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Winter microbial carbon metabolism and community composition in Alaskan tundra soil
http://udini.proquest.com/view/winter-microbial-carbon-metabolism-goid:230770144/
C. Microbes in Cold Environments
In 1975, Morita proposed that psychrophilic organisms (bacteria, in parti cular) be defined as those that have a optimal growth temperature below 15°C, a maximal growth temperature of about 20°C and are capable of growth at or below 0°C. He suggested that any organism that was capable of growth in near freezing temperatures but had growth optima above 15°C should be referred to a psychrotrophs. However, due to the arbitrary nature of the growth temperature cutoffs, the strict definition is often ignored and the term psychrophile applied to any organism living in extreme cold envi ronments (Feller and Gerday, 2003) .
Sub - zero environmen ts, including sea ice, deep - sea water and permafrost, have been the target of many studies that are attempting to define the lower thermal range for microbial life. So far, most organisms isolated from extreme cold environments are bacteria, but a few psyc hrophilic yeasts have also been isolated. However, it is not clear whether these organisms are simply surviving or if they are actually able to grow at temperatures below freezing. Bacteria isolated from permafrost are able to grow very slowly below 0°C in pure culture (Vishnivetskaya et al., 2000; Panikov and Sizova, 2007) , but only a few studies have measured in situ growth in permafrost (Rivkina et al., 2000; Gilichinsky et al., 2003; Rivkina et al., 2004) . However, permafrost and arctic tundra soil differ in their temperature regimes; where permafrost is continually frozen, putting intense pressure on microbes to adapt to life in frozen conditions if they are to remain acti ve, tundra soil cycles between frozen and thawed states annually, releasing the pressure to adapt if only a microbe
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can survive. Thus, it is unclear whether winter - active microbial communities in tundra soil are specially adapted to frozen conditions or wh ether they are simply a hardy subset of the summer community. I hypothesized that the winter microbial community would be different from the summer community both functionally and genetically. In Chapter 3 , I use BrdU - labelling coupled with T - RFLP analysis
to compare the active and bulk bacterial communities at a genetic level. Chapters 4 & 5 address the functional aspects of winter vs. summer microbial communities.
Though few studies have directly measured winter - active microbial communities in Arctic tund ra, extensive studies have been conducted on microbial communities in frozen alpine tundra soil in Colorado (Lipson and Monson, 1998; Lipson et al., 1999, 2000; Lipson et al., 2002; Schadt et al., 2003; Schmidt and L ipson, 2004; Lipson, 2007) . The alpine differs from the Arctic in that it receives both more snow and more sun throughout the winter. However, vegetation types and nutrient cycling patterns are similar to those of Arctic tundra, making the Colorado alpine a useful ecosystem for comparison. One of the more interesting aspects of microbial community composition in the alpine is that bacteria dominate during the summer and fungi during the winter (Lipson et al., 2002; S chadt et al., 2003) . One might hypothesize that fungi would be less suited to winter conditions because of their filamentous nature. As described earlier, microbial activity in frozen soil takes place in micron - thin water films. Fungi may have difficulty fitting into the water films and thus accessing nutrients, etc. On the other hand, fungi may be able to bridge water films in a way a single - celled bacteria cannot; by maintaining channels of
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unfrozen fluid through their hyphae, fungi can translocate nutri ents from one area to another without having to wait for diffusion (Panikov and Sizova, 2007) .
Though I expected to find that microbial communities would vary between seasons, we had already shown that between summer 2004 and spring 2005, microbial communities in tussock and shrub organic soil remained fairly constant at the phyla level. In fact, native vegetation type proved to be a much stronger influence than season, as tussock and shrub communities were quite different from each other.