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. (Conant et al., 2011)

Soil enzymatic activities and microbial biomass carbon (Cmic) are considered to be
two important soil biological activities influenced by oil contamination occurring in the soil
ecosystem. This study focused on changes in the soil microbial community enzymatic
activities as a result of the potential inhibitory effects of hydrocarbon contamination. The
relationship between hydrocarbons (kerosene and diesel), Cmic and enzymatic activity
(dehydrogenase and phosphatase) was evaluated in three amended soil types collected from
different areas (Fresh Boyndie, Insch and Brechin) in Aberdeenshire (UK). Results showed
that hydrocarbon contamination inhibited enzymatic activities in all the amended soil
samples. The extent of inhibition increased significantly with increasing levels of
hydrocarbons, and varied with the incubation period. Insch soil had high Cmic values and
high numbers of heterotrophic bacteria CFU, but it had the lowest dehydrogenase and
phosphatase activities of all three soils. Brechin soils had the highest phosphatase activity.
Results also showed that both Insch and Brechin soils had similar numbers of culturable
hydrocarbon degrader bacteria across all soil treatments with the exception of kerosene
treatments, while Brechin soils had the highest culturable numbers of hydrocarbon
degrading fungi across all three soil treatments with the exception of incubated control and
kerosene treatments. There were generally strong positive relationships in non-treated
samples between bacterial heterotrophs and hydrocarbon degrading bacteria in all three
soils. Both incubated Insch and Brechin soil treatments exhibited a strong correlation
between fungal heterotrophs and hydrocarbon degraders. However, non-incubated Insch
and Brechin soils had a weak relationship between fungal heterotrophs and degrading
fungi. Hydrocarbons in soils provide a source of carbon for microbial growth and this helps
to explain the high variation in fungal data between soils which may be associated with
different microbial communities in each soil. (Alrrunman et al., 2015)

Precipitation directly alters N cycling via its impact on soil water availability, erosion,
and leaching, and indirectly alters N cycling by influencing plant N uptake as well as plant
productivity. Increases in precipitation, specifically in large rain events, might lead to large
leaching and runoff events (Nearing et al. 2005), which could increase N loss and decrease
N retention. For example, nitrate is readily leached down the soil profile in arid ecosystems,
which could lead to N accumulation beneath the reach of plant roots (Jobaggy et al. 2001,
Walvoord et al. 2003). Changes in the distribution of rainfall can lead to large rain events
(Easterling et al. 2000). These large events and their associated runoff can redistribute N
among tree types and from the intercrown space to intercrown vegetated areas leading to
the loss of N beneath tree canopies (Wilcox 1994, Reid et al. 1999). In water-limited
ecosystems, N uptake from the soil is limited by water availability, thus when precipitation
and soil water availability increase, plant N uptake from the soil and overall plant
productivity also increase leading to a decline in soil N availability (Stark and Firestone
1995, Austin 2002, Knapp et al. 2008). Similarly, N can accumulate during periods of
drought. When soils are dry, decreased plant transpiration results in decreased N uptake by
roots and increased N accumulation in soils (Stark and Firestone 1995, Weltzin et al. 2003)
(Cregger et al., 2014)
 Increased precipitation generally increases microbial biomass and fungal:bacterial
ratio. Few studies report responses to reduced precipitation but the effects likely counter
those of increased precipitation. Altered precipitation regimes have also been found to alter
microbial community composition but broader generalizations are difficult to make.
Changes in event size and frequency influences invertebrate activity and density with
cascading impacts on the soil food web, which will likely impact carbon and nutrient pools.
The long-term implications for biogeochemical cycling are inconclusive but several studies
suggest that increased aridity may cause decoupling of carbon and nutrient cycling.
(Nielsen et al., 2015)

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. (Suseela et al., 2012)

The temperature sensitivity of soil organic matter (SOM) decomposition has been a
crucial topic in global change research, yet remains highly uncertain. One of the
contributing factors to this uncertainty is the lack of understanding about the role of
rhizosphere priming effect (RPE) in shaping the temperature sensitivity. Using a novel
continuous 13C-labeling method, we investigated the temperature sensitivity of RPE and its
impact on the temperature sensitivity of SOM decomposition.We observed an overall
positive RPE. The SOM decomposition rates in the planted treatments increased 17–163%
above the unplanted treatments in three growth chamber experiments including two plant
species, two growth stages, and two warming methods. More importantly, warming by 5
1C increased RPE up to threefold, hence, the overall temperature sensitivity of SOM
decomposition was consistently enhanced (Q10 values increased 0.3–0.9) by the presence
of active rhizosphere. In addition, the proportional contribution of SOM decomposition to
total soil respiration was increased by soil warming, implying a higher temperature
sensitivity of SOM decomposition than that of autotrophic respiration. Our results, for the
first time, clearly demonstrated that root–soil interactions play a crucial role in shaping the
temperature sensitivity of SOM decomposition. Caution is required for interpretation of any
previously determined temperature sensitivity of SOM decomposition that omitted
rhizosphere effects using either soil incubation or field root-exclusion. More attention
should be paid to RPE in future experimental and modeling studies of SOM
decomposition.(Zhu & Cheng, 2011)

Sudden changes in soil moisture, such as those resulting from drought and subsequent
rain events, can induce a physiological stress on bacteria due to the rapid change in water
potential (Harris 1981). Some bacteria may be able to tolerate these stresses because they
possess certain adaptive traits, like a thicker cell wall to withstand osmotic pressure (Kieft
et al. 1987; Schimel et al. 2007). However, tolerating stress also requires an investment in
resources (Schimel et al. 2007) such that taxa investing in this stress-tolerance strategy
should have reduced relative fitness (including as a result of competitive interactions) when
stress events are rare. (Evans & Wallenstein, 2014)
 Our objective was to assess the effects of long-term continuous grazing on soil enzyme
activities in relation to shifts in plant litter attributes and soil resources in an arid
ecosystem, considering both spatial and temporal variations. Grazing led to reduced grass
cover, decreasing plant litter mass with increasing soluble phenolics, and reduced
phosphatases, ß-glucosidase and microbial biomass-C at PCP. A localized nutrient input
from animal excreta seems to promote microbial biomass-C, alkaline phosphatase and
dehydrogenase activities but only at IC from the site with high grazing intensity. Plant
heterogeneous distribution, plant litter quantity and quality, nutrient inputs from grazers
and seasonal variation in soil moisture, also affecting soil resources and microbial biomass,
modulate soil enzyme responses to long-term grazing in the arid Patagonian Monte (Olivera
et al., 2014)

Grasslands comprise 85% of Southern Patagonia land area and play a critical role in
the global carbon cycle. We evaluated seasonal dynamics to identify differences in soil
respiration rates between contrasting grasslands across a climate gradient (rainfall), long
term grazing intensity (moderate and high stocking rates) and land uses (silvopastoral
system, primary forest and grassland). Soil respiration varied from 0.09 g CO2 h_1 m_2 in
winter to a maximum of 1.43 g CO2 h_1 m_2 in spring. We found that the soil respiration
rate was 30% higher in moderately grazed grasslands than in heavily grazed grasslands.
Landuse changes showed that soil respiration followed the order silvopastoral system >
native forest > grassland. While almost all plant and soil variables had a significant effect
on soil respiration, soil carbon concentration, litter cover and depth and bare soil cover
were the main factors explaining 78 e 83% of the variance in soil respiration. Soil
respiration rates were correlated strongly to air and soil temperatures and to a lesser extent
with mean monthly rainfall and soil volumetric water content. The information provided in
the present work about soil respiration is essential to estimate carbon balance for a range of
important and widespread ecosystems in Patagonia (Peri et al., 2015)

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 1C on four sampling dates in tussock, intertussock, shrub organic, and
shrub mineral soils at Toolik Lake, Alaska. Potential activities of N-acetyl
glucosaminidase, b-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 situ b-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. (Wallentein et al., 2009)

Previous studies that have been performed at different scales have indicated that
enzyme distribution in space is highly heterogeneous. This review summarizes the current
knowledge regarding the extent of this spatial variability and the factors that contribute to
its establishment. The distribution of enzymes is spatially heterogeneous at various scales.
Spatial autocorrelation has been recorded for the activity of hydrolytic and oxidative
enzymes in the range of 10s of centimeters to a scale of meters when considering areas
ranging from square meters to hectares; however, enzyme molecules are unevenly
distributed even across areas of several centimeters, exhibiting spatial autocorrelation in the
range of 10s of millimeters. At these scales, patches of nutrients and microbial biomass,
plant roots, or colonies of specific microorganisms seem to induce high enzyme activity. In
such areas, the activity of multiple enzymes is often increased. At larger scales, variation in
microbial biomass, effect of individual trees, and actual soil moisture may shape enzyme
distribution. Additionally, when studying a range of several square kilometers, the
dominant vegetation, land use type, and soil physicochemical properties affect enzyme
distribution. Because these factors change with time (soil moisture content, in particular),
the distribution of soil enzymes and decomposition rates are probably highly dynamic.
(Baldrian, 2014)

Researchers agree that climate change factors such as rising atmospheric [CO2] and
warming will likely interact to modify ecosystem properties and processes. However, the
response of the microbial communities that regulate ecosystem processes is less
predictable. We measured the direct and interactive effects of climatic change on soil
fungal and bacterial communities (abundance and composition) in a multifactor climate
change experiment that exposed a constructed old-field ecosystem to different atmospheric
CO2 concentration (ambient,_300 ppm), temperature (ambient, _3°C), and precipitation
(wet and dry) might interact to alter soil bacterial and fungal abundance and community
structure in an old-field ecosystem. We found that (i) fungal abundance increased in
warmed treatments; (ii) bacterial abundance increased in warmed plots with elevated
atmospheric [CO2] but decreased in warmed plots under ambient atmospheric [CO2]; (iii)
the phylogenetic distribution of bacterial and fungal clones and their relative abundance
varied among treatments, as indicated by changes in 16S rRNA and 28S rRNA genes; (iv)
changes in precipitation altered the relative abundance of Proteobacteria and
Acidobacteria, where Acidobacteria decreased with a concomitant increase in the
Proteobacteria in wet relative to dry treatments; and (v) changes in precipitation altered
fungal community composition, primarily through lineage specific changes within a
recently discovered group known as soil clone group I. Taken together, our results indicate
that climate change drivers and their interactions may cause changes in bacterial and fungal
overall abundance; however, changes in precipitation tended to have a much greater effect
on the community composition. These results illustrate the potential for complex
community changes in terrestrial ecosystems under climate change scenarios that alter
multiple factors simultaneously (Castro et al., 2010)
 stimulate microbial decomposition of soil carbon, producing a positive feedback to
rising global temperatures1,2. Although field experiments document an initial increase in
the loss of CO2 from soils in response to warming, in line with these predictions, the
carbon dioxide loss from soils tends to decline to control levels within a few years3–5. This
attenuation response could result from changes in microbial physiological properties with
increasing temperature, such as a decline in the fraction of assimilated carbon that is
allocated to growth, termed carbon-use efficiency6. Here we explore these mechanisms
using a microbial-enzyme model to simulate the responses of soil carbon to warming by 5
_C. We find that declines in microbial biomass and degradative enzymes can explain the
observed attenuation of soil-carbon emissions in response to warming. Specifically,
reduced carbon-use efficiency limits the biomass of microbial decomposers and mitigates
the loss of soil carbon. However, microbial adaptation or a change in microbial
communities could lead to an upward adjustment of the efficiency of carbon use,
counteracting the decline in microbial biomass and accelerating soil-carbon loss. We
conclude that the soil-carbon response to climate warming depends on the efficiency of soil
microbes in using carbon (Allison et al., 2010).

The Arrhenius equation was used to determine the activation energy (Ea) and the
temperature coefficient (Q10) for all enzymes. The values of Ea and Q10 for each enzyme
differed among soils, although in general the differences were small, especially for those
enzymes that act on substrates of low molecular weight. In terms of the values of Ea and
Q10 and the differences established among soils, the results obtained for those enzymes
that act on substrates of high molecular weight differed most from those corresponding to
the other enzymes. Thus the lowest Ea and Q10 values corresponded to BAA-protease, and
the highest values to CM-cellulase and casein-protease. Except for catalase in one of the
soils, the values of Ea and Q10 for the oxidoreductases were similar to those of most of the
hydrolases. In general, the effect of temperature appeared to be more dependent on the type
of enzyme than on the characteristics of the soil. Determination of the activity of an enzyme
at different temperatures not only provides the optimal range of temperature for enzyme
activity, but also allows estimation of thermodynamic parameters, e.g. the activation energy
(Ea). The Ea for enzyme-catalyzed reactions is lower than for reactions not catalyzed by
enzymes, because enzymes act by lowering the energy barrier that must be surmounted
before the reaction can take place (Juma and Tabatabai, 1988). Determination of enzyme
activity at different temperatures also allows the value of the temperature coefficient, or
Q10, to be obtained. This coefficient indicates the increase in the rate of reaction for
increases in temperature of 10 1C. Enzyme-catalyzed reactions are less sensitive to
temperature changes than their uncatalyzed counterparts. Thus, although the uncatalyzed
reaction rate may double for every increase of 10 1C, the enzyme catalyzed reaction rate
generally increases by a factor of less than two (Tabatabai, 1982). The value of Q10 is not
constant, and varies for different enzymes depending on the activation energy and on the
temperature at which it is obtained (Lehninger, 1978). There are many reports of the effects
of incubation temperature on the activity of hydrolytic enzymes, although most of these
studies have involved finding out the optimum temperature for activity (Geller and
Ginzburg, 1979; Kanazawa and Miyashita, 1986; Trevors, 1984a; Kandeler, 1990; Schinner
and von Mersi, 1990; Nannipieri et al., 1991). Furthermore, most published studies on the
effect of the incubation temperature on the activity of soil enzymes refer to hydrolytic
enzymes, and very few are concerned with oxidoreductase enzymes such as dehydrogenase
or catalase. In any case, studies on thermodynamic parameters are very scarce and in fact,
for many soil enzymes no studies of their thermodynamic parameters have been carried out.
This is surprising considering the importance of these parameters, which are intrinsic
characteristics of soil enzymes under any particular set of environmental conditions.
(Trasar- Cepeda, 2007)

Decomposition of soil organic matter (SOM) is mediated by microbial extracellular

hydrolytic enzymes (EHEs). Thus, given the large amount of carbon (C) stored as SOM, it
is imperative to understand how microbial EHEs will respond to global change (and
warming in particular) to better predict the links between SOM and the global C cycle.

Here, we measured the Michaelis–Menten kinetics [maximal rate of velocity (Vmax) and

half-saturation constant (Km)] of five hydrolytic enzymes involved in SOM degradation

(cellobiohydrolase, b-glucosidase, b-xylosidase, a-glucosidase, and N-acetyl-b-D-
glucosaminidase) in five sites spanning a boreal forest to a tropical rainforest. We tested the
specific hypothesis that enzymes from higher latitudes would show greater temperature
sensitivities than those from lower latitudes. We then used our data to parameterize a
mathematical model to test the relative roles of Vmax and Km temperature sensitivities in
SOM decomposition. We found that both Vmax and Km were temperature sensitive, with
Q10 values ranging from 1.53 to 2.27 for Vmax and 0.90 to 1.57 for Km. The Q10 values
for the Km of the cellulose-degrading enzyme b-glucosidase showed a significant (P =
0.004) negative relationship with mean annual temperature, indicating that enzymes from
cooler climates can indeed be more sensitive to temperature. Our model showed that Km
temperature sensitivity can offset SOM losses due to Vmax temperature sensitivity, but the
offset depends on the size of the SOM pool and the magnitude of Vmax. Overall, our
results suggest that there is a local adaptation of microbial EHE kinetics to temperature and
that this should be taken into account when making predictions about the responses of C

cycling to global change. the Michaelis–Menten constant (Km) increases more strongly

with increasing temperature in cold-adapted enzymes than in warmadapted enzymes

(Hochachka & Somero, 2002; Johns & Somero, 2004; Somero, 2004; Dong & Somero,
2009; Huestis et al., 2009). Km is the substrate concentration at half-maximal enzymatic
velocity (Vmax), and is indicative of the affinity an enzyme has for its substrate (German et
al., 2011b). Therefore, an increase in Km indicates a decrease in overall enzyme function.
In fact, an increase in Km can counteract an increase in enzyme Vmax under warming
conditions, thereby reducing the temperature sensitivity of decomposition in soils
(Davidson et al., 2006). The temperature sensitivity of enzymes may be a function of ‘local
adaptation’ (Williams, 1966; Belotte et al., 2003) of organisms to a prevailing temperature
regime across season or latitude. Although tests of local adaptation have been difficult to
design for soil microbial communities (Belotte et al., 2003; Reed & Martiny, 2007), a
‘natural experiment’ using a latitudinal gradient may be one way to test for local adaptation
of microbial physiology. Such an experiment could be particularly valuable for

understanding the Michaelis–Menten kinetics of microbial enzymes, as has been done for

invertebrate enzymes (Johns & Somero, 2004; Dong & Somero, 2009; Huestis et al., 2009).
(German et al., 2012).