Repeated freeze–thaw cycling can reduce the microbial
biomass by as much as one-third (Larsen et al. 2002,
Pesaro et al. 2003), and basal respiration by up to 30%
(Schimel and Clein 1996). Damage is usually greater
when microbes are active and growing (Schimel and
Clein 1996). When soils experience multiple freeze–thaw
cycles, most of the vulnerable biomass appears to be lost
in the first cycle and microbes that survive the first cycle
are resistant to additional events (Larsen et al. 2002,
Walker et al. 2006). Such results suggest that different
groups of microbes have different growth strategies
when it comes to responding to freezing. In contrast to
drought, one might hypothesize that bacteria are more
tolerant than fungi because the mycelial growth form
should be sensitive to the physical disruption associated
with freezing. Multiple freeze/thaw cycles appear to shift
communities toward increased bacterial dominance
(Nieminen and Setala 2001, Larsen et al. 2002) as
bacteria survive while eukaryote and archaeal populations
are reduced (Pesaro et al. 2003). In contrast to
these studies, the gram negative Pseudomonas/Alcaligenes
group appears to be sensitive to freeze/thaw
(Skogland et al. 1988), although Pseudomonas paucimobilis
was tolerant of a first freeze/thaw cycle but only
sensitive to a second (Morley et al. 1983), suggesting it
may have consumed the resources needed for acclimation
in surviving the first freeze cycle. Although fungi are
vulnerable to freezing, they sometimes increase under
prolonged frozen conditions (Schadt et al. 2003, Sjursen
et al. 2005). Since fungi are generally tolerant of low
water potentials and are able to bridge gaps to tap
spatially separated resources, this wouldn’t be surprising.
There is little information about how low temperatures
affect microbial community structure in nature.
The one relatively complete picture of microbial winter
dynamics is from the Colorado alpine where the
microbial community cycles between a summertime,
bacteria-dominated community and a winter, fungidominated
community (Schadt et al. 2003); this community
shift produces a flush of nutrients that support
springtime plant growth (Schmidt and Lipson 2004).
However, in the alpine, with thick snow cover, soil
temperatures are never extremely low and the shifts in
microbial communities are thought to be driven more by
changes in available substrates—root exudates and
other fresh, simple compounds in the summer, and
more complex and recalcitrant litter-derived materials in
the winter (Schmidt and Lipson 2004, Schmidt et al.
2007). Although this work has been cited as characterizing
‘‘tundra’’ (Neufeld and Mohn 2005), arctic and
alpine tundra differ in many ways and it is quite possible
that arctic communities have different patterns of
microbial community dynamics.
Whereas surviving drought appears to be physiologically
expensive, microbes appear generally able to do
so. Freezing appears to be a greater challenge, possibly
involving more actual mortality, although many of the
direct physiological costs of surviving dry and freezing
conditions appear similar in magnitude. The ecophysiological
costs of both, and their effects on functional
community dynamics, need to be better constrained at
the whole-ecosystem scale.