|Soil respiration rate; Growth season; Temperature; Soil
moisture; Sloping pasture; Heihe river basin
|Soil surface CO2 efflux is one of the major pathways by which CO2
fixed by terrestrial plants is released into the atmosphere . Given the
controversy over its potential role in amplifying global warming, soil
surface CO2 efflux has recently been the subject of intensive study [2-5].
In the face of impending global warming, increases in soil respiration
are likely to mediate progressively lower rates of carbon sequestration
[2,6-8]. Despite its obvious importance to carbon cycle processes, soil
respiration has proven to be extremely difficult to quantify accurately.
Like many other soil processes, respiration exhibits both great spatial
heterogeneity, particularly at small spatial scales, and great temporal
variability on diurnal, seasonal and inter-annual time scales [9-15].
|Sloping pasture is the main pasture types on the mountain watershed
of the Heihe River basin, and they cover about 28.27% of the total land
area. Understanding of the pasture efflux of CO2 from the soil surface
is a key component of the carbon balance of its ecosystem. Quantifying
this flux and understanding the factors that underlie the temperature
and soil moisture variation in its magnitude are fundamental to our
understanding of the behavior of the ecosystem as a whole and to
our ability to predict the likely consequences of climate change .
Up to now, a large number of studies have been carried out on the
relationship between the dynamics of soil CO2 flux and related factors
[16-19]. However, little has been reported on the sloping pasture soil
respiration intensity at regional scale. This study describes the changes
in CO2 flux of sloping pasture soil and the temporal differences during
the growing season under different environment conditions were
explored in this study. The spatial variations of soil CO2 flux and its
relation to the environmental elements such as soil water content, and
temperature were discussed. The results help the scientific community’s
understanding of carbon exchange mechanisms between soil and
atmosphere and the source-sink changes of the terrestrial ecosystem.
|Materials and Methods
|The experimental site is located in the Pailugou watershed in the Xishui Forest Farm of Sunan County in Gansu Province. The basin
covers an area of 2.95 km2, with 55% grassland and 40% forest land. The
study area has a high, cold, semiarid and sub-humid mountain forest
and grassland climate, with mean annual temperature of 0.5°C, mean
annual precipitation of 435 mm, and annual potential evaporation of
21051 mm . Sloping pasture mainly occurs in the mountain forest
grassland zone at elevation of 2,500~3,000 m and occupies about
28.27% of the Qilian Mountain region’s total area. The main plant
species are Carex, Achnatherum inebrians, Polygonum viviparum,
Oxytropis ochrocephala, Achnatherum splendens. Soil type in the region
is mountain grey cinnamon soil, with a depth of about 1 m. Hence, the
study of CO2 flux of alpine meadow soil at this altitude will contribute
to a better understanding of the feedback effect of soil CO2 emission
flux on the climate and land use changes under high altitude and low
|Soil surface CO2 efflux measurements
|Soil surface CO2 effluxes were measured on early April to late October
2010 with a LiCor 6400 gas exchange analyzer with soil respiration
chamber attachment (LiCor Inc, Lincoln, NE, USA). The 6400 is a
closed system infrared gas analyzer that measures soil respiration. We
installed five PVC collars (0.008 m2) randomly located in each 5m×5m
plot 2 days. Collars were imbedded approximately 2 cm into the soil
and left in place throughout the measurement period. We did not begin
measurements until 1 week after installation to minimize the effects of
disturbance from collar installation and removed live vegetation inside the chamber collars at least 24 hr before measurements to minimize
influences of soil disturbance and root injury on the measurements.
|Soil temperature and moisture measurements
|Soil temperature and moisture were determined at each soil CO2
efflux measurement location and time. Soil moisture within the top 80
mm of soil was measured with a cable tester (1502C Metallic TDR Cable
Tester, Tektronix, Inc., Beaverton, OR) and dedicated Time-Domain
Reflectometry (TDR) steel probes (8 cm long) inserted vertically from
the soil surface. Temperature was measured at a depth of 0.1 m from
the soil surface with a copper-constantan thermocouple mounted in an
|The relationship between soil CO2 flux and temperature was
analyzed using the statistical analysis software SPSS 13.0 for Windows
and the dynamical curve was drawn using the Origin Pro 8.0.
|The diurnal variation in soil CO2 efflux
|Soil CO2 efflux showed an asymmetric diurnal pattern, with a minimum between 0300H and 0700H (local time) and a maximum
in the early afternoon (13:00H-16:00H). Soil CO2 efflux followed the
increasing trend of soil temperature in the morning, but then leveled
off with slight fluctuations, while soil temperature continued to increase
in the afternoon. From evening to early morning of the next day, soil
CO2 efflux followed, with few fluctuations, the declining trend of
soil temperature. In growth season, Heihe basin mountainous area
sloping pasture soil CO2 efflux daily variation assumes the following
characteristic: In the evening maintains at the low level, Minimum
value about 6:00, Starts in 7:00~8:30 to elevate,14:00 about maximizing,
16:00~18:30 to drop gradually, The entire process assumes the single
peak curve. Different month, Soil breath speed daily variation existence
remarkable difference, displays starts in the soil breath speed to elevate,
the maximizing time is different. From May to October, the diurnal
variation of soil respiration was low at night, the lowest at 07:00 H,
06:30 H, 05:30 H, 06:00 H and 07:00 H, rise rapidly between 07:00 H
and 08:30 H, and then descended between 16:00 H and 18:30 H. The
maximum soil CO2 efflux appeared at 15:00 H, 14:30 H, 14:30 H, 13:30
H, 14:00 H and 15:00 H (Figure 1).
|In different periods of growth, soil CO2 efflux diurnal variation
existence remarkable difference (Figure 2) The mean daily soil respiration rate between 0.31~2.58 μmol·m-2·s-1 in May, 0.78~4.85
μmol·m-2·s-1 in June, 4.61~6.98 μmol·m-2·s-1 in July, and then gradually
descended at 2.37~6.26 μmol·m-2·s-1 in August, 3.47~4.23 μmol·m-2·s-1
in September, 1.61~4.21 μmol·m-2·s-1 in October.
|The seasonal variation in soil CO2 efflux
|The continuously observed results of CO2 flux of sloping pasture
soil in the Heihe river basin (Figure 3, Table 1) showed that associated
with the diurnal variations, soil CO2 flux was low in the initial growing
stage (May) but gradually increased in June, reached a maximum value
8.49 μmol·m-2·s-1 in July to August (Figure 3) and started to decrease
in September, the entire process change tendency assumes the single
|Effects of soil temperature and moisture on soil CO2 efflux
|Non-linear regression analysis showed that the exponential model can better describe the relationship between CO2 flux of sloping pasture
soil in the Heihe river basin and soil temperature at 15 cm depth as
shown in figure 4a. Statistical analysis revealed that the correlation
between CO2 flux of alpine meadow soil (S) and soil temperature was
significant (P<0.001, n=25), and its regression relation is as follows:
|The variation trends of CO2 flux of sloping pasture soil in the Heihe
river basin and soil water content were not consistent figure 4b. When
soil water content was low, the variations in soil CO2 flux and soil water
content were almost synchronous, i.e. CO2 flux increased with increase
in soil water content, but when soil content increased to a certain level
the increase in soil CO2 flux became slow. Analytical results show that
the Boltzmann model can better describe the relationship between CO2
flux of sloping pasture soil in the Heihe river basin and soil moisture
(P<0.001, n=25), and its regression relation is as follows:
|S=5.54-5.03/ (1+exp ((x-8.38)/1.34)) (R2=0.98)
|Simultaneous stepwise ranking of variables’ effects on soil
|Soil temperature and soil moisture, as well as their interaction
showed effects on changes in soil CO2 efflux. Using the stepwise
regression process in SPSS, all variables were tested simultaneously
for their relative contribution to explaining variance in soil CO2
efflux. These results were similar to those obtained using simple linear
regression with individual factors: soil temperature was positively
related to soil CO2 efflux and explained about 46.8% of its variance; soil
moisture was weakly positively related to soil CO2 efflux and explained
about 15.7% of its variance; All of the variables combined explained
about 62% of the variation in annual CO2 efflux in 2010.
|Soil CO2 efflux showed diurnal and seasonal changes (Figures 1,2)
The measurements of soil CO2 efflux presented here for the sloping
pasture sites were in the range of 0.31-6.98 μmol·m-2·s-1. This result is higher than one of 2.32-5.70 μmol·m-2·s-1 reported by Michael  for
grassland in northern Ontario of Canada. Raich and Schlessinger 
summarizes large amounts of data showed that average Q10 of grassland
around 2.4, but for forest soils, Liu Shaohui and Fang jinyun 
believes that global average Q10 is 1.57. This study indicated that the
Q10 value of sloping pasture soil is 2.16, And LI Linhao  in sheep
grass grassland research results showed that Q10 value is 2.0~3.0 (to
temperatures for based on), slightly than tropical dilute tree grassland
in North Australia  and high grass grassland in North America 
they think this phenomenon attributed to research locations of latitude
location partial high, because Q10 value in cold region higher than
warm region . This study site and the leymus chinensis grassland
study sites at the same temperate zone, and temperature ranges of the
experimental period commensurate with each other so their Q10 values
|While multiple factors contribute to the differences in measured
efflux rates, the generally low soil moisture and high soil temperature
at our site are likely to be the two major factors which determined the
magnitude of the soil CO2 efflux at the Heihe river basin. Jensen et al.
 measured soil-surface CO2 efflux over two days at 8 locations in a
Pinus radiata D. Don forest in New Zealand using a dynamic chamber
method (portable infra-red CO2 analyser). Their results showed no
apparent diurnal pattern in CO2 efflux, which may have been a result
of a lack in variation in soil temperature (at 150 mm depth) and the
high soil moisture (close to field capacity during the measurements).
However, Davidson et al.  reported a diurnal trend resembling the
temperature pattern. Kutsch & Kappen’s  measurements in crop
fields showed a diurnal trend of CO2 efflux similar to ours, except that
their diurnal maximum occurred later (about 16:00 H).
|When simultaneously considered, soil temperature and soil
moisture explained 46.8 and 15.7, respectively, of the variance in soil
CO2 efflux on our sites. Thus our results indicate that soil temperature
as a single factor explains the greatest amount of variance in soil CO2
efflux observed within and across sites and over seasons in the Heihe
river basin (R2=0.88, P<0.0001). Our findings are consistent with reports that cite a strong relationship between soil temperature and soil
CO2 efflux [30,31].
|Generally, soil moisture limits soil CO2 efflux at either extremely
high or low moisture levels [30-32]. In agreement with the results of
Davidson et al. , at the Heihe river basin the single factor of soil
moisture was correlated (R2=0.98, P<0.0001, N=25) with CO2 efflux,
and the soil CO2 efflux increased slow at high soil moisture contents
(>15%). This effect at high soil moisture may also be related to the
availability of O2 in the soil pore space, which affects microbial activity.
From laboratory and theoretical studies some researchers have found
that high water content can impede diffusion of O2 into the soil, which
in turn impedes decomposition and CO2 production [33-34].
|Soil-surface CO2 efflux measurements were made on sloping
pasture of Heihe river basin from April to October 2010. During the
growing season, the diurnal variation of pasture soil respiration in
the mountain watershed of the Heihe River valley was low at night,
with lowest appears at 7:00, 6:30, 5:30, 5:00, 6:00 and 7:00 from May
to October, and started to rise rapidly during 7:00~8:30, and then
descend during 16:00~18:30. The maximum soil CO2 efflux appears at
15:00, 14:30, 14:30, 13:30, 14:00 and 15:00. The maximum of average
soil CO2 efflux occurred in July and August, and the second was in
May and September, and the third was in April and October and it was
basically consistent in April and October. The diurnal mean soil-surface
CO2 efflux of sloping pasture stand ranged from 0.31 μmol·m-2·s-1 to
6.98 μmol·m-2·s-1, and the Q10 value is 2.16. The positive relationship
between soil CO2 efflux and soil temperature that was observed is well
documented. And the soil CO2 efflux showed a Boltzmann correlation
with soil water content.
|This work was funded by the National Natural Science Foundation of
China (91025002; 30970492; 31270482; 41271037), National Natural Science
Foundation of Gansu Province (1107RJZA089) and the National Key Technology
R&D Program (2012BAC08B05). The authors would like to express their deep gratitude to the anonymous reviewers for their valuable suggestions that greatly
improved the manuscript.
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