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and larch plantation
Kai Yang1, Jiaojun Zhu1,*, Min Zhang1,2, Qiaoling Yan1 and
Abstract
Aims increasing soil depth in the two types of stands. The ratios of MBC to
Natural secondary forest (NSF) and larch plantation are two of the SOC (MBC/SOC) and MBN to TN (MBN/TN) were also significantly
predominant forest types in Northeast China. However, how the lower in the LOP than in the NSF. Moreover, the values of MBC,
two types of forests compare in sustaining soil quality is not well un- MBC/SOC, and MBN/TN significantly varied with time and followed
derstood. This study was conducted to determine how natural sec- a similar pattern during the growing season, all with an apparent
ondary forest and larch plantation would differ in soil microbial peak in summer. Our results indicate that NSF is better in sustaining
biomass and soil organic matter quality. soil microbial biomass and nutrients than larch plantation in the tem-
perate Northeast China. This calls for cautions in large-scale conver-
Methods
sions of the native forests to coniferous plantations as a forest
Microbial biomass carbon (MBC), microbial biomass nitrogen (MBN),
management practice on concerns of sustaining soil productivity.
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
Keywords: microbial biomass carbon d microbial biomass
and biological measurements in the montane region of eastern Liaon-
nitrogen d soil quality d temperate forests
ing Province, Northeast China, during the growing season of 2008 in
stands of NSF and Larix olgensis plantation (LOP). Received: 24 March 2010 Revised: 21 June 2010 Accepted: 9 July
2010
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
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.
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176 Journal of Plant Ecology
sequestration capacity with conversion of natural forests to are studies demonstrating that the maximum values of micro-
plantations and/or other land uses (Chen and Li 2003; Shi bial biomass may occur in summer (Gallardo and Schlesinger
et al. 2009; Wang and Yang 2007; Yang et al. 2007). 1994; Zhong and Makeschin 2006) or spring (Diaz-Raviña et al.
There is a widespread perception that plantations are 1995; Zhu and Carreiro 2004) or show no significant seasonal
generally inferior to naturally regenerated forest stands in nu- variations (Bauhus and Barthel 1995). The variable results
trient cycling and soil quality (Burton et al. 2007; Liu et al. suggest that seasonal variations in soil microbial biomass
1998); while this view is supported by a large body of literature probably depend on the specific forest ecosystems and climatic
(Behera and Sahani 2003; Kasel and Bennett 2007; Yan et al. factors.
2008; Zhang et al. 2009), there also exist contradictory exper- In this study, the concentrations of microbial biomass C and
imental results. For example, the study of Li et al. (2005) shows N, soil organic C, total N and the relationships between soil or-
that there is no significant difference in total soil organic car- ganic matter and microbial biomass in the 0- to 15-cm and 15-
bon (SOC) between a pine plantation (5.59 6 0.09 kg m 2) to 30-cm soil layers were compared between the stands of NSF
and a secondary mixed forest (5.68 6 0.16 kg m 2) in Puerto and LOP in the montane region of eastern Liaoning Province,
Rico. Our previous study also indicates that NSF and Northeast China. The specific objectives of this study were to
Larix olgensis plantation (LOP) differ in chemical composi- determine: (1) how NSFs and larch plantations would differ
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-
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
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.
0
spring summer autumn spring summer autumn
*
60 *
Soil MBC, MBN, MBC/SOC and MBN/TN ratios
(mg kg-1)
There were significant effects of forest types and seasons on in the NSF in each season. This may suggest that the NSF
the ratios of MBC/SOC and MBN/TN. In addition, the interac- stands would be of advantage to sustain soil fertility.
tion between forest type and soil layer significantly influenced Many factors have been suggested to explain the effects of
the MBC/SOC ratio. The MBC/SOC ratio was ;0.0130 in the vegetation type on microbial biomass in soils (Hackl et al.
NSF stands and 0.0115 in the LOP stands in the 0- to 15-cm 2004). For example, differences in the quantity and quality
layer and 0.0143 in the NSF stands and 0.0085 in the LOP of substrate inputs via varying litter and root types and asso-
stands in the 15- to 30-cm layer, across seasons. The MBN ciated nutrient specificity can be crucial drivers to influence
to TN ratio, MBN/TN, was 0.0142 in the NSF stands and the soil microbial biomass (Feng et al. 2009; Jin et al. 2010).
0.0110 in the LOP stands for the 0- to 15-cm layer, and It is well known that soil microbial biomass greatly depends
0.0137 in the NSF stands and 0.0098 in the LOP stands for on soil organic matter as substrate; a decrease in SOC causes
the 15- to 30-cm layer, respectively, across seasons (Fig. 3). reduction in soil microbial biomass (Chen et al. 2005). Thus,
The ratios of MBC/SOC and MBN/TN varied significantly the higher MBC and MBN in the NSF stands than that in
across sampling seasons both in NSF and LOP stands, which LOP stands are mainly attributable to the greater availability
were higher in summer than in spring and autumn. of organic matter in NSF stands. This is evident from the
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
1.0 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 Journal of Plant Ecology
significant positive correlations between soil microbial biomass alterations of soil conditions (Sparling 1992). This value is also
and soil organic matter, in agreement with the finding by used as for comparison of soil quality parameter across soils
Wang and Wang (2007). A decline of soil organic matter in with different organic matter contents (Joergensen and Scheu
the LOP stands as compared with the NSF stands may be 1999). The MBC/SOC ratio can reflect the relationship and
explained by: (1) loss of soil organic matter due to removal interactions between the MBC and mineral SOC (Insam and
of slash and plant residues during site preparation and (2) poor Domsch 1988). In other words, the MBC/SOC ratio is inter-
litter quantity and quality, such as pine litter in larch planta- preted as substrate available and the portion of total soil carbon
tion containing a large amount of recalcitrant compounds, lead immobilized in microbial cells. In this, the ratios of MBC/SOC
to a lower rate of decomposition and consequently slower and MBN/TN were lower in the LOP stands than in the NSF
transformation of particulate organic matter into mineral soil stands in both 0- to 15-cm and 15- to 30-cm soil layers. Similar
component (Leckie et al. 2004). results have been previously founds by Liu et al. (1998) and
The concentrations of MBC in this study (192–776 mg kg-1) Wang and Wang (2007). Relatively lower MBC/SOC ratio
fall well within the range observed in temperate forest soils in the LOP stands may be ascribed to the inhibition of microbial
(Vance et al. 1987b; Zhong and Makeschin 2006), whereas immobilization, whereas a relatively higher MBC/SOC ratio in
the concentrations of MBN (22–58 mg kg 1) are lower than the NSF stands may be explainable by more diversified organic
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