Changes to soil freezing dynamics with climate change can modify ecosystem carbon and nutrient losses. Soil freezing is influenced strongly by both air temperature and insulation by the snowpack, and ...it has been hypothesized that winter climate warming may lead to increased soil freezing as a result of reduced snowpack thickness. I used weather station data to explore the relationships between winter air temperature, precipitation and soil freezing for 31 sites in Canada, ranging from the temperate zone to the high Arctic. Inter-annual climate variation and associated soil temperature variation over the last 40 years were examined and used to interpolate the effects of projected climate change on soil freezing dynamics within sites using linear regression models. Annual soil freezing days declined with increasing mean winter air temperature despite decreases in snow depth and cover, and reduced precipitation only increased annual soil freezing days in the warmest sites. Annual soil freeze-thaw cycles increased in both warm and dry winters, although the effects of precipitation were strongest in sites that experience low mean winter precipitation. Overall, it was projected that by 2050, changes in winter temperature will have a much stronger effect on annual soil freezing days and freeze-thaw cycles than changes in total precipitation, with sites close to but below freezing experiencing the largest changes in soil freezing days. These results reveal that experimental data relevant to the effects of climate changes on soil freezing dynamics and changes in associated soil physical and biological processes are lacking.
ABSTRACT
Winter is a key driver of individual performance, community composition, and ecological interactions in terrestrial habitats. Although climate change research tends to focus on performance ...in the growing season, climate change is also modifying winter conditions rapidly. Changes to winter temperatures, the variability of winter conditions, and winter snow cover can interact to induce cold injury, alter energy and water balance, advance or retard phenology, and modify community interactions. Species vary in their susceptibility to these winter drivers, hampering efforts to predict biological responses to climate change. Existing frameworks for predicting the impacts of climate change do not incorporate the complexity of organismal responses to winter. Here, we synthesise organismal responses to winter climate change, and use this synthesis to build a framework to predict exposure and sensitivity to negative impacts. This framework can be used to estimate the vulnerability of species to winter climate change. We describe the importance of relationships between winter conditions and performance during the growing season in determining fitness, and demonstrate how summer and winter processes are linked. Incorporating winter into current models will require concerted effort from theoreticians and empiricists, and the expansion of current growing‐season studies to incorporate winter.
Soil microbial responses to climate warming in temperate regions may interact with the effects of increased atmospheric N deposition. In addition, the combined effects of these factors on microbial ...activity during the plant growing season may differ from the effects over winter, when reduced plant soil C inputs and soil freezing can alter microbial nutrient availability and demand. We examined seasonal changes in soil extracellular enzyme activity (EEA), microbial biomass C and N, and soil fungal and bacterial content in a warming and N addition experiment in a temperate old field. For EEA, we examined both hydrolases (organic C degrading enzymes, a chitinase and phosphatase) and ligninases (phenol oxidase and peroxidase). While both hydrolase and ligninase activities exhibited significant seasonal variation, EEA was unresponsive to the experimental treatments. Microbial biomass C increased with warming year round, however, and microbial biomass N increased with N addition but only over summer. Despite increased microbial biomass in response to warming, phosphatase was the only enzyme that exhibited a significant change in specific activity (enzyme activity per unit of microbial biomass) in response to warming. Likewise, soil fungal and bacterial biomass varied seasonally, but treatment effects on these variables were minimal. Overall, while the effects of N addition on microbial N varied seasonally, microbial responses were relatively insensitive to the warming and N addition treatments in our experiment. This insensitivity was unexpected given the large treatment effects on plant productivity and soil N dynamics documented during the same time frame in the field experiment.
Although freeze–thaw cycles can alter soil physical properties and microbial activity, their overall impact on soil functioning remains unclear. This review addresses the effects of freeze–thaw ...cycles on soil physical properties, microorganisms, carbon and nutrient dynamics, trace gas losses and higher organisms associated with soil. I discuss how the controlled manipulation of freeze–thaw cycles has varied widely among studies and propose that, despite their value in demonstrating the mechanisms of freeze–thaw action in soils, many studies of soil freeze–thaw cycles have used cycle amplitudes, freezing rates and minimum temperatures that are not relevant to temperature changes across much of the soil profile in situ. The lack of coordination between the timing of soil collection and the season for which freeze–thaw cycles are being simulated is also discussed. Suggested improvements to future studies of soil freeze–thaw cycles include the maintenance of realistic temperature fluctuations across the soil profile, soil collection in the appropriate season and the inclusion of relevant surface factors such as plant litter in the fall or excess water in the spring. The implications of climate change for soil freeze–thaw cycles are addressed, along with the need to directly assess how changes in soil freeze–thaw cycle dynamics alter primary production.
Assays for extracellular enzyme activity (EEA) have become a common tool for studying soil microbial responses in climate change experiments. Nevertheless, measures of potential EEA, which are ...conducted under controlled conditions, often do not account for the direct effects of climate change on EEA that occur as a result of the temperature and moisture dependence of enzyme activity in situ. Likewise, the indirect effects of climate on EEA in the field, that occur via effects on microbial enzyme producers, must be assessed in the context of potential changes in plant and soil faunal communities. Here, EEA responses to warming and altered precipitation in field studies are reviewed, with the goal of evaluating the role of EEA in enhancing our understanding of soil and ecosystem responses to climate change. Seasonal and interannual variation in EEA responses to climate change treatments are examined, and potential interactions with elevated atmospheric CO2, increased atmospheric N deposition and changes in disturbance regimes are also explored. It is demonstrated that in general, soil moisture manipulations in field studies have had a much greater influence on potential EEA than warming treatments. However, these results may simply reflect the low magnitude of soil warming achieved in many field experiments. In addition, changes in plant species composition over the longer term in response to warming could strongly affect EEA. Future challenges involve extending studies of potential EEA to address EEA responses to climate change in situ, and gaining further insights into the mechanisms, such as enzyme production, stabilization and turnover, that underlie EEA responses.
► Climate change can have a strong influence on soil EEA, both directly and indirectly. ► In general, soil moisture manipulations in field studies have had a much greater influence on potential EEA than warming treatments. ► Future challenges involve addressing EEA responses to climate change in situ. ► Further insights into mechanisms underlying potential EEA responses (i.e. turnover) are required.
Increased atmospheric nitrogen (N) deposition and climate warming are both anticipated to influence the N dynamics of northern temperate ecosystems substantially over the next century. In field ...experiments with N addition and warming treatments, cumulative treatment effects can be important for explaining variation in treatment effects on N dynamics over time; however, comparisons between data collected in the early vs. later years potentially can be confounded with interactions between treatment effects and inter-annual variation in environmental conditions or other factors. We compared the short-term versus long-term effects of N addition and warming on net N mineralization and N leaching in a grass-dominated old field using in situ soil cores. We added new N addition and warming plots (3 years old) to an existing field experiment (16 years old), which enabled comparison of the treatment effects at both time scales while controlling for potential inter-annual variation in other factors. For net N mineralization, there was a significant interaction between plot age and N addition over the growing season, and for extractable inorganic N there was a significant interaction between plot age and warming over winter. In both cases, the directions of the treatment effects differed among old and new plots. Moreover, the responses in the new plots differed from the responses observed previously when the 16-year-old plots had been new. These results demonstrate how inter-annual variation in responses, independent from cumulative treatment effects, can play an important role in interpreting long-term effects on soil N cycling in global change field experiments.
There is limited understanding of the combined effects of discrete climate extremes and chronic environmental changes on ecosystem processes and functioning. We assessed the interactions of extreme ...drought timing (45 days, in spring or summer) and nitrogen (N) addition in a full factorial field experiment in a Leymus chinensis-dominated meadow steppe in northeast China. We evaluated the resistance and recovery of the grassland (calculated in terms of aboveground biomass) to these two drought events. The spring drought reduced aboveground biomass by 28% in the unfertilized plots and by 33% in the fertilized plots, and the effects persisted during the subsequent post-drought period within the same growing season; however, the summer drought had no significant influence on aboveground biomass. Although there were no significant interactive effects between drought timing and N addition, we observed a potential trend of N addition increasing the proportion of aboveground biomass suppressed by spring drought but not summer drought. Moreover, the drought resistance of the aboveground biomass was positively correlated with the response of the belowground biomass to drought. One year after the extreme drought events, the spring drought effects on aboveground and belowground biomass were negligible. Our results indicate that the drought sensitivity of productivity likely depends on the phenological and morphological traits of the single highly dominant species (Leymus chinensis) in this meadow steppe.
In annual grasslands that experience a mediterranean-type climate, the synchrony between plant senescence and peak solar radiation over summer results in high litter sun exposure. We examined the ...decomposition of both shaded and sun-exposed litter over summer and inferred the effects of photodegradation from changes in mass loss and litter chemistry. The carry-over effects of summer litter exposure on wet season decomposition were also assessed, and the attenuation of photodegradation with litter layer thickness was used to estimate the proportion of grass litter lignin susceptible to photodegradation under different treatments of a factorial global change experiment. Over summer, mass loss from grass and forb litter exposed to ambient sunlight ranged from 8% to 10%, whereas lignin decreased in grass litter by approximately 20%. After one year of decomposition, mass losses from grass leaves exposed to sunlight over summer were more than double the mass losses from summer-shaded leaves. When shade litter layer thickness was varied, mass losses over summer for all treatments were also approximately 8%; however, lignin decreased significantly only in the low shade treatments (0-64 g m⁻² of shade litter). Aboveground production of annual grasses nearly quadrupled in response to the combined effects of N addition, elevated atmospheric CO₂, increased precipitation and warming. The estimated proportion of grass litter lignin experiencing full photodegradation ranged from 100% under ambient conditions to 31-62% in plots receiving the combined global change treatments. These results reveal an important role of sun exposure over summer in accelerating litter decomposition in these grasslands and provide evidence that future changes in the quantity of litter deposition may modulate the influence of photodegradation integrated across the litter layer.
Background and aims
Climate warming and atmospheric nitrogen deposition are both expected to exert strong influences on soil organic matter in northern temperate ecosystems over the next century, but ...it is unclear to what extent these effects may be revealed by short-term field experiments. We compared the short-term (1–2 year) versus long-term (14–15 year) effects of nitrogen and warming on soil organic matter, while controlling for the effects of interannual environmental variability.
Methods
We added the new nitrogen addition and warming plots (short-term) to a pre-existing nitrogen and warming field experiment (long-term) in a grass-dominated field. We used soil density fractionation and size fractionation to separate the soil organic matter into the fine free light fraction, the coarse free light fraction and the occluded light fraction. We analyzed the quality of each soil organic matter fraction using Fourier-transform infrared (FTIR) spectroscopy.
Results
While the free light fraction decreased by 14% with nitrogen addition in the short-term plots, contrary to our prediction, it increased by 12% in the long-term plots. Likewise, in the long-term plots, the occluded light fraction increased by 58% with nitrogen addition when combined with warming. Organic matter quality generally did not differ among treatments for the different density and size fractions.
Conclusion
Our results demonstrate a divergence in the directions of long-term nitrogen addition and warming responses of soil organic matter fractions from those of short-term responses.
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