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Forest Ecology and Management 260 (2010) 422–428

Contents lists available at ScienceDirect

Forest Ecology and Management

journal homepage: www.elsevier.com/locate/foreco

Changes in soil P chemistry as affected by conversion of natural secondary forests

to larch plantations
Kai Yang a,b , Jiao-Jun Zhu a,∗ , Qiao-Ling Yan a , Osbert Jianxin Sun c
a
Qingyuan Experimental Station of Forest Ecology, Institute of Applied Ecology, Chinese Academy of Sciences, 72 Wenhua Road, Shenyang 110016, China
b
Graduate University of Chinese Academy of Sciences, Beijing 100039, China
c
MOE Key Laboratory for Silviculture and Conservation, Institute of Forestry and Climate Change Research, Beijing Forestry University, Beijing 100083, China

a r t i c l e i n f o a b s t r a c t

Article history: To assess the impact of conversion of native forests to monocultural larch plantations on soil chemical
Received 21 January 2010 properties, we compared the total and various fractions of soil phosphorus (P) and acid phosphatase
Received in revised form 28 April 2010 activity (APA) between natural secondary forests (NSF) and Larix olgensis plantations (LOP) on a montane
Accepted 29 April 2010
forest site in eastern Liaoning Province, Northeast China. We found that the concentrations of total P
(TP), inorganic P, and iron-bound P (Fe-P) were significantly higher, and the concentrations of microbial
Keywords:
biomass P (MBP), sodium bicarbonate-extractable organic P (NaHCO3 -Po), and APA were significantly
Acid phosphatase activity
lower, in the LOP stands than in the NSF stands; whilst organic P, sodium bicarbonate-extractable inor-
Larch plantation
Microbial biomass
ganic P (NaHCO3 -Pi), aluminum-bound P (Al-P) and calcium-bound P (Ca-P) were comparable between
Natural secondary forest the two forest types. Our study also showed that the ratios of MBP/TP, NaHCO3 -Pi/TP, NaHCO3 -Po/TP, and
Northeast China APA significantly varied with time during the growing season. Moreover, the concentrations of NaHCO3 -
Soil P chemistry Pi, NaHCO3 -Po, and MBP had significant (P < 0.01) and positive linear relationships with APA. Overall,
results from this study suggest that conversion of native forests to larch plantations in the region is more
likely to cause compositional change in soil P than to result in reduction in overall P availability.
Crown Copyright © 2010 Published by Elsevier B.V. All rights reserved.

1. Introduction
Mixed broadleaved-Korean pine (Pinus koraiensis Sieb. Et Zucc)
forest is one of the most important regional climax forest types
in Northeast China, of which more than 70% have become natu-
ral secondary forests after a century of timber exploitation and
historical natural disturbances (Zhu et al., 2007). To meet the grow-
ing demands for timber, extensive areas of these secondary forests
have been replaced by plantations predominantly of larch species
(Zhu et al., 2008). In the eastern Liaoning Province of Northeast
China, for example, there are about 200,000 ha of Larix olgensis
Henry plantations in the montane regions (Liu et al., 2007); these
plantations intersperse with the remnant natural forests to form
mosaic of plantation/natural secondary forest landscapes. Given
the extensive coverage and economic value, larch plantations have
in recent years attracted much attention in their role for contribut-
ing to ecosystem functioning as well as timber production. Impacts
of larch plantations on soil properties and nutrient cycling have
been a particular research interest for concerns on sustainable for-
est productivity.
∗ Corresponding author. Tel.: +86 24 83970342; fax: +86 24 83970300.
E-mail address: jiaojunzhu@iae.ac.cn (J.-J. Zhu).
Many studies have demonstrated that changes in land cover can
influence soil biological properties and nutrient cycling (Yeates et
al., 1997; Oberson et al., 2001; Sharma et al., 2004). However, the
directions of the influences can vary depending on the type of land
cover change. For example, it is reported that land cover change
from grassland to conifers results in a reduction in soil microbial
biomass, rate of soil respiration, and phosphatase activity (Chen et
al., 2000). Similarly, changes from native forests to larch planta-
tions have been found to result in significant decline in soil C and N
concentrations (Chen and Li, 2003). In contrast, a conversion from
native savanna to rice–pasture rotations causes improvement in
biological properties (Gijsman et al., 1997). In general, conversion
of natural forests to plantations has been found to result in reduc-
tion in soil nutrient availability and deterioration in microbiological
properties (Wang and Wang, 2007; Yan et al., 2008).
Compared with other major nutrients, P is known to be the
least mobile element and to have low availability to plants under
most soil conditions. The availability of P for plant growth depends
on complex geochemical and biochemical processes (Walker and
Syers, 1976). Land cover change may affect P inputs and outputs,
resulting in an altered P balance and changes in P forms (Ross et
al., 1999), and alter soil P transformation by modifying soil phys-
ical, chemical and biological properties, especially soil microbial
processes (Chen et al., 2008). Soil microorganisms play a key role
in facilitating P transformations through mineralization of P from

0378-1127/$ – see front matter. Crown Copyright © 2010 Published by Elsevier B.V. All rights reserved.
doi:10.1016/j.foreco.2010.04.038
 Author's personal copy
K. Yang et al. / Forest Ecology and Management 260 (2010) 422–428
organic sources, synthesis and release of organic P, and solubiliza-
tion of barely soluble P forms (Oberson et al., 2001). Microbes also
have the ability to rapidly immobilize considerable amounts of P
when labile C is readily available (Bünemann et al., 2004).
Most P utilized by plants is in inorganic forms (Marschner,
1995). Organic P accounts for about 15–80% of total P (Condron
and Tiessen, 2005); the organically bound P needs to be mineralized
to become available for plant uptake and utilization. Phosphatase
activity has been considered to be an important driver in this min-
eralization process and to reflect organic P mineralization potential
of soils (Chen, 2003; Criquet et al., 2004). However, the studies of
Dilly and Nannipieri (2001) and Olander and Vitousek (2000) show
that increased P availability depresses phosphatase activity in for-
est soils. Therefore the relationship between phosphatase activity
and soil P availability still remains a controversial issue.
Knowledge of the seasonal variation in soil P fractions is
important for understanding the dynamics of P availability and
mineralization in soils (Magid and Nielsen, 1992). Seasonal
changes in soil P fractions may reflect P uptake by plants and
mineralization–immobilization processes. The fluctuation in P frac-
tions during the growing season, particularly soil P transformation
process, is considered an important factor in controlling P avail-
ability and ultimately in influencing plant nutrition (Eviner et al.,
2006). Several studies on temperate ecosystems have shown a con-
siderable seasonal variation of P fractions (Chen et al., 2003; Styles
and Coxon, 2007; Zhao et al., 2009). However, there also appear
inconsistent findings (e.g. Tate et al., 1991; Magid and Nielsen,
1992; Fabre et al., 1996). For instance, the maximum values of soil
bicarbonate-extractable inorganic P and bicarbonate-extractable
organic P have been found to occur in summer (Zhao et al., 2009) or
winter (Fabre et al., 1996), or there lacks significant seasonal vari-
ation in soil resin-extractable inorganic P (Tate et al., 1991). The
variable results suggest that seasonal variation in soil P fractions
might be dependent of ecosystem types and climatic conditions.
In this study, we compared various forms of soil P between
natural secondary forests and larch plantations in the montane
region of eastern Liaoning Province of Northeast China, with aims
to understand how the two contrasting forest types may differ in
soil P properties. Detailed measurements were made on soil total
P (TP), various fractions of soil P (i.e. organic P, inorganic P, sodium
bicarbonate-extractable organic P, sodium bicarbonate-extractable
inorganic P, aluminum-bound P, iron-bound P, and calcium-bound
P), and the factors affecting P cycling and transformation (i.e. micro-
bial biomass P and acid phosphatase activity). The objectives of this
study were to determine: (1) how natural secondary forests and
larch plantations would differ in the quantity and compositional
structure of soil P; and (2) how seasonally variable the various P
fractions would be in forests of the temperate climate. Our study
was to test the hypothesis that forest conversion from native forests
to larch plantations would reduce P availability and result in sub-
stantive compositional changes of soil P.
2. Materials and methods
2.1. Site description and experimental design
The study was conducted at the Qingyuan Experimental Station
of Forest Ecology of Institute of Applied Ecology, Chinese Academy
of Sciences. The station is located in a mountainous region in
the eastern Liaoning Province, Northeast China (latitude 41◦ 51 N,
longitude 124◦ 54 E, elevation 500–1100 m above sea level). The cli-
mate of the region is a continental monsoon type with a humid and
rainy summer, and a cold and dry winter. Mean annual air tempera-
ture varies between 3.9 and 5.4 ◦ C, and the maximum and minimum
temperatures are 36.5 and −37.6 ◦ C. Annual precipitation fluctu-
423
ates between 700 and 850 mm, with more than 80% falling during
June–August. The growing season is from early April to late October
(Zhu et al., 2007). The brown forest soil belongs to Udalfs according
to the second edition of U.S. Soil Taxonomy (1999). The soil is a clay
loamy soil (sand: 25.6%, silt: 51.2%, clay: 23.2%).
The study site was originally occupied by primary mixed
broadleaved-Korean pine forests until 1930s and subsequently sub-
jected to decades of unregulated timber removal. A large fire that
occurred in the early 1950s completely cleared off the original
forests and the site was replaced by a mixture of naturally regen-
erating broadleaved native tree species. Since 1960s, patches of
the secondary natural forests were cleared for larch plantations.
Our sample plots were set up on three stands of natural secondary
forests and three stands of larch plantations of the age 16–44 years.
The six stands were selected from separate small watersheds over
an area of 100 ha, with experimental plots set up on sites of slightly
variable altitudes (from 568 to 730 m above sea level) and on gen-
tle slopes of variable aspects as strict controls of altitude and slope
aspect proven highly difficult because of the fragmented landscape
in the region. Larch plantations often occur in small patches among
the natural secondary forest stands in our study area. Where possi-
ble, we set up the larch plantation plots close to, but not necessarily
adjacent with, the natural secondary forest sites to avoid complica-
tion by variation in topographical features. Overall, the six stands
have the same topographical features and are on soils developed
from the same parental material, i.e. granite gneiss. Soil depth of the
study site varied from 60 to 80 cm and was comparable between
LOP and NSF stands. However, the two types of stands differed in
the thickness of the litter layer and A-horizon soil layer; the litter
layer depth was 4.4–6.0 cm in the LOP stands and 2.7–5.6 cm in the
NSF stands, whilst the A-horizon soil layer depth was 6.5–8.5 cm in
the LOP stands and 8–10 cm in the NSF stands; the AB-horizon soil
layer depth was about 10 cm; and the B-horizon soil depth varied
between 30 and 40 cm.
In each of the stands, three 20 m × 20 m plots were laid out in
September 2006. The plots of natural secondary forests consisted of
Juglans mandshurica, Quercus mongolica, Acer mono, Fraxinus rhyn-
chophylla and Ulmus macrocarpa in the tree layer, A. triflorun, A.
tegmentosum, A. mandshricum and Syringa amurensis in the under-
story component, and Cardamine leucantha, Allium monanthum,
Arisaema amurense, and Polygonatum involucratum in the herbage
component. The plots of larch plantations contain Acer tegmento-
sum, A. pseudo-sieboldianum, Schisandra chinensis, Syringa wolfi and
Acanthopanax senticosus in the shrub layer and Cardamine leucan-
tha, Rubia sylvatica, Spuriopimpinella brachycarpa in the herbage
layer.
2.2. Soil sampling
Soil samples were collected during the growing season in spring
(April), summer (July) and autumn (September) of 2008. Soil cores
of 5 cm in diameter were taken at nine random locations on each
plot after removing litters, and each soil core was separated into
0–15 and 15–30 cm layers. The nine samples of the same layer on
each plot were mixed as a composite sample, which was further
divided into three sets of sub-samples. One set of the sub-samples
was processed to pass through a 2 mm sieve, and stored at 4 ◦ C for
measurements of microbial biomass P (MBP) and acid phosphatase
activity (APA); one was air-dried and passed through a 1 mm
sieve for analysis of soil pH, sodium bicarbonate-extractable inor-
ganic P (NaHCO3 -Pi) and sodium bicarbonate-extractable organic
P (NaHCO3 -Po); the remaining sub-sample was air-dried, and
homogenized and passed through a 0.25 mm sieve for analyses of
soil organic C (SOC), total N (TN), TP, organic P and fractions of inor-
ganic P. The basic soil physical and chemical properties, seasonal
variations in soil water content (%, v/v) in the natural secondary
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424 K. Yang et al. / Forest Ecology and Management 260 (2010) 422–428

Table 1
Soil physiochemical characteristics in the natural secondary forest (NSF) and the Larix olgensis plantation (LOP).

Soil depth (cm) Forest type SOC (g kg−1 ) TN (g kg−1 ) C/N pH Soil bulk density (g cm−3 )

NSF 50.45 ± 4.25 a

4.21 ± 0.52 12.3 ± 0.2 a
5.82 ± 0.03 a
1.32 ± 0.03
0–15
LOP 34.70 ± 1.86 3.20 ± 0.17 10.9 ± 0.1 5.55 ± 0.05 1.31 ± 0.01

NSF 23.42 ± 2.63 2.18 ± 0.32 11.1 ± 0.3a 5.91 ± 0.05a 1.45 ± 0.03
15–30
LOP 23.97 ± 1.90 2.38 ± 0.16 10.0 ± 0.1 5.71 ± 0.04 1.49 ± 0.02

SOC: soil organic C; TN: soil total N. Values shown in tables are means ± standard errors (n = 3).
a
Indicate significant differences between the natural secondary forest and the Larix olgensis plantation of the corresponding depth at P = 0.05 levels.

forest stands and larch plantations are listed in Tables 1 and 2,

respectively.

2.3. Soil analyses

Soil pH was estimated on a 1:2.5 soil–water mixture. Soil bulk

density samples were measured at each soil depth using a known
volume metal container. Bulk density measurements were per-
formed by drying the soil at 105 ◦ C for 12 h and then reweighing
the samples. Gravimetric soil water content was calculated from
mass loss after drying for 12 h at 105 ◦ C in 0–15 and 15–30 cm
of soil layers, respectively. The SOC and TN were analyzed by dry
combustion on a Vario EL III elemental analyzer (Germany). The
TP was determined following H2 SO4 –HClO4 digestion (Olsen and
Sommers, 1982). Organic P was determined by ignition method of
Saunders and Williams (1955), and inorganic P was computed as
the difference between the TP and organic P.
Soil MBP was determined by the chloroform fumigation extrac-
tion (soil: extractant [0.5 mol L−1 NaHCO3 ] ratio 1:20) method. The
MBP was estimated from the relationship EP /KEP , where EP is the
difference between inorganic P extracted from fumigated and unfu-
Fig. 1. Soil total P (TP), organic P (Po), and inorganic P (Pi) in the natural secondary
migated soils and KEP = 0.40 (Brookes et al., 1982). The correction
forest (NSF) and the Larix olgensis plantation (LOP) at different seasons (standard
for chloroform released P that was absorbed by soil colloids during errors from 3 samples shown by vertical bars).
extraction was made by adding 25 mg P kg−1 soil during extraction
and then corrected for its recovery (Brookes et al., 1982). Soil APA
was assayed by the method of Tabatabai (1994) at pH 6.5. Soil acid 2.4. Statistical analyses
phosphatase activity was assayed at pH 6.5 rather than at the soil
pH because of: (1) that the difference in soil pH between the NSF All reported values are expressed on air-dry soil weight basis.
stands and the LOP stands is about 0.2–0.3 units apart; and (2) that Statistical analyses were performed by using the SPSS 11.5 for Win-
the soil pH value of 6.5 is optimum for acid phosphatase activity. dows. One-way ANOVA was performed by soil layer, and LSD’s
The NaHCO3 -Pi and NaHCO3 -Po were determined by Bowman (least significant difference) test was applied post hoc to distin-
and Cole (1978) method. The fractionation of inorganic P in soils guish the differences of soil physiochemical properties between the
was determined by a modified Chang and Jackson procedure (as NSF and the LOP. Repeated measures analysis of variance (ANOVA)
described by Petersen and Corey, 1966). Soil samples were sequen- was used to test the effects of forest types on various P variables
tially extracted with 0.5 mol L−1 NH4 F for aluminum-bound P (Al-P) of the same soil layer across the growing season. When there were
determination, 0.1 mol L−1 NaOH for iron-bound P (Fe-P), and significant interactions between the forest type and sampling sea-
0.5 mol L−1 H2 SO4 for calcium-bound P (Ca-P). All phosphate ions son, one-way ANOVA was used to test the effects of vegetation
were determined by the ascorbic acid molybdenum antimony spec- type on soil P variables by sampling season and the effects of sam-
trophotometric method. pling season on soil P variables by forest types. Data were tested
for normality and homogeneity of error variances to determine
whether transformations were needed. Pearson linear correlations
Table 2 were used to assess the relationships between acid phosphatase
Seasonal variations in soil water content (%, v/v) in the natural secondary forest activities and active soil P fractions (including MBP, NaHCO3 -Po
(NSF) and the Larix olgensis plantation (LOP). and NaHCO3 -Pi).
Soil depth (cm) Sampling season Forest type

NSF LOP 3. Results

Spring 30.4 ± 1.2a 27.5 ± 0.6
0–15 Summer 34.8 ± 1.1a 30.9 ± 0.8 3.1. Total P, organic P, and inorganic P fractions
Autumn 26.7 ± 0.9a 21.7 ± 1.3

Spring 24.0 ± 0.5 22.9 ± 1.2

Concentrations of total P and inorganic P were significantly
15–30 Summer 25.2 ± 1.9 25.0 ± 0.7 higher in the LOP stands than in the NSF stands in both 0–15 and
Autumn 19.6 ± 0.8 18.5 ± 1.2 15–30 cm soil layers (Fig. 1, P < 0.05), whilst the organic P concen-
Values shown in tables are means ± standard errors (n = 3). tration did not vary between the NSF and LOP stands each sampling
a
Indicate significant differences between the natural secondary forest and the season (Fig. 1). Sampling season had no significant effect on soil
Larix olgensis plantation in same season of the corresponding depth at P = 0.05 levels. total P, inorganic P and organic P. Across the three seasons, soil
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K. Yang et al. / Forest Ecology and Management 260 (2010) 422–428 425

Fig. 3. Soil microbial biomass P (MBP), sodium bicarbonate-extractable organic P

Fig. 2. Soil Al-P, Fe-P and Ca-P in natural secondary forest (NSF) and Larix olgensis (NaHCO3 -Po) and sodium bicarbonate-extractable inorganic P (NaHCO3 -Pi) in the
plantation (LOP) at different seasons (standard errors from 3 samples shown by natural secondary forest (NSF) and the Larix olgensis plantation (LOP) at different
vertical bars). seasons (standard errors from 3 samples shown by vertical bars).

organic P accounted for 53–64% of total P in both types of stands,
with the NSF stands having higher ratios of organic P to total P than
LOP stands (Table 3).
For inorganic P fractions, the NSF stands had significantly lower
Fe-P concentration than the LOP stands. Specifically, the average
concentrations of soil Fe-P in the NSF stands were 51.3 mg kg−1
lower in the 0–15 cm soil layer and 35.2 mg kg−1 lower in the
15–30 cm soil layer, respectively, than in the LOP stands across
seasons (Table 3 and Fig. 2). There were no significant difference
between NSF and LOP for Al-P and Ca-P concentrations in spring
and autumn. Effects of forest type on Al-P and Ca-P concentrations
were significant in summer in the 0–15 cm soil layer (P < 0.05). The
NSF stands had significantly higher Al-P concentration and lower
Ca-P concentration than the LOP stands. All the inorganic P fractions
were lower in the 15–30 cm soil layer than in the 0–15 cm soil layer
for both forest types (Fig. 2). The seasonal average concentrations
of the inorganic P fractions followed a descending order of: Fe-
P > Ca-P > Al-P. There was no significant effect of sampling season
on concentrations of Al-P, Fe-P and Ca-P (Fig. 2).
3.2. Soil MBP, NaHCO3 -Po, and NaHCO3 -Pi
MBP and NaHCO3 -Po concentrations were higher in the NSF
stands than in the LOP stands in the 0–15 cm soil layer in spring
and summer, and decreased with soil depth (Fig. 3). In the 0–15 cm
soil layer, the concentrations of MBP and NaHCO3 -Po averaged
across seasons were 14.2 and 4.2 mg kg−1 higher in the NSF stands
than in the LOP stands, respectively (Table 3). The NaHCO3 -Pi con-
centration did not differ between the NSF and LOP stands across
seasons in both 0–15 and 15–30 cm soil layers. There was a signifi-
cant effect of sampling season on MBP in both soil layers in the LOP
stands, but not in the NSF stands. In the LOP stands, MBP concen-
tration was higher in summer than in spring and autumn, with the
maximum difference being 11.1 mg kg−1 in the 0–15 cm soil layer
and 5.2 mg kg−1 in the 15–30 cm soil layer, respectively (Fig. 3).
There was also a significant seasonal variation in soil NaHCO3 -
Po in the both types of stands in the two soil layers; significantly
higher NaHCO3 -Po concentrations were also observed in summer
than in spring and autumn (Fig. 3) (P < 0.05). In the NSF stands,
the maximum seasonal difference in NaHCO3 -Po concentration
was 13.9 mg kg−1 in the 0–15 cm soil layer and 10.7 mg kg−1 in
the 15–30 cm soil layer, respectively; whilst in the LOP stands,
the maximum seasonal difference in NaHCO3 -Po concentration
was 3.0 mg kg−1 in the 0–15 cm soil layer and 3.6 mg kg−1 in the
15–30 cm soil layer, respectively. The NaHCO3 -Pi concentrations
were relatively stable across seasons in both NSF and LOP stands.
Effects of forest type on MBP/TP and NaHCO3 -Po/TP ratios were
significant across seasons in both soil layers (Fig. 4): in the 0–15 cm
soil layer, the MBP/TP was 5.5% in the NSF stands and 2.6% in the LOP
stands; where as in the 15–30 cm soil layer, the MBP/TP was 1.8% in
the NSF stands and 1.4% in the LOP stands. The NaHCO3 -Po/TP was
2.8% in the NSF stands and 1.6% in the LOP stands in the 0–15 cm
layer, and 1.7% in the NSF stands and 1.0% in the LOP stands in the
15–30 cm layer, respectively, across seasons (Fig. 4). The difference
in NaHCO3 -Pi/TP between the NSF and LOP stands was dependent
on the sampling season. The NaHCO3 -Pi/TP was generally higher in
the NSF stands than in the LOP stands. The MBP/TP and NaHCO3 -
Po/TP showed the similar seasonal patterns as MBP and NaHCO3 -Po

Table 3
The average (n = 9) of soil P fractions (mg kg−1 ) and acid phosphatase activities (mg kg−1 h−1 ) in the natural secondary forest (NSF) and the Larix olgensis plantation (LOP)
across seasons.

Soil depth (cm) Forest type TP Po Pi Al-P Fe-P Ca-P MBP NaHCO3 -Po NaHCO3 -Pi APA

NSF 741 475 272 17.8 84.9 70.0 40.3 19.8 10.2 818
0–15
LOP 1025 543 481 15.2 136.2 78.3 26.1 15.6 10.1 673

NSF 627 376 251 13.9 68.0 53.5 12.5 10.1 4.5 448
15–30
LOP 897 495 402 11.6 103.2 53.4 11.8 8.3 5.2 277

TP: total P; Po: organic P; Pi: inorganic P; MBP: microbial biomass P; NaHCO3 -Po: sodium bicarbonate-extractable organic P; NaHCO3 -Pi: sodium bicarbonate-extractable
inorganic P; APA: acid phosphatase activity.
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426 K. Yang et al. / Forest Ecology and Management 260 (2010) 422–428
Fig. 4. Percent (%) of various active P fractions of soils in the natural secondary
forest (NSF) and the Larix olgensis plantation (LOP) at different seasons (standard
errors from 3 samples shown by vertical bars).
Fig. 5. Seasonal variation in acid phosphatase activity (APA) of soils in the natural
secondary forest (NSF) and the Larix olgensis plantation (LOP) (standard errors from
3 samples shown by vertical bars).
in the two types of stands. There was a significant effect of sam-
pling season on NaHCO3 -Pi/TP in both soil layers in the NSF stands
(P < 0.05), but not in the LOP stands.
3.3. Soil APA
The value of APA was significantly greater in the NSF stands
than in the LOP regardless of soil depth and sampling season (with
an exception of the APA in the 0–15 cm soil layer in autumn),
and was higher in the 0–15 cm soil layer than in the 15–30 cm
soil layer (Fig. 5). The average value of soil APA in the NSF
stands was 145 mg kg−1 h−1 greater in the 0–15 cm soil layer, and
171 mg kg−1 h−1 greater in the 15–30 cm soil layer, respectively,
than in the LOP stands across seasons (Table 3). There was a signif-
icant effect of sampling season on APA in the 15–30 cm soil layer,
with the maximum APA found in autumn, and the minimum in
summer for both forest types. The maximum seasonal differences
in APA were 189 mg kg−1 h−1 in the NSF stands and 124 mg kg−1 h−1
in the LOP stands, respectively.
3.4. Relationships between active soil P fractions and APA
APA was positively correlated with MBP based on a combined
dataset for both forest types (r = 0.822, P < 0.01, n = 36). Significant
and positive correlations were also found for APA-NaHCO3 -Po and
APA-NaHCO3 -Pi (Fig. 6). The two forest types did not differ in the
relationships of APA with MBP, NaHCO3 -Po, and NaHCO3 -Pi.
Fig. 6. Relationship between acid phosphatase activity and microbial biomass
P (A), sodium bicarbonate-extractable organic P (NaHCO3 -Po) (B) and sodium
bicarbonate-extractable inorganic P (NaHCO3 -Pi) (C) for the natural secondary forest
and the Larix olgensis plantation in three seasons (n = 36, i.e. three plots × two soil
depths × three seasons × two forest types). *, ** Significant at P < 0.05 and P < 0.01
respectively.
4. Discussion
This study has shown that the two forest types differed markedly
in soil P chemistry. We found that the concentrations of total P, inor-
ganic P, and Fe-P were significantly higher, and the concentrations
of MBP, NaHCO3 -Po, and APA were significantly lower, in the LOP
stands than in the NSF stands; whilst concentrations of organic P,
NaHCO3 -Pi, Al-P, and Ca-P were comparable between the two forest
types. The finding of generally greater amount of soil P in larch plan-
tation than in natural secondary stand was contrary to our initial
expectation of a reduced soil P availability with land cover change
from native forests to coniferous plantations.
Many factors may attribute to differences between different for-
est types in soil P. For example, differences in the quantity of plant
litter inputs and chemical composition of the litter may be crucial
drivers to alter the soil P concentration (Ordóñez et al., 2008; Redel
et al., 2008). In addition, plant species composition and root type
can also influence soil P status in the ecosystem scale (Grierson
and Adams, 2000). In this study, the difference in total P between
the two forest types may be explained firstly by more abundant
understory shrubs in the LOP (personal observation) and secondly
by greater organic matter input as decaying roots and slash when
original forest trees were cleared for planting L. olgensis. It has been
shown by Liu et al. (1998) that the role of the shrub species and
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K. Yang et al. / Forest Ecology and Management 260 (2010) 422–428
understory vegetation is very important in sustaining soil fertility
in larch plantations. The organic matter inputs from leave decom-
positions can also affect P cycling in the soil (Wood et al., 2009).
The results of this study also revealed seasonal patterns of total
P, organic P and inorganic P for both two forest types, i.e. the con-
centrations of total P, organic P and inorganic P were relatively
stable over time in both NSF and LOP stands. This result was consis-
tent with previous studies in temperate forest ecosystems by Fabre
et al. (1996).
Since P as orthophosphate is largely the preferred source for
plant P uptake, knowledge of the inorganic P fractions within
soils is essential to understanding P bioavailability (McDowell and
Stewart, 2006). It should be noted that the low pH values of the
soils (5.55–5.91) might also influence the behavior of P due to P
binding by Al and Fe in acid soils (Richter et al., 2006). The high
proportion of Fe-P in the two forest types in our study was consis-
tent with previous research in Chinese fir forest soils (Chen, 2003).
The relative distribution of inorganic P fractions for the two for-
est types in decreasing order was Fe-P, Ca-P and Al-P in our study,
which differed with the finding of Barroso and Nahas (2005) that
the percentages of the P fractions decreased in the order of Fe-P,
Al-P and Ca-P in Brazilian soils. Al-P content in the two forest types
was lower than that in Brazilian soils because the former is a less
weathered soil. In the current study, the concentrations of inorganic
P fractions were significantly affected by forest types, especially
the Fe-P fraction. Different phosphate-solubilizing microorganisms
between the NSF and LOP stands may be responsible for Fe-P con-
centrations (Barroso and Nahas, 2005). Furthermore, the Fe oxide
concentration could dominate the mechanisms governing values of
Fe-P in temperate forest ecosystems (Darke and Walbridge, 2000).
The results support the viewpoint that the soil inorganic P content
varies with different forest ecosystems (Redel et al., 2008). Gener-
ally, Al-P increases with declining soil pH (Chen, 2003). But, in our
study, Al-P concentration was not significant difference between
the NSF and LOP stands, albeit significantly lower soil pH in the
LOP stands than that in the NSF stands. The difference in soil pH
between the NSF and LOP stands (0.2–0.3) could be too small to
impose significant effects.
Soil microbial biomass is an important component in most ter-
restrial ecosystems. A small change in microbial biomass could have
a major impact on plant nutrient availability, at least in the short-
term. Furthermore, the P held in soil microbial biomass is a readily
available P source for plants. In this study, soil MBP in the LOP was
lower than in the NSF stands, supporting the finding of Wang and
Wang (2007) that soil microbial biomass in pure coniferous planta-
tions was lower than in natural broadleaved forests. Differences in
the quantity and quality of organic matter returned to soil between
the NSF and LOP stands, especially labile organic matter (includ-
ing microbial biomass C), could be responsible for changes in soil
MBP. On the other hand, changes in MBP between the NSF and
the LOP stands observed in this study could also be influenced by
microclimatic modifications, such as decreases in soil water con-
tent due to increased evaporation or hydraulic redistribution in
the LOP stands. The results of a greater NaHCO3 -Po concentration
in the NSF stands could be explained by faster microbial decom-
position of plant litters in this forest type. Differences in the active
soil P fractions (including MBP and NaHCO3 -Po) between the NSF
and LOP stands implicate that soil organic P status might deteri-
orate following conversion of natural secondary forests to larch
plantations.
We found that the ratios of MBP/TP, NaHCO3 -Po/TP and
NaHCO3 -Pi/TP were generally higher in soils of the NSF stands
than of the LOP stands across seasons, and that MBP/TP
and NaHCO3 -Po/TP were significantly higher in summer than
in spring or autumn, which suggest that it is important to
take temporal variations into consideration in the sampling
427
strategy. The value of soil bioavailable P depends on the fac-
tors such as mineralization–immobilization of organic P, and
adsorption–desorption and precipitation–dissolution of inorganic
P (Frossard et al., 2000). Recently, Styles and Coxon (2007) shows
that labile inorganic P concentration follows a distinct seasonal
trend of winter maximum and summer minimum in western Ire-
land. However, we found that the NaHCO3 -Po/TP ratio was higher
in summer and lower in spring or autumn in our study, contrary to
the findings of Chen et al. (2003) and Styles and Coxon (2007). This
may be because the NaHCO3 -Po fraction was affected by microbial
biomass P and plant uptake (Fabre et al., 1996). Additionally, cli-
matic factors such as soil and air temperatures, soil water content
can also influence the seasonal dynamics of MBP and NaHCO3 -Po
fraction (Magid and Nielsen, 1992).
Phosphatase activity plays an important role in P bioavailabil-
ity. It has been suggested as an indicator of P availability in soils
and soil quality (Chen, 2003). Soil acid phosphatase activities in this
study falls well within the range of the values reported in other for-
est ecosystems (Chen et al., 2000). Low soil APA activity observed
in LOP stands than those in NSF stands were related to decrease
soil organic matter and microbial biomass. We found that seasonal
variations in acid phosphatase activities were small for both for-
est types and the maximum acid phosphatase activities in NSF and
LOP occurred in autumn. Criquet et al. (2004) also reported a sea-
sonal increase in the acid phosphatase activities during the cold
period in an evergreen oak litter of the French Mediterranean area.
In this study, acid phosphatase activities were significantly corre-
lated with MBP, consistent with the finding of Šnajdr et al. (2008)
that soil acid phosphatase positively correlated with total micro-
bial biomass in Quercus petraea forest soil. In our study, APA was
also positively correlated with NaHCO3 -Po and NaHCO3 -Pi, which
is consistent with the previous finding by Grierson and Adams
(2000), but differed with findings of Dilly and Nannipieri (2001)
and Olander and Vitousek (2000). These contradicting findings may
attribute to differences in the concentration of NaHCO3 -Pi because
the concentrations of NaHCO3 -Pi in the range of 3.4–11.8 mg kg−1
in this study were much lower than in other studies (e.g. Dick,
1994).
In summary, results from this study showed that the contents
of soil MBP and NaHCO3 -Po were significantly higher in the native
forest stands than in the larch plantation, whilst there was no sig-
nificant difference in NaHCO3 -Pi between the two forest types,
indicating that the native forests are better in maintaining soil
organic P fertility than larch plantations. Our results also suggest
that conversion of native forests to larch plantations in the region
is more likely to cause compositional change in soil P than to result
in reduction in overall P availability.
Acknowledgements
This research was supported by the 11th-Five-Year National
Science and Technology Research Projects (2006BAD03A0401,
2006BAD03A0903) and the CAS/SAFEA International Partner-
ship Program for Creative Research Teams (KZCX2-YW-445) and
National Non-commercial Forests Project (200804027-05).
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