Geoderma 402 (2021) 115273

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Geoderma
journal homepage: www.elsevier.com/locate/geoderma

Effects of fire on soil respiration and its components in a Dahurian larch

(Larix gmelinii) forest in northeast China: Implications for forest ecosystem
carbon cycling
Tongxin Hu, Binqing Zhao, Fei Li, Xu Dou, Haiqing Hu *, 1, Long Sun *, 1
Key Laboratory of Sustainable Forest Ecosystem Management-Ministry of Education, College of Forestry, Northeast Forestry University, 26 Hexing Road, Harbin 150040,
China

A R T I C L E I N F O A B S T R A C T

Handling Editor: Jan Willem Van Groenigen Fire plays a critical part in regulating soil carbon (C) budgets in forest ecosystems. However, few studies have
focused on the effects of fire on soil respiration (Rs) and its heterotrophic (Rh) and autotrophic (Ra) components.
Keywords: In this study, we examined the response of Rs, Rh, and Ra to fire and the driving factors that affect Rs after a fire
Fire disturbance disturbance in a Dahurian larch (Larix gmelinii) forest in northeastern China. Fire disturbance significantly
Soil respiration
increased Rh by approximately 24% and significantly reduced Ra by approximately 66% in burned plots
Soil heterotrophic respiration
compared to unburned plots. The ratio of Rh to Rs significantly increased from 0.57 in the unburned plots to 0.79
Soil autotrophic respiration
Larix gmelinii in the burned plots. Fire disturbance also led to cumulative Rs C efflux increasing by approximately 10% and 28%
Cold temperature forest during the growing season and the period of spring freeze–thaw cycles (FTC) in the burned plots. Furthermore,
using structural equation modelling, we showed that T5 (soil temperature at 5 cm depth) was the main abiotic
factor affecting Rs in both unburned and burned plots. Fine root biomass (FR) was the dominant biotic driver of
Rs in the control plots, whereas microbial biomass carbon (MBC) was the dominant biotic driver of Rs in burned
plots. Our findings demonstrated a significant discrepancy in the response of Rs components to fire disturbance.
Forest fires can cause considerable losses of soil C in a cold temperate forest ecosystem, which suggests that
proactive management of forest regeneration should be carried out after forest fire disturbances.

1. Introduction
Forest fires, governed by biomass availability, landscape, meteoro­
logical conditions, and human activity, are critical terrestrial distur­
bance factors that shape the forest ecosystem (Abatzoglou et al., 2018;
Collins et al., 2017; Lacroix et al., 2019; Pausas and Ribeiro, 2017).
Forest ecosystems play a dominant role in the global carbon (C) cycle
and act predominantly as sinks for atmospheric carbon dioxide (CO2)
(Pan et al., 2011). Soil respiration (Rs) in forest ecosystems is a critical
process in C cycling that regulates regional climate and determines the
net C balance in terrestrial ecosystems (IPCC, 2013; Zhou et al., 2016).
However, recent studies have shown that forest fires can significantly
and rapidly affect the Rs of forest ecosystems and their C feedback to the
atmospheric system (Chen et al., 2019; Hu et al., 2017a; Plaza-Álvarez
et al., 2017). Extreme climatic events, such as the El Niño effect and
extreme drought (Seidl et al., 2017; Siegert et al., 2001), and
anthropogenic activities, such as urban expansion and intense land use
for agricultural and livestock activities (Abatzoglou and Williams, 2016;
Balch et al., 2017; Celso et al., 2018), contribute to an ongoing increase
in forest fire occurrence, which may, in turn, disrupt the global C cycle
(Santín et al., 2016; Walker et al., 2019). Thus, quantifying the effect of
forest fires on Rs and its driving factors is key to lowering the un­
certainties associated with C sink estimates in forest ecosystems
(Kasischke and Stocks, 2000; Liu et al., 2014; Schimel and Baker, 2002).
Soil respiration is the sum of soil autotrophic respiration (Ra) from
plant metabolic activity and soil heterotrophic respiration (Rh) from the
decomposition of soil organic matter by microbes (Davidson and Jans­
sens, 2006; Chen et al., 2016). Soil respiration is the second-largest
source of C efflux (80–98 Pg C⋅yr− 1) in terrestrial ecosystems, and
global Rh from soils has been estimated to be 53–57 Pg C⋅yr− 1 (Bond-
Lamberty and Thomson, 2010). Slight changes in Rs and its components
may influence global C balance (Sihi et al., 2018). Although several

* Corresponding authors.
E-mail addresses: htxhtxapple@sina.com (T. Hu), huhq@nefu.edu.cn (H. Hu), sunlong365@126.com (L. Sun).
1
Contributed equally to this work.

https://doi.org/10.1016/j.geoderma.2021.115273
Received 30 September 2020; Received in revised form 27 May 2021; Accepted 1 June 2021
Available online 10 June 2021
0016-7061/© 2021 Elsevier B.V. All rights reserved.
T. Hu et al.
studies have been carried out on the effect of fire disturbances on Rs,
there is still no consensus on the results and the regulatory mechanisms
(Yang et al., 2020). The reason for the lack of consensus is that varying
fire characteristics such as duration, burn severity or season, varying
ecosystem traits, climate differences, and C pools in soils make it diffi­
cult to obtain similar results. A recent study observed that the Rh:Rs ratio
increased significantly from 0.54 to 0.63 between 1990 and 2014 due to
global climate change (Bond-Lamberty et al., 2018). If the soil Rh:Rs
ratio continues to increase, the forest ecosystem becomes a source of C
efflux into the atmosphere (Mayer et al., 2017). Thus, the remaining
challenge is to understand the variations that occur in Rs and the
dominant factors that affect Rs and its components Rh and Ra after a fire
disturbance (Song et al., 2019). In particular, the response of Rh:Rs to
forest fires might be the key factor that influences the amount of net C
stored in forest ecosystems (Bond-Lamberty et al., 2004b; Wang et al.,
2014a). However, previous studies have mainly focused on the effects of
fire disturbance on Rs, and only a few studies have focused on the
variation in Rh and Ra after fire disturbance (Hu et al., 2017a; Pourreza
et al., 2014; Song et al., 2019).
Soil respiration and its components can be severely disturbed by fires
and are potentially controlled by variations in soil environmental factors
(Luo and Zhou, 2006; Pereira et al., 2016). For instance, forest fires can
affect Rs by decreasing vegetation cover and increasing albedo, which
can increase soil temperatures and litter decomposition rates (Throop
et al., 2017). Moreover, higher temperatures after fire disturbance drive
the increase in Rs and its components accumulated C efflux during the
spring freeze–thaw cycles (FTC) period (Hu et al., 2017b). In addition,
high severity fires can increase soil hydrophobicity, which may indi­
rectly control the components of Rs by reducing soil moisture infiltration
and increasing surface runoff (O’Donnell et al., 2009). Although soil
temperature and soil water content account for a considerable propor­
tion of the variation in Rs after fire disturbance, a large proportion of the
post-fire variations in Rs are not fully understood, and the mechanisms
through which fire disturbances affect Rs remain unclear (Li et al.,
2018a, 2018b; Moyano et al., 2013). Thus, it is necessary to further
study the influence of biotic factors (e.g. soil microbial activities and
plant roots) on Rs after fire disturbance.
Several studies have indicated that fire affects Rs directly by affecting
soil microbial activities and plant roots, especially fine roots (<2 mm)
(Holden et al., 2016; Kitchen et al., 2009; Uribe et al., 2013; Wang et al.,
2012), and indirectly by changing soil pH, soil nutrient availability, and
substrate quality, which are related to soil microorganisms and fine
roots (Makita et al., 2016; Throop et al., 2017). The extent to which fire
disturbance affects these factors depends on the burn severity and
duration of the fire (Certini, 2005). Several studies have shown that high
temperatures during a fire can reduce soil microbial biomass, thus
inhibiting soil microbial activities in the short term after a fire (Dooley
and Treseder, 2012; Ludwig et al., 2018). Others have found that fire
releases alkaline ions in ash, which increases soil pH and provides
greater availability of nutrients after the fire disturbance, an effect that
could promote soil microbial activities in the long term (Barreiro et al.,
2016; Stirling et al., 2019). Similarly, fires with high burn severity can
significantly reduce fine root biomass and inhibit the physiological ac­
tivities of plant root organisms (Kitchen et al., 2009; Vergani et al.,
2017). However, some studies have found that an increase in the
availability of soil ammonia and nitrate nitrogen through the burning of
plant tissue can promote the rapid recovery of vegetation, thus leading
to the recovery of fine root biomass in a short period after a fire (Day
et al., 2006). The complex responses of soils, microbes, and vegetation to
fire disturbance lead to uncertainties in estimating post-fire Rs levels
(Tufekcioglu et al., 2010). Therefore, it is of great ecological significance
to consider fully the relationship between Rs and the changes in soil
biotic factors, nutrient availability, and soil microclimate after a fire to
reveal the mechanisms through which fire disturbance affects Rs.
In the present study, we selected a forest of Dahurian larch (Larix
gmelinii), which accounts for approximately 30% of timber production in
2
Geoderma 402 (2021) 115273
China (Hu et al., 2019). This forest type is mainly located along the
southern edge of the Eurasian boreal forests (a cold-temperature forest
transition zone in northeast China) and is extremely sensitive to climate
change (Li et al., 2020b). The specific objectives of this study were (1) to
explore the pattern of change of Rs and its components (Rh and Ra) in a
cold, temperate forest ecosystem one year after a fire disturbance, and
(2) to elucidate the abiotic and biotic driving factors of Rs one year after
a fire disturbance. Based on the findings of previous studies, we
hypothesised that (1) compared with the unburned control plots, there
are differences in the response of Rh and Ra to fire disturbances, which
leads to a post-fire increase in the Rh:Rs ratio in burned plots; (2) a fire
will change the dominant biotic and abiotic factors that regulate Rs in
burned plots; and (3) a fire will drive the increase in accumulated C
efflux of Rs, Rh, and Ra during the FTC period in burned plots.
2. Material and methods
2.1. Site description
The present study was conducted at the Forest Ecosystem Research
Station on the Maoer Mountain in Heilongjiang Province, northeast
China (45◦ 20–45◦ 25 N, 127◦ 30–127◦ 34 E; 400 m above sea level)
(Fig. 1). Heilongjiang Province has the highest incidence of forest fires in
northeastern China, and the fire season in this area is mainly from March
to June and September to November each year, with a total of 1302 fires
occurring between 2000 and 2020, destroying an area of 5.13 × 104 ha
yr− 1.
The region experiences a continental monsoon climate with dry and
cold winters. During the sampling period (May 2017–Apr 2018), the
mean annual air temperature was 6.3 ◦ C, the mean maximum and
minimum air temperatures were 30.5 ◦ C and − 38.2 ◦ C, respectively, and
the mean total annual precipitation was 560.1 (Fig. 2). The snow lasted
for 154 days (4 Nov. 2017 to 7 Apr. 2018), with a maximum snowpack
depth of 31.2 cm and a mean snowpack depth of 14.1 cm; snow started
to melt at the end of March. The parent material is granite bedrock, and
the soil is classified as a Haplumbrepts in the United States Soil Tax­
onomy (Soil Survey Staff, 2014). The forest is a Dahurian larch (Larix
gmelinii (Rupr.) Kuzen) plantation (>80% total basal area), which was
established in 1985 (32 years old), is a mixed forest with naturally re­
generated tree species, such as Betula platyphylla and Populus davidiana.
The dominant herb species observed in the area were Anisodus acu­
tangulus, Adenocaulon himalaicum, Dryopteris crassirhizoma, and Aegopo­
dium alpestre.
In April 2016, forest fires caused by lightning occurred at the Maoer
Mountain Forest Ecosystem Research Station. The fires lasted one day,
and approximately 500 ha of forest was burned, which provided an
opportunity for us to study the effects of fire disturbances on Rs and its
components. Burn severity was estimated to be moderate based on the
depth of the burned organic soil, the consumption of the aboveground
biomass, tree mortality, and bark char height – characteristics that help
us understand the extent of fire damage to a forest (Keeley, 2009). In the
burned area, approximately 47% of the understory shrubs were burned,
approximately 37% of the trees died, the bark char height was 1.7–2.5
m, approximately 65% of the organic layer was burned, and the ash
layer was 3.4–4.2 cm. Moreover, a substantial amount of charcoal was
left on the burned area ground surface. Herbaceous vegetation recov­
ered rapidly three months after the fire disturbance, and the species
were the same as those in the unburned areas. Some Dahurian larch tree
seeds germinated and regenerated one year after the fire disturbance.
We selected three replicate plots in the burned area to conduct our
investigation and selected the nearby unburned area as the control. Six
experimental plots were selected in the burned and unburned areas for
our study in May 2016. The size of each plot was 400 m2 (20 m × 20 m),
and the distance between the plots was>200 m to ensure the indepen­
dence of data acquisition. All replicate plots had a low slope gradient
(5%–8%). The diameter at breast height (DBH), tree height, canopy
T. Hu et al. Geoderma 402 (2021) 115273

Fig. 1. The location and plot arrangement of the research area in the Maoer mountains, northeastern China.

cover, depth of the organic layer (OL), and bulk density in the control
and burned plots are shown in Table 1. All sampling points (SP) and
trenched sampling points (TSP) were established in May 2016 (Fig. 1),
approximately 12 months before the measurement of Rs and its com­
ponents, to ensure that the effects of the trenching process on Rs and its
components could be avoided (Jiao and Wang, 2019; Mayer et al.,
2017).

2.2. Dividing the study period into specified periods

Fig. 2. Temporal changes in mean daily air temperature (Ta), soil temperature
at 5 cm depth (T5), and precipitation during the measurement period from May
2017 to April 2018.
The study period was divided into four periods based on the mean air
temperature (Ta) and soil temperature at 5 cm depth (T5) (Fig. 2) (Piao
et al., 2007; Wang et al., 2011, 2013). The winter period (8 Nov.
2017–31 Mar. 2018) was defined as the period in which the mean Ta
remained below 0 ◦ C for at least consecutive days. The FTC period (1
Apr. 2018–13 Apr. 2018) in spring began at a daily maximum Ta of
above 0 ◦ C (i.e., the beginning of the snowmelt) and ended when the
daily minimum T5 was above 0 ◦ C (i.e., when the snow had completely
melted). The non-growing season (8 Nov. 2017–13 Apr. 2018) included
winter and the FTC periods. The growing season (1 May 2017–7 Nov.
2017; 14 Apr. 2018–30 Apr. 2018) was defined as the period from the
end of the spring FTC to the beginning of winter.
2.3. Partitioning sampling points for estimation of Rs and its components

A trenching method was used to separate the soil Rh and Ra (Fig. S1).
Five sampling points (SP) were randomly selected for Rs measurement,
and three trenched sampling points (TSP) for Rh measurement were
Table 1
selected in each plot (Fig. 1). A total of 30 sampling points (5 soil rings ×
Basic information of the research plots. Values are means (±standard deviation)
6 replicate plots) were used to measure Rs in burned plots and in un­
of three replicates (n = 3) in both the burned and control plots.
burned control plots, and a total of 18 trenched sampling points (three
Plot type DBH (cm) Height of tree Canopy OL Bulk soil rings × six replicate plots) were used to measure the Rh in burned
(m) density
cover (cm) (g cm− 3)
and unburned control plots. We excavated 20 cm wide and 80 cm deep
(%) (approximately the bottom of the root zone) trenches around the
trenching sampling points, each encompassing an area of 1.5 × 1.5 m
Burned 12.96 ± 11.83 ± 72 ± 6.2 ± 0.58 ±
plots 7.18a 2.71a 10a 0.6a 0.08a (Fig. S1a). All roots within the trenches were severed, and a plastic
Control 13.87 ± 12.17 ± 90 ± 10.5 ± 0.82 ± lining was installed to inhibit root and mycorrhizal in-growth. The
plots 7.83a 2.32a 16b 1.0b 0.01b trenches were then refilled according to their original soil profile, and
Note: DBH = diameter at breast height; OL = depth of the organic layer. Mean the surface vegetation was also removed in trenched sampling points
values in a column that have the different lowercase letters indicate that the (Fig. S1b).
corresponding variable differed significantly of the two plot types at the p = 0.05 A polyvinylchloride (PVC) soil collar (inner diameter 20 cm and
level. height 10 cm) was inserted 3 cm deep into the soil in each plot for Rs

3
T. Hu et al.
measurement. The surface vegetation in the trenched sampling points
was removed during the study period. The Ra was calculated as the
difference in soil CO2 effluxes measured from the trenched and un­
treated sampling points in the respective areas.
2.4. Field measurement and laboratory analysis
Soil CO2 effluxes were measured using a Li-8100 portable automatic
measuring system for soil C flux (LI-COR Biosciences, Inc., Lincoln, NE,
USA). Soil CO2 effluxes were measured on the 15th of each month from
May 2017 to April 2018, for a total of 12 times during the study period.
The measurement cycle lasted approximately 2 min for each R or Rh
ring. All 48 (30 non-trenched soil rings for Rs + 18 trenched soil rings for
Rh) measurements were conducted between 9:00 and 11:00 am. The T5
and soil volumetric water content (W5) at 5 cm depth near the PVC rings
were measured simultaneously using a temperature probe (p/
n8100–201; LI-COR Biosciences, Lincoln, NE, USA) and the ECH20
model EC-5 soil volumetric water content sensor (p/n 8100202;
Decagon Devices, Inc., Pullman, WA, USA), respectively.
Soil samples were collected once a month, and the soil sample
collection time was consistent with that of soil CO2-efflux measure­
ments. Soil samples were taken from 0 to 10 cm with a 5-cm-diameter
tube auger and 10 cm away from the PVC ring. Twelve soil samples
collected from the same soil layer in each plot were combined to form a
composite sample. Samples were kept cool (4 ◦ C) and sieved (2 mm
sieve) within a few days. Ammonium-nitrogen (NH4-N) and nitrate-
nitrogen (NO3-N) were extracted from 5 g of soil incubated in 50 ml
2 M KCl for 1 h (Li et al., 2018a, 2018b). The extracts were filtered using
Whatman 42 (Whatman International Ltd., Maidstone, Kent, UK) and
analysed for NH4-N and NO3-N concentrations (mg kg− 1) using a BRAN
+ LUEBBE-AA3 autoanalyzer (Seal Analytical, Germany). Soil pH (soil:
water 1:2.5) was measured in fresh samples using a PHS-3S pH meter
(Sartorius, Germany).
Fine root (<2 mm) biomass (FR) was determined according to the
method described by Rewald et al. (2012). Tree roots were distinguished
from other root segments by root colour, odour, resilience (to breakage),
presence of root resin, and woody structure, using a stereomicroscope
(10–40x), before tree roots were further separated into living and dead
fine roots. Dead fine roots were separated from living ones based on
their colour and lustre, resilience (to breakage), smell, and appearance
of phloem (Brassard et al., 2013). Subsequently, all FR samples were
dried at 70 ◦ C for 48 h and weighed (±0.1 mg) to determine the FR mass
per hectare.
Soil samples used for the determination of soil organic carbon (SOC)
were dried at 40 ◦ C and then ball-milled until they passed through a 200-
mm-mesh sieve. Ball milling was performed in a planetary agate ball
mill (QM-3SP04, Nanjing, China) under ambient conditions for 2 min at
300 rpm (Yapiyev et al., 2018). All the milled soil samples were analysed
for SOC concentration (g kg− 1) using a Multi N/C 2100 s (Analytik Jena,
Germany) elemental analyser.
Soil microbial biomass carbon (MBC) values were determined using
the chloroform fumigation-extraction method (Brookes et al., 1985). For
this, 20 g of fresh soil samples was extracted using 0.5 mol L-1 of po­
tassium sulphate (K2SO4) (at a ratio of 1:4) and by shaking for 1 h at 200
rpm. A further 20 g of fresh soil was extracted using the same procedure,
but after being fumigated with chloroform for 24 h in the dark. Dissolved
organic carbon (DOC) content in both fumigated and unfumigated ex­
tracts was analysed using Multi N/C 2100 s (Analytik Jena, Germany).
The MBC was calculated as follows:
MBC = EC /0.45 (1)
where EC is the difference between the fumigated and unfumigated DOC
content (mg kg− 1), and 0.45 is the conversion factor for MBC (Brookes
et al., 1985).
4
Geoderma 402 (2021) 115273
2.5. Data analysis and modelling for soil C efflux
Data were processed and analysed using R statistical software
version 3.5.2 (R Core Team 2018). A two-way analysis of covariance
(ANOVA) was performed with fire disturbance (F), measurement date
(D), and their interaction effects (F × D) on soil properties (T5, W5, FR,
MBC, soil pH, NH4-N, NO3-N, and SOC) and Rs and its components (Rh
and Ra). A three-way ANOVA was performed with fire disturbance (F),
measurement date (D), trenching effect (TE), and their interaction ef­
fects on soil temperature (T5) (℃), and soil water content (W5) (%) of at
5 cm depth in sampling points and trenched sampling points. All sta­
tistical analyses were performed at a significance level of 0.05.
Structural equation modelling (SEM) was used to determine the
direct and indirect effects of abiotic (T5, W5, soil pH, NH4-N, NO3-N,
SOC) and biotic factors (FR and MBC) on Rs. The SEM model was
calculated using the “lavann” package in R (Rosseel, 2012). In a stepwise
procedure, pathways and variables were removed or added to the model
to obtain the absolute fit of the best model; Akaike’s information crite­
rion (AIC), chi-squared test results (in SEM, a reliable model is indicated
by a low χ 2 and a model p-value > 0.05), and the comparative fit index
(CFI) served as measures for the overall model fit. Only significant re­
lationships (p < 0.05) were maintained in the model.
An exponential model and the temperature sensitivity (Q10) of Rs and
its components (Rh and Ra) were used to describe the relationship be­
tween Rs and soil temperature, which was determined by fitting the
exponential function (Eq. (2) and Eq. (3) below) for the burned and
unburned control plots (Hu et al., 2017a; Lloyd and Taylor, 1994):
SR = αe(βT5 ) (2)
Q10 = e10β (3)
where SR is the measured Rs and its components (Rh and Ra), T5 is the
soil temperature (◦ C) at 5 cm, α and β are regression coefficients, e is the
natural number, and Q10 is the factor by which Rs and its components
increase during a temperature increase of 10 ◦ C.
Previous studies have indicated that soil temperature and soil water
content are the two main abiotic variables that affect Rs and its com­
ponents (Cable et al., 2012; Hursh et al., 2017; Song et al., 2019), and we
previously developed an exponential model to describe the effects of soil
temperature and soil water content on Rs and its components (Rh and Ra)
(Hu et al., 2017a). Logarithmic transformation of Rs and its components
(Rh and Ra) was required to achieve linearity and homoscedasticity.
Here, we selected the best-fitted model based on R2 and AIC. The
regression model is given by Eq. (4):
Ln(SR) = α + β × T5 + ε × W5 + ω × T5 × W5 (4)
where Ln (SR) is the logarithmic transformation of Rs and its com­
ponents (Rs or Rh) that was applied to achieve linearity and homosce­
dasticity, T5 is the soil temperature at 5 cm (◦ C), W5 is the soil water
content at 5 cm (%), T5 × W5 is the interaction effect of T5 and W5, and α,
β, ε, and ω are regression coefficients. A stepwise regression procedure
was performed to remove insignificant terms (p = 0.05).
The accumulated C efflux (g C⋅m− 2) of daily SR (Rs or Rh) was esti­
mated based on the hourly emissions calculated by Eq. (5):
∑24
∑ j=1 SR(i,j) × 12 × 3600
SR(i) = (5)
1, 000, 000
∑ ∑
where SR(i) is the daily emission of Rs or Rh; SR(i,j) is the daily
emission of Rs or Rh at moment j (1 h) on day i, 12 is the molecular weight
of C, and the figure of 3600 is a constant value (unit: second) based on
the Campbell Scientific data logger (Campbell Scientific, Inc., Utah,
USA) to record soil temperature and soil water content every 60 min
from May 2017 to April 2018. The annual C efflux (g C⋅m− 2 yr− 1) of Rs
and Rh was calculated by adding up the daily emissions. The sum of C
T. Hu et al. Geoderma 402 (2021) 115273

efflux in the growing and non-growing seasons is the annual C efflux.

3. Results

3.1. Effect of fire on soil abiotic and biotic factors

The T5, W5, soil pH, NH4-N, and MBC values showed a significant
seasonal variation pattern during the year-long field experiment (Fig. 3
a, b, c, e, g; Table 2). However, the NO3-N, SOC, and FR content did not
show such a significant seasonal variation pattern over the study period
(Fig. 3 d, f, h; Table 2). The mean values of T5 in the control and burned
plots were 6.13 ± 0.64 ◦ C and 7.97 ± 1.30 ◦ C, respectively. The mean
values of W5 in the control and burned plots were 34.35 ± 2.56% and
27.45 ± 3.84%, respectively. We found that T5 was significantly higher
in the burned plots than in the controls, whereas W5 in the burned plots
was significantly lower than that in the controls (Table 2). The trenching
approach had no significant effect on T5 and W5 between the trenched
and non-trenched sampling points (Fig. S2; Table S1).
The mean values of soil pH in the control and burned plots were 4.95
± 0.38 and 5.48 ± 0.42, respectively; the mean values of NO3-N in the
control and burned plots were 31.10 ± 13.38 mg kg− 1 and 35.85 ±
11.33 mg kg− 1; the mean values of NH4-N in the control and burned
plots were 100.63 ± 37.78 mg kg− 1, and 140.49 ± 38.86 mg kg− 1,
respectively; the mean values of SOC in the control and burned plots
were 142.04 ± 18.54 g kg− 1, and 128.27 ± 17.68 g kg− 1, respectively;
the mean values of MBC in the control and burned plots were 398.70 ±
169.32 mg kg− 1, and 588.02 ± 259.67 mg kg− 1, respectively; the mean
values of FR biomass in the control and burned plots were 0.59 ± 0.53 t
ha− 1 and 0.41 ± 0.37 t ha− 1, respectively. The soil pH, NH4-N, and MBC
were significantly higher in the burned plots than in the control plots
(Table 2). However, the SOC of the burned plots was significantly lower
than that of the control plots (Table 2). There was no significant dif­
ference in soil NO3-N content between the control and burned plots
(Table 2). Although the FR biomass in the burned plots was approxi­
mately 31% lower than in the control plots, the difference was not sig­
nificant (Table 2).
During the spring FTC period, the mean values of soil T5 in the
control and burned plots were 3.39 ± 0.93 ◦ C and 7.43 ± 1.87 ◦ C,
respectively; the mean values of W5 in the control and burned plots were
38.55 ± 3.02% and 30.15 ± 4.47%, respectively; the mean values of soil
pH in the control and burned plots were 4.60 ± 0.07 and 5.90 ± 0.05,
respectively; the mean values of NO3-N in the control and burned plots
were 44.64 ± 27.39 mg kg− 1 and 45.81 ± 8.11 mg kg− 1; the mean values
of NH4-N in the control and burned plots were 86.50 ± 70.03 mg kg− 1,
and 203.98 ± 30.65 mg kg− 1, respectively; the mean values of SOC in
the control and burned plots were 138.28 ± 17.69 g kg− 1, and 124. 79 ±
16.73 g kg− 1, respectively; the mean values of MBC in the control and
burned plots were 406.56 ± 58.13 mg kg− 1, and 767.66 ± 58.13 mg
kg− 1, respectively; the mean values of FR biomass in the control and
burned plots were 0.57 ± 0.06 t ha− 1 and 0.93 ± 0.42 t ha− 1, respec­
tively. Compared with the control plots, the soil T5, pH, MBC, and FR
were significantly higher in the burned plots during the FTC period (p <
0.05). There was no significant difference in soil W5, NO3-N, NH4-N, and
SOC between the control and burned plots during the FTC period (p >
0.05).

3.2. Effect of fire disturbance on Rs and its components

Like T5, the temporal dynamics of Rs, Rh, and Ra also showed a
significantly varied seasonal pattern during the experimental period,
with the highest values in the summer and the lowest values in winter
(Fig. 4, Table 2). The mean values of Rs in the control and burned plots
were 1.84 ± 0.37 µmol CO2⋅m− 2 s− 1 and 1.53 ± 0.27 µmol CO2⋅m− 2 s− 1,
respectively. The mean values of Rs in the control plots were signifi­
cantly higher by approximately 20% (p < 0.05) than those in the burned
plots (Fig. 4a, Table 2).
Fig. 3. Temporal changes of (a) soil temperature at the 5 cm depth (T5), (b) soil
volumetric water content at 5 cm depth (W5), (c) soil pH of 0–10 cm depth soil,
(d) soil nitrate-nitrogen (NO3-N) concentration of the 0–10 cm depth soil, (e)
soil ammonium-nitrogen (NH4-N) concentration of the 0–10 cm depth soil, (f)
soil organic carbon (SOC) concentration of the 0–10 cm depth soil, (g) soil
microbial biomass C (MBC) concentration of the 0–10 cm depth soil, (h) fine
roots biomass (FR) in control and burned plots. Vertical bars are standard de­
viations of the mean (n = 3).

5
T. Hu et al. Geoderma 402 (2021) 115273

Table 2
Results (F-values) of repeated-measures analysis of variance of the effects of fire disturbance (F), measurement date (D), and their interaction on soil temperature at 5
cm depth (T5; ℃) , soil volumetric water content at 5 cm depth (W5; %), nitrate-nitrogen (NO3-N; mg kg− 1), ammonium-nitrogen (NH4-N; mg kg− 1), soil organic carbon
(SOC; g kg− 1), microbial biomass carbon (MBC, mg kg− 1); soil pH, fine roots biomass (FR, t ha− 1); soil total respiration (Rs; µmol CO2⋅m− 2 s− 1), soil heterotrophic
respiration (Rh; µmol CO2⋅m− 2 s− 1), soil autotrophic respiration (Ra; µmol CO2⋅m− 2 s− 1) (n = 36).
F D F×D

Df F ratio p-value Df F ratio p-value Df F ratio p-value

T5 1 39.98 ** ↑ 11 508.36 ** 11 3.57 **

W5 1 57.89 ** ↓ 11 3.23 ** 11 2.21 *
NO3-N 1 2.86 n.s. 11 1.63 n.s. 11 0.9 n.s.
NH4-N 1 51.91 ** ↑ 11 7.43 ** 11 5.17 **
SOC 1 11.82 ** ↓ 11 1.94 n.s. 11 0.92 n.s.
MBC 1 48.48 ** ↑ 11 14.16 ** 11 4.45 **
Soil pH 1 46.62 ** ↑ 11 3.67 ** 11 1.32 n.s.
FR 1 3.66 n.s. 11 1.83 n.s. 11 2.1 *
Rs 1 9.24 ** ↓ 11 64.01 ** 11 4.01 **
Rh 1 10.02 ** ↑ 11 52.08 ** 11 9.84 **
Ra 1 39.13 ** ↓ 11 9.84 ** 11 5.25 **
Rh: Rs 1 54.82 **↑ 11 2.669 ** 11 0.30 n.s.

Note: n.s., *, and ** represents non-significant, p < 0.05, p < 0.01, respectively; “↑” indicates that fire disturbance causes a significant increase in this variable; “↓”
indicates that fire disturbance causes a significant decrease in this variable; “n.s” indicates that the fire disturbance has no significant effect on this variable.

The mean Rh values in the control and burned plots were 1.01 ± 0.17
µmol CO2⋅m− 2 s− 1, respectively, and 1.25 ± 0.28 µmol CO2⋅m− 2 s− 1,
respectively. The mean Rh in the burned plots was significantly higher
by approximately 24% than in the control plots (p < 0.05) (Fig. 4b,
Table 2). The mean Ra in the control and burned plots was 0.82 ± 0.33
µmol CO2⋅m− 2 s− 1 and 0.28 ± 0.14 µmol CO2⋅m− 2 s− 1, respectively. The
Ra in the burned plots was significantly lower (approximately 66%) than
in the control plots (p < 0.05) (Fig. 4c, Table 2). The mean values of Rh:
Rs ratio of the control and burned plots was 0.57 ± 0.14 and 0.79 ± 0.10,
respectively. The Rh:Rs in the burned plots was significantly higher by
approximately 39% than that in the control plots (p < 0.05) (Fig. S3,
Table 2).
3.3. Relationship between Rs and soil properties
The factors showing a significant correlation with Rs in either the
control or burned plots were introduced in the SEM models (Fig. 5,
Fig. 6, Fig. S4). In the control plots, Rs was significantly correlated with
T5 (α = 0.819, p < 0.01) and FR (α = 0.293, p < 0.01) (Fig. 5a). The SOC
was significantly correlated (α = 0.382, p < 0.05) with FR in the control
plots (Fig. 5a). In the burned plots, Rs was also significantly correlated
with T5 (α = 0.569, p < 0.01) and MBC (α = 0.377, p < 0.01) (Fig. 5b).
We found that FR was significantly positively correlated with NH4-N (α
= 0.552, p < 0.01) but was significantly negatively correlated with T5 (α
= -0.680, p < 0.01) in burned plots. The MBC was significantly corre­
lated with NH4-N (α = 0.387, p < 0.01) in the burned plots (Fig. 5b).
3.4. Modelling and C-efflux estimation of Rs and its components
The exponential regression models with T5 as a single controlling
factor for Rs, Rh, and Ra were significant (p < 0.05) for both the control
and burned plots and explained approximately 82%–85%, 81%–82%,
and 43%–67% of the variation in Rs, Rh, and Ra, respectively (Fig. 6).
However, only Rh showed a significant linear relationship with W5 in the
burned plots, whereas Rs, Rh, and Ra did not have a significant linear
relationship with W5 in the control plots (Fig. 6). T5 had a significant
effect on Rs in both the control and burned plots. In contrast, W5 had no
significant effect on Rs in either the control or burned plots. The Q10 of
Rs, Rh, and Ra in the burned plots was 22%, 21%, and 42% lower than in
the control plots (T5 in Fig. 6).
The empirical model containing both T5 and W5 improved the fitting
of the observed Rh in burned plots compared to that of the exponential
model with only T5 (Table 3). The growing season, non-growing season,
winter period, spring FTC period, and annual C effluxes of Rs, Rh, and Ra
in the control and burned plots are shown in Table 4. Compared with the
control plots, the cumulative C efflux of Rs during the annual, growing,
and spring FTC periods increased by approximately 5%, 10%, and 27%,
respectively, in the burned plots. However, compared with the control
plots, the cumulative C efflux of Rs during the non-growing season
decreased by approximately 28% in the burned plots (Table 4). The ratio
of the cumulative C efflux of Rs during the spring FTC period to the non-
growing season cumulative C efflux of Rs was 11% in the control plots,
while the proportion increased to 19% in the burned plots.
Compared with the control plots, the cumulative C efflux of Rh
during the annual, growing season, non-growing season, and spring FTC
period increased by approximately 34%, 37%, 19%, and 19%, respec­
tively (Table 4). Compared with that in the control plots, the cumulative
C efflux of Ra during the annual, growing season, and non-growing
season decreased by approximately 31%, 22%, and 79%, respectively
(Table 4). However, the cumulative C efflux of Ra during the spring FTC
period increased by approximately 37% in the burned plots (Table 4).
4. Discussion
4.1. Effect of fire disturbance on soil respiration and its components
Consistent with our first hypothesis, there were differences in the
responses of Rs components to fire disturbances. In our study, Rs and Ra
significantly decreased after a fire, whereas Rh and Rh:Rs ratios increased
significantly. The significant decrease in Rs was mainly driven by a
significant decrease in Ra after a fire. However, our results disagree with
our second hypothesis in terms of abiotic factors. Fire did not cause a
shift in the dominant abiotic factors that regulated Rs. Our results
indicated that soil temperature was the dominant abiotic factor for Rs
variation in both the control and burned plots, which was consistent
with previous findings (Hu et al., 2017a; Sawamoto et al., 2000; Uribe
et al., 2013). Although previous studies have shown that low soil water
content is an important constraining factor in forest soil organic matter
decomposition after fire disturbance, it did not act as a controlling factor
for Rs in our study, either in control plots or burned plots (Holden et al.,
2015; Muñoz-Rojas et al., 2016). Other studies have indicated that soil
water content could be a significant controlling factor for Rs only under
extremely dry conditions (Rey et al., 2011; Sun et al., 2019; Zhou et al.,
2019). However, consistent with our second hypothesis, the dominant
biotic factors that regulate Rs shifted after fire disturbance. According to
our SEM results, FR was a dominant biotic factor for Rs variation in
unburned control plots, whereas MBC was the dominant biotic factor for
Rs variation after a fire. These results were mainly attributed to the
difference in the Rs components’ (Rh and Ra) responses to the fire. Soil
autotrophic respiration is an important component of soil respiration in

6
T. Hu et al.
Fig. 4. Variations of (a) total soil respiration rate (Rs), (b) soil heterotrophic
respiration (Rh), and (c) soil autotrophic respiration (Ra) in control and burned
plots. Vertical bars are standard deviations of the mean (n = 3).
unburned control plots, as it originates mainly from the root system. If
root biomass declines in response to fire disturbance, Ra decreases at the
stand level. Soil heterotrophic respiration, mainly from the decompo­
sition of organic material by free-living microbes, was significantly
higher in the burned plots. If the MBC increases in response to fire
disturbance, Rh increases at the stand level.
Studies have shown that global warming has driven the increase in
Rh:Rs ratio due to rising soil organic carbon losses during the last de­
cades (Bond-Lamberty et al., 2018; Crowther et al., 2016; Melillo et al.,
2017; Nottingham et al., 2020). We found that the Rh:Rs ratio increased
from 0.57 to 0.79 after a fire, which was consistent with previous
findings that Rh accounts for approximately 71%–93% of the Rs after
fires of varying severity (Bond-Lamberty et al., 2004b; Hu et al., 2017a).
The decrease in Ra can be mainly attributed to the decrease in FR after a
fire, which is consistent with previous findings that Ra decreases at the
stand level when the root biomass declines in response to fire distur­
bance (Czimczik et al., 2006; Makita et al., 2016; Smirnova et al., 2008).
Previous studies have indicated that higher temperature and NH4-N
content after a fire is mainly attributed to the volatilisation of organic N
7
Geoderma 402 (2021) 115273
during combustion and its condensation after deposition to the cool soil
layer, and the increased N mineralisation, which had a significantly
positive correlation with FR after a fire (Grogan et al., 2000; Hu et al.,
2019; Pellegrini et al., 2018; Smithwick et al., 2005). However, in our
study, the FR of the burned plots was still lower than in the unburned
control plots. Fire not only stimulates an increase in soil temperature
(Throop et al., 2017), it also decreases the soil water content through
evapotranspiration and reduces the soil’s water-holding capacity (Badía
et al., 2017). This is because the organic layer, characterised by a high
water-holding capacity, has been destroyed (Ebel 2012). This effect, in
turn, indirectly suppresses root production (Bai et al., 2010; Nishar
et al., 2017). Decreased soil water availability can inhibit root growth
and carbon allocation capacity due to reduced plant photosynthesis (Niu
et al., 2008). We also found that SOC was the dominant factor for FR
growth in the control plots, although they did not exhibit a significant
correlation one year after the fire. The finding that SOC content
decreased significantly after a fire indicated that the decrease in the
substrate supply might be another reason for the limited post-fire FR
growth. In addition, the decrease in FR biomass may also be caused by
the incorporation of charcoal, which is a by-product of incomplete
combustion that influences the below-ground ecosystem (Hart and
Luckai, 2013; Hernandez-Soriano et al., 2016). Several studies have
found that charcoal absorbs toxic compounds (e.g. phenols, dioxin,
furan, and benzoic acid) and delays the N mineralisation process, which
may indirectly suppress fine root development after a fire (Bryanin and
Makoto, 2017; Wang et al., 2017; Wardle et al., 1998). Nearly 50% of
the carbohydrates from photosynthesis are supposed to be transported
to the roots, and approximately 40% of the total assimilated C is used in
root respiration, which is the main component of Ra in the forest
ecosystem (Bond-Lamberty et al., 2004a; Hopkins et al., 2013). There­
fore, the effect of fire on Ra should not be ignored, as it is critical to
evaluate the changes in soil respiration components after fire
disturbance.
The higher Rh in burned plots may be driven by the post-fire increase
in MBC. Previous studies mainly attributed the increase in MBC to the
increase in nutrient availability, decomposition of additional organic
matter input from aboveground plant residue, and changes in soil
microclimate (Bárcenas-Moreno et al., 2016; Fernández-García et al.,
2019). Our SEMs also showed that MBC had a significantly positive
correlation with NH4-N after a fire. An increase in NH4-N one year after a
fire has been observed in several studies (Smithwick et al., 2005; Wan
and Luo, 2001; Wang et al., 2014c). Such an increase is beneficial to soil
microbial activity because microbes use NH4-N for their growth (Dooley
and Treseder, 2012; Nordin et al., 2004). Like NH4-N, NO3-N is another
important form of inorganic nitrogen in the soil. Previous studies have
also indicated that post-fire soil NO3-N content is generally higher than
pre-fire levels (DeLuca et al., 2006; Smithwick et al., 2005; Wang et al.,
2014c). However, our results showed that the NO3-N levels did not differ
significantly between the burned and control plots, which may be due to
the leaching effect and uptake by microbes after the fire disturbance
(Koyama et al., 2012).
Several studies have indicated that there may be an immediate and
significant increase in SOC after a fire (Czimczik et al., 2005; Li et al.,
2020a; Sawamoto et al., 2000; Zhao et al., 2012). However, we found
that SOC in burned plots was significantly lower than in unburned
control plots one year after the fire. In conjunction with our findings that
there was higher Rh and MBC in burned plots, this suggests that the
decrease in SOC in the burned plots might be caused by the consumption
of SOC by soil microorganisms for their own growth after the fire
disturbance (Chen et al., 2019). Kuzyakov (2006) reported that a change
in Ra has little effect on the variation in SOC stock, whereas a change in
Rh could substantially alter the SOC pool size and atmospheric CO2
concentration (Moinet et al., 2016). If the fire continues to increase Rh,
this will lead to the loss of more SOC into the atmosphere in the form of
CO2 (Martínez-García et al., 2017). Current research has also found that
litter decomposition is not the main driving force for the fixation of the
T. Hu et al.
organic C pool; instead, fine roots are the main driving force for C
sequestration in the forest ecosystem (Bryanin and Makoto, 2017; Wu
et al., 2018). Therefore, fire disturbance may further lead to increased
soil heterotrophic respiration and decreased soil carbon sequestration
capacity one year later, which will have an important impact on the
underground carbon cycle of forest ecosystems (Singh et al., 2008; Song
et al., 2019).
4.2. Effect of fire disturbance on Q10 and FTC period
Compared with the control plots, the Q10 of Ra was 42% lower in the
burned plots. Wan and Luo (2003) found that Q10 not only reflects Ra
sensitivity to temperature, but also expresses the combined response to
fluctuations in temperature, root biomass, soil water content conditions,
and substrate quality. Fire destroys root structures and causes the loss of
8
Geoderma 402 (2021) 115273
Fig. 5. Structural equation models using soil
abiotic and biotic factors as predictors of soil
respiration (Rs) for unburned control plots (a)
and burned plots (b). Solid lines represent sig­
nificant paths (p < 0.05), and the dashed lines
represent insignificant paths (p > 0.05). FR, fine
root biomass; MBC, microbial biomass carbon;
T5, soil temperature at 5 cm depth; NH4-N,
ammonium-nitrogen; W5, soil water content at 5
cm depth; SOC, soil organic carbon; CFI,
comparative fit index; Chisq, chi-square test; AIC,
Akaike’s information criterion.
the labile fraction of SOC in the form of CO2 into the atmosphere, which
reduces the ability of fine roots to absorb nutrients and the amount of
organic carbon available to microorganisms (Bryanin and Makoto,
2017; Ulery et al., 2017). Fire has a much more significant effect on the
temperature sensitivity of Ra than that of Rh in forest ecosystems, which
highlights the importance of Ra in regulating the Q10 of Rs (Atkin et al.,
2000; Boone et al., 1998; Song et al., 2019). The decomposition of soil
organic matter increases the availability of readily available substrates,
resulting in high temperature sensitivity. In contrast, the low substrate
availability may lead to a cancelling effect, which would suppress the
intrinsic temperature dependency of SOC decomposition, leading to
lower Q10 values (Conant et al., 2011; Gershenson et al., 2010; Hu et al.,
2017a).
Consistent with our third hypothesis, fire disturbance led to a pulse
increase in cumulative C efflux from Rs and its components during the
T. Hu et al. Geoderma 402 (2021) 115273

Fig. 6. Relationship between soil total respiration (Rs) and its components (Rh and Ra) and soil temperature (T5) and soil water content (W5) at 5 cm depth in control
and burned plots. Dashed lines represent the fitting model in control plots, solid lines represent the fitting model in burned plots. “n.s.” indicates “not significant.”

efflux in burned plots. Currently, the continuous decrease in forested

Table 3
areas caused by fire (Giglio et al., 2018; Liu et al., 2019) may cause soil
Summary of the best-fit regression models of soil respiration against soil tem­
carbon pools to release more CO2 into the atmosphere during the
perature and soil water content.
freeze–thaw cycle period, which has an uncertain impact on the carbon
Plot type Model α β ε ω AIC R2 cycle of regional forest ecosystems (Brooks et al., 2005; Meigs et al.,
Burned plots Ln(Rs) − 0.497 0.071 n.s. n.s. 39.95 0.82 2009).
Ln(Rh) 0.411 0.073 − 0.044 n.s. 36.67 0.86
Control plots Ln(Rs) − 0.474 0.095 n.s. n.s. 44.11 0.85
Ln(Rh) − 1.116 0.098 n.s. n.s. 55.85 0.81 4.3. Implications for forest ecosystem carbon cycling after fire disturbance
Note: Ln(SR) = α + β × T5 + ε × W5 + ω × T5 × W5. Where, SR is the soil total
Although the effect of fire on overall Rs has been studied, its effects
respiration (Rs) or soil (Rh); T5 is the soil temperature at 5 cm depth (◦ C), W5 is
the soil water content at 5 cm depth (%), and T5 × W5 is the interaction effect of on Rs components, that is, Ra and Rh, and the mechanisms involved are
T5 and W5. “n.s.” this symbol of the model was not significant in model fitting (at still poorly understood (Song et al., 2019). Elucidating the mechanisms
the p = 0.05 level), α, β, ε, ω are regression coefficients. Akaike Information of the effects of fire disturbance on Ra and Rh is vital for understanding C
Criterion (AIC), and determination coefficient (R2) are given. cycling one year after a fire (Kobziar, 2007). We demonstrated that Rh
increased significantly one year after fire disturbance while Ra decreased
spring FTC period one year after the fire, which was likely to cause significantly. In addition, there were major changes in soil temperature,
considerable loss of soil C in this area. During the spring FTC, MBC and soil water content, NH4-N, MBC, soil pH, SOC, and FR (not significant) in
FR were significantly higher in the burned plots than in the control plots. a cold temperate forest. This is the first field evidence from a cold
These results are mainly attributed to the significantly higher post-fire temperate forest ecosystem that reveals the discrepancy in the responses
temperature in burned areas, which led to the melting of snow cover of Rs components to fire disturbance. In conjunction with the significant
in this area and, combined with soil nutrient content in the burned areas, increase in the Rh:Rs ratio and the potential increase in C efflux during
promoted microbial growth and rhizosphere activities (Brooks et al., the FTC period, our findings emphasise that forest fires have a high
2005; Li et al., 2010). A previous study found that the C efflux of Rs in the potential to cause loss of forest soil C into the atmosphere against the
spring FTC period accounts for an important proportion in the non- background of increasing frequency of high severity fire and longer fire
growing season (Wang et al., 2014b). This is consistent with our result seasons. To reduce fire-related soil C loss and limit the increase in soil
that the C efflux of Rs accounted for 19% of the non-growing season C temperature via shading, proactive management of forest regeneration,
including afforestation and corresponding tending measurements,

Table 4
Estimates of the specific period cumulative CO2 efflux (g C m− 2) from soil respiration (Rs) and its components (Rh and Ra) in control and burned plots.
Growing season Non-growing season Winter period Spring FTC period Annual

Control plots Burned plots Control plots Burned plots Control plots Burned plots Control plots Burned plots Control plots Burned plots

Rs 461.80 508.37 78.38 56.45 69.89 45.63 8.49 10.82 540.18 564.82
Rh 253.68 346.96 40.95 48.56 36.49 43.23 4.47 5.33 294.63 395.52
Ra 208.13 161.41 37.42 7.89 33.41 2.40 4.02 5.49 245.55 169.30

Note: FTC = freeze–thaw cycle, the “Non-growing season” includes the “Winter period” and the “Spring FTC period”.

9
T. Hu et al.
should be carried out after forest fires.
5. Conclusions
In conclusion, our data demonstrate that Rs components exhibit a
discrepancy in their response to fire disturbance in cold temperate for­
ests in China. Forest fires significantly increased Rh and significantly
decreased the Ra. The fires also significantly increased the Rh:Ra ratio
from 0.57 in the unburned plots to 0.79 one year after fire disturbance,
and this was caused by post-fire changes in the abiotic and biotic factors
related to Rs components. We found that forest fires caused soil carbon
pools to release more CO2 into the atmosphere during the FTC. There­
fore, proactive management of forest regeneration should be undertaken
after fire forest disturbance. In addition, this one-year field study
covered a relatively short time span after a fire disturbance. Further
studies are needed to explore the long-term change patterns of Rs and its
components after fire disturbance. In particular, quantifying the effects
of vegetation restoration and by-products (such as charcoal or pyrolytic
carbon) after fire disturbance on Rs and its components is critical to
reveal whether the forest ecosystem plays the role of a carbon source or
carbon sink after fire disturbance and to improve the regional carbon
cycle model.
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Acknowledgements
This research was financially supported by the National Natural
Science Foundation (No. 32001324) and Fundamental Research Funds
for the Central Universities (No. 2572019BA03). We thank the Northern
Forest Fire Management Key Laboratory of the State Forestry and
Grassland Bureau and the National Innovation Alliance of Wildland Fire
Prevention and Control Technology of China for supporting this
research. The database can be downloaded using the following reference
and doi: Hu TX, Sun L (2021). Data for “Effects of fire on soil respiration
and its components in a Dahurian larch (Larix gmelinii) forest in north­
east China: implications for forest ecosystem carbon cycling” [Data set].
V1. CREA. http://doi.org/10.5281/zenodo.4574684 under the Creative
Commons Attribution-NonCommercial 4.0 License.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.geoderma.2021.115273.
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