Snow depth, soil freezing and nitrogen cycling in a northern hardwood forest landscape

Increases in soil freezing associated with decreases in snow cover have been identified as a significant disturbance to nitrogen (N) cycling in northern hardwood forests. We created a range of soil freezing intensity through snow manipulation experiments along an elevation gradient at the Hubbard Brook Experimental Forest (HBEF) in the White Mountains, NH USA in order to improve understanding of the factors regulating freeze effects on nitrate (NO₃ ^?) leaching, nitrous oxide (N₂O) flux, potential and in situ net N mineralization and nitrification, microbial biomass carbon (C) and N content and respiration, and denitrification. While the snow manipulation treatment produced deep and persistent soil freezing at all sites, effects on hydrologic and gaseous losses of N were less than expected and less than values observed in previous studies at the HBEF. There was no relationship between frost depth, frost heaving and NO₃ ^? leaching, and a weak relationship between frost depth and winter N₂O flux. There was a significant positive relationship between dissolved organic carbon (DOC) and NO₃ ^? concentrations in treatment plots but not in reference plots, suggesting that the snow manipulation treatment mobilized available C, which may have stimulated retention of N and prevented treatment effects on N losses. While the results support the hypothesis that climate change resulting in less snow and more soil freezing will increase N losses from northern hardwood forests, they also suggest that ecosystem response to soil freezing disturbance is affected by multiple factors that must be reconciled in future research.     

Climate Variation and Soil Carbon and Nitrogen Cycling Processes in a Northern Hardwood Forest

We exploited the natural climate gradient in the northern hardwood forest at the Hubbard Brook Experimental Forest (HBEF) to evaluate the effects of climate variation similar to what is predicted to occur with global warming over the next 50–100 years for northeastern North America on soil carbon (C) and nitrogen (N) cycle processes. Our objectives were to (1) characterize differences in soil temperature, moisture and frost associated with elevation at the HBEF and (2) evaluate variation in total soil (TSR) and microbial respiration, N mineralization, nitrification, denitrification, nitrous oxide (N₂O) flux, and methane (CH₄) uptake along this gradient. Low elevation sites were consistently warmer (1.5–2.5°C) and drier than high elevation sites. Despite higher temperatures, low elevation plots had less snow and more soil frost than high elevation plots. Net N mineralization and nitrification were slower in warmer, low elevation plots, in both summer and winter. In summer, this pattern was driven by lower soil moisture in warmer soils and in winter the pattern was linked to less snow and more soil freezing in warmer soils. These data suggest that N cycling and supply to plants in northern hardwood ecosystems will be reduced in a warmer climate due to changes in both winter and summer conditions. TSR was consistently faster in the warmer, low elevation plots. N cycling processes appeared to be more sensitive to variation in soil moisture induced by climate variation, whereas C cycling processes appeared to be more strongly influenced by temperature.     

Nitrate and dissolved organic carbon mobilization in response to soil freezing variability

Reduced snowpack and associated increases in soil freezing severity resulting from winter climate change have the potential to disrupt carbon (C) and nitrogen (N) cycling in soils. We used a natural winter climate gradient based on elevation and aspect in a northern hardwood forest to examine the effects of variability in soil freezing depth, duration, and frequency on the mobilization of dissolved organic carbon (DOC) and nitrate (NO₃ ^?) in soils over the course of 2 years. During a winter with a relatively thin snowpack, soils at lower elevation sites experienced greater freezing and especially variable freeze/thaw cycles, which in turn led to greater leaching of DOC from the organic horizon during the following growing season. In contrast to several previous field manipulation studies, we did not find changes in soil solution NO₃ ^? concentrations related to soil freezing variables. Our results are consistent with a soil matrix disturbance from freezing and thawing which increases leachable C. These results build upon previous laboratory experiments and field manipulations that found differing responses of DOC and NO₃ ^? following soil freezing, suggesting that mobilization of labile C may suppress NO₃ ^? losses through microbial immobilization of N. This research highlights the importance of studying natural variation in winter climate and soil freezing and how they impact soil C and N retention, with implications for surface water runoff quality.     

Winter climate change implications for decomposition in northeastern forests: comparisons of sugar maple litter with herbivore fecal inputs

Forests in northeastern North America are influenced by varying climatic and biotic factors; however, there is concern that rapid changes in these factors may lead to important changes in ecosystem processes such as decomposition. Climate change (especially warming) is predicted to increase rates of decomposition in northern latitudes. Warming in winter may result in complex effects including decreased levels of snow cover and an increased incidence of soil freezing that will effect decomposition. Along with these changes in climate, moose densities have also been increasing in this region, likely affecting nutrient dynamics. We measured decomposition and N release from ¹⁵N‐labeled sugar maple leaf litter and moose feces over 20 months in reference and snow removal treatment (to induce soil freezing) plots in two separate experiments at the Hubbard Brook Experimental Forest in New Hampshire, USA. Snow removal/soil freezing decreased decomposition of maple litter, but stimulated N transfer to soil and microbial biomass. Feces decomposed more rapidly than maple litter, and feces N moved into the mineral soil more than N derived from litter, likely due to the lower C : N ratio of feces. Feces decomposition was not affected by the snow removal treatment. Total microbial biomass (measured as microbial N and C) was not significantly affected by the treatments in either the litter or feces plots. These results suggest that increases in soil freezing and/or large herbivore populations, increase the transfer rate of N from plant detritus or digested plants into the mineral soil. Such changes suggest that altering the spatial and temporal patterns of soil freezing and moose density have important implications for ecosystem N cycling.     

Thaw depth determines reaction and transport of inorganic nitrogen in valley bottom permafrost soils

Nitrate (NO₃^–) export coupled with high inorganic nitrogen (N) concentrations in Alaskan streams suggests that N cycles of permafrost‐influenced ecosystems are more open than expected for N‐limited ecosystems. We tested the hypothesis that soil thaw depth governs inorganic N retention and removal in soils due to vertical patterns in the dominant N transformation pathways. Using an in situ, push–pull method, we estimated rates of inorganic N uptake and denitrification during snow melt, summer, and autumn, as depth of soil–stream flowpaths increased in the valley bottom of an arctic and a boreal catchment. Net NO₃^– uptake declined sharply from snow melt to summer and decreased as a nonlinear function of thaw depth. Peak denitrification rate occurred during snow melt at the arctic site, in summer at the boreal site, and declined as a nonlinear function of thaw depth across both sites. Seasonal patterns in ammonium (NH₄⁺) uptake were not significant, but low rates during the peak growing season suggest uptake that is balanced by mineralization. Despite rapid rates of hydrologic transport during snow melt runoff, rates of uptake and removal of inorganic N tended to exceed water residence time during snow melt, indicating potential for retention of N in valley bottom soils when flowpaths are shallow. Decreased reaction rates relative to water residence time in subsequent seasons suggest greater export of inorganic N as the soil–stream flowpath deepens due to thawing soils. Using seasonal thaw as a proxy for longer term deepening of the thaw layer caused by climate warming and permafrost degradation, these results suggest increasing potential for export of inorganic N from permafrost‐influenced soils to streams.     

Plant-microbe competition for soil amino acids in the alpine tundra: effects of freeze-thaw and dry-rewet events

Amino acids have been shown to be a potentially significant N source for the alpine sedge, Kobresia myosuroides. We hypothesised that freeze-thaw and dry-rewet events allow this plant species increased access to amino acids by disrupting microbial cells, which decreases the size of competing microbial populations, but increases soil amino acid concentrations. To test this hypothesis, we characterized freeze-thaw and dry-rewet events in the field and simulated them in laboratory experiments on plant-soil microcosms. In one experiment, ¹⁵N,¹³C-[2]-glycine was added to microcosms that had previously been subjected to a freeze-thaw or dry-rewet event, and isotopic concentrations in the plant and microbial fractions were compared to non-stressed controls. Microbial biomass and uptake of the labeled glycine were unaffected by the freezing and drying treatments, but microbial uptake of ¹⁵N was lower in the two warmer treatments (dry-rewet and summer control) then in the two colder treatments (freeze-thaw and fall control). Plant uptake of glycine-¹⁵N was decreased by climatic disturbance, and uptake in plants that had been frozen appeared to be dependent on the severity of the freeze. The fact that intact glycine was absorbed by the plants was confirmed by near equal enrichment of plant tissues in ¹³C and ¹⁵N. Plants under optimal conditions recovered 3.5% of the added ¹⁵N and microbes recovered 5.0%. The majority of the ¹³C and ¹⁵N label remained in a non-extractable fraction in the bulk soil. To better understand the isolated influences of environmental perturbations on soil amino acid pools and population sizes of amino-acid utilizing microbes, separate experiments were performed in which soils, alone, were subjected to drying and rewetting or freezing and thawing. Potential respiration of glycine and glutamate (substrate-induced respiration; SIR) by the soil microbial communities was unaffected by a single freeze-thaw event. Glycine SIR was decreased slightly (∼10%) by the most extreme drying treatment, but glutamate SIR was not significantly affected. Freezing lowered the concentration of water-extractable amino acids while drying increased their concentration. We interpret the surprising former result as either a decrease in proteolytic activity in frozen soils relative to amino acid uptake, or a stimulation in microbial uptake by physical nutrient release from the soil. We conclude that climatic disturbance does not provide opportunities for increased amino acid uptake by K. myosuroides, but that this plant competes well for amino acid N under non-stressed conditions, especially when soils are warm. We also note that this alpine tundra microbial community's high resistance to freeze-thaw and dry-rewet events is novel and contrasts with studies in other ecosystems. Received: 24 February 1997 / Accepted: 28 August 1997     

Intensive measurement of nitrous oxide emissions from a corn–soybean–wheat rotation under two contrasting management systems over 5 years

No‐tillage (NT), a practice that has been shown to increase carbon sequestration in soils, has resulted in contradictory effects on nitrous oxide (N₂O) emissions. Moreover, it is not clear how mitigation practices for N₂O emission reduction, such as applying nitrogen (N) fertilizer according to soil N reserves and matching the time of application to crop uptake, interact with NT practices. N₂O fluxes from two management systems [conventional (CP), and best management practices: NT + reduced fertilizer (BMP)] applied to a corn (Zea mays L.), soybean (Glycine max L.), winter‐wheat (Triticum aestivum L.) rotation in Ontario, Canada, were measured from January 2000 to April 2005, using a micrometeorological method. The superimposition of interannual variability of weather and management resulted in mean monthly N₂O fluxes ranging from − 1.9 to 61.3 g N ha⁻¹ day⁻¹. Mean annual N₂O emissions over the 5‐year period decreased significantly by 0.79 from 2.19 kg N ha⁻¹ for CP to 1.41 kg N ha⁻¹ for BMP. Growing season (May–October) N₂O emissions were reduced on average by 0.16 kg N ha⁻¹ (20% of total reduction), and this decrease only occurred in the corn year of the rotation. Nongrowing season (November–April) emissions, comprised between 30% and 90% of the annual emissions, mostly due to increased N₂O fluxes during soil thawing. These emissions were well correlated (r²= 0.90) to the accumulated degree‐hours below 0 °C at 5 cm depth, a measure of duration and intensity of soil freezing. Soil management in BMP (NT) significantly reduced N₂O emissions during thaw (80% of total reduction) by reducing soil freezing due to the insulating effects of the larger snow cover plus corn and wheat residue during winter. In conclusion, significant reductions in net greenhouse gas emissions can be obtained when NT is combined with a strategy that matches N application rate and timing to crop needs.     

Evidence of enhanced freezing damage in treeline plants during six years of CO2 enrichment and soil warming

Climate change and elevated atmospheric CO₂ levels could increase the vulnerability of plants to freezing. We analyzed tissue damage resulting from naturally occurring freezing events in plants from a long–term in situ CO₂ enrichment (+ 200 ppm, 2001–2009) and soil warming (+ 4°C since 2007) experiment at treeline in the Swiss Alps (Stillberg, Davos). Summer freezing events caused damage in several abundant subalpine and alpine plant species in four out of six years between 2005 and 2010. Most freezing damage occurred when temperatures dropped below –1.5°C two to three weeks after snow melt. The tree Larix decidua and the dwarf shrubs Vaccinium myrtillus and Empetrum hermaphroditum showed more freezing damage under experimentally elevated CO₂ and/or temperatures than under control conditions. Soil warming induced a 50% die‐back of E. hermaphroditum during a single freezing event due to melting of the protective snow cover. Although we could not identify a clear mechanism, we relate greater freezing susceptibility to a combination of advanced plant phenology in spring and changes in plant physiology. The climate record since 1975 at the treeline site indicated a summer warming by 0.58°C/decade and a 3.5 days/decade earlier snow melt, but no significant decrease in freezing events during the vegetation period. Therefore, in a warmer climate with higher CO₂ levels but constant likelihood of extreme weather events, subalpine and alpine plants may be more susceptible to freezing events, which may partially offset expected enhanced growth with global change. Hence, freezing damage should be considered when predicting changes in growth of alpine plants or changes in community composition under future atmospheric and climate conditions.     

Effects of mild winter freezing on soil nitrogen and carbon dynamics in a northern hardwood forest

Overwinter and snowmelt processes are thought to be critical to controllersof nitrogen (N) cycling and retention in northern forests. However, therehave been few measurements of basic N cycle processes (e.g.mineralization, nitrification, denitrification) during winter and littleanalysis of the influence of winter climate on growing season N dynamics.In this study, we manipulated snow cover to assess the effects of soilfreezing on in situ rates of N mineralization, nitrification and soilrespiration, denitrification (intact core, C₂H₂ – based method),microbial biomass C and N content and potential net N mineralization andnitrification in two sugar maple and two yellow birch stands with referenceand snow manipulation treatment plots over a two year period at theHubbard Brook Experimental Forest, New Hampshire, U.S.A. The snowmanipulation treatment, which simulated the late development of snowpackas may occur in a warmer climate, induced mild (temperatures >–5 °C) soil freezing that lasted until snowmelt. The treatmentcaused significant increases in soil nitrate (NO₃ ^–)concentrations in sugar maple stands, but did not affect mineralization,nitrification, denitrification or microbial biomass, and had no significanteffects in yellow birch stands. Annual N mineralization and nitrificationrates varied significantly from year to year. Net mineralization increasedfrom 12.0 g N m^–2 y^–1 in 1998 to 22 g N m^–2 y^–1 in 1999 and nitrification increased from 8 g N m^–2 y^–1 in 1998 to 13 g N m^–2 y^–1 in 1999.Denitrification rates ranged from 0 to 0.65 g N m^–2 y^–1. Ourresults suggest that mild soil freezing must increase soil NO₃ ^– levels by physical disruption of the soil ecosystem and not by direct stimulation of mineralization and nitrification. Physical disruption canincrease fine root mortality, reduce plant N uptake and reduce competitionfor inorganic N, allowing soil NO₃ ^– levels to increase evenwith no increase in net mineralization or nitrification.     

Effects of freeze–thaw on C and N release from soils below different vegetation in a montane system: a laboratory experiment

Unstable snow cover and more frequent freeze–thaw events have been predicted for montane areas in southern Norway, where stable winters are common today. These systems are important contributors to the flux of carbon (C) and nitrogen (N) to air and water. Here we quantify and compare the effects of freeze–thaw on C and N release from soils collected below Calluna, Molinia or Sphagnum. Intact organic soil cores were subjected to four different freeze–thaw regimes for four consecutive 2‐week periods: (1) slow cycling (SC) with one long freezing event during each 2‐week period, (2) fast cycling (FC) with four short freezing events during each 2‐week period, (3) permanent frost (PF) and (4) permanent thaw (PT). The freezing temperature was −5 °C and the thawing temperature was 5 °C. Before start of treatment, at the end of each 2‐week period, and during postincubation periods, carbon dioxide (CO₂) emission as well as leachable dissolved organic C (DOC), dissolved organic N (DON), ammonium (NH₄), nitrate (NO₃) and absorbance at 254 nm were measured. In soils from all three vegetations, PF increased the release of CO₂, DOC, DON and NH₄ compared with PT. SC caused some scattered effects whereas FC only resulted in some increase in NO₃ release below Molinia. Generally, the emission of CO₂ and leaching of DOC, DON and NH₄ increased in the following order: Sphagnum < Calluna < Molinia. The release of NO₃ was greatest below Calluna. Our data suggest that vegetation cover and composition seem at least as important as increased soil frost for future winter fluxes of CO₂, DOC, DON and dissolved inorganic N (DIN) from the soil to air and water. The freezing period needs to be sufficiently long to give significant effects.     

Long‐term increase in snow depth leads to compositional changes in arctic ectomycorrhizal fungal communities

Many arctic ecological processes are regulated by soil temperature that is tightly interconnected with snow cover distribution and persistence. Recently, various climate‐induced changes have been observed in arctic tundra ecosystems, e.g. shrub expansion, resulting in reduction in albedo and greater C fixation in aboveground vegetation as well as increased rates of soil C mobilization by microbes. Importantly, the net effects of these shifts are unknown, in part because our understanding of belowground processes is limited. Here, we focus on the effects of increased snow depth, and as a consequence, increased winter soil temperature on ectomycorrhizal (ECM) fungal communities in dry and moist tundra. We analyzed deep DNA sequence data from soil samples taken at a long‐term snow fence experiment in Northern Alaska. Our results indicate that, in contrast with previously observed responses of plants to increased snow depth at the same experimental site, the ECM fungal community of the dry tundra was more affected by deeper snow than the moist tundra community. In the dry tundra, both community richness and composition were significantly altered while in the moist tundra, only community composition changed significantly while richness did not. We observed a decrease in richness of Tomentella, Inocybe and other taxa adapted to scavenge the soil for labile N forms. On the other hand, richness of Cortinarius, and species with the ability to scavenge the soil for recalcitrant N forms, did not change. We further link ECM fungal traits with C soil pools. If future warmer atmospheric conditions lead to greater winter snow fall, changes in the ECM fungal community will likely influence C emissions and C fixation through altering N plant availability, fungal biomass and soil‐plant C‐N dynamics, ultimately determining important future interactions between the tundra biosphere and atmosphere.     

A reciprocal transplant experiment within a climatic gradient in a semiarid shrub-steppe ecosystem: effects on bunchgrass growth and reproduction, soil carbon, and soil nitrogen

We investigated the effect of climate change on Poa secunda Presl. and soils in a shrub‐steppe ecosystem in south‐eastern Washington. Intact soil cores containing P. secunda were reciprocally transplanted between two elevations. Plants and soils were examined, respectively, 4.5 and 5 years later. The lower elevation (310 m) site is warmer (28.5 °C air average monthly maximum) and drier (224 mm yr^?1) than the upper elevation (844 m) site (23.5 °C air average monthly maximum, 272 mm yr^?1). Observations were also made on undisturbed plants at both sites. There was no effect of climate change on plant density, shoot biomass, or carbon isotope discrimination in either transplanted plant population. The cooler, wetter environment significantly reduced percent cover and leaf length, while the warmer, drier environment had no effect. Warming and drying reduced percent shoot nitrogen, while the cooler, wetter environment had no effect. Culm density was zero for the lower elevation plants transplanted to the upper site and was 10.3 culms m^?2 at the lower site. There was no effect of warming and drying on the culm density of the upper elevation plants. Culm density of in situ lower elevation plants was greater than that of the in situ upper elevation plants. Warming and drying reduced total soil carbon 32% and total soil nitrogen 40%. The cooler, wetter environment had no effect on total soil C or N. Of the C and N that was lost over time, 64% of both came from the particulate organic matter fraction (POM, > 53 µ m). There was no effect of warming and drying on the upper population of P. secunda while exposing the lower population to the cooler, wetter environment reduced reproductive effort and percent cover. With the warmer and drier conditions that may develop with climate change, total C and N of semiarid soils may decrease with the active fraction of soil C also rapidly decreasing, which may alter ecosystem diversity and function.     

Growth performance of an inland population of Plantago maritima in response to nitrogen and salinity

The inland distribution of Plantago maritima in Ireland and Britain is strongly western and generally associated with base-rich soils or with flushes in more acidic upland soils. Plants from an eastern Irish salt marsh and from an inland population growing on shallow calcareous soils in east County Clare, Ireland, were grown in culture solutions with a range of seawater dilutions (10–100%) and nitrogen (2.8–140 ppm N) concentrations added as ammonium nitrate. A low nitrogen supply (2.8 ppm N) resulted in very low dry matter production but with increased nitrogen, plant growth increased, even of inland population plants in 50% seawater. Some inland plants survived 100% seawater, but growth was much reduced. The relatively high salinity tolerance of this inland population is discussed.Nomenclature follows Flora Europaea (Tutin et al., 1964–80) for angiosperms, Watson (1968) for bryophytes and Hawksworth et al. (1980) for lichens.Thanks are due to the Central Marine Services Unit and Michael Coughlan, Microbiology Department at University College Galway for Mullaghmore soil nitrogen analyses.     

Consistent effects of nitrogen amendments on soil microbial communities and processes across biomes

Ecosystems worldwide are receiving increasing amounts of reactive nitrogen (N) via anthropogenic activities with the added N having potentially important impacts on microbially mediated belowground carbon dynamics. However, a comprehensive understanding of how elevated N availability affects soil microbial processes and community dynamics remains incomplete. The mechanisms responsible for the observed responses are poorly resolved and we do not know if soil microbial communities respond in a similar manner across ecosystems. We collected 28 soils from a broad range of ecosystems in North America, amended soils with inorganic N, and incubated the soils under controlled conditions for 1 year. Consistent across nearly all soils, N addition decreased microbial respiration rates, with an average decrease of 11% over the year‐long incubation, and decreased microbial biomass by 35%. High‐throughput pyrosequencing showed that N addition consistently altered bacterial community composition, increasing the relative abundance of Actinobacteria and Firmicutes, and decreasing the relative abundance of Acidobacteria and Verrucomicrobia. Further, N‐amended soils consistently had lower activities in a broad suite of extracellular enzymes and had decreased temperature sensitivity, suggesting a shift to the preferential decomposition of more labile C pools. The observed trends held across strong gradients in climate and soil characteristics, indicating that the soil microbial responses to N addition are likely controlled by similar wide‐spread mechanisms. Our results support the hypothesis that N addition depresses soil microbial activity by shifting the metabolic capabilities of soil bacterial communities, yielding communities that are less capable of decomposing more recalcitrant soil carbon pools and leading to a potential increase in soil carbon sequestration rates.     

Reduced Snow Cover Increases Wintertime Nitrous Oxide (N<Subscript>2</Subscript>O) Emissions from an Agricultural Soil in the Upper U.S. Midwest

Throughout most of the northern hemisphere, snow cover decreased in almost every winter month from 1967 to 2012. Because snow is an effective insulator, snow cover loss has likely enhanced soil freezing and the frequency of soil freeze–thaw cycles, which can disrupt soil nitrogen dynamics including the production of nitrous oxide (N₂O). We used replicated automated gas flux chambers deployed in an annual cropping system in the upper Midwest US for three winters (December–March, 2011–2013) to examine the effects of snow removal and additions on N₂O fluxes. Diminished snow cover resulted in increased N₂O emissions each year; over the entire experiment, cumulative emissions in plots with snow removed were 69% higher than in ambient snow control plots and 95% higher than in plots that received additional snow (P < 0.001). Higher emissions coincided with a greater number of freeze–thaw cycles that broke up soil macroaggregates (250–8000 µm) and significantly increased soil inorganic nitrogen pools. We conclude that winters with less snow cover can be expected to accelerate N₂O fluxes from agricultural soils subject to wintertime freezing.     

Methane consumption by montane soils: implications for positive and negative feedback with climatic change

We report here three years of field observations of methane uptake, averaging 1.2 mg CH₄ m^–2 d^–1 in montane meadow soils. Surface soil moisture influenced diffusion of substrate while in deeper soil, where methane oxidation was maximum, moisture influenced both diffusion and microbial activity. Microbial oxidation of methane was maximum at an intermediate level of soil moisture, at this site at about 25% moisture by weight (50% water holding capacity). Laboratory incubations also showed inhibition below 20% moisture. These results provide in situ characterization of moisture limitation of methanotroph activity and evidence that soil drying may diminish the methane sink strength. The microbial limitation to methane consumption at low soil moisture provides a mechanism for positive feedback between methane flux and climate warming, as suggested by ice core data (Blunier et al. 1993; Chappellaz et al. 1990; Stauffer et al. 1985).     

Snow depth manipulation and its influence on soil frost and water dynamics in a northern hardwood forest

Climate change will likelyresult in warmer winter temperatures leading toless snowfall in temperate forests. Thesechanges may lead to increases in soil freezingbecause of lack of an insulating snow cover andchanges in soil water dynamics during theimportant snowmelt period. In this study, wemanipulated snow depth by removing snow for twowinters, simulating the late development of thesnowpack as may occur with global warming, toexplore the relationships between snow depth,soil freezing, soil moisture, and infiltration.We established four sites, each with two pairedplots, at the Hubbard Brook Experimental Forest(HBEF) in New Hampshire, U.S.A. and instrumentedall eight plots with soil and snow thermistors,frost tubes, soil moisture probes, and soillysimeters. For two winters, we removed snowfrom the designated treatment plots untilFebruary. Snow in the reference plots wasundisturbed. The treatment winters (1997/1998 and1998/1999) were relatively mild, withtemperatures above the seasonal norm and snowdepths below average. Results show the treatedplots accumulated significantly less snow andhad more extensive soil frost than referenceplots. Snow depth was a strong regulator ofsoil temperature and frost depth at all sites.Soil moisture measured by time domainreflectometry probes and leaching volumescollected in lysimeters were lower in thetreatment plots in March and April compared tothe rest of the year. The ratio of leachatevolumes collected in the treatment plots tothat in the reference plots decreased as thesnow ablation seasons progressed. Our data showthat even mild winters with low snowfall,simulated by snow removal, will result inincreased soil freezing in the forests at theHBEF. Our results suggest that a climate shifttoward less snowfall or a shorter duration ofsnow on the ground will produce increases insoil freezing in northern hardwood forests.Increases in soil freezing will haveimplications for changes in soil biogeochemicalprocesses.     

Bird effects on organic processes in soils from five microhabitats on a nunatak with and without breeding snow petrels in Dronning Maud Land, Antarctica

Five microhabitat types with varying degrees of bird influence were examined. Soils were collected from open polygons, under mosses and bird nests on a nunatak with breeding snow petrels (Pagodroma nivea) and from open polygons and under mosses on a non-bird nunatak. Nutrient levels (total N and P, nitrate, nitrite and ammonia), moisture levels and δ ¹⁵N values were determined and the organic processes of nitrogen fixation (acetylene reduction) and soil respiration (CO₂ flux) were examined. Nests represented the most favourable microhabitat type for soil respiration having the highest nutrient levels and most favourable temperature and moisture regimes. The soils under mosses were also favourable and appear to act as a nutrient sink for nutrients originating from the nests. The open polygons were the least favourable for biological activity. There was little nitrogen fixation in any of the soils except for the soils under mosses from the non-bird nunatak. Fixation is possibly limited in favourable microhabitat types on the bird nunatak by high nitrogen levels. These results were confirmed by the δ ¹⁵N results, which had high values typical of a seabird signal in the soils from the bird nunatak and values near zero, typical of soils containing fixed nitrogen, on the non-bird nunatak. Received: 3 March 1997 / Accepted: 30 March 1998     

N uptake and growth responses to sub-lethal freezing in the grass Poa pratensis L.

Background and aims

Climate warming has the potential to increase both the exposure and vulnerability of grass roots to frost in temperate regions by reducing snow cover and altering the timing of cold acclimation. Despite a strong research focus on the direct effects of freezing on grass mortality, the direct sub-lethal effects of freezing on grass performance have not been well-characterized. We examined sub-lethal responses of the grass Poa pratensis to variation in the timing, severity, rate and length of freezing.

Methods

We assessed short term root functional responses (¹⁵N uptake) and longer term plant growth responses to freezing administered both under controlled conditions in a refrigerated incubator, and in the field by manipulating snow and litter cover.

Results

In fall and spring, ¹⁵N uptake declined in response to 1?day of freezing down to ?10?°C or to 3?days of freezing at ?5?°C, whereas in winter, ¹⁵N uptake was insensitive to freezing. Long term growth responses were similar, with reduced growth only occurring for grasses frozen for 3?days at ?5?°C in spring, but not for grasses frozen in fall or winter. Snow and litter removal intensified soil freezing over winter, but did not significantly affect plant growth.

Conclusions

Our results demonstrate that while P. pratensis is relatively tolerant to frost damage over winter, it may be vulnerable to sub-lethal frost effects in fall, and particularly in spring. These sub-lethal effects occur at temperatures approximately 15–20?°C warmer than the published LT₅₀ values for this species.