Effect of seasonal freeze–thaw cycle on net nitrogen mineralization of soil organic layer in the subalpine/alpine forests of western Sichuan,China

Nitrogen mineralization, a main way that soil organic nitrogen converts to mineral nitrogen, is one of the key processes in soil nitrogen cycle. The mineral nitrogen has an important role in plant growth in the growing season. It has been widely accepted that soil freezing in winter can kill a number of microorganisms, weakening soil nitrogen mineralization. However, more and more recent studies have documented that soil microorganisms still have high activity during the deep freezing period, and obvious nitrogen mineralization in winter. Seasonal freeze–thaw cycle is a common phenomenon in the subalpine/alpine forest region, which may have a strong effect on soil ecological processes. Furthermore, the changing pattern of seasonal freeze–thaw cycles might have a significant influence on soil nitrogen mineralization in this region in the scenarios of global warming. As yet, little attention has been given to nitrogen mineralization of soil organic layer as affected by changed seasonal freeze–thaw pattern, although the increasing studies have demonstrated that winter warming might give strong effects on the litter decomposition and microbial activity in the subalpine/alpine forest regions. Therefore, a method of intact soil core incubation in combination with natural environmental gradient was employed by transferring forest soils from 3582 m (A₁) of altitude to 3298 m (A₂) of altitude and 3023 m (A₃) of altitude in the subalpine/alpine forests of western Sichuan, respectively. The amounts and rates of net nitrogen mineralization in soil organic layer were measured. The incubation period included the growing season and the freeze–thaw season from May 24, 2010 to April 19, 2011. The results suggested that significant net nitrogen mineralization was only observed in soil organic layer at low altitude (A₃) during the whole incubation period. Forest soils at higher altitudes (A₁ and A₂) showed obvious soil nitrogen immobilization. In comparison with the growing season which showed remarkable nitrogen immobilization characteristic, the freeze–thaw season showed obvious nitrogen mineralization at lower altitudes (A₂ and A₃). In contrast, the nitrogen immobilization amounts at high altitude (A₁) in freeze–thaw period were less than those in the growing season. Besides, the maximum of net nitrogen mineralization amounts and rates at high altitude (A₁) in soil organic layer mainly occurred in the late stage of growing season and the onset of freezing, soil nitrogen mineralization at the middle altitude (A₂) mainly occurred in the onset of freezing and the deep freezing period, while the highest amount and rate of net nitrogen mineralization at low altitude (A₃) occurred in the early stage of thawing and the late stage of growing season. Furthermore, the amount and rate of soil net nitrogen mineralization during the freeze–thaw season were increasing with the decrease of altitude, which correlated with soil freeze–thaw cycle and freezing process at different altitudes. These results indicated that increasing soil temperature in the future could not only significantly enhance soil nitrogen mineralization in the freeze–thaw season, but also improve soil nitrogen mineralization by increasing freeze–thaw cycle times and shortening freeze–thaw period. However, the processes were significantly influenced by soil micro-environment of subalpine/alpine forest regions.     

Soil biogeochemistry during the early spring in low arctic mesic tundra and the impacts of deepened snow and enhanced nitrogen availability

Air temperature freeze–thaw cycles often occur during the early spring period directly after snowmelt and before budbreak in low arctic tundra. This early spring period may be associated with nitrogen (N) and carbon (C) loss from soils as leachate or as trace gases, due to the detrimental impact of soil freeze–thaw cycles and a developing active layer on soil microorganisms. We measured soil and microbial pools of C and N in early spring during a period of fluctuating air temperature (ranging from ?4 to +10°C) and in midsummer, in low arctic birch hummock tundra. In addition we measured N₂O, CH₄ and CO₂ production in the early spring. All of these biogeochemical variables were also measured in long-term snowfence (deepened snow) and N-addition plots to characterize climate-change related controls on these variables. Microbial and soil solution pools of C and N, and trace gas production varied among the five early spring sample dates, but only marginally and no more than among sample dates in midsummer. N-addition greatly elevated N₂O fluxes, indicating that although denitrifiers were present their activity during early spring was strongly limited by N-availability, but otherwise trace gas production was very low in early spring. The later thaw, warmer winter and colder spring soil temperatures resulting from deepened snow did not significantly alter N pools or rates in early spring. Together, our results indicate strong stability in microbial and soil solution C and N pool sizes in the early spring period just after snowmelt when soil temperatures are close to 0°C (?1.5 to +5°C). A review of annual temperature records from this and other sites suggests that soil freeze–thaw cycles are probably infrequent in mesic tundra in early spring. We suggest that future studies concerned with temperature controls on soil and microbial biogeochemistry should focus not on soil freeze–thaw cycles per se, but on the rapid and often stepped increases in soil temperature that occur under the thawing snowpack.     

Impact of snowpack decrease on net nitrogen mineralization and nitrification in forest soil of northern Japan

Winter climate change is an important environmental driver that alters the biogeochemical processes of forest soils. The decrease in snowpack amplifies soil freeze–thaw cycles and decreases the snowmelt water supply to soil. This study examined how snow decrease affects nitrogen (N) mineralization and nitrification in forest soil in northern Japan by conducting an in situ experimental snowpack manipulation experiment and a laboratory incubation of soil with different moisture, temperature and freeze–thaw magnitudes. For the incubation studies, surface mineral soil (0–10 cm) was collected from a cool-temperate natural mixed forest and incubated using the resin core method during the winter. In the field, there were two treatments: 50 and 100 % snow removal and control plots. The increase in the soil freeze–thaw cycle increased net N mineralization and marginally decreased the net nitrification in soil. The dissolved organic carbon (DOC) and DOC/DON ratio in soil increased with the decrease in snowpack especially during the snow melt period. These results suggested that the change in substrate quality by the increase in freeze–thaw cycles caused the significant enhancement of microbial ammonium production in soil. The lower soil moisture and higher gross immobilization of inorganic N by soil microbes may be maintaining the slow net nitrification and low nitrate leaching in freeze–thaw cycles with less snowpack. The results indicate that winter climate change would strongly impact N biogeochemistry through the increase in ammonium availability in soil for plants and microbes, whereas it would be unlikely that nitrate loss from surface soil would be enhanced.     

Effects of experimental warming of air,soil and permafrost on carbon balance in Alaskan tundra

The carbon (C) storage capacity of northern latitude ecosystems may diminish as warming air temperatures increase permafrost thaw and stimulate decomposition of previously frozen soil organic C. However, warming may also enhance plant growth so that photosynthetic carbon dioxide (CO₂) uptake may, in part, offset respiratory losses. To determine the effects of air and soil warming on CO₂ exchange in tundra, we established an ecosystem warming experiment – the Carbon in Permafrost Experimental Heating Research (CiPEHR) project – in the northern foothills of the Alaska Range in Interior Alaska. We used snow fences coupled with spring snow removal to increase deep soil temperatures and thaw depth (winter warming) and open‐top chambers to increase growing season air temperatures (summer warming). Winter warming increased soil temperature (integrated 5–40 cm depth) by 1.5 °C, which resulted in a 10% increase in growing season thaw depth. Surprisingly, the additional 2 kg of thawed soil C m^?2 in the winter warming plots did not result in significant changes in cumulative growing season respiration, which may have been inhibited by soil saturation at the base of the active layer. In contrast to the limited effects on growing‐season C dynamics, winter warming caused drastic changes in winter respiration and altered the annual C balance of this ecosystem by doubling the net loss of CO₂ to the atmosphere. While most changes to the abiotic environment at CiPEHR were driven by winter warming, summer warming effects on plant and soil processes resulted in 20% increases in both gross primary productivity and growing season ecosystem respiration and significantly altered the age and sources of CO₂ respired from this ecosystem. These results demonstrate the vulnerability of organic C stored in near surface permafrost to increasing temperatures and the strong potential for warming tundra to serve as a positive feedback to global climate change.     

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.     

Frost fatigue response to simulated frost drought using a centrifuge in Acer mono Maxim

In the coldest part of winter, water uptake is blocked by the frozen soil and frozen stems known as ‘frost drought’ causing severe embolisms in woody plants. Frost drought in stems was simulated in a centrifuge by a synergy between freeze–thaw cycles and the different tensions induced by changing the rotation speed. Frost fatigue was defined as a reduction of embolism resistance after a freeze–thaw cycle and determined from ‘vulnerability curves’, which showed percent losses of conductivity vs tension (positive value) or xylem pressure (negative value). Different tensions combined with a controlled freeze–thaw cycle were induced to investigate the effects on frost resistance over the course of year. During the growing season, Acer mono Maxim. developed significant frost fatigue, and a significant positive correlation was found between frost fatigue response and exogenous tension. During the dormant season, A. mono showed very high embolism resistance to frost drought, even under a tension of 2 MPa. When the exogenous tension was increased to 3 MPa while the stem was frozen, significant frost fatigue occurred. Longer freezing times had more serious effects on frost fatigue in A. mono. A hypothesis was raised that at the same low temperature, the severer the drought (higher tension) when stems were frozen, the higher frost fatigue response would be induced.     

Importance of recent shifts in soil thermal dynamics on growing season length, productivity, and carbon sequestration in terrestrial high-latitude ecosystems

In terrestrial high‐latitude regions, observations indicate recent changes in snow cover, permafrost, and soil freeze–thaw transitions due to climate change. These modifications may result in temporal shifts in the growing season and the associated rates of terrestrial productivity. Changes in productivity will influence the ability of these ecosystems to sequester atmospheric CO₂. We use the terrestrial ecosystem model (TEM), which simulates the soil thermal regime, in addition to terrestrial carbon (C), nitrogen and water dynamics, to explore these issues over the years 1960–2100 in extratropical regions (30–90°N). Our model simulations show decreases in snow cover and permafrost stability from 1960 to 2100. Decreases in snow cover agree well with National Oceanic and Atmospheric Administration satellite observations collected between the years 1972 and 2000, with Pearson rank correlation coefficients between 0.58 and 0.65. Model analyses also indicate a trend towards an earlier thaw date of frozen soils and the onset of the growing season in the spring by approximately 2–4 days from 1988 to 2000. Between 1988 and 2000, satellite records yield a slightly stronger trend in thaw and the onset of the growing season, averaging between 5 and 8 days earlier. In both, the TEM simulations and satellite records, trends in day of freeze in the autumn are weaker, such that overall increases in growing season length are due primarily to earlier thaw. Although regions with the longest snow cover duration displayed the greatest increase in growing season length, these regions maintained smaller increases in productivity and heterotrophic respiration than those regions with shorter duration of snow cover and less of an increase in growing season length. Concurrent with increases in growing season length, we found a reduction in soil C and increases in vegetation C, with greatest losses of soil C occurring in those areas with more vegetation, but simulations also suggest that this trend could reverse in the future. Our results reveal noteworthy changes in snow, permafrost, growing season length, productivity, and net C uptake, indicating that prediction of terrestrial C dynamics from one decade to the next will require that large‐scale models adequately take into account the corresponding changes in soil thermal regimes.     

Effects of soil freeze–thaw cycles differ between experimental plant communities

Soil freeze–thaw cycles (FTC) influence nutrient cycling, but their consequences for productivity and composition of vegetation are not well investigated. Ongoing global warming will increase the recurrence of FTC in cool-temperate and high-latitude regions.Here, we report on the above- and belowground biomass production as well as the nitrogen nutrition of two common vegetation types, grassland and heath, after more frequent FTC in a controlled field experiment in Central Europe. Furthermore, we analyze the duration of the observed effects. Five FTC were induced by buried heating wires in addition to three naturally occurring FTC during winter 2005/06. More frequent FTC significantly increased aboveground production of experimental grassland early in the following growing season. However, no reaction was found for experimental heath within the first growing season. Biomass production of heath communities dropped significantly and C/N ratio increased in the freeze–thaw treated plots in the second year after the manipulation, whereas production in the grassland communities was no longer affected significantly, except for an increase in C/N ratio. This response can at least partly be explained by changes in nutrient availability, as plant available nitrate increased in the manipulated grassland plots and decreased in the manipulated heath plots.The results show the high ecological importance of climate changes during winter, with the outcomes differing strongly between contrasting vegetation types. Furthermore, we show that short term climatic events can cause long-lasting effects, sometimes emerging in the vegetation only after considerable time lags (here: one growing season).     

冬季升温对高山生态系统碳氮循环过程的影响

全球温度升高是目前面临的重要环境问题,但存在明显的季节差异性,即冬季升温幅度显著高于夏季的季节非对称性趋势,这在高纬度和高海拔地区更加显著。冬季升温会直接影响积雪覆盖与冰冻层厚度,并引起冻融交替循环的增加,而冬季植物处于休眠状态,这会直接影响土壤中有效氮的吸收与损失,引起土壤有效氮可利用性的变化。然而,关于冬季增温对后续生长季节植物活动、土壤碳氮循环过程的影响等方面的研究仍存在诸多不确定。综述了冬季升温对积雪覆盖与冻融交替循环改变对高山生态系统物质循环的影响,以及冬季升温对土壤碳氮循环、微生物与酶活性的影响,并由此引起的植物物候期、群落结构、生产与养分循环与凋落物分解等生理、生态过程方面的研究进展。在未来的研究中,应针对不同生态系统特点选择合适的冬季增温方式,加强非极地苔原地区关于冬季升温的研究,注重关注冬季升温对植物-土壤微生物之间反馈作用的影响,重点关注冬季升温对生态系统的延滞效应。     

Absence of soil frost affects plant-soil interactions in temperate grasslands

Background and aims

Intermittently frozen ground in winter is expected to disappear over large areas in the temperate zone due to ongoing climate warming. The lack of soil frost influences plant soil interactions and needs to be studied in more detail.

Methods

Winter soil frost was avoided by belowground heating wires in a field experiment over two subsequent winters in a temperate grassland. Soil respiration, soil nitrogen availability and plant performance (aboveground biomass, root length at two depth levels, greenness, nutrient content) were compared between “no-frost” and reference plots which underwent repeated freeze-thaw cycles in both winters.

Results

Soil respiration increased in the “no-frost” treatment during the warming phase (+291 %). N-availability in the upper 10 cm of the soil profile was not affected, possibly due to increased plant N accumulation during winter (+163 %), increased plant N concentration (+18 %) and increased biomass production (+31.5 %) in the growing season. Translocation of roots into deeper soil layers without changes in total root length in response to the “no-frost” treatment, however, may be a sign of nutrient leaching.

Conclusions

The cumulative effect on carbon cycling due to warmer soils therefore depends on the balance between increased winter carbon loss due to higher soil biotic activity and enhanced plant productivity with higher nutrient accumulation in the growing season.     

Net nitrogen mineralization and leaching in response to warming and nitrogen deposition in a temperate old field: the importance of winter temperature

While climate warming can increase plant N availability over the growing season by increasing rates of N mineralization, increased N mineralization over winter at a time when plant roots are largely inactive, coupled with an increased frequency of soil freeze–thaw cycles, may increase soil N leaching losses. We examined changes in soil net N mineralization and N leaching in response to warming and N addition (6 g m^?2 year^?1) in a factorial experiment conducted in a temperate old field. We used two warming treatments, year-round and winter-only warming, to isolate the effects of winter warming on soil N dynamics from the year-round warming effects. We estimated net N mineralization using in situ soil cores with resin bags placed at the bottom to catch throughput, and we measured N leaching using lysimeters located below the plant rooting zone at a depth of 50 cm. There were minor effects of warming on changes in soil extractable N and resin N in the soil cores over winter. Nevertheless, the overall effects of both warming and N addition on net N mineralization (the sum of changes in soil extractable N and resin N) were not significant over this period. Likewise, there were no significant treatment effects on the concentration of N in leachate collected below the plant rooting zone. However, in response to winter warming, net N mineralization over summer was approximately double that of both the ambient and year-round warming treatments. This result demonstrates a potentially large and unexpected effect of winter warming on soil N availability in this old field system.     

Interactive effects of soil warming and fertilization on root production, mortality, and longevity in a Norway spruce stand in Northern Sweden

The effects of soil warming and nitrogen availability on root production, longevity and mortality were studied using minirhizotrons in irrigation (C), fertilized (F), heated (H), and heated‐fertilized (HF) plots in a Norway spruce stand in northern Sweden from October 1996 to October 1997. Irrigation was included in all treatment plots. Heating cables were used to maintain the soil temperature in heated plots at 5°C above that in unheated plots during the growing season. A Kaplan–Meier approach was used to estimate the longevity of fine roots and Cox proportional hazards regression to analyze the effects of the H, F, and HF treatments on the risk of root mortality. The proportion of annual root length production contributed by winter–spring production amounted to 52% and 49% in heated plots and heated‐fertilized plots, respectively. The annual root length production in C plots was significantly higher than in other treatments, while the HF treatment gave significantly greater production compared with the F treatment. The risk of mortality (hazard ratio) relative to C plots was higher in H plots (358%) and F plots (191%). The interaction between heating and fertilizing was strongly significant. The increase in the risk of root mortality in combined fertilization and heating (103%) was lower than that in the H or F plots. The results show that nitrogen addition combined with warmer temperatures decreases the risk of root mortality, and fine root production is a function of the length of the growing season. In the future, fertilization combined with the warmer temperatures expected to follow predicted climatic change may increase root production in boreal forests at low fertility sites.     

Impact of winter climate change on nitrogen biogeochemistry in forest ecosystems: A synthesis from Japanese case studies

Nitrogen (N) is a critical ecological and environmental indicator under changing environments. The impact of winter climate change on N biogeochemical processes in forest ecosystems has gained increasing recognition. Decreasing snowfall has caused a decrease in the heat insulation properties of the snowpack, resulting in an increase in the frequency and magnitude of freezing and thawing cycles in surface soil, where biological processes are most active. Here I synthesize recent research findings from integrated field observations and experiments conducted in northern Japan and compare these results with previous research outcomes from other regions to identify current research gaps and develop the next research agenda to further advance our understanding of this complex problem. Japanese case studies indicated that net ammonium production (ammonification) was mostly dominant in terms of available soil N fertility in cold environments and was sensitive to the increase in soil freezing and thawing cycles because of the decreased snowpack. On the other hands, nitrate dynamics were more stable or conservative than those of ammonium. The soil characteristics (i.e., N pool and microbial activities) were significant explanatory factors of the responses of soil N dynamics and N leakage among different soils to increased freezing–thawing cycles at watershed and national scale. This synthesis indicates that winter climate change had significant impacts on soil N biogeochemistry (such as soil N pool size and microbial N transformation) during the winter and snowmelt season and also during the following growing season. Several research gaps and possible research topics (path dependency and soil microbial community composition) are also presented by synthesizing the current research findings. Further field experiments and observations quantifying the pools and fluxes of inorganic N with modeling analysis under freeze–thaw environments would contribute to increase the understandings of N transformation processes under winter climate change.     

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 affects growing‐season soil microbial biomass and activity in northern hardwood forests

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.     

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.     

Effects of lengthened growing season and soil warming on the phenology and physiology of Polygonum bistorta

The phenological and physiological responses of arctic tundra plant species are key to predicting their survival in a warmer climate. One of the consequences of a warmer climate in the Arctic will be a longer growing season. We examined the effects of lengthened growing season and soil warming on the widely distributed forb, Polygonum bistorta L. Three treatments were established near Toolik Lake, Alaska in 1995 and 1996: extended season, extended season with soil warming, and an unmanipulated control. The season was extended by removing the snow load in the spring and keeping the treatments free of snow in the autumn. The spring snow removal extended the snow‐free period over that of controls by 8 d in 1995 and 24 d in 1996. As a result, the number of accumulated soil thaw days and consequently the depth of soil thaw increased on the treatment plots. Polygonum bistorta responded to the treatments by becoming active earlier and senescing earlier, resulting in a growth period of similar duration to that of the controls. Leaf size and leaf number were unaffected by the treatments, as were leaf photosynthetic assimilation rates and nutrient concentrations. The results indicate that internal constraints limit the response of this species to lengthened growing season, suggesting that it is a determinant or periodic species. With climate warming, this periodic growth will put P. bistorta at a competitive disadvantage relative to plants that can respond to lengthened growing season.     

Soil Respiration along Environmental Gradients in Olympic National Park

Although mountainous landscapes dominate large areas of the Earth, our understanding of how elevation and aspect influence soil respiration in complex mountainous terrain is very limited. Therefore, we measured soil respiration throughout the growing season in 1999 and 2000 at 11 forested sites in Olympic National Park, Washington, USA along elevation-climatic gradients. The study sites ranged from temperate rain forest to alpine forests near tree line. Soil temperature was a significant predictor of soil respiration at all sites, and soil moisture explained additional variability at three sites (R2 from 0.42 to 0.90, P ≤ 0.01). Soil temperatures at the highest-elevation sites were 4.5°C cooler than those at the lowest elevation, but there were no relationships between soil respiration rates at a given temperature and elevation or mean annual temperature that would indicate acclimation of soil respiration to the cooler temperatures at high-elevation sites. Experimental urea additions (1.0 and 2.0 g N m-2 y-1) made at seven of the sites had no consistent effect on soil respiration. Total soil carbon dioxide (CO2) efflux during the growing season (May-September) varied from 0.34 to 0.75 kg C/m2 and was greater at low-elevation sites with warmer soil temperatures and longer growing seasons. Elevation and the length of the frost-free season could both be used to predict growing season (r2 = 0.53) and annual (r2 = 0.81) soil CO2 efflux for the 10 sites located in steep mountainous terrain. Significant correlations also existed with mean annual temperature. These results suggest that warmer soils and a longer snow-free season associated with climatic warming could cause the mountainous ecosystems of the Olympic peninsula to evolve increasing amounts of CO2 from all elevations and aspects.     

Snow melt stimulates ecosystem respiration in Arctic ecosystems

Cold seasons in Arctic ecosystems are increasingly important to the annual carbon balance of these vulnerable ecosystems. Arctic winters are largely harsh and inaccessible leading historic data gaps during that time. Until recently, cold seasons have been assumed to have negligible impacts on the annual carbon balance but as data coverage increases and the Arctic warms, the cold season has been shown to account for over half of annual methane (CH₄) emissions and can offset summer photosynthetic carbon dioxide (CO₂) uptake. Freeze–thaw cycle dynamics play a critical role in controlling cold season CO₂ and CH₄ loss, but the relationship has not been extensively studied. Here, we analyze freeze–thaw processes through in situ CO₂ and CH₄ fluxes in conjunction with soil cores for physical structure and porewater samples for redox biogeochemistry. We find a movement of water toward freezing fronts in soil cores, leaving air spaces in soils, which allows for rapid infiltration of oxygen‐rich snow melt in spring as shown by oxidized iron in porewater. The snow melt period coincides with rising ecosystem respiration and can offset up to 41% of the summer CO₂ uptake. Our study highlights this important seasonal process and shows spring greenhouse gas emissions are largely due to production from respiration instead of only bursts of stored gases. Further warming is projected to result in increases of snowpack and deeper thaws, which could increase this ecosystem respiration dominate snow melt period causing larger greenhouse gas losses during spring.     

Frost damage and winter nitrogen uptake by the grass Poa pratensis L.: consequences for vegetative versus reproductive growth

Frost damage can decrease nitrogen uptake by grasses over winter, and it can also decrease biomass production over the following growing season. However, it is not clear to what extent reduced nitrogen uptake over winter decreases grass production, or whether is it merely a symptom of root damage. We examined the growth response of the grass Poa pratensis L. (Kentucky bluegrass) to variation in the timing of freezing and nitrogen availability over winter in London, Ontario, Canada. All tillers were transplanted into untreated soil in early spring, and at peak seed maturation, root, shoot, and reproductive biomass were measured. There was an interaction between freezing and increased winter nitrogen availability, whereby nitrogen addition increased tiller biomass under ambient temperatures, but decreased tiller biomass in combination with a late winter freeze. The nitrogen response of ambient temperature tillers occurred primarily via increased seed production, whereas for frozen tillers seed production was generally absent. Our results support the hypothesis that nitrogen uptake over winter can increase growing season productivity in P. pratensis, but also demonstrate that increased nitrogen availability increases tiller vulnerability to frost. These results have important implications for grass responses to the alteration of soil freezing dynamics with climate change.