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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]

Arctic soils contain large amounts of organic matter due to very slow rates of detritus decomposition. The first step in decomposition results from the activity of extracellular enzymes produced by soil microbes. We hypothesized that potential enzyme activities are low relative to the large stocks of organic matter in Arctic tundra soils, and that enzyme activity is low at in situ temperatures. We measured the potential activity of six hydrolytic enzymes at 4 and 20 °C on four sampling dates in tussock, intertussock, shrub organic, and shrub mineral soils at Toolik Lake, Alaska. Potential activities of N-acetyl glucosaminidase, β-glucosidase, and peptidase tended to be greatest at the end of winter, suggesting that microbes produced enzymes while soils were frozen. In general, enzyme activities did not increase during the Arctic summer, suggesting that enzyme production is N-limited during the period when temperatures would otherwise drive higher enzyme activity in situ. We also detected seasonal variations in the temperature sensitivity ( Q10) of soil enzymes. In general, soil enzyme pools were more sensitive to temperature at the end of the winter than during the summer. We modeled potential in situβ-glucosidase activities for tussock and shrub organic soils based on measured enzyme activities, temperature sensitivities, and daily soil temperature data. Modeled in situ enzyme activity in tussock soils increased briefly during the spring, then declined through the summer. In shrub soils, modeled enzyme activities increased through the spring thaw into early August, and then declined through the late summer and into winter. Overall, temperature is the strongest factor driving low in situ enzyme activities in the Arctic. However, enzyme activity was low during the summer, possibly due to N-limitation of enzyme production, which would constrain enzyme activity during the brief period when temperatures would otherwise drive higher rates of decomposition. [ABSTRACT FROM AUTHOR]

There is some evidence that microbes inhabiting permanently frozen environments are capable of cell maintenance and growth under in situ conditions. In contrast, Arctic tundra surface soils that are only frozen during the winter have exhibited CO2 respiration under frozen conditions. This could result from maintenance metabolism fuelled by recycling cell material, but does necessarily indicate growth, which requires the synthesis of new cell material. We used 13C-glucose to trace C into membrane lipids and 5-bromo-3-deoxyuridine to monitor DNA synthesis in a laboratory microcosm study on Arctic tundra soils. Organisms were not equally active: fungi incorporated more C than any other group. Although some bacteria were growing, Gram(+) bacteria were almost completely inactive. Shrub tundra microbes were more active in early winter than tussock microbes, incorporating more C and synthesizing more DNA. In late winter, the C incorporation pattern reversed, although DNA production was similar. We demonstrated for the first time that microbes in frozen tundra soils synthesized new cell membranes and DNA, processes fundamentally associated with growth. That microbes can grow during Arctic winters may have important implications for C cycle modelling under a changing climate. [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]

Rapid climate change in the Arctic is altering microbial structure and function, with important consequences for the global ecosystem. Emerging evidence suggests organisms in higher trophic levels may also influence microbial communities, but whether warming alters these effects is unclear. Wolf spiders are dominant Arctic predators whose densities are expected to increase with warming. These predators have temperature-dependent effects on decomposition via their consumption of fungal-feeding detritivores, suggesting they may indirectly affect the microbial structure as well. To address this, we used a fully factorial mesocosm experiment to test the effects of wolf spider density and warming on litter microbial structure in Arctic tundra. We deployed replicate litter bags at the surface and belowground in the organic soil profile and analyzed the litter for bacterial and fungal community structure, mass loss, and nutrient characteristics after 2 and 14 months. We found there were significant interactive effects of wolf spider density and warming on fungal but not bacterial communities. Specifically, higher wolf spider densities caused greater fungal diversity under ambient temperature but lower fungal diversity under warming at the soil surface. We also observed interactive treatment effects on fungal composition belowground. Wolf spider density influenced surface bacterial composition, but the effects did not change with warming. These findings suggest a widespread predator can have indirect, cascading effects on litter microbes and that effects on fungi specifically shift under future expected levels of warming. Overall, our study highlights that trophic interactions may play important, albeit overlooked, roles in driving microbial responses to warming in Arctic terrestrial ecosystems., Importance: The Arctic contains nearly half of the global pool of soil organic carbon and is one of the fastest warming regions on the planet. Accelerated decomposition of soil organic carbon due to warming could cause positive feedbacks to climate change through increased greenhouse gas emissions; thus, changes in ecological dynamics in this region are of global relevance. Microbial structure is an important driver of decomposition and is affected by both abiotic and biotic conditions. Yet how activities of soil-dwelling organisms in higher trophic levels influence microbial structure and function is unclear. In this study, we demonstrate that predicted changes in abundances of a dominant predator and warming interactively affect the structure of litter-dwelling fungal communities in the Arctic. These findings suggest predators may have widespread, indirect cascading effects on microbial communities, which could influence ecosystem responses to future climate change., Competing Interests: The authors declare no conflict of interest.

Molecular ecology is poised to tackle a host of interesting questions in the coming years. The Arctic provides a unique and rapidly changing environment with a suite of emerging research needs that can be addressed through genetics and genomics. Here we highlight recent research on boreal and tundra ecosystems and put forth a series of questions related to plant and microbial responses to climate change that can benefit from technologies and analytical approaches contained within the molecular ecologist's toolbox. These questions include understanding (i) the mechanisms of plant acquisition and uptake of N in cold soils, (ii) how these processes are mediated by root traits, (iii) the role played by the plant microbiome in cycling C and nutrients within high-latitude ecosystems and (iv) plant adaptation to extreme Arctic climates. We highlight how contributions can be made in these areas through studies that target model and nonmodel organisms and emphasize that the sequencing of the Populus and Salix genomes provides a valuable resource for scientific discoveries related to the plant microbiome and plant adaptation in the Arctic. Moreover, there exists an exciting role to play in model development, including incorporating genetic and evolutionary knowledge into ecosystem and Earth System Models. In this regard, the molecular ecologist provides a valuable perspective on plant genetics as a driver for community biodiversity, and how ecological and evolutionary forces govern community dynamics in a rapidly changing climate., (© 2015 John Wiley & Sons Ltd.)

Global climate change is already having significant impacts on arctic and alpine ecosystems, and ongoing increases in temperature and altered precipitation patterns will affect the strong seasonal patterns that characterize these temperature-limited systems. The length of the potential growing season in these tundra environments is increasing due to warmer temperatures and earlier spring snow melt. Here, we compare current and projected climate and ecological data from 20 Northern Hemisphere sites to identify how seasonal changes in the physical environment due to climate change will alter the seasonality of arctic and alpine ecosystems. We find that although arctic and alpine ecosystems appear similar under historical climate conditions, climate change will lead to divergent responses, particularly in the spring and fall shoulder seasons. As seasonality changes in the Arctic, plants will advance the timing of spring phenological events, which could increase plant nutrient uptake, production, and ecosystem carbon (C) gain. In alpine regions, photoperiod will constrain spring plant phenology, limiting the extent to which the growing season can lengthen, especially if decreased water availability from earlier snow melt and warmer summer temperatures lead to earlier senescence. The result could be a shorter growing season with decreased production and increased nutrient loss. These contrasting alpine and arctic ecosystem responses will have cascading effects on ecosystems, affecting community structure, biotic interactions, and biogeochemistry., (© 2014 John Wiley & Sons Ltd.)

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.