Journal of

Plant Ecology Soil microbial biomass carbon and

VOLUME 3, NUMBER 3,
PAGES 175–182 nitrogen in forest ecosystems of
SEPTEMBER 2010

doi: 10.1093/jpe/rtq022 Northeast China: a comparison

between natural secondary forest
Advanced Access published
on 4 August 2010

available online at
www.jpe.oxfordjournals.org
and larch plantation
Kai Yang1, Jiaojun Zhu1,*, Min Zhang1,2, Qiaoling Yan1 and

Downloaded from http://jpe.oxfordjournals.org by Osbert Sun on August 19, 2010

Osbert Jianxin Sun3
1
Qingyuan Experimental Station of Forest Ecology, Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang
110016, China
2
Graduate School, Chinese Academy of Sciences, Beijing 100039, China
3
MOE Key Laboratory for Silviculture and Conservation, Beijing Forestry University, Beijing 100083, China
*Correspondence address. Institute of Applied Ecology, Chinese Academy of Sciences, 72 Wenhua Road,
Shenyang 110016, China. Tel: 86-24-83970342; Fax: 86-24-83970342. E-mail: jiaojunzhu@iae.ac.cn

Abstract

Aims
Natural secondary forest (NSF) and larch plantation are two of the
predominant forest types in Northeast China. However, how the
two types of forests compare in sustaining soil quality is not well un-
derstood. This study was conducted to determine how natural sec-
ondary forest and larch plantation would differ in soil microbial
biomass and soil organic matter quality.
Methods
Microbial biomass carbon (MBC), microbial biomass nitrogen (MBN),
soil organic carbon (SOC) and total nitrogen (TN) in the 0- to 15-cm
and 15- to 30-cm soil layers were investigated by making chemical
and biological measurements in the montane region of eastern Liaon-
ing Province, Northeast China, during the growing season of 2008 in
stands of NSF and Larix olgensis plantation (LOP).
Important Findings
We found that soil MBC and MBN were significantly lower in the
LOP than in the NSF. Both MBC and MBN declined significantly with
increasing soil depth in the two types of stands. The ratios of MBC to
SOC (MBC/SOC) and MBN to TN (MBN/TN) were also significantly
lower in the LOP than in the NSF. Moreover, the values of MBC,
MBC/SOC, and MBN/TN significantly varied with time and followed
a similar pattern during the growing season, all with an apparent
peak in summer. Our results indicate that NSF is better in sustaining
soil microbial biomass and nutrients than larch plantation in the tem-
perate Northeast China. This calls for cautions in large-scale conver-
sions of the native forests to coniferous plantations as a forest
management practice on concerns of sustaining soil productivity.
Keywords: microbial biomass carbon d microbial biomass
nitrogen d soil quality d temperate forests
Received: 24 March 2010 Revised: 21 June 2010 Accepted: 9 July
2010

China, NSFs account for as much as 70% of the regional forests

INTRODUCTION
Natural secondary forests (NSFs) widely occur in temperate
regions around the world (Wang and Yang 2007). They are
formed through natural regeneration following stand-replacing
disturbances of primary forests by anthropogenic activities or
by extreme natural events (Zhu and Liu 2007). In Northeast
(Hao et al. 2000), of which large areas have been turned into
larch plantations for fast timber production since the 1950s
(Wang et al. 2006). However, concerns for biological
conservation of forest resources and mitigation of climate
change in the context of global change have created large inter-
ests in assessing the changes of soil quality and carbon

Ó The Author 2010. Published by Oxford University Press on behalf of the Institute of Botany, Chinese Academy of Sciences and the Botanical Society of China.
All rights reserved. For permissions, please email: journals.permissions@oxfordjournals.org
176
sequestration capacity with conversion of natural forests to
plantations and/or other land uses (Chen and Li 2003; Shi
et al. 2009; Wang and Yang 2007; Yang et al. 2007).
There is a widespread perception that plantations are
generally inferior to naturally regenerated forest stands in nu-
trient cycling and soil quality (Burton et al. 2007; Liu et al.
1998); while this view is supported by a large body of literature
(Behera and Sahani 2003; Kasel and Bennett 2007; Yan et al.
2008; Zhang et al. 2009), there also exist contradictory exper-
imental results. For example, the study of Li et al. (2005) shows
that there is no significant difference in total soil organic car-
bon (SOC) between a pine plantation (5.59 6 0.09 kg m 2)
and a secondary mixed forest (5.68 6 0.16 kg m 2) in Puerto
Rico. Our previous study also indicates that NSF and
Larix olgensis plantation (LOP) differ in chemical composi-
tion of soil P but not in overall P availability in eastern
Liaoning Province of Northeast China (Yang et al. 2010), where
L. olgensis is an introduced species and known to have a slow
rate of litter decomposition than the tree species in the natural
forests (Liu et al. 1998). A conversion from natural forest to
coniferous plantation is generally found to reduce SOC stock
(Guo and Gifford 2002) and result in a decline in soil organic
matter quality (Behera and Sahani 2003; He et al. 2006).
Soil microbial biomass, which can be either a source or sink
of available nutrients, plays a critical role in nutrient transfor-
mation in terrestrial ecosystems (Singh et al. 1989). Any
change in the microbial biomass may affect soil organic matter
turnover. Thus, the soil microbial activity has a direct influ-
ence on ecosystem stability and fertility (Smith et al. 1993).
In general, microbial biomass can be used for assessing soil
quality of different types of vegetation (Groffman et al.
2001; Zeng et al. 2009) as well as for evaluating soil perturba-
tion and restoration (Ross et al. 1982; Smith and Paul 1990).
Forest types influence soil microbial biomass and activities
by determining the quantity and quality of organic matter
inputs (Hackl et al. 2004; Xu et al. 2008). Shifts in plant com-
munity composition may affect SOC dynamics as a result of
changes in the amount and chemical composition of plant res-
idues returned to the soil (Jin et al. 2010), which in turn influ-
ences the pool size and activity of the soil microbial biomass
(Kasel and Bennett 2007).
Besides forest types, seasonal variations of temperature,
rainfall, plant development, and organic matter accumulation
from litterfall also have great influences on soil microbial bio-
mass (Chen et al. 2005; Devi and Yadava 2006; Maithani et al.
1996; Tonon et al. 2005). Fluctuations in the size of soil micro-
bial biomass during the growing season are considered an im-
portant factor in controlling the turnover of soil carbon and
nitrogen. Therefore, information on seasonal variations in soil
microbial biomass is needed to improve our understanding on
soil nutrient transformation and availability. In temperate for-
ests, however, several studies on the seasonal dynamics of soil
microbial biomass in recent decades have reported inconsis-
tent findings (e.g. Bauhus and Barthel 1995; Gallardo and
Schlesinger 1994; Zhu and Carreiro 2004). For instance, there
Journal of Plant Ecology
are studies demonstrating that the maximum values of micro-
bial biomass may occur in summer (Gallardo and Schlesinger
1994; Zhong and Makeschin 2006) or spring (Diaz-Raviña et al.
1995; Zhu and Carreiro 2004) or show no significant seasonal
variations (Bauhus and Barthel 1995). The variable results
suggest that seasonal variations in soil microbial biomass
probably depend on the specific forest ecosystems and climatic
factors.
In this study, the concentrations of microbial biomass C and
N, soil organic C, total N and the relationships between soil or-
ganic matter and microbial biomass in the 0- to 15-cm and 15-
to 30-cm soil layers were compared between the stands of NSF
and LOP in the montane region of eastern Liaoning Province,
Northeast China. The specific objectives of this study were to
determine: (1) how NSFs and larch plantations would differ
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in soil microbial biomass and soil organic matter quality and
(2) how soil microbial biomass would vary seasonally for the
two forest types of forests under temperate climatic conditions.
We hypothesize that the stands of LOP would have markedly
lower soil microbial biomass and activity, hence slower nutrient
cycling than the NSFs because of the slower rate of litter decom-
position in L. olgensis than in the tree species in the NSFs.
MATERIALS AND METHODS
Site description and experimental design
The study was conducted at Qingyuan Experimental Station of
Forest Ecology of Institute of Applied Ecology, Chinese Acad-
emy of Sciences. The station is located in a mountainous region
in the eastern Liaoning Province, Northeast China (lati-
tude 41°51’N, longitude 124°54’E, elevation 500–1100 m
above sea level). The climate of the region is a continental
monsoon type with a humid and rainy summer and a cold
and snowy winter. Mean annual air temperature varies be-
tween 3.9 and 5.4°C with the minimum of 37.6°C in January
and the maximum of 36.5°C in July. The mean annual
precipitation ranges between 700 and 850 mm, of which
80% rain falls in June, July and August. The frost-free period
lasts for 130 days on average, with an early frost in October and
late frost in April (Zhu et al. 2007). The mean monthly temper-
ature and precipitation during the growing season of 2008 are
shown in Fig. 1. The soil type is a typical brown forest soil with
a thickness of 60–80 cm. The brown forest soil belongs to
Udalfs according to the second edition of US Soil Taxonomy
(1999).
The study site was originally occupied by primary mixed
broadleaved-Korean pine (Pinus koraiensis Sieb. et Zucc.) for-
ests until 1930s and subsequently subjected to decades of un-
regulated timber removal. A large fire in the early 1950s
cleared off the entire forests and the site was gradually replaced
by a mixture of naturally regenerating broadleaved native tree
species. Since 1960s, patches of the secondary natural forests
were cleared and replaced by larch (L. olgensis or Larix keampferi
[Lamb.] Carr.) plantations (Wang and Yang 2007).
Yang et al. | Soil microbial C and N in temperate forests 177

Precipitation Temperature GmbH, Hanau, Germany). Soil MBC and MBN were deter-
25 300
Mean air temperature ( )
mined by fumigation–extraction method (Brookes et al.

Precipitation (mm) 1
20 250 1985; Vance et al. 1987a). For each plot, three out of six sub-
200 samples (each 10.0 g fresh soil) were fumigated with ethanol-
15
150 free chloroform for 24 h at 25°C in an evacuated extractor. The
10
100 remaining samples were treated as control. Fumigated and
5 50 non-fumigated soils were extracted with 40 ml 0.5 mol l 1
K2SO4 (soil:extractant = 1:4) and shaken for 1 h on a reciprocal
0 0
Apr May Jun Jul Aug Sep Oct shaker. The extracts were filtered using Whatman No.42 filter
paper with diameter 7 cm and frozen stored at 15°C prior to
Figure 1: mean monthly air temperature and precipitation in the analysis. The total organic carbon and nitrogen in the extracts
study area in 2008 in eastern Liaoning Province, Northeast China.
were measured using a Multi N/C 3000 analyzer (Elementar
Analysensysteme GmbH). Soil pH was estimated on a 1:2.5
Our sample plots were set up on three stands of NSF and soil–water mixture. Gravimetric soil water content was calcu-
three stands of LOP of the age 16–44 years. The six stands lated from mass loss after drying for 12 h at 105°C sepa-

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have a similar topographical feature and the underlying soils rately for the 0- to 15 and 15- to 30-cm soil layers. The
are developed from the same parental materials. In each of forest floor litter was determined using six mesh traps (each
the stands, three 20 3 20-m plots were laid out in September an area of 1.0 m2) installed 0.5 m above the soil surface.
2006. The NSF plots consisted of the tree layer, the under- The fine root (<2mm in diameter) biomass was estimated from
story component and the herbage component. The tree layer eight soil cores (each an area of 28.3 cm2) in each plot. (Table 2)
includes Juglans mandshurica Maxim., Quercus mongolica All data are expressed on oven-dry (105°C) soil weight basis.
Fischer ex Ledebour, Acer mono Maxim., Fraxinus rhyncho- MBC was calculated as:
phylla Hance and Ulmus macrocarpa Hance; the understory
MBC = EC =kEC ;
component includes Acer triflorun Kom., Acer tegmentosum
Maxim, Asarum hetelotopoides var. mandshricum Maxim. and where EC = (organic C extracted from fumigated soils)
Syringa amurensis Rupr; and the herbage component includes (organic C extracted from non-fumigated soils) and
Cardamine leucantha (Tausch) OE Schulz, Allium monanthum kEC = 0.45 (Wu et al. 1990).
Maxim., Arisaema amurense Maxim., and Polygonatum involu- MBN was calculated as:
cratum Maxim. The LOP plots contain the shrub layer and the MBN = EN =kEN ;
herbage layer. The shrub layer includes A. tegmentosum, Acer
pseudo-sieboldianum (Pax.) Kom., Schisandra chinensis (Turcz.) where EN = (total N extracted from fumigated soils) (total N
Bail., Syringa wolfi Schneid. and Acanthopanax senticosus (Rupr. extracted from non-fumigated soils) and kEN = 0.54 (Brookes
et Maxim.) Harms., and the herbage layer includes et al. 1985).
C. leucantha, Rubia sylvatica Nakai and Spuriopimpinella brachy-
Statistical analysis
carpa (Kom.) Kitag.
The statistical analyses were conducted with SPSS 11.5.
Soil sampling and chemical analysis A three-way analysis of variance (ANOVA) was used to test
In spring, summer and autumn, mineral soil samples at 0- to the effects of forest type, sampling season, and soil depth on
15-cm and 15- to 30-cm depths were collected on 25 April, 18 soil microbial biomass and soil chemical properties. Pearson’s
July and 19 September 2008, respectively. The experimental correlation analysis was used to determine whether there were
design did not include winter season because of the snow cover significant interrelationships among the measured properties
and frozen soils, which made the winter sampling impractical. of the soils.
Litter layer was removed before mineral soil sampling. Nine
soil samples were randomly collected on each plot using
a stainless cylinder with 5 cm diameter. The samples collected
RESULTS
from each stand were mixed and homogenized by compositing The SOC and TN concentrations
soils at the same depth from three plots. The composited soils Concentration of SOC and C/N ratio differed significantly be-
were divided into two parts: one was sieved to pass through tween NSF stands and LOP stands (P < 0.05). Moreover, the
a 2-mm mesh immediately and stored at 4°C until analysis concentration of SOC in the 0- to 15-cm layer of the NSF stands
for the estimation of microbial biomass carbon (MBC) and mi- was significantly higher than that in the LOP stands (Table 1).
crobial biomass nitrogen (MBN), the other air-dried and However, there was no significant difference in TN concentra-
passed through a 0.25-mm sieve for SOC and total nitrogen tion between the NSF and LOP stands (Table 1). Concentra-
(TN). tions of SOC and TN decreased with depth and did not
The SOC and TN were analyzed by dry combustion on differ among the seasons (Table 1). The C/N ratios of two soil
a Vario EL III elemental analyzer (Elementar Analysensysteme layers (0–15 cm and 15- to 30-cm depth for the two forests)
178 Journal of Plant Ecology

Table 1: three-way ANOVA on SOC, TN, C/N and soil water contents in the 0- to 15 and 15- to 30-cm soil layers in the NSF and the LOP in
eastern Liaoning Province, Northeast China

Soil depth (cm) Season Forest type SOC (g kg 1) TN (g kg 1) C/N Soil water content (%) pH

0–15 Spring NSF 50.5 6 4.2 4.2 6 0.4 12.3 6 0.3 30.4 6 1.2 5.82 6 0.03
LOP 34.7 6 1.8 3.2 6 0.2 10.9 6 0.2 27.5 6 0.6 5.55 6 0.05
Summer NSF 49.8 6 3.9 3.9 6 0.4 13.1 6 0.3 34.8 6 1.1 nd
LOP 37.0 6 2.4 3.3 6 0.3 11.4 6 0.2 30.9 6 0.8 nd
Autumn NSF 46.7 6 4.6 3.7 6 0.5 12.9 6 0.2 26.7 6 0.9 nd
LOP 36.8 6 2.0 3.2 6 0.2 11.4 6 0.1 21.7 6 1.3 nd
15-30 Spring NSF 23.4 6 2.8 2.2 6 0.3 11.1 6 0.2 24.0 6 0.5 5.91 6 0.05
LOP 24.0 6 1.9 2.4 6 0.1 10.0 6 0.2 22.9 6 1.2 5.71 6 0.04
Summer NSF 22.5 6 2.0 2.0 6 0.2 11.6 6 0.2 25.2 6 1.9 nd
LOP 23.4 6 1.9 2.2 6 0.1 10.6 6 0.1 25.0 6 0.7 nd
Autumn NSF 25.8 6 3.2 2.2 6 0.3 11.7 6 0.2 19.6 6 0.8 nd

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LOP 25.5 6 0.9 2.4 6 0.1 10.4 6 0.1 18.5 6 1.2 nd
Three-way ANOVA results
Forest type * ns ** ** /
Sampling season ns ns ** ** /
Depth ** ** ** ** /
Forest 3 season ns ns ns ns /
Forest 3 depth * ns ns ** /
Season 3 depth ns ns ns ns /
Forest 3 season 3 depth ns ns ns ns /

ns denotes not significant; nd denotes not determined. Values shown in Table 1 are means 6 standard error (n = 3).
*
P < 0.05, **P < 0.01.

Table 2: forest litterfall (t hm 2 y 1) and fine root biomass (g m 2) 0-15 cm 15-30 cm

1000
to 30-cm soil depth for the NSF and the LOP in eastern Liaoning
NSF
Province, Northeast China 800 LOP
MBC concentration
(mg kg-1)

Forest type Forest litterfall Fine root biomass 600

NSF 2.99 295.3 400

LOP 1.15 114.4 200

0
spring summer autumn spring summer autumn

ranged from 10.0 to 13.1 and were significantly (P < 0.01)

80
affected by sampling seasons.
MBN concentration

*
60 *
Soil MBC, MBN, MBC/SOC and MBN/TN ratios
(mg kg-1)

Results from three-way ANOVA indicated that both forest 40 *

types and sampling seasons had significant effects on MBC
20
concentration; the soil MBN was significantly affected by forest
types, but not the sampling season. The MBC and MBN con- 0
centrations were also significantly influenced by soil layers spring summer autumn spring summer autumn
Season
(P < 0.01). In general, concentrations of MBC and MBN
showed higher values in the NSF stands than in the LOP stands
Figure 2: MBC and MBN in the soils of NSF and LOP in different sea-
across seasons (Fig. 2, P < 0.01). The values of soil MBC and sons in eastern Liaoning Province, Northeast China. Vertical bars in-
MBN were 30–38% and 18–34% lower in the 0- to 15-cm dicate standard errors of three stands.
layer and 27–37% and 7–28% lower in the 15- to 30-cm layer,
respectively, in the LOP stands than in the NSF stands across concentration was higher in summer than in spring and au-
the three seasons (Fig. 2). There was significant seasonal var- tumn at both soil depths. Soil MBN did not display apparent
iation in soil MBC for both forest types (P < 0.05); soil MBC seasonal patterns (Fig. 2).
Yang et al. | Soil microbial C and N in temperate forests 179

There were significant effects of forest types and seasons on
the ratios of MBC/SOC and MBN/TN. In addition, the interac-
tion between forest type and soil layer significantly influenced
the MBC/SOC ratio. The MBC/SOC ratio was ;0.0130 in the
NSF stands and 0.0115 in the LOP stands in the 0- to 15-cm
layer and 0.0143 in the NSF stands and 0.0085 in the LOP
stands in the 15- to 30-cm layer, across seasons. The MBN
to TN ratio, MBN/TN, was 0.0142 in the NSF stands and
0.0110 in the LOP stands for the 0- to 15-cm layer, and
0.0137 in the NSF stands and 0.0098 in the LOP stands for
the 15- to 30-cm layer, respectively, across seasons (Fig. 3).
The ratios of MBC/SOC and MBN/TN varied significantly
across sampling seasons both in NSF and LOP stands, which
were higher in summer than in spring and autumn.
in the NSF in each season. This may suggest that the NSF
stands would be of advantage to sustain soil fertility.
Many factors have been suggested to explain the effects of
vegetation type on microbial biomass in soils (Hackl et al.
2004). For example, differences in the quantity and quality
of substrate inputs via varying litter and root types and asso-
ciated nutrient specificity can be crucial drivers to influence
the soil microbial biomass (Feng et al. 2009; Jin et al. 2010).
It is well known that soil microbial biomass greatly depends
on soil organic matter as substrate; a decrease in SOC causes
reduction in soil microbial biomass (Chen et al. 2005). Thus,
the higher MBC and MBN in the NSF stands than that in
LOP stands are mainly attributable to the greater availability
of organic matter in NSF stands. This is evident from the

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Relationships between soil organic matter and
1500
NSF LOP
microbial biomass y = 11.658x - 6.453

MBC (mg kg )
-1
2
R = 0.777**
There was a strong positive correlation of SOC and MBC 1000
(R2 = 0.78, n = 36, Fig. 4) or MBN (R2 = 0.83, n = 36,
500
Fig. 4) and of TN and MBC (R2 = 0.66, n = 36, Fig. 5) or
MBN (R2 = 0.77, n = 36, Fig. 5) based on a combined data 0
0.0 20.0 40.0 60.0 80.0
set for both forest types. The two forest types did not differ sig-
-1
nificantly in the relationships of SOC with MBC or MBN and of SOC (g kg )
TN with MBC or MBN.
MBN (mg kg ) 80.0 y = 0.874x + 4.453
-1

2
60.0 R = 0.834**
DISCUSSION 40.0
Results in this study indicate that the two stand types differ 20.0
markedly in soil microbial biomass. The concentrations of 0.0
MBC and MBN were significantly lower in the LOP stands than 0.0 20.0 40.0 60.0 80.0
-1
SOC (g kg )

Figure 4: relationship between SOC and MBC or MBN for the NSF
0-15 cm 15-30 cm
2.5 and the LOP in eastern Liaoning Province, Northeast China (n = 36,
NSF i.e. three stands 3 two soil depths 3 three seasons 3 two stand types).
2.0 LOP
*
MBC/ SOC

1.5 * NSF LOP

(%)

1500 y = 128.92x + 6.7935

MBC (mg kg )

1.0 2
-1

R = 0.665**
1000
0.5
500
0.0
spring summer autumn spring summer autumn 0
0.0 2.0 4.0 6.0 8.0
2.5 TN (g kg )
-1

2.0 * *
80.0 y = 10.006x + 4.4454
MBN (mg kg )
MBN/ TN

1.5 *
-1

2
* 60.0 R = 0.766**
(%)

1.0 40.0

0.5 20.0
0.0
0.0 0.0 2.0 4.0 6.0 8.0
spring summer autumn spring summer autumn
-1
Season TN (g kg )

Figure 3: the ratios of MBC to SOC and MBN to TN in the soils of NSF Figure 5: relationship between soil TN and MBC or MBN for the NSF
and LOP in different seasons in eastern Liaoning Province, Northeast and the LOP in eastern Liaoning Province, Northeast China (n = 36, i.e.
China. Vertical bars indicate standard errors of three stands. three stands 3 two soil depths 3 three seasons 3 two stand types).
180
significant positive correlations between soil microbial biomass
and soil organic matter, in agreement with the finding by
Wang and Wang (2007). A decline of soil organic matter in
the LOP stands as compared with the NSF stands may be
explained by: (1) loss of soil organic matter due to removal
of slash and plant residues during site preparation and (2) poor
litter quantity and quality, such as pine litter in larch planta-
tion containing a large amount of recalcitrant compounds, lead
to a lower rate of decomposition and consequently slower
transformation of particulate organic matter into mineral soil
component (Leckie et al. 2004).
The concentrations of MBC in this study (192–776 mg kg-1)
fall well within the range observed in temperate forest soils
(Vance et al. 1987b; Zhong and Makeschin 2006), whereas
the concentrations of MBN (22–58 mg kg 1) are lower than
those in the temperate humid forest soils (Diaz-Raviña et al.
1995; Gómez-Luna et al. 2009). The low MBN from our study
site may be due to the rapid rate of N mineralization and N
leaching in the temperate secondary forest soils (Li et al. 2001).
The two stand types in this study exhibited a significant sea-
sonal variation in soil MBC, but soil MBN was less variable.
Seasonal variations of soil microbial biomass reflect the degree
of immobilization–mineralization of soil carbon and nitrogen.
A decrease in soil microbial biomass can result in mineraliza-
tion of nutrients, whereas an increase in microbial biomass
may lead to immobilization of nutrients (McGill et al. 1986).
In both NSF and LOP stands, MBC concentration was higher
in summer than in spring and autumn. This finding is consis-
tent with some of the previous studies in temperate forest
regions (Bohlen et al. 2001). However, the patterns of seasonal
variations in MBC in our study differ from that reported for
subtropical humid forest, in which the highest values of micro-
bial biomass are observed during the winter and the lowest
during the rainy season (Maithani et al. 1996). It has been sug-
gested that soil moisture is important in determining MBC of
temperate forest soils and is a major factor controlling the mi-
crobial biomass (Diaz-Raviña et al. 1995). In this study, the sea-
sonal dynamics of soil water content coincided with the
seasonal variations in rainfall of the study area, which was
higher in summer and lower in spring and autumn. The sea-
sonal variations in soil moisture may be responsible for the var-
iation in MBC in both types of stands, as soil water content was
significantly related to MBC concentrations (R2 = 0.74 and P <
0.01 in the NSF stands; R2 = 0.58 and P < 0.01 in the LOP
stands). Chen et al. (2003) also found similar relationships be-
tween soil water content and MBC in pine plantations in the
subtropical zone. In addition, the mean monthly temperature
was 21°C in summer and 10°C in spring or autumn in our
study area. Thus, the higher MBC in summer may well be re-
lated to the high temperature in July. A combination of in-
creased soil moisture, temperature and plant root activity in
summer all favor soil microbial biomass activities and higher
microbial biomass in the temperate forest ecosystem.
Soil MBC/SOC ratio, or microbial quotient, has been widely
used as an indicator of future changes in organic matter due to
Journal of Plant Ecology
alterations of soil conditions (Sparling 1992). This value is also
used as for comparison of soil quality parameter across soils
with different organic matter contents (Joergensen and Scheu
1999). The MBC/SOC ratio can reflect the relationship and
interactions between the MBC and mineral SOC (Insam and
Domsch 1988). In other words, the MBC/SOC ratio is inter-
preted as substrate available and the portion of total soil carbon
immobilized in microbial cells. In this, the ratios of MBC/SOC
and MBN/TN were lower in the LOP stands than in the NSF
stands in both 0- to 15-cm and 15- to 30-cm soil layers. Similar
results have been previously founds by Liu et al. (1998) and
Wang and Wang (2007). Relatively lower MBC/SOC ratio
in the LOP stands may be ascribed to the inhibition of microbial
immobilization, whereas a relatively higher MBC/SOC ratio in
the NSF stands may be explainable by more diversified organic
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substrate production as a result of mixed species.
In this study, the ratios of MBC/SOC and MBN/TN varied with
season, attaining higher values in summer and lower in spring or
autumn. The seasonal average of the MBC/SOC ratio is 1.18%
lower than the values reported for other temperate forests (1.8–
2.9%; Vance et al. 1987b). Furthermore, the seasonal average
of MBN/TN ratio (1.22%) is also lower than that of temperate
forest soils (1.6–3.0%; Zhong and Makeschin 2006), resulting
possibly from low soil organic matter (Jones et al. 2008).
In conclusion, our results suggest that the NSF is better in
sustaining the microbial biomass and soil organic matter qual-
ity than larch plantation in Northeast China. This calls for cau-
tions in large-scale conversions of the native forests to
coniferous plantations as a forest management practice on
concerns of sustainable soil productivity.
FUNDING
The Academy of Sciences / State Administration of Foreign
Experts Affairs (CAS/SAFEA) International Partnership Pro-
gram for Creative Research Teams (KZCX2-YW-445); National
Forestry Scientific Support Program (2006BAD03A0903
and 2006BAD03A0401); Non-commercial Forest Project
(200804027-05).
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