Winter is an important period for ecological processes in northern regions; however, compared to other seasons, the impacts of winter climate on ecosystems are poorly understood. In this review we evaluate the influence of winter climate on carbon dynamics based on the current state of knowledge and highlight emerging topics and future research challenges. Studies that have addressed this topic include plot-scale snow cover manipulation experiments that alter soil temperatures, empirical investigations along natural climatic gradients, laboratory temperature incubation experiments aimed at isolating influential factors in controlled environments, and time series of climate and carbon data that evaluate long-term natural variation and trends. Combined, these studies have demonstrated how winter climate can influence carbon in complex ways that in some cases are consistent across studies and in other cases are difficult to predict. Despite advances in our understanding, there is a great need for studies that further explore: (i) carry-over effects from one season to another, (ii) ecosystem processes in the fall-winter and winter-spring shoulder seasons, (iii) the impacts of extreme events, (iv) novel experimental approaches, and (v) improvements to models to include ecological effects of winter climate. We also call for the establishment of an international winter climate change research network that enhances collaboration and coordination among studies, which could provide a more thorough understanding of how the snow-covered period influences carbon cycling, thereby improving our ability to predict future responses to climate change.

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 (2mm pore size) or excluded root ingrowth (50m pore size) for up to 29months 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.

Short-term N2O emission occurs in relation to snowmelt within seasonally frozen soil. To understand the effects of changing winter climates on the N2O flux, snow cover manipulation experiments are useful. In Japan, snow cover manipulation is practiced by farmers to improve agricultural yield and is executed either by applying a broadcast of blackish agent onto the snow cover, which leads to faster snow-melting thereby extending the crop-growing season, or by snow cover removal/re-accumulation, leading to an enhanced soil frost depth for weed management. Implementation of these practices involves using an amount of fossil fuel, in addition to influencing soil-derived N2O emissions, therefore, the load factors of snow cover management practices per unit area of agricultural field were estimated in this study. Field data including micrometeorological conditions, ground surface flux of N2O, and amount of fossil fuel consumed during machinery operation for management practices, were obtained at two sites in Hokkaido over 2 years (2008-2010). Fuel consumption for the field spreading was found to be unexpectedly small (0.017 Mg CO2 eq ha(-1)). It was therefore suggested that acceleration of snowmelt may have the potential to reduce net greenhouse gas emissions if the agent used is a low-degradable C-rich material, such as charcoal. For soil frost control, the fossil fuel consumption during a set of snow cover removal/re-accumulation (estimated as 0.052 Mg CO2 eq ha(-1)) is discussed, together with the relationship between possible mechanisms causing stimulation of N2O production in frozen soil and inherent large differences in N2O flux among sites.

Decomposition is a key process in carbon and nutrient cycling. However, little is known about its response to altered winter soil temperature regimes in boreal forests. Here, the impact of soil frost on cellulose decomposition over 1 yr and soil biotic activity (bait-lamina sticks) over winter, in spring, and in summer was investigated using a long-term (9-yr) snow-cover manipulation experiment in a boreal Picea abies forest. The experiment consisted of the treatments: snow removal, increased insulation, and ambient control. The snow removal treatment caused longer and deeper soil frost (minimum temperature - 8.6 degrees C versus - 1.4 degrees C) at 10 cm soil depth in comparison with control, while the increased insulation treatment resulted in nearly no soil frost during winter. Annual cellulose decomposition rates were reduced by 46% in the snow removal manipulation in comparison with control conditions. Increased insulation had no significant effect on decomposition. The decomposition was mainly driven by microorganisms, as no significant difference was observed for containers enclosed with a 44-mu m and a 1-mm mesh. Soil biotic activity was slightly increased by both the snow removal and the increased insulation treatment in comparison with control conditions over winter. However, this effect disappeared over spring and summer. We conclude that soil frost can have strong effects on decomposition in boreal ecosystems. Further studies should investigate to which degree the observed reduction in decomposition due to reduced snow cover in winter slows or even offsets the expected increase in decomposition rates with global warming.

Snow regimes affect biogeochemistry of boreal ecosystems and are altered by climate change. The effects on plant communities, however, are largely unexplored despite their influence on relevant processes. Here, the impact of snow cover on understory community composition and below-ground production in a boreal Picea abies forest was investigated using a long-term (8-year) snow cover manipulation experiment consisting of the treatments: snow removal, increased insulation (styrofoam pellets), and control. The snow removal treatment caused longer (118 vs. 57 days) and deeper soil frost (mean minimum temperature -5.5 vs. -2.2 degrees C) at 10 cm soil depth in comparison to control. Understory species composition was strongly altered by the snow cover manipulations; vegetation cover declined by more than 50% in the snow removal treatment. In particular, the dominant dwarf shrub Vaccinium myrtillus (-82%) and the most abundant mosses Pleurozium schreberi (-74%) and Dicranum scoparium (-60%) declined strongly. The C:N ratio in V. myrtillus leaves and plant available N in the soil indicated no altered nitrogen nutrition. Fine-root biomass in summer, however, was negatively affected by the reduced snow cover (-50%). Observed effects are attributed to direct frost damage of roots and/or shoots. Besides the obvious relevance of winter processes on plant ecology and distribution, we propose that shifts in the vegetation caused by frost damage may be an important driver of the reported alterations in biogeochemistry in response to altered snow cover. Understory plant performance clearly needs to be considered in the biogeochemistry of boreal systems in the face of climate change.

Climate change may cause a decrease in snow cover in northern latitudes. This, on the other hand, may result in more severe soil frost even in areas where it is not common at present, and may lead to increased stress on the tree canopy. We studied the effects of snow removal and consequent changes in soil frost and water content on the physiology of Norway spruce (Picea abies [L] Karst.) needles and implications on root biomass. The study was conducted at a 47-year-old Norway spruce stand in eastern Finland during the two winters of 2005/06 and 2006/07. The treatments in three replicates were: (i) natural snow accumulation and melting (CTRL), (ii) artificial snow removal during the winter (OPEN), and (id) the same as OPEN, but the ground was insulated in early spring to delay soil thawing (FROST). In spite of the deeper soil frost in the OPEN than in the CTRL treatment, soil warming in spring occurred at the same time, whereas soil warming in the FROST was delayed by 2 and 1.5 months in 2006 and 2007, respectively. The soil water content was affected by snow manipulations, being at a lower level in the OPEN and FROST than CTRL in spring and early summer. The physiological measurements of the needles (e.g. starch, carbon and nitrogen content and apoplastic electrical resistance) showed differences between soil frost treatments. The differences were mostly seen between the CTRL and FROST, but also in the case of the starch content in early spring 2007 between the CTRL and OPEN. The needle responses in the FROST were more evident after the colder winter of 2006. The physiological changes seemed to be related to the soil temperature and water content in the early growing season rather than to the wintertime soil temperature. No difference was found in the fine root (diameter <2 mm) biomass between the treatments assessed in 2007. In the future, conditions similar to the OPEN treatment may be more common than at present in areas experiencing a thick snow cover. The present experiment took place over the course of two years. It is possible that whenever thin snow cover occurs yearly, the reduced starch content during the early spring may be reflected in the tree growth itself as a result of reduced energy reserves. (C) 2011 Elsevier B.V. All rights reserved.

Background: Winter conditions are changing considerably due to climate change. Resulting alterations in the frequency of soil freeze-thaw cycles (FTCs) are ecologically important. Aim: We quantified the impact of winter soil-warming pulses on the community structure of temperate plant communities. Methods: The cover of vascular plant species in two vegetation types, each at three diversity levels, was recorded 1 year before to 3 years after an FTC-manipulation that added five additional FTCs. Changes in species abundance patterns (Bray-Curtis similarity) were analysed by linear mixed effect models. Results: Communities exposed to additional FTCs showed less change in their species abundance patterns than the reference plots. Community development in the grassland differed between the FTC-manipulation and the reference plots in the first growing season after the FTC-manipulation, but such effects disappeared over time, whereas the divergence from the reference plots in the dwarf-shrub heath started in the second year after the FTC-manipulation and effects grew over time. Responses to FTCs were related to growth forms: some grasses increased after the FTC-manipulation, whereas the cover of dwarf shrubs was reduced. There was less change in species abundance distributions in the more diverse communities with legumes present. Conclusions: Winter climate change is a critical driver of temperate ecosystems. Short-term climatic events can have long-term implications on the structure of ecosystems. Community composition regulates alterations in the development and competitive balance of plant communities caused by soil warming pulses.

This study considered how climate change affects the accumulation of snow, the soil moisture and soil frost at sites without tree cover in boreal conditions in Finland (60 degrees-70 degrees N). An increase of 4.5 degrees C in annual mean temperature and 20% in annual precipitation were assumed for Finland by the year 2100 according to A2 emission scenario. Along with climate, the soil type of the permanent inventory plots of the Finnish National Forest Inventory was used. Soil and climate data were combined by using a process-based ecosystem model. Calculations were done for four periods: current climate (1971-2000), near future (2001-2020), mid-term future (2021-2050) and long-term future (2071-2100). According to our simulations, the average monthly duration and depth of snow decreased over the simulation period. However, the increasing precipitation may locally increase the snow depths in the mid-term calculations. In the autumn and winter, the average volumetric soil moisture content slightly increased in southern Finland during the near future, but decreased towards the end of the century, but still remained on a higher level than presently. In northern Finland, the soil moisture in the autumn and winter increased by the end of this century. In the summertime soil moisture decreased slightly regardless of the region. Throughout Finland, the length and the depth of soil frost decreased by the end of the century. In the south, the reduction in the depth was largest in the autumn and spring, while in the mid-winter it remained relatively deep in the middle of the century. In the north, the depth tended to increase during the first two calculation periods, in sonic areas, even during the third calculation period (2071-2100) due to reduced insulation effects of snow during cold spells. The wintertime increase in soil moisture and reduced soil frost may be reflected to reduced carrying capacity of soil for timber harvesting.