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Previously-frozen stores of organic carbon (C) are now subject to decomposition due to a warming Arctic climate and associated permafrost thaw; however, estimates of the amount of greenhouse gases (GHG) that may be released are not well constrained. Knowing more about the functions of the extant permafrost microbial community will inform this knowledge gap. The exploration of microbial functional traits may be useful to elucidate the relationship between microbial diversity and ecosystem function. We characterized the community traits and functional diversity of the bacterial and Archaeal component of the microbial community from three depths of permafrost, as well as the organic and mineral horizons of the seasonally-thawed active layer, by assessing 'substrate-use richness,' 'substrate preference,' 'growth rate,' 'and substrate specific growth rate.' We measured the microbial community response to 31 substrates with an EcoPlate (Biolog, Inc.) assay at three incubation temperatures (1, 10, and 20 °C) using a kinetic approach, and modeled the microbial response to each substrate with a modified logistic growth function. We hypothesized that the permafrost communities would be selected for high functional potential and activity at cold temperatures. Rather, we found that the permafrost community did not have a higher functional diversity or activity at 1 °C than the organic active layer soils. In addition, permafrost communities increased their growth rates with increasing temperature, indicating that the highest incubation temperature (20 °C) was below their temperature optimum for growth. As predicted, the permafrost communities did exhibit temperature dependent substrate preferences. Thus, permafrost microbial communities did not appear to be selected for higher metabolism and the ability to use a broad suite of substrates at low temperatures, which suggests that they may have limited function immediately following thaw when temperatures are near 0 °C. However, changes in community composition or additional permafrost warming will increase the functional capabilities of permafrost microbes to decompose the C stored in those soils. [ABSTRACT FROM AUTHOR]

While much is known about the factors that control each component of the terrestrial nitrogen (N) cycle, it is less clear how these factors affect total N availability, the sum of organic and inorganic forms potentially available to microorganisms and plants. This is particularly true for N-poor ecosystems such as drylands, which are highly sensitive to climate change and desertification processes that can lead to the loss of soil nutrients such as N. We evaluated how different climatic, abiotic, plant and nutrient related factors correlate with N availability in semiarid Stipa tenacissima grasslands along a broad aridity gradient from Spain to Tunisia. Aridity had the strongest relationship with N availability, suggesting the importance of abiotic controls on the N cycle in drylands. Aridity appeared to modulate the effects of pH, plant cover and organic C (OC) on N availability. Our results suggest that N transformation rates, which are largely driven by variations in soil moisture, are not the direct drivers of N availability in the studied grasslands. Rather, the strong relationship between aridity and N availability could be driven by indirect effects that operate over long time scales (decades to millennia), including both biotic (e.g. plant cover) and abiotic (e.g. soil OC and pH). If these factors are in fact more important than short-term effects of precipitation on N transformation rates, then we might expect to observe a lagged decrease in N availability in response to increasing aridity. Nevertheless, our results suggest that the increase in aridity predicted with ongoing climate change will reduce N availability in the Mediterranean basin, impacting plant nutrient uptake and net primary production in semiarid grasslands throughout this region. [ABSTRACT FROM AUTHOR]

Abstract: Recent evidence suggests that soil organic matter (SOM) is largely composed of microbial products rather than plant compounds that resist decomposition. The chemical transformation of leaf litter components during decomposition is critical in controlling SOM formation. Plant leaf litter tends to decompose faster in its native environment than when it is placed under other vegetation types. This home-field advantage (HFA) suggests that decomposer communities are specialized to most efficiently degrade the litter found in their native environment, possibly through the production of specific enzymes that degrade unique compounds within that litter. Could this affect the degree to which leaf litter chemistry is altered during decomposition? We used pyrolysis-molecular beam mass spectrometry (py-MBMS) to analyze whether the chemistry of aspen and lodgepole pine litter was altered to a greater degree when decomposed in its home environment compared to an away environment. We had previously reported a 4% HFA for pine litter decomposition rates in this reciprocal experiment, and attributed that effect to differences in decomposer communities. Our high-resolution analysis revealed that litter chemistry also changed to a greater extent in its home environment. The changes in litter chemistry were more pronounced for the more recalcitrant pine litter, suggesting that decomposer community specialization is more important for recalcitrant litter. The accumulation of microbial products and microbially-transformed plant components resulted in an overall convergence in litter chemistry as decomposition proceeded, but the imprints of both initial litter chemistry and decomposer communities remained evident. The detection of new compounds in decomposed litter and the HFA effect on litter chemistry suggest that decomposer communities affect both the rate at which individual compounds within litter are decomposed, and the chemical nature of compounds that are incorporated into SOM. [Copyright &y& Elsevier]

Soil microbial communities may be able to rapidly respond to changing environments in ways that change community structure and functioning, which could affect climate-carbon feedbacks. However, detecting microbial feedbacks to elevated CO2 (eCO2) or warming is hampered by concurrent changes in substrate availability and plant responses. Whether microbial communities can persistently feed back to climate change is still unknown. We overcame this problem by collecting microbial inocula at subfreezing conditions under eCO2 and warming treatments in a semi-arid grassland field experiment. The inoculant was incubated in a sterilised soil medium at constant conditions for 30 days. Microbes from eCO2 exhibited an increased ability to decompose soil organic matter (SOM) compared with those from ambient CO2 plots, and microbes from warmed plots exhibited increased thermal sensitivity for respiration. Microbes from the combined eCO2 and warming plots had consistently enhanced microbial decomposition activity and thermal sensitivity. These persistent positive feedbacks of soil microbial communities to eCO2 and warming may therefore stimulate soil C loss. [ABSTRACT FROM AUTHOR]

As the earth system changes in response to human activities, a critical objective is to predict how biogeochemical process rates (e.g. nitrification, decomposition) and ecosystem function (e.g. net ecosystem productivity) will change under future conditions. A particular challenge is that the microbial communities that drive many of these processes are capable of adapting to environmental change in ways that alter ecosystem functioning. Despite evidence that microbes can adapt to temperature, precipitation regimes, and redox fluctuations, microbial communities are typically not optimally adapted to their local environment. For example, temperature optima for growth and enzyme activity are often greater than in situ temperatures in their environment. Here we discuss fundamental constraints on microbial adaptation and suggest specific environments where microbial adaptation to climate change (or lack thereof) is most likely to alter ecosystem functioning. Our framework is based on two principal assumptions. First, there are fundamental ecological trade-offs in microbial community traits that occur across environmental gradients (in time and space). These trade-offs result in shifting of microbial function (e.g. ability to take up resources at low temperature) in response to adaptation of another trait (e.g. limiting maintenance respiration at high temperature). Second, the mechanism and level of microbial community adaptation to changing environmental parameters is a function of the potential rate of change in community composition relative to the rate of environmental change. Together, this framework provides a basis for developing testable predictions about how the rate and degree of microbial adaptation to climate change will alter biogeochemical processes in aquatic and terrestrial ecosystems across the planet. [ABSTRACT FROM AUTHOR]

Climate models project that precipitation patterns will likely intensify in the future, resulting in increased duration of droughts and increased frequency of large soil rewetting events, which are stressful to the microorganisms that drive soil biogeochemical cycling. Historical conditions can affect contemporary microbial responses to environmental factors through the persistence of abiotic changes or through the selection of a more tolerant microbial community. We examined how a history of intensified rainfall would alter microbial functional response to drying and rewetting events, whether this historical legacy was mediated through altered microbial community composition, and how long community and functional legacies persisted under similar conditions. We collected soils from a long-term field manipulation (Rainfall Manipulation Plot Study) in Kansas, USA, where rainfall variability was experimentally amplified. We measured respiration, microbial biomass, fungal:bacterial ratios and bacterial community composition after collecting soils from the field experiment, and after subjecting them to a series of drying-rewetting pulses in the lab. Although rainfall history affected respiration and microbial biomass, the differences between field treatments did not persist throughout our 115-day drying-rewetting incubation. However, soils accustomed to more extreme rainfall did change less in response to lab moisture pulses. In contrast, bacterial community composition did not differ between rainfall manipulation treatments, but became more dissimilar in response to drying-rewetting pulses depending on their previous field conditions. Our results suggest that environmental history can affect contemporary rates of biogeochemical processes both through changes in abiotic drivers and through changes in microbial community structure. However, the extremity of the disturbance and the mechanism through which historical legacies occur may influence how long they persist, which determines the importance of these effects for biogeochemical cycling. [ABSTRACT FROM AUTHOR]

Microbial decomposition of soil organic matter produces a major flux of CO2 from terrestrial ecosystems and can act as a feedback to climate change. Although climate-carbon models suggest that warming will accelerate the release of CO2 from soils, the magnitude of this feedback is uncertain, mostly due to uncertainty in the temperature sensitivity of soil organic matter decomposition. We examined how warming and altered precipitation affected the rate and temperature sensitivity of heterotrophic respiration ( Rh) at the Boston-Area Climate Experiment, in Massachusetts, USA. We measured Rh inside deep collars that excluded plant roots and litter inputs. In this mesic ecosystem, Rh responded strongly to precipitation. Drought reduced Rh, both annually and during the growing season. Warming increased Rh only in early spring. During the summer, when Rh was highest, we found evidence of threshold, hysteretic responses to soil moisture: Rh decreased sharply when volumetric soil moisture dropped below ~15% or exceeded ~26%, but Rh increased more gradually when soil moisture rose from the lower threshold. The effect of climate treatments on the temperature sensitivity of Rh depended on the season. Apparent Q10 decreased with high warming (~3.5 °C) in spring and fall. Presumably due to limiting soil moisture, warming and precipitation treatments did not affect apparent Q10 in summer. Drought decreased apparent Q10 in fall compared to ambient and wet precipitation treatments. To our knowledge, this is the first field study to examine the response of Rh and its temperature sensitivity to the combined effects of warming and altered precipitation. Our results highlight the complex responses of Rh to soil moisture, and to our knowledge identify for the first time the seasonal variation in the temperature sensitivity of microbial respiration in the field. We emphasize the importance of adequately simulating responses such as these when modeling trajectories of soil carbon stocks under climate change scenarios. [ABSTRACT FROM AUTHOR]

The response of soil organic matter ( OM) decomposition to increasing temperature is a critical aspect of ecosystem responses to global change. The impacts of climate warming on decomposition dynamics have not been resolved due to apparently contradictory results from field and lab experiments, most of which has focused on labile carbon with short turnover times. But the majority of total soil carbon stocks are comprised of organic carbon with turnover times of decades to centuries. Understanding the response of these carbon pools to climate change is essential for forecasting longer-term changes in soil carbon storage. Herein, we briefly synthesize information from recent studies that have been conducted using a wide variety of approaches. In our effort to understand research to-date, we derive a new conceptual model that explicitly identifies the processes controlling soil OM availability for decomposition and allows a more explicit description of the factors regulating OM decomposition under different circumstances. It explicitly defines resistance of soil OM to decomposition as being due either to its chemical conformation ( quality) or its physico-chemical protection from decomposition. The former is embodied in the depolymerization process, the latter by adsorption/desorption and aggregate turnover. We hypothesize a strong role for variation in temperature sensitivity as a function of reaction rates for both. We conclude that important advances in understanding the temperature response of the processes that control substrate availability, depolymerization, microbial efficiency, and enzyme production will be needed to predict the fate of soil carbon stocks in a warmer world. [ABSTRACT FROM AUTHOR]

Abstract: Arctic soil microorganisms remain active at ecologically relevant rates in frozen soils. We used bromodeoxyuridine (BrdU) labeling and terminal restriction fragment length polymorphism (T-RFLP) analysis of 16S rRNA gene amplicons to examine active bacterial communities in two Alaskan tundra soils collected in summer and winter of 2005. Active community T-RFLP profiles were compared to total community profiles to determine if active bacteria were a subset of the total community. In shrub soils, active bacteria communities differed in composition between summer and winter, and winter-active bacterial taxa were not detected in the total community, suggesting that they are likely rare within the overall community. In contrast, tussock tundra soil contained more bacterial taxa that were active in both summer and winter and also represented a large portion of the total community. Using in silico digest of a sequence library from this site, we attempted to identify the dominant organisms in our samples. Our previous research suggested that the total microbial community was stable throughout the year, but this new study suggests that the active community is more dynamic seasonally. In general, only a subset of the total community was growing at a given time. This temporal niche partitioning may contribute to the high diversity of microbial communities in soils. Understanding which taxa contribute to microbial function under different conditions is the next frontier in microbial ecology and linking composition to biogeochemical cycling. [ABSTRACT FROM AUTHOR]

The biogeochemical cycles of carbon (C), nitrogen (N), and phosphorus (P) are fundamental to life on Earth. Because organisms require these elements in strict proportions, the cycles of C, N, and P are coupled at molecu- lar to global scales through their effects on the biochemical reactions controlling primary production, respira- tion, and decomposition. The coupling of the C, N, and P cycles constrains organismal responses to climatic and atmospheric change, suggesting that present-day estimates of climate warming through the year 2100 are conservative. N and P supplies constrain C uptake in the terrestrial biosphere, yet these constraints are often not incorporated into global-scale analyses of Earth's climate. The inclusion of coupled biogeochemical cycles is critical to the development of next-generation, global-scale climate models. [ABSTRACT FROM AUTHOR]

Altered surface ultraviolet-B (UV-B) radiation resulting from a combination of factors that include changes in stratospheric ozone concentrations, cloud cover, and aerosol conditions may affect litter decomposition and, thus, terrestrial nutrient cycling on a global scale. Although litter decomposition rates vary across biomes, patterns of decomposition suggest that UV-B radiation accelerates litter decay in xeric environments where precipitation is infrequent. However, under more frequent precipitation regimes where litter decay rates are characteristically high, the effect of UV-B radiation on litter decomposition has not been fully elucidated. To evaluate this association between moisture regime and UV-B exposure, a litter decomposition experiment was designed for aspen ( Populus tremuloides) leaf litter, where conditions that influence both abiotic (photodegradation) and biotic (microbial) processes could be manipulated quantitatively. We found that experimentally increasing UV-B exposure (0, 7.4, and 11.2 kJ m−2 day−1, respectively) did not consistently increase litter decomposition rates across simulated precipitation frequencies of 4, 12, and 24 days. Instead, a UV-B exposure of 11.2 kJ m−2 day−1 resulted in a 13% decrease in decomposition rates under the 4-day precipitation frequency, but an increase of 80% under the 24-day frequency. Furthermore, the same UV-B dose increased litter decomposition rates under the 24-day precipitation frequency by 78% even in conditions where microbial activity was suppressed. Therefore, under more xeric conditions, greater exposure to UV-B radiation increased decomposition rates, presumably through photodegradation. In contrast, when decomposition was not moisture-limited, greater UV-B exposure slowed decomposition rates, most likely from the resulting inhibition of microbial activity. Ultimately, these experimental results highlight UV-B radiation as a potential driver of decomposition, as well as indicate that both the direction and magnitude of the UV-B effect is dependent on moisture availability, a factor that may change according to future patterns in global precipitation. [ABSTRACT FROM AUTHOR]

Soil microbes are among the most abundant and diverse organisms on Earth. Although microbial decomposers, particularly fungi, are important mediators of global carbon and nutrient cycling, the functional roles of specific taxa within natural environments remain unclear. We used a nucleotide-analog technique in soils from the Harvard Forest to characterize the fungal taxa that responded to the addition of five different carbon substrates—glycine, sucrose, cellulose, lignin, and tannin-protein. We show that fungal community structure and richness shift in response to different carbon sources, and we demonstrate that particular fungal taxa target different organic compounds within soil microcosms. Specifically, we identified eleven taxa that exhibited changes in relative abundances across substrate treatments. These results imply that niche partitioning through specialized resource use may be an important mechanism by which soil microbial diversity is maintained in ecosystems. Consequently, high microbial diversity may be necessary to sustain ecosystem processes and stability under global change. [ABSTRACT FROM AUTHOR]

Extracellular enzymes are the proximate agents of organic matter decomposition and measures of these activities can be used as indicators of microbial nutrient demand. We conducted a global-scale meta-analysis of the seven-most widely measured soil enzyme activities, using data from 40 ecosystems. The activities of β-1,4-glucosidase, cellobiohydrolase, β-1,4- N-acetylglucosaminidase and phosphatase g−1 soil increased with organic matter concentration; leucine aminopeptidase, phenol oxidase and peroxidase activities showed no relationship. All activities were significantly related to soil pH. Specific activities, i.e. activity g−1 soil organic matter, also varied in relation to soil pH for all enzymes. Relationships with mean annual temperature (MAT) and precipitation (MAP) were generally weak. For hydrolases, ratios of specific C, N and P acquisition activities converged on 1 : 1 : 1 but across ecosystems, the ratio of C : P acquisition was inversely related to MAP and MAT while the ratio of C : N acquisition increased with MAP. Oxidative activities were more variable than hydrolytic activities and increased with soil pH. Our analyses indicate that the enzymatic potential for hydrolyzing the labile components of soil organic matter is tied to substrate availability, soil pH and the stoichiometry of microbial nutrient demand. The enzymatic potential for oxidizing the recalcitrant fractions of soil organic material, which is a proximate control on soil organic matter accumulation, is most strongly related to soil pH. These trends provide insight into the biogeochemical processes that create global patterns in ecological stoichiometry and organic matter storage. [ABSTRACT FROM AUTHOR]

Microorganisms have a variety of evolutionary adaptations and physiological acclimation mechanisms that allow them to survive and remain active in the face of environmental stress. Physiological responses to stress have costs at the organismal level that can result in altered ecosystem-level C, energy, and nutrient flows. These large-scale impacts result from direct effects on active microbes' physiology and by controlling the composition of the active microbial community. We first consider some general aspects of how microbes experience environmental stresses and how they respond to them. We then discuss the impacts of two important ecosystem-level stressors, drought and freezing, on microbial physiology and community composition. Even when microbial community response to stress is limited, the physiological costs imposed on soil microbes are large enough that they may cause large shifts in the allocation and fate of C and N. For example, for microbes to synthesize the osmolytes they need to survive a single drought episode they may consume up to 5% of total annual net primary production in grassland ecosystems, while acclimating to freezing conditions switches Arctic tundra soils from immobilizing N during the growing season to mineralizing it during the winter. We suggest that more effectively integrating microbial ecology into ecosystem ecology will require a more complete integration of microbial physiological ecology, population biology, and process ecology. [ABSTRACT FROM AUTHOR]

Fungal and bacterial community structure in tussock, intertussock and shrub organic and mineral soils at Toolik Lake, Alaska were evaluated. Community structure was examined by constructing clone libraries of partial 16S and 18S rRNA genes. The soil communities were sampled at the end of the growing season in August 2004 and just after the soils thawed in June 2005. The communities differed greatly between vegetation types, although tussock and intertussock soil communities were very similar at the phyla level. The communities were relatively stable between sample dates at the phyla and subphyla levels, but differed significantly at finer phylogenetic scales. Tussock and intertussock bacterial communities were dominated by Acidobacteria, while shrub soils were dominated by Proteobacteria. These results appear consistent with previous work demonstrating that shrub soils contain an active, bioavailable C fraction, while tussock soils are dominated by more recalcitrant substrates. Tussock fungi communities had higher proportions of Ascomycota than shrub soils, while Zygomycota were more abundant in shrub soils. Recent documentation of increasing shrub abundance in the Arctic suggests that soil microbial communities and their functioning are likely to be altered by climate change. [ABSTRACT FROM AUTHOR]