquisition (increasing BG:NAG ratio) and lower active microbial biomass, providing some evidence that heterotrophic microbial N demand controls the ammonium pool for nitrifiers. However, higher total N deposition increased ƒ nitrified , including for high soil C sites predicted to have low ƒ nitrified , which decreased the role of soil C as a predictor of ƒ nitrified . Notably, the drop in contemporary atmospheric N deposition rates during the 2020 COVID-19 pandemic did not weaken the effect of N deposition on relationships between soil C and ƒ nitrified . Our results suggest that N deposition can disrupt the relationship between soil C and net N transformations, with this change potentially explained by weaker microbial competition for N. Therefore, past N inputs and soil C should be used together to predict N dynamics across terrestrial ecosystems., (© 2024 John Wiley & Sons Ltd.)

Street trees support climate resiliency through a variety of pathways, such as offsetting urban heat and attenuating storm water runoff. While urban trees in arid and semiarid ecosystems have been shown to take up water from irrigation, it is unknown where street trees in mesic cities obtain their water. In this study, we use natural abundance stable isotopes to estimate the proportional sources of water taken up by Acer platanoides street trees in Boston, Massachusetts, United States, including precipitation, irrigation, groundwater, and wastewater. We use Bayesian multisource mixing models to estimate water sources by comparing the natural abundance isotopic ratios of hydrogen and oxygen across potential water sources with water extracted from tree stem samples. We find that during the summer of 2021, characterized by anomalously high rainfall, street trees predominantly utilized water from precipitation. Precipitation accounted for 72.3 % of water extracted from trees sampled in August and 65.6 % from trees sampled in September. Of the precipitation taken up by street trees, most water was traced back to large storm events in July, with July rainfall alone accounting for up to 84.0 % of water found within street trees. We find strong relationships between canopy cover fractions and the proportion of precipitation lost to evapotranspiration across the study domain, supporting the conclusion that tree planting initiatives result in storm water mitigation benefits due to utilization of water from precipitation by urban vegetation. However, while the mature trees studied here currently support their water demand from precipitation, the dependency of street trees on precipitation in mesic cities may lead to increased water stress in a changing climate characterized by a higher frequency and severity of drought., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Global proliferation of forest edges through anthropogenic land-use change and forest fragmentation is well documented, and while forest fragmentation has clear consequences for soil carbon (C) cycling, underlying drivers of belowground activity at the forest edge remain poorly understood. Increasing soil C losses via respiration have been observed at rural forest edges, but this process was suppressed at urban forest edges. We offer a comprehensive, coupled investigation of abiotic soil conditions and biotic soil activity from forest edge to interior at eight sites along an urbanization gradient to elucidate how environmental stressors are linked to soil C cycling at the forest edge. Despite significant diverging trends in edge soil C losses between urban and rural sites, we did not find comparable differences in soil % C or microbial enzyme activity, suggesting an unexpected decoupling of soil C fluxes and pools at forest edges. We demonstrate that across site types, soils at forest edges were less acidic than the forest interior (p < 0.0001), and soil pH was positively correlated with soil calcium, magnesium and sodium content (adj R 2  = 0.37), which were also elevated at the edge. Compared to forest interior, forest edge soils exhibited a 17.8 % increase in sand content and elevated freeze-thaw frequency with probable downstream effects on root turnover and decomposition. Using these and other novel forest edge data, we demonstrate that significant variation in edge soil respiration (adj R 2  = 0.46; p = 0.0002) and C content (adj R 2  = 0.86; p < 0.0001) can be explained using soil parameters often mediated by human activity (e.g., soil pH, trace metal and cation concentrations, soil temperature), and we emphasize the complex influence of multiple, simultaneous global change drivers at forest edges. Forest edge soils reflect legacies of anthropogenic land-use and modern human management, and this must be accounted for to understand soil activity and C cycling across fragmented landscapes., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Temperate forests are threatened by urbanization and fragmentation, with over 20% (118,300 km 2 ) of U.S. forest land projected to be subsumed by urban land development. We leveraged a unique, well-characterized urban-to-rural and forest edge-to-interior gradient to identify the combined impact of these two land use changes-urbanization and forest edge creation-on the soil microbial community in native remnant forests. We found evidence of mutualism breakdown between trees and their fungal root mutualists [ectomycorrhizal (ECM) fungi] with urbanization, where ECM fungi colonized fewer tree roots and had less connectivity in soil microbiome networks in urban forests compared to rural forests. However, urbanization did not reduce the relative abundance of ECM fungi in forest soils; instead, forest edges alone led to strong reductions in ECM fungal abundance. At forest edges, ECM fungi were replaced by plant and animal pathogens, as well as copiotrophic, xenobiotic-degrading, and nitrogen-cycling bacteria, including nitrifiers and denitrifiers. Urbanization and forest edges interacted to generate new "suites" of microbes, with urban interior forests harboring highly homogenized microbiomes, while edge forest microbiomes were more heterogeneous and less stable, showing increased vulnerability to low soil moisture. When scaled to the regional level, we found that forest soils are projected to harbor high abundances of fungal pathogens and denitrifying bacteria, even in rural areas, due to the widespread existence of forest edges. Our results highlight the potential for soil microbiome dysfunction-including increased greenhouse gas production-in temperate forest regions that are subsumed by urban expansion, both now and in the future.

Winter at high latitudes is characterized by low temperatures, dampened light levels and short photoperiods which shape ecological and evolutionary outcomes from cells to populations to ecosystems. Advances in our understanding of winter biological processes (spanning physiology, behaviour and ecology) highlight that biodiversity threats (e.g. climate change driven shifts in reproductive windows) may interact with winter conditions, leading to greater ecological impacts. As such, conservation and management strategies that consider winter processes and their consequences on biological mechanisms may lead to greater resilience of high altitude and latitude ecosystems. Here, we use well-established threat and action taxonomies produced by the International Union of Conservation of Nature-Conservation Measures Partnership (IUCN-CMP) to synthesize current threats to biota that emerge during, or as the result of, winter processes then discuss targeted management approaches for winter-based conservation. We demonstrate the importance of considering winter when identifying threats to biodiversity and deciding on appropriate management strategies across species and ecosystems. We confirm our expectation that threats are prevalent during the winter and are especially important considering the physiologically challenging conditions that winter presents. Moreover, our findings emphasize that climate change and winter-related constraints on organisms will intersect with other stressors to potentially magnify threats and further complicate management. Though conservation and management practices are less commonly considered during the winter season, we identified several potential or already realized applications relevant to winter that could be beneficial. Many of the examples are quite recent, suggesting a potential turning point for applied winter biology. This growing body of literature is promising but we submit that more research is needed to identify and address threats to wintering biota for targeted and proactive conservation. We suggest that management decisions consider the importance of winter and incorporate winter specific strategies for holistic and mechanistic conservation and resource management., Competing Interests: Cooke is the Editor in Chief of the journal Conservation Physiology. The paper was handled at arms length from Cooke as per COPE guidelines to ensure integrity of the editorial process., (© The Author(s) 2023. Published by Oxford University Press and the Society for Experimental Biology. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Nitrogen (N) availability relative to plant demand has been declining in recent years in terrestrial ecosystems throughout the world, a phenomenon known as N oligotrophication. The temperate forests of the northeastern U.S. have experienced a particularly steep decline in bioavailable N, which is expected to be exacerbated by climate change. This region has also experienced rapid urban expansion in recent decades that leads to forest fragmentation, and it is unknown whether and how these changes affect N availability and uptake by forest trees. Many studies have examined the impact of either urbanization or forest fragmentation on nitrogen (N) cycling, but none to our knowledge have focused on the combined effects of these co-occurring environmental changes. We examined the effects of urbanization and fragmentation on oak-dominated (Quercus spp.) forests along an urban to rural gradient from Boston to central Massachusetts (MA). At eight study sites along the urbanization gradient, plant and soil measurements were made along a 90 m transect from a developed edge to an intact forest interior. Rates of net ammonification, net mineralization, and foliar N concentrations were significantly higher in urban than rural sites, while net nitrification and foliar C:N were not different between urban and rural forests. At urban sites, foliar N and net ammonification and mineralization were higher at forest interiors compared to edges, while net nitrification and foliar C:N were higher at rural forest edges than interiors. These results indicate that urban forests in the northeastern U.S. have greater soil N availability and N uptake by trees compared to rural forests, counteracting the trend for widespread N oligotrophication in temperate forests around the globe. Such increases in available N are diminished at forest edges, however, demonstrating that forest fragmentation has the opposite effect of urbanization on coupled N availability and demand by trees., (© 2023 John Wiley & Sons Ltd.)

As urbanization and forest fragmentation increase around the globe, it is critical to understand how rates of respiration and carbon losses from soil carbon pools are affected by these processes. This study characterizes soils in fragmented forests along an urban to rural gradient, evaluating the sensitivity of soil respiration to changes in soil temperature and moisture near the forest edge. While previous studies found elevated rates of soil respiration at temperate forest edges in rural areas compared to the forest interior, we find that soil respiration is suppressed at the forest edge in urban areas. At urban sites, respiration rates are 25% lower at the forest edge relative to the interior, likely due to high temperature and aridity conditions near urban edges. While rural soils continue to respire with increasing temperatures, urban soil respiration rates asymptote as temperatures climb and soils dry. Soil temperature- and moisture-sensitivity modeling shows that respiration rates in urban soils are less sensitive to rising temperatures than those in rural soils. Scaling these results to Massachusetts (MA), which encompasses 0.25 Mha of the urban forest, we find that failure to account for decreases in soil respiration rates near urban forest edges leads to an overestimate of growing-season soil carbon fluxes of >350,000 Mg C. This difference is almost 2.5 times that for rural soils in the analogous comparison (underestimate of <143,000 Mg C), even though rural forest area is more than four times greater than urban forest area in MA. While a changing climate may stimulate carbon losses from rural forest edge soils, urban forests may experience enhanced soil carbon sequestration near the forest edge. These findings highlight the need to capture the effects of forest fragmentation and land use context when making projections about soil behavior and carbon cycling in a warming and increasingly urbanized world., (© 2022 John Wiley & Sons Ltd.)

The productivity of ecosystems and their capacity to support life depends on access to reactive nitrogen (N). Over the past century, humans have more than doubled the global supply of reactive N through industrial and agricultural activities. However, long-term records demonstrate that N availability is declining in many regions of the world. Reactive N inputs are not evenly distributed, and global changes-including elevated atmospheric carbon dioxide (CO 2 ) levels and rising temperatures-are affecting ecosystem N supply relative to demand. Declining N availability is constraining primary productivity, contributing to lower leaf N concentrations, and reducing the quality of herbivore diets in many ecosystems. We outline the current state of knowledge about declining N availability and propose actions aimed at characterizing and responding to this emerging challenge.

Background: With a lack of United States federal policy to address climate change, cities, the private sector, and universities have shouldered much of the work to reduce carbon dioxide (CO 2 ) and other greenhouse gas emissions. This study aims to determine how landcover characteristics influence the amount of carbon (C) sequestered and respired via biological processes, evaluating the role of land management on the overall C budget of an urban university. Boston University published a comprehensive Climate Action Plan in 2017 with the goal of achieving C neutrality by 2040. In this study, we digitized and discretized each of Boston University's three urban campuses into landcover types, with C sequestration and respiration rates measured and scaled to provide a University-wide estimate of biogenic C fluxes within the broader context of total University emissions., Results: Each of Boston University's three highly urban campuses were net sources of biogenic C to the atmosphere. While trees were estimated to sequester 0.6 ± 0.2 kg C m -2 canopy cover year -1 , mulch and lawn areas in 2018 emitted C at rates of 1.7 ± 0.4 kg C m -2  year -1 and 1.4 ± 0.4 kg C m -2  year -1 , respectively. C uptake by tree canopy cover, which can spatially overlap lawn and mulched landcovers, was not large enough to offset biogenic emissions. The proportion of biogenic emissions to Scope 1 anthropogenic emissions on each campus varied from 0.5% to 2%, and depended primarily on the total anthropogenic emissions on each campus., Conclusions: Our study quantifies the role of urban landcover in local C budgets, offering insights on how landscaping management strategies-such as decreasing mulch application rates and expanding tree canopy extent-can assist universities in minimizing biogenic C emissions and even potentially creating a small biogenic C sink. Although biogenic C fluxes represent a small fraction of overall anthropogenic emissions on urban university campuses, these biogenic fluxes are under active management by the university and should be included in climate action plans.

Climate models project higher growing-season temperatures and a decline in the depth and duration of winter snowpack throughout many north temperate ecosystems over the next century. A smaller snowpack is projected to induce more frequent soil freeze/thaw cycles in winter in northern hardwood forests of the northeastern United States. We measured the combined effects of warmer growing-season soil temperatures and increased winter freeze/thaw cycles on rates of leaf-level photosynthesis and transpiration (sap flow) of red maple (Acer rubrum) trees in a northern hardwood forest at the Climate Change Across Seasons Experiment at Hubbard Brook Experimental Forest in New Hampshire. Soil temperatures were warmed 5°C above ambient temperatures during the growing season and soil freeze/thaw cycles were induced in winter to mimic the projected changes in soil temperature over the next century. Relative to reference plots, growing-season soil warming increased rates of leaf-level photosynthesis by up to 85.32 ± 4.33%, but these gains were completely offset by soil freeze/thaw cycles in winter, suggesting that increased freeze/thaw cycles in winter over the next 100 yr will reduce the effect of warming on leaf-level carbon gains. Soil warming in the growing season increased rates of transpiration per kilopascal of vapor pressure deficit (VPD) by up to 727.39 ± 0.28%, even when trees were exposed to increased frequency of soil freeze/thaw cycles in the previous winter, which could influence regional hydrology in the future. Using climate projections downscaled from the Coupled Model Intercomparison Project, we project increased rates of whole-season transpiration in these forests over the next century by 42-61%. We also project 52-77 additional days when daily air temperatures will be above the long-term average daily maximum during the growing season at Hubbard Brook. Together, these results show that projected changes in climate across both the growing season and winter are likely to cause greater rates of water uptake and have no effect on rates of leaf-level carbon uptake by trees, with potential ecosystem consequences for hydrology and carbon cycling in northern hardwood forests., (© 2020 by the Ecological Society of America.)

Winter air temperatures are rising faster than summer air temperatures in high-latitude forests, increasing the frequency of soil freeze/thaw events in winter. To determine how climate warming and soil freeze/thaw cycles affect soil microbial communities and the ecosystem processes they drive, we leveraged the Climate Change across Seasons Experiment (CCASE) at the Hubbard Brook Experimental Forest in the northeastern United States, where replicate field plots receive one of three climate treatments: warming (+5°C above ambient in the growing season), warming in the growing season + winter freeze/thaw cycles (+5°C above ambient +4 freeze/thaw cycles during winter), and no treatment. Soil samples were taken from plots at six time points throughout the growing season and subjected to amplicon (rDNA) and metagenome sequencing. We found that soil fungal and bacterial community composition were affected by changes in soil temperature, where the taxonomic composition of microbial communities shifted more with the combination of growing-season warming and increased frequency of soil freeze/thaw cycles in winter than with warming alone. Warming increased the relative abundance of brown rot fungi and plant pathogens but decreased that of arbuscular mycorrhizal fungi, all of which recovered under combined growing-season warming and soil freeze/thaw cycles in winter. The abundance of animal parasites increased significantly under combined warming and freeze/thaw cycles. We also found that warming and soil freeze/thaw cycles suppressed bacterial taxa with the genetic potential for carbon (i.e., cellulose) decomposition and soil nitrogen cycling, such as N fixation and the final steps of denitrification. These new soil communities had higher genetic capacity for stress tolerance and lower genetic capacity to grow or reproduce, relative to the communities exposed to warming in the growing season alone. Our observations suggest that initial suppression of biogeochemical cycling with year-round climate change may be linked to the emergence of taxa that trade-off growth for stress tolerance traits., (Copyright © 2020 Garcia, Templer, Sorensen, Sanders-DeMott, Groffman and Bhatnagar.)

Frameworks for limiting ecosystem exposure to excess nutrients and acidity require accurate and complete deposition budgets of reactive nitrogen (Nr). While much progress has been made in developing total Nr deposition budgets for the U.S., current budgets remain limited by key data and knowledge gaps. Analysis of National Atmospheric Deposition Program Total Deposition (NADP/TDep) data illustrates several aspects of current Nr deposition that motivate additional research. Averaged across the continental U.S., dry deposition contributes slightly more (55%) to total deposition than wet deposition and is the dominant process (>90%) over broad areas of the Southwest and other arid regions of the West. Lack of dry deposition measurements imposes a reliance on models, resulting in a much higher degree of uncertainty relative to wet deposition which is routinely measured. As nitrogen oxide (NO x ) emissions continue to decline, reduced forms of inorganic nitrogen (NH x  = NH 3  + NH 4 + ) now contribute >50% of total Nr deposition over large areas of the U.S. Expanded monitoring and additional process-level research are needed to better understand NH x deposition, its contribution to total Nr deposition budgets, and the processes by which reduced N deposits to ecosystems. Urban and suburban areas are hotspots where routine monitoring of oxidized and reduced Nr deposition is needed. Finally, deposition budgets have incomplete information about the speciation of atmospheric nitrogen; monitoring networks do not capture important forms of Nr such as organic nitrogen. Building on these themes, we detail the state of the science of Nr deposition budgets in the U.S. and highlight research priorities to improve deposition budgets in terms of monitoring and flux measurements, leaf- to regional-scale modeling, source apportionment, and characterization of deposition trends and patterns., (Published by Elsevier B.V.)

Direct quantification of terrestrial biosphere responses to global change is crucial for projections of future climate change in Earth system models. Here, we synthesized ecosystem carbon-cycling data from 1,119 experiments performed over the past four decades concerning changes in temperature, precipitation, CO 2 and nitrogen across major terrestrial vegetation types of the world. Most experiments manipulated single rather than multiple global change drivers in temperate ecosystems of the USA, Europe and China. The magnitudes of warming and elevated CO 2 treatments were consistent with the ranges of future projections, whereas those of precipitation changes and nitrogen inputs often exceeded the projected ranges. Increases in global change drivers consistently accelerated, but decreased precipitation slowed down carbon-cycle processes. Nonlinear (including synergistic and antagonistic) effects among global change drivers were rare. Belowground carbon allocation responded negatively to increased precipitation and nitrogen addition and positively to decreased precipitation and elevated CO 2 . The sensitivities of carbon variables to multiple global change drivers depended on the background climate and ecosystem condition, suggesting that Earth system models should be evaluated using site-specific conditions for best uses of this large dataset. Together, this synthesis underscores an urgent need to explore the interactions among multiple global change drivers in underrepresented regions such as semi-arid ecosystems, forests in the tropics and subtropics, and Arctic tundra when forecasting future terrestrial carbon-climate feedback.

Rising winter air temperature will reduce snow depth and duration over the next century in northern hardwood forests. Reductions in snow depth may affect soil bacteria and fungi directly, but also affect soil microbes indirectly through effects of snowpack loss on plant roots. We incubated root exclusion and root ingrowth cores across a winter climate-elevation gradient in a northern hardwood forest for 29 months to identify direct (i.e., winter snow-mediated) and indirect (i.e., root-mediated) effects of winter snowpack decline on soil bacterial and fungal communities, as well as on potential nitrification and net N mineralization rates. Both winter snowpack decline and root exclusion increased bacterial richness and phylogenetic diversity. Variation in bacterial community composition was best explained by differences in winter snow depth or soil frost across elevation. Root ingrowth had a positive effect on the relative abundance of several bacterial taxonomic orders (e.g., Acidobacterales and Actinomycetales). Nominally saprotrophic (e.g., Saccharomycetales and Mucorales) or mycorrhizal (e.g., Helotiales, Russalales, Thelephorales) fungal taxonomic orders were also affected by both root ingrowth and snow depth variation. However, when grouped together, the relative abundance of saprotrophic fungi, arbuscular mycorrhizal fungi, and ectomycorrhizal fungi were not affected by root ingrowth or snow depth, suggesting that traits in addition to trophic mode will mediate fungal community responses to snowpack decline in northern hardwood forests. Potential soil nitrification rates were positively related to ammonia-oxidizing bacteria and archaea abundance (e.g., Nitrospirales, Nitrosomondales, Nitrosphaerales). Rates of N mineralization were positively and negatively correlated with ectomycorrhizal and saprotrophic fungi, respectively, and these relationships were mediated by root exclusion. The results from this study suggest that a declining winter snowpack and its effect on plant roots each have direct effects on the diversity and abundance of soil bacteria and fungal communities that interact to determine rates of soil N cycling in northern hardwood forests.

Changes in growing season climate are often the foci of research exploring forest response to climate change. By contrast, little is known about tree growth response to projected declines in winter snowpack and increases in soil freezing in seasonally snow-covered forest ecosystems, despite extensive documentation of the importance of winter climate in mediating ecological processes. We conducted a 5-year snow-removal experiment whereby snow was removed for the first 4-5 weeks of winter in a northern hardwood forest at the Hubbard Brook Experimental Forest in New Hampshire, USA. Our results indicate that adverse impacts of reduced snowpack and increased soil freezing on the physiology of Acer saccharum (sugar maple), a dominant species across northern temperate forests, are accompanied by a 40 ± 3% reduction in aboveground woody biomass increment, averaged across the 6 years following the start of the experiment. Further, we find no indication of growth recovery 1 year after cessation of the experiment. Based on these findings, we integrate spatial modeling of snowpack depth with forest inventory data to develop a spatially explicit, regional-scale assessment of the vulnerability of forest aboveground growth to projected declines in snowpack depth and increased soil frost. These analyses indicate that nearly 65% of sugar maple basal area in the northeastern United States resides in areas that typically experience insulating snowpack. However, under the RCP 4.5 and 8.5 emissions scenarios, we project a 49%-95% reduction in forest area experiencing insulating snowpack by the year 2099 in the northeastern United States, leaving large areas of northern forest vulnerable to these changes in winter climate, particularly along the northern edge of the region. Our study demonstrates that research focusing on growing season climate alone overestimates the stimulatory effect of warming temperatures on tree and forest growth in seasonally snow-covered forests., (© 2018 John Wiley & Sons Ltd.)

Human societies depend on an Earth system that operates within a constrained range of nutrient availability, yet the recent trajectory of terrestrial nitrogen (N) availability is uncertain. Examining patterns of foliar N concentrations and isotope ratios (δ 15 N) from more than 43,000 samples acquired over 37 years, here we show that foliar N concentration declined by 9% and foliar δ 15 N declined by 0.6-1.6‰. Examining patterns across different climate spaces, foliar δ 15 N declined across the entire range of mean annual temperature and mean annual precipitation tested. These results suggest declines in N supply relative to plant demand at the global scale. In all, there are now multiple lines of evidence of declining N availability in many unfertilized terrestrial ecosystems, including declines in δ 15 N of tree rings and leaves from herbarium samples over the past 75-150 years. These patterns are consistent with the proposed consequences of elevated atmospheric carbon dioxide and longer growing seasons. These declines will limit future terrestrial carbon uptake and increase nutritional stress for herbivores.

Excess nitrogen and phosphorus ("nutrients") loadings continue to affect ecosystem function and human health across the U.S. Our ability to connect atmospheric inputs of nutrients to aquatic end points remains limited due to uncoupled air and water quality monitoring. Where connections exist, the information provides insights about source apportionment, trends, risk to sensitive ecosystems, and efficacy of pollution reduction efforts. We examine several issues driving the need for better integrated monitoring, including: coastal eutrophication, urban hotspots of deposition, a shift from oxidized to reduced nitrogen deposition, and the disappearance of pristine lakes. Successful coordination requires consistent data reporting; collocating deposition and water quality monitoring; improving phosphorus deposition measurements; and filling coverage gaps in urban corridors, agricultural areas, undeveloped watersheds, and coastal zones.

Atmospheric deposition of nitrogen (N) is a major input of N to the biosphere and is elevated beyond preindustrial levels throughout many ecosystems. Deposition monitoring networks in the United States generally avoid urban areas in order to capture regional patterns of N deposition, and studies measuring N deposition in cities usually include only one or two urban sites in an urban-rural comparison or as an anchor along an urban-to-rural gradient. Describing patterns and drivers of atmospheric N inputs is crucial for understanding the effects of N deposition; however, little is known about the variability and drivers of atmospheric N inputs or their effects on soil biogeochemistry within urban ecosystems. We measured rates of canopy throughfall N as a measure of atmospheric N inputs, as well as soil net N mineralization and nitrification, soil solution N, and soil respiration at 15 sites across the greater Boston, Massachusetts area. Rates of throughfall N are 8.70±0.68kgNha -1 yr -1 , vary 3.5-fold across sites, and are positively correlated with rates of local vehicle N emissions. Ammonium (NH 4 + ) composes 69.9±2.2% of inorganic throughfall N inputs and is highest in late spring, suggesting a contribution from local fertilizer inputs. Soil solution NO 3 - is positively correlated with throughfall NO 3 - inputs. In contrast, soil solution NH 4 + , net N mineralization, nitrification, and soil respiration are not correlated with rates of throughfall N inputs. Rather, these processes are correlated with soil properties such as soil organic matter. Our results demonstrate high variability in rates of urban throughfall N inputs, correlation of throughfall N inputs with local vehicle N emissions, and a decoupling of urban soil biogeochemistry and throughfall N inputs., (Copyright © 2017 Elsevier B.V. All rights reserved.)

Climate models project an increase in mean annual air temperatures and a reduction in the depth and duration of winter snowpack for many mid and high latitude and high elevation seasonally snow-covered ecosystems over the next century. The combined effects of these changes in climate will lead to warmer soils in the growing season and increased frequency of soil freeze-thaw cycles (FTCs) in winter due to the loss of a continuous, insulating snowpack. Previous experiments have warmed soils or removed snow via shoveling or with shelters to mimic projected declines in the winter snowpack. To our knowledge, no experiment has examined the interactive effects of declining snowpack and increased frequency of soil FTCs, combined with soil warming in the snow-free season on terrestrial ecosystems. In addition, none have mimicked directly the projected increase in soil FTC frequency in tall statured forests that is expected as a result of a loss of insulating snow in winter. We established the Climate Change Across Seasons Experiment (CCASE) at Hubbard Brook Experimental Forest in the White Mountains of New Hampshire in 2012 to assess the combined effects of these changes in climate on a variety of pedoclimate conditions, biogeochemical processes, and ecology of northern hardwood forests. This paper demonstrates the feasibility of creating soil FTC events in a tall statured ecosystem in winter to simulate the projected increase in soil FTC frequency over the next century and combines this projected change in winter climate with ecosystem warming throughout the snow-free season. Together, this experiment provides a new and more comprehensive approach for climate change experiments that can be adopted in other seasonally snow-covered ecosystems to simulate expected changes resulting from global air temperature rise.

Molecular hydrogen (H 2 ) is an atmospheric trace gas with a large microbe-mediated soil sink, yet cycling of this compound throughout ecosystems is poorly understood. Measurements of the sources and sinks of H 2 in various ecosystems are sparse, resulting in large uncertainties in the global H 2 budget. Constraining the H 2 cycle is critical to understanding its role in atmospheric chemistry and climate. We measured H 2 fluxes at high frequency in a temperate mixed deciduous forest for 15 months using a tower-based flux-gradient approach to determine both the soil-atmosphere and the net ecosystem flux of H 2 . We found that Harvard Forest is a net H 2 sink (-1.4 ± 1.1 kg H 2  ha -1 ) with soils as the dominant H 2 sink (-2.0 ± 1.0 kg H 2  ha -1 ) and aboveground canopy emissions as the dominant H 2 source (+0.6 ± 0.8 kg H 2  ha -1 ). Aboveground emissions of H 2 were an unexpected and substantial component of the ecosystem H 2 flux, reducing net ecosystem uptake by 30% of that calculated from soil uptake alone. Soil uptake was highly seasonal (July maximum, February minimum), positively correlated with soil temperature and negatively correlated with environmental variables relevant to diffusion into soils (i.e., soil moisture, snow depth, snow density). Soil microbial H 2 uptake was correlated with rhizosphere respiration rates (r = 0.8, P < 0.001), and H 2 metabolism yielded up to 2% of the energy gleaned by microbes from carbon substrate respiration. Here, we elucidate key processes controlling the biosphere-atmosphere exchange of H 2 and raise new questions regarding the role of aboveground biomass as a source of atmospheric H 2 and mechanisms linking soil H 2 and carbon cycling. Results from this study should be incorporated into modeling efforts to predict the response of the H 2 soil sink to changes in anthropogenic H 2 emissions and shifting soil conditions with climate and land-use change., (© 2016 John Wiley & Sons Ltd.)

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; R 2  = 0.25-0.46) than to root biomass, snow or soil frost, or winter soil temperature (R 2  < 0.10). Root ingrowth was positively related to soil frost (P < 0.01; R 2  = 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., (© 2016 by the Ecological Society of America.)

The respiratory release of carbon dioxide (CO 2 ) from soil is a major yet poorly understood flux in the global carbon cycle. Climatic warming is hypothesized to increase rates of soil respiration, potentially fueling further increases in global temperatures. However, despite considerable scientific attention in recent decades, the overall response of soil respiration to anticipated climatic warming remains unclear. We synthesize the largest global dataset to date of soil respiration, moisture, and temperature measurements, totaling >3,800 observations representing 27 temperature manipulation studies, spanning nine biomes and over 2 decades of warming. Our analysis reveals no significant differences in the temperature sensitivity of soil respiration between control and warmed plots in all biomes, with the exception of deserts and boreal forests. Thus, our data provide limited evidence of acclimation of soil respiration to experimental warming in several major biome types, contrary to the results from multiple single-site studies. Moreover, across all nondesert biomes, respiration rates with and without experimental warming follow a Gaussian response, increasing with soil temperature up to a threshold of ∼25 °C, above which respiration rates decrease with further increases in temperature. This consistent decrease in temperature sensitivity at higher temperatures demonstrates that rising global temperatures may result in regionally variable responses in soil respiration, with colder climates being considerably more responsive to increased ambient temperatures compared with warmer regions. Our analysis adds a unique cross-biome perspective on the temperature response of soil respiration, information critical to improving our mechanistic understanding of how soil carbon dynamics change with climatic warming., Competing Interests: The authors declare no conflict of interest.

Nitrogen (N) deposition is impacting the services that ecosystems provide to humanity. However, the mechanisms determining impacts on the N cycle are not fully understood. To explore the mechanistic underpinnings of N impacts on N cycle processes, we reviewed and synthesised recent progress in ecosystem N research through empirical studies, conceptual analysis and model simulations. Experimental and observational studies have revealed that the stimulation of plant N uptake and soil retention generally diminishes as N loading increases, while dissolved and gaseous losses of N occur at low N availability but increase exponentially and become the dominant fate of N at high loading rates. The original N saturation hypothesis emphasises sequential N saturation from plant uptake to soil retention before N losses occur. However, biogeochemical models that simulate simultaneous competition for soil N substrates by multiple processes match the observed patterns of N losses better than models based on sequential competition. To enable better prediction of terrestrial N cycle responses to N loading, we recommend that future research identifies the response functions of different N processes to substrate availability using manipulative experiments, and incorporates the measured N saturation response functions into conceptual, theoretical and quantitative analyses., (© 2016 John Wiley & Sons Ltd/CNRS.)

Urban areas are the dominant source of U.S. fossil fuel carbon dioxide (FFCO2) emissions. In the absence of binding international treaties or decisive U.S. federal policy for greenhouse gas regulation, cities have also become leaders in greenhouse gas reduction efforts through climate action plans. These plans focus on anthropogenic carbon flows only, however, ignoring a potentially substantial contribution to atmospheric carbon dioxide (CO2) concentrations from biological respiration. Our aim was to measure the contribution of CO2 efflux from soil respiration to atmospheric CO2 fluxes using an automated CO2 efflux system and to use these measurements to model urban soil CO2 efflux across an urban area. We find that growing season soil respiration is dramatically enhanced in urban areas and represents levels of CO2 efflux of up to 72% of FFCO2 within greater Boston's residential areas, and that soils in urban forests, lawns, and landscaped cover types emit 2.62 ± 0.15, 4.49 ± 0.14, and 6.73 ± 0.26 μmolCO2 m(-2) s(-1), respectively, during the growing season. These rates represent up to 2.2 times greater soil respiration than rates found in nearby rural ecosystems in central Massachusetts (MA), a potential consequence of imported carbon amendments, such as mulch, within a general regime of landowner management. As the scientific community moves rapidly towards monitoring, reporting, and verification of CO2 emissions using ground based approaches and remotely-sensed observations to measure CO2 concentrations, our results show that measurement and modeling of biogenic urban CO2 fluxes will be a critical component for verification of urban climate action plans., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Carbonyl sulfide (OCS), the most abundant sulfur gas in the atmosphere, has a summer minimum associated with uptake by vegetation and soils, closely correlated with CO2. We report the first direct measurements to our knowledge of the ecosystem flux of OCS throughout an annual cycle, at a mixed temperate forest. The forest took up OCS during most of the growing season with an overall uptake of 1.36 ± 0.01 mol OCS per ha (43.5 ± 0.5 g S per ha, 95% confidence intervals) for the year. Daytime fluxes accounted for 72% of total uptake. Both soils and incompletely closed stomata in the canopy contributed to nighttime fluxes. Unexpected net OCS emission occurred during the warmest weeks in summer. Many requirements necessary to use fluxes of OCS as a simple estimate of photosynthesis were not met because OCS fluxes did not have a constant relationship with photosynthesis throughout an entire day or over the entire year. However, OCS fluxes provide a direct measure of ecosystem-scale stomatal conductance and mesophyll function, without relying on measures of soil evaporation or leaf temperature, and reveal previously unseen heterogeneity of forest canopy processes. Observations of OCS flux provide powerful, independent means to test and refine land surface and carbon cycle models at the ecosystem scale.

Background and Aims: Many individual studies have shown that the timing of leaf senescence in boreal and temperate deciduous forests in the northern hemisphere is influenced by rising temperatures, but there is limited consensus on the magnitude, direction and spatial extent of this relationship., Methods: A meta-analysis was conducted of published studies from the peer-reviewed literature that reported autumn senescence dates for deciduous trees in the northern hemisphere, encompassing 64 publications with observations ranging from 1931 to 2010., Key Results: Among the meteorological measurements examined, October temperatures were the strongest predictors of date of senescence, followed by cooling degree-days, latitude, photoperiod and, lastly, total monthly precipitation, although the strength of the relationships differed between high- and low-latitude sites. Autumn leaf senescence has been significantly more delayed at low (25° to 49°N) than high (50° to 70°N) latitudes across the northern hemisphere, with senescence across high-latitude sites more sensitive to the effects of photoperiod and low-latitude sites more sensitive to the effects of temperature. Delays in leaf senescence over time were stronger in North America compared with Europe and Asia., Conclusions: The results indicate that leaf senescence has been delayed over time and in response to temperature, although low-latitude sites show significantly stronger delays in senescence over time than high-latitude sites. While temperature alone may be a reasonable predictor of the date of leaf senescence when examining a broad suite of sites, it is important to consider that temperature-induced changes in senescence at high-latitude sites are likely to be constrained by the influence of photoperiod. Ecosystem-level differences in the mechanisms that control the timing of leaf senescence may affect both plant community interactions and ecosystem carbon storage as global temperatures increase over the next century., (© The Author 2015. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Leaf traits are closely associated with nutrient use by plants and can be utilized as a proxy for nutrient cycling processes. However, open questions remain, in particular regarding the variability of leaf traits within and across seasonally dry tropical forests. To address this, we considered six leaf traits (specific area, thickness, dry matter content, N content, P content and natural abundance (15)N) of four co-occurring tree species (two that are not associated with N2-fixing bacteria and two that are associated with N2-fixing bacteria) and net N mineralization rates and inorganic N concentrations along a precipitation gradient (537-1036 mm per year) in the Yucatan Peninsula, Mexico. Specifically we sought to test the hypothesis that leaf traits of dominant plant species shift along a precipitation gradient, but are affected by soil N cycling. Although variation among different species within each site explains some leaf trait variation, there is also a high level of variability across sites, suggesting that factors other than precipitation regime more strongly influence leaf traits. Principal component analyses indicated that across sites and tree species, covariation in leaf traits is an indicator of soil N availability. Patterns of natural abundance (15)N in foliage and foliage minus soil suggest that variation in precipitation regime drives a shift in plant N acquisition and the openness of the N cycle. Overall, our study shows that both plant species and site are important determinants of leaf traits, and that the leaf trait spectrum is correlated with soil N cycling.

Ecological research is increasingly concentrated at particular locations or sites. This trend reflects a variety of advantages of intensive, site-based research, but also raises important questions about the nature of such spatially delimited research: how well does site based research represent broader areas, and does it constrain scientific discovery? We provide an overview of these issues with a particular focus on one prominent intensive research site: the Hubbard Brook Experimental Forest (HBEF), New Hampshire, USA. Among the key features of intensive sites are: long-term, archived data sets that provide a context for new discoveries and the elucidation of ecological mechanisms; the capacity to constrain inputs and parameters, and to validate models of complex ecological processes; and the intellectual cross-fertilization among disciplines in ecological and environmental sciences. The feasibility of scaling up ecological observations from intensive sites depends upon both the phenomenon of interest and the characteristics of the site. An evaluation of deviation metrics for the HBEF illustrates that, in some respects, including sensitivity and recovery of streams and trees from acid deposition, this site is representative of the Northern Forest region, of which HBEF is a part. However, the mountainous terrain and lack of significant agricultural legacy make the HBEF among the least disturbed sites in the Northern Forest region. Its relatively cool, wet climate contributes to high stream flow compared to other sites. These similarities and differences between the HBEF and the region can profoundly influence ecological patterns and processes and potentially limit the generality of observations at this and other intensive sites. Indeed, the difficulty of scaling up may be greatest for ecological phenomena that are sensitive to historical disturbance and that exhibit the greatest spatiotemporal variation, such as denitrification in soils and the dynamics of bird communities. Our research shows that end member sites for some processes often provide important insights into the behavior of inherently heterogeneous ecological processes. In the current era of rapid environmental and biological change, key ecological responses at intensive sites will reflect both specific local drivers and regional trends.

We found that up to 52 ± 17% of residential litterfall carbon (C) and nitrogen (N; 390.6 kg C and 6.5 kg N ha(-1) yr(-1)) is exported through yard waste removed from the City of Boston, which is equivalent to more than half of annual N outputs as gas loss (i.e. denitrification) or leaching. Our results show that removing yard waste results in a substantial decrease in N inputs to urban areas, which may offset excess N inputs from atmospheric deposition, fertilizer application and pet waste. However, export of C and N via yard waste removal may create nutrient limitation for some vegetation due to diminished recycling of nutrients. Removal of leaf litter from residential areas disrupts nutrient cycling and residential yard management practices are an important modification to urban biogeochemical cycling, which could contribute to spatial heterogeneity of ecosystems that are either N limited or saturated within urban ecosystems., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Atmospheric inputs of N and S in bulk deposition (open collectors) and throughfall (beneath canopy collectors) were measured in and adjacent to two Class 1 wilderness areas of the northeastern US. In general, atmospheric S inputs followed our expectations with throughfall S fluxes increasing with elevation in the White Mountains, New Hampshire and throughfall S fluxes being greater in coniferous than deciduous stands in both sites. In contrast, throughfall N fluxes decreased significantly with elevation. Throughfall NO3 (-) fluxes were greater in coniferous than deciduous stands of Lye Brook, Vermont, but were greater in deciduous than coniferous stands of the White Mountains. We found overlap in the range of values for atmospheric N inputs between our measurements and monitoring data [National Atmospheric Deposition Program (NADP) and Clean Air Status and Trends Network (CASTNET)] for wet and total (wet + dry) deposition at Lye Brook. However, our measurements of total S deposition in the White Mountains and bulk (wet) deposition at both Lye Brook and the White Mountains were significantly lower than NADP plus CASTNET, and NADP data, respectively. Natural abundance (18)O in throughfall and bulk deposition were not significantly different, suggesting that there was no significant biological production of [Formula: see text] via nitrification in the canopy. NO3 (-) concentrations in streams were low and had natural abundance (18)O values consistent with microbial production, demonstrating that atmospheric N is being biologically transformed while moving through these watersheds and that these forested watersheds are unlikely to be N saturated.

Understanding the responses of terrestrial ecosystems to global change remains a major challenge of ecological research. We exploited a natural elevation gradient in a northern hardwood forest to determine how reductions in snow accumulation, expected with climate change, directly affect dynamics of soil winter frost, and indirectly soil microbial biomass and activity during the growing season. Soils from lower elevation plots, which accumulated less snow and experienced more soil temperature variability during the winter (and likely more freeze/thaw events), had less extractable inorganic nitrogen (N), lower rates of microbial N production via potential net N mineralization and nitrification, and higher potential microbial respiration during the growing season. Potential nitrate production rates during the growing season were particularly sensitive to changes in winter snow pack accumulation and winter soil temperature variability, especially in spring. Effects of elevation and winter conditions on N transformation rates differed from those on potential microbial respiration, suggesting that N-related processes might respond differently to winter climate change in northern hardwood forests than C-related processes., (© 2014 John Wiley & Sons Ltd.)

The depth and duration of snow pack is declining in the northeastern United States as a result of warming air temperatures. Since snow insulates soil, a decreased snow pack can increase the frequency of soil freezing, which has been shown to have important biogeochemical implications. One of the most notable effects of soil freezing is increased inorganic nitrogen losses from soil during the following growing season. Decreased nitrogen retention is thought to be due to reduced root uptake, but has not yet been measured directly. We conducted a 2-year snow-removal experiment at Hubbard Brook Experimental Forest in New Hampshire, USA to determine the effects of soil freezing on root uptake and leaching of inorganic nitrogen simultaneously. Snow removal significantly increased the depth of maximal soil frost by 37.2 and 39.5 cm in the first and second winters, respectively (P < 0.001 in 2008/2009 and 2009/2010). As a consequence of soil freezing, root uptake of ammonium declined significantly during the first and second growing seasons after snow removal (P = 0.023 for 2009 and P = 0.005 for 2010). These observed reductions in root nitrogen uptake coincided with significant increases in soil solution concentrations of ammonium in the Oa horizon (P = 0.001 for 2009 and 2010) and nitrate in the B horizon (P < 0.001 and P = 0.003 for 2009 and 2010, respectively). The excess flux of dissolved inorganic nitrogen from the Oa horizon that was attributable to soil freezing was 7.0 and 2.8 kg N ha(-1) in 2009 and 2010, respectively. The excess flux of dissolved inorganic nitrogen from the B horizon was lower, amounting to 1.7 and 0.7 kg N ha(-1) in 2009 and 2010, respectively. Results of this study provide direct evidence that soil freezing reduces root nitrogen uptake, demonstrating that the effects of winter climate change on root function has significant consequences for nitrogen retention and loss in forest ecosystems., (© 2014 John Wiley & Sons Ltd.)

Projections of future changes in land carbon (C) storage using biogeochemical models depend on accurately modeling the interactions between the C and nitrogen (N) cycles. Here, we present a framework for analyzing N limitation in global biogeochemical models to explore how C-N interactions of current models compare to field observations, identify the processes causing model divergence, and identify future observation and experiment needs. We used a set of N-fertilization simulations from two global biogeochemical models (CLM-CN and O-CN) that use different approaches to modeling C-N interactions. On the global scale, net primary productivity (NPP) in the CLM-CN model was substantially more responsive to N fertilization than in the O-CN model. The most striking difference between the two models occurred for humid tropical forests, where the CLM-CN simulated a 62% increase in NPP at high N addition levels (30 g N m(-2) yr(-1)), while the O-CN predicted a 2% decrease in NPP due to N fertilization increasing plant respiration more than photosynthesis. Across 35 temperate and boreal forest sites with field N-fertilization experiments, we show that the CLM-CN simulated a 46% increase in aboveground NPP in response to N, which exceeded the observed increase of 25%. In contrast, the O-CN only simulated a 6% increase in aboveground NPP at the N-fertilization sites. Despite the small response of NPP to N fertilization, the O-CN model accurately simulated ecosystem retention of N and the fate of added N to vegetation when compared to empirical (15) N tracer application studies. In contrast, the CLM-CN predicted lower total ecosystem N retention and partitioned more losses to volatilization than estimated from observed N budgets of small catchments. These results point to the need for model improvements in both models in order to enhance the accuracy with which global C-N cycle feedbacks are simulated., (© 2013 John Wiley & Sons Ltd.)

Due to projected increases in winter air temperatures in the northeastern USA over the next 100 years, the snowpack is expected to decrease in depth and duration, thereby increasing soil exposure to freezing air temperatures. To evaluate the potential physiological responses of sugar maple (Acer saccharum Marsh.) to a reduced snowpack, we measured root injury, foliar cation and carbohydrate concentrations, woody shoot carbohydrate levels, and terminal woody shoot lengths of trees in a snow manipulation experiment in New Hampshire, USA. Snow was removed from treatment plots for the first 6 weeks of winter for two consecutive years, resulting in lower soil temperatures to a depth of 50 cm for both winters compared to reference plots with an undisturbed snowpack. Visibly uninjured roots from trees in the snow removal plots had significantly higher (but sub-lethal) levels of relative electrolyte leakage than trees in the reference plots. Foliar calcium: aluminum (Al) molar ratios were significantly lower, and Al concentrations were significantly higher, in trees from snow removal plots than trees from reference plots. Snow removal also reduced terminal shoot growth and increased foliar starch concentrations. Our results are consistent with previous research implicating soil freezing as a cause of soil acidification that leads to soil cation imbalances, but are the first to show that this translates into altered foliar cation pools, and changes in soluble and structural carbon pools in trees. Increased soil freezing due to a reduced snowpack could exacerbate soil cation imbalances already caused by acidic deposition, and have widespread implications for forest health in the northeastern USA.

Effects of anthropogenic nitrogen (N) deposition and the ability of terrestrial ecosystems to store carbon (C) depend in part on the amount of N retained in the system and its partitioning among plant and soil pools. We conducted a meta-analysis of studies at 48 sites across four continents that used enriched 15N isotope tracers in order to synthesize information about total ecosystem N retention (i.e., total ecosystem 15N recovery in plant and soil pools) across natural systems and N partitioning among ecosystem pools. The greatest recoveries of ecosystem 15N tracer occurred in shrublands (mean, 89.5%) and wetlands (84.8%) followed by forests (74.9%) and grasslands (51.8%). In the short term (< 1 week after 15N tracer application), total ecosystem 15N recovery was negatively correlated with fine-root and soil 15N natural abundance, and organic soil C and N concentration but was positively correlated with mean annual temperature and mineral soil C:N. In the longer term (3-18 months after 15N tracer application), total ecosystem 15N retention was negatively correlated with foliar natural-abundance 15N but was positively correlated with mineral soil C and N concentration and C:N, showing that plant and soil natural-abundance 15N and soil C:N are good indicators of total ecosystem N retention. Foliar N concentration was not significantly related to ecosystem 15N tracer recovery, suggesting that plant N status is not a good predictor of total ecosystem N retention. Because the largest ecosystem sinks for 15N tracer were below ground in forests, shrublands, and grasslands, we conclude that growth enhancement and potential for increased C storage in aboveground biomass from atmospheric N deposition is likely to be modest in these ecosystems. Total ecosystem 15N recovery decreased with N fertilization, with an apparent threshold fertilization rate of 46 kg N x ha(-1) x yr(-1) above which most ecosystems showed net losses of applied 15N tracer in response to N fertilizer addition.

Leaf-level responses to increases in atmospheric carbon dioxide (CO(2)) concentrations could have large implications for water and carbon cycles. We investigated whether stomatal density, guard cell length, and intrinsic water use efficiency (iWUE) of 27 individual trees growing at the Arnold Arboretum in Boston, Massachusetts have responded to changing environmental conditions over the last 100 years. We examined leaves from 74 herbarium specimens collected from three genera-Acer (maples), Quercus (oaks), and Carpinus (hornbeams)-from 1893 to 2006. During this period, global average atmospheric CO(2) concentrations increased by approximately 29% (86 ppm), and temperatures in Boston increased by 1.8°C. Stomatal density and guard cell length were negatively correlated in oaks and hornbeams. Although stomatal density declined and guard cell length increased over time, the changes were not dependent on the magnitude of changes in CO(2) concentrations. Intrinsic WUE did not change significantly over time. Our findings suggest that iWUE may not respond to changes in CO(2) concentrations over the lifetimes of individual trees, possibly because of compensating changes in stomatal density and guard cell size. We provide an example of a method that can enable researchers to differentiate between genetic and plastic responses to global change in long-lived trees.

Ratios of nitrogen (N) isotopes in leaves could elucidate underlying patterns of N cycling across ecological gradients. To better understand global-scale patterns of N cycling, we compiled data on foliar N isotope ratios (delta(15)N), foliar N concentrations, mycorrhizal type and climate for over 11,000 plants worldwide. Arbuscular mycorrhizal, ectomycorrhizal, and ericoid mycorrhizal plants were depleted in foliar delta(15)N by 2 per thousand, 3.2 per thousand, 5.9 per thousand, respectively, relative to nonmycorrhizal plants. Foliar delta(15)N increased with decreasing mean annual precipitation and with increasing mean annual temperature (MAT) across sites with MAT >or= -0.5 degrees C, but was invariant with MAT across sites with MAT < -0.5 degrees C. In independent landscape-level to regional-level studies, foliar delta(15)N increased with increasing N availability; at the global scale, foliar delta(15)N increased with increasing foliar N concentrations and decreasing foliar phosphorus (P) concentrations. Together, these results suggest that warm, dry ecosystems have the highest N availability, while plants with high N concentrations, on average, occupy sites with higher N availability than plants with low N concentrations. Global-scale comparisons of other components of the N cycle are still required for better mechanistic understanding of the determinants of variation in foliar delta(15)N and ultimately global patterns in N cycling.

Humid tropical forests are generally characterized by the lack of nitrogen (N) limitation to net primary productivity, yet paradoxically have high potential for N loss. We conducted an intensive field experiment with 15 NH 4 and 15 NO 3 additions to highly weathered tropical forest soils in Puerto Rico to determine the relative importance of N retention and loss mechanisms. Over one-half of all the NH 4 + produced was rapidly converted to NO 3 - via the process of gross nitrification. During the first 24 hours, plant roots took up 28% of the inorganic N produced, dominantly as NH 4 + , and were a greater sink for N than soil microbial biomass. Soil microbes were not a significant sink for added 15 NH 4 + or 15 NO 3 - during the first 24 hours, and only for 15 NH 4 + after 7 days. Patterns of microbial community composition, as determined by terminal restriction fragment length polymorphism analysis (TRFLP), were weakly but significantly correlated with nitrification and denitrification to N 2 O. Rates of dissimilatory NO 3 - reduction to NH 4 + (DNRA) were high in this forest, accounting for up to 25% of gross NH 4 + production and 35% of gross nitrification. DNRA was a major sink for NO 3 - , which may have contributed to the lower rates of N 2 O and leaching losses. Despite considerable N conservation via DNRA and plant NH 4 + uptake, the fate of ∼45% of the NO 3 - produced and 4% of the NH 4 + produced were not measured in our fluxes, suggesting that other important pathways for N retention and loss (e.g., denitrification to N 2 ) are important in this system. The high proportion of mineralized N that was rapidly nitrified and the fates of that NO 3 - highlight the key role of gross nitrification as a proximate control on N retention and loss in humid tropical forest soils. Furthermore, our results demonstrate the importance of the coupling between DNRA and plant uptake of NH 4 + as a potential N-conserving mechanism within tropical forests., (© 2008 by the Ecological Society of America.)

Watersheds within the Catskill Mountains, New York, receive among the highest rates of nitrogen (N) deposition in the northeastern United States and are beginning to show signs of N saturation. Despite similar amounts of N deposition across watersheds within the Catskill Mountains, rates of soil N cycling and N retention vary significantly among stands of different tree species. We examined the potential use of delta (15)N of plants and soils as an indicator of relative forest soil N cycling rates. We analyzed the delta (15)N of foliage, litterfall, bole wood, surface litter layer, fine roots and organic soil from single-species stands of American beech (Fagus grandifolia), eastern hemlock (Tsuga canadensis), red oak (Quercus rubra), and sugar maple (Acer saccharum). Fine root and organic soil delta (15)N values were highest within sugar maple stands, which correlated significantly with higher rates of net mineralization and nitrification. Results from this study suggest that fine root and organic soil delta (15)N can be used as an indicator of relative rates of soil N cycling. Although not statistically significant, delta (15)N was highest within foliage, wood and litterfall of beech stands, a tree species associated with intermediate levels of soil N cycling rates and forest N retention. Our results show that belowground delta (15)N values are a better indicator of relative rates of soil N cycling than are aboveground delta (15)N values.