Pedosphere 25(4): 489–500, 2015

ISSN 1002-0160/CN 32-1315/P

c 2015 Soil Science Society of China
Published by Elsevier B.V. and Science Press

Soil Enzyme Activities on Eroded Slopes in the Sichuan

Basin, China
NIE Xiaojun1 , ZHANG Jianhui2,∗ and GAO Han1
1 Land Research Integrated Monitoring and Sustainable Utilization in Central China, Field Science Observation & Research Base of

the Ministry of Land and Resources of China, School of Surveying and Land Information Engineering, Henan Polytechnic University,
Jiaozuo 454003 (China)
2 Key Laboratory of Mountain Surface Processes and Ecological Regulation, Institute of Mountain Hazards and Environment, Chinese

Academy of Sciences, Chengdu 610041 (China)

(Received December 15, 2014; revised May 21, 2015)

ABSTRACT
Determining how soil erosion affects enzyme activity may enhance our understanding of soil degradation on eroded agricultural
landscapes. This study assessed the changes in enzyme activity with slope position and erosion type by selecting water and tillage
erosion-dominated slopes and performing analyses using the 137 Cs technique. The 137 Cs data revealed that soil loss occurred in the
upper section of the two eroded slope types, while soil accumulation occurred in the lower section. The invertase activity increased
downslope and exhibited a pattern similar to the 137 Cs data. The spatial patterns of urease and alkaline phosphatase activities were
similar to the 137 Cs inventories on the water and tillage erosion-dominated slopes, respectively. On both the eroded slope types, the
invertase activity and soil organic carbon content were correlated, but no correlation was observed between the alkaline phosphatase
activity and total phosphorus content. Nevertheless, the urease activity was correlated with the total nitrogen content only on the
water erosion-dominated slopes. The enzyme activity-to-microbial biomass carbon ratios indicated high activities of invertase and
urease but low activity of phosphatase on the water erosion-dominated slopes compared with the tillage erosion-dominated slopes.
Both the invertase activity and the invertase activity-to-microbial biomass carbon ratio varied with the slope position. Changes in the
urease activity-to-microbial biomass carbon ratio were significantly affected by the erosion type. These suggested that the dynamics of
the invertase activity were linked to soil redistribution on the two eroded slope types, whereas the dynamics of the urease and alkaline
phosphatase activities were associated with soil redistribution only on the water or tillage erosion-dominated slopes, respectively. The
erosion type had an obvious effect on the activities of invertase, urease and alkaline phosphatase. Soil redistribution might influence
the involvement of urease in the N cycle and alkaline phosphatase in the P cycle. Thus, enzyme activity-to-microbial biomass ratios
may be used to better evaluate microbiological activity in eroded soils.
Key Words: 137 Cs technique, microbial biomass C, N cycle, P cycle, soil organic C, soil redistribution, tillage erosion, water erosion

Citation: Nie X J, Zhang J H, Gao H. 2015. Soil enzyme activities on eroded slopes in the Sichuan Basin, China. Pedosphere. 25(4):
489–500.

Soil erosion is a primary cause of land degrada-
tion in mountainous and hilly areas of China, such as
the Qinghai-Tibet Plateau, the Loess Plateau, and the
black soil region in Northeast China (Zhao et al., 2013;
Li et al., 2014; Liu et al., 2014). Effective measures
such as afforestation programs and land conversion
programs on sloping landscapes have been undertaken
to combat erosion-induced soil degradation in these a-
reas. However, in the hilly areas with intense farming,
such as the Sichuan Basin, which is known as China’s
breadbasket, soil erosion is still a serious impediment
to the development of sustainable agriculture. Inten-
sive soil erosion mostly occurs on sloping agricultur-
al landscapes in the Sichuan Basin. Soil erosion from
sloping landscapes has been estimated to account for
60%–80% of the total soil erosion in the region (Chen et
∗ Corresponding author. E-mail: zjh@imde.ac.cn.
al., 2000). To minimize the risk of water erosion, most
of the hillslopes in the region have been apportioned
by local farmers into slope segments with different gra-
dients. Unfortunately, this agricultural practice has led
to intensive tillage erosion. Field evidence from diffe-
rent regions of the world has emphasized that water
and tillage erosion processes are two major soil erosion
processes on tilled sloping landscapes (Van Oost et al.,
2006; Garcı́a-Orenes et al., 2009; Mandal and Sharda,
2013; Lieskovský and Kenderessy, 2014). Moreover, the
impacts of tillage and water erosion on temperate agri-
cultural soils are of approximately the same order of
magnitude (Van Oost et al., 2006). Currently, the nega-
tive effect of tillage erosion on soil quality is drawing
more attention in the Sichuan Basin.
Among the available soil quality indicators, soil or-
490
ganic carbon (SOC), nitrogen (N) and phosphorus (P)
have been widely studied in eroded soils. There is in-
creasing evidence that water erosion accelerates the
losses of SOC, N and P from soils (Polyakov and Lal
2004; Nie et al., 2013a) and that tillage erosion main-
ly increases the spatial variability of these nutrients
(Heckrath et al., 2005). Studies conducted previously
in the Sichuan Basin further demonstrated that the
effects of water and tillage erosion on the spatial dis-
tribution of SOC, N and P on sloping landscapes are
different due to selective soil transport by water and
complete soil disturbance by tillage (Zhang et al., 2006;
Ni et al., 2007; Cerdà et al., 2007, 2009, 2010; Haile
and Fetene, 2012; Prokop and Porȩba, 2012). Howe-
ver, little is known about the effect of soil erosion on
soil microbial properties, such as enzyme activity, in
relation to nutrient dynamics in the Sichuan Basin.
Soil enzyme activity is a more vital contributor to
soil quality than soil nutrients (Dick, 1997) because
all biochemical transformations that occur in soil de-
pend on or are related to the presence of enzymes
(Tabatabai, 1994). The impact of soil erosion on en-
zyme activity has been addressed in degraded soils
(Garcı́a and HernÁndez, 1997; Moreno-de las Heras,
2009; Park et al., 2014), thus enhancing our under-
standing of the biochemical processes of soil degrada-
tion by erosion. Among the available soil enzyme pa-
rameters, invertase, urease and phosphatase activities
are regarded as three desirable indicators of the sup-
plies of SOC, N and P, respectively. Invertase partici-
pates in the C cycle by catalyzing the hydrolysis of
sucrose into glucose and fructose. Urease is involved in
the N cycle and catalyzes the hydrolysis of urea into
CO2 and NH3 , which is of particular interest because
urea is an important N fertilizer. In the P cycle, phos-
phatase catalyzes the hydrolysis of both organic phos-
phate esters and phosphoric anhydrides into inorganic
P. In the Sichuan Basin of China, the losses of SOC,
N and P induced by soil erosion are threatening food
security (Su et al., 2010). To ensure crop production,
farmers are accustomed to increasing fertilizer use to
offset soil nutrient losses. However, the effect of long-
term increases in fertilization on crop production is
unknown for eroded slopes. The conventional practice
accelerates N and P pollution of the local aquatic en-
vironment. Conservation measures considering soil en-
zymes would likely contribute to the development of
sustainable agriculture in the Sichuan Basin. In this
context, there is a need to assess the effect of soil ero-
sion on the activity of soil enzymes such as invertase,
urease and phosphatase in relation to C, N and P cy-
cling.
X. J. NIE et al.
Cesium-137 (137 Cs) is a well-established soil ero-
sion tracer (Walling and Quine, 1991), and determina-
tion of erosion (less 137 Cs in test sites than in reference
sites) or deposition (more 137 Cs in test sites than in re-
ference sites) can be performed by comparing 137 Cs in-
ventories of test sites with those of reference sites. Fur-
thermore, 137 Cs inventories in eroded soil are inversely
proportional to the erosion degree. The relationships
between 137 Cs and soil chemical properties (e.g., C, N
and P) are well known (Zhang et al., 2006; Ni et al.,
2007; Nie et al., 2013a; Li et al., 2014), supporting the
use of the 137 Cs technique as a valuable tool for asses-
sing the effect of soil erosion on soil quality.
In this study, we used the 137 Cs technique to
evaluate erosion-enzyme relationships in eroded soils
of the Sichuan Basin, China, aiming 1) to examine
the dynamics of soil invertase, urease and alkaline
phosphatase activities on water and tillage erosion-
dominated landscapes and 2) to elucidate the impacts
of water and tillage erosion on soil invertase, urease
and alkaline phosphatase activities. Because water ero-
sion may easily liberate more organic substrates from
soil aggregates to increase enzymatic catalysis (Berhe
et al., 2012), we hypothesized that the activities of
soil invertase, urease and alkaline phosphatase would
be higher on water erosion-dominated landscapes than
tillage erosion-dominated landscapes.
MATERIALS AND METHODS
Study area
This study was conducted in Jianyang County,
Sichuan Province, in the Sichuan Basin of the sou-
thwestern part of China (30◦ 04 28 –30◦ 39 00 N,
104◦ 11 34 –104◦ 53 36 E). The study area is typical
of the hilly areas of Sichuan (400–587 m above sea
level), exhibiting a humid subtropical climate with a
mean annual temperature of 17 ◦ C and a mean annual
rainfall of 872 mm, 90% of which occurs between May
and October. The soils in the study area, derived from
purple mudstone and sandstone of the Jurassic Age,
are classified as Orthic Regosols in the FAO soil classi-
fication system. The soils are clay loam in texture (27%
clay, 29% silt and 44% sand) and generally contain less
than 20 g kg−1 organic matter.
Hoes are the predominant tillage implement in this
area. One major tillage operation is carried out per
year, starting at the bottom of the slope and mo-
ving upslope step by step; however, within every step,
the tillage direction is downslope (i.e., always grading
downward). Due to long-term intensive hoeing, the up-
per parts of the slope are characterized by a thin soil
SOIL ENZYME ACTIVITIES ON ERODED SLOPES
layer underlain by bedrock, whereas the deeper soils
occur in the lower parts. Cropping in the area involves
a wheat (Triticum aestivum L.) and maize (Zea mays
L.)/sweet potato (Ipomoea batatas) rotation. Fertilizers
(330 kg N ha−1 year−1 as urea and ammonium bicar-
bonate, and 166 kg P2 O5 ha−1 year−1 as superphos-
phate) are uniformly applied on the cultivated slopes
and residue management is also uniformly implemen-
ted on the slopes in the study area. The wheat and
maize residues are cut by hand as low as possible and
removed; the remaining residues, approximately 10 cm
in height, are mixed into the tillage layer through ma-
nual hoeing. In sweet potato crops, all of the residues
are removed from the field.
Soil sampling
Two gentle (4% and 7%) slopes (44 and 53 m in
length, respectively) and two steep (19% and 26%)
slopes (22 and 17 m in length, respectively) were selec-
ted for soil sampling. According to the soil erosion mo-
dels reported by Zhang et al. (1990, 2004), water ero-
sion, accounting for 67%–71% of the total erosion rate,
was the dominant erosion process on the 4% and 7%
gentle slopes, which are hereafter referred to as WES-I
and WES-II, respectively, whereas tillage erosion was
responsible for 66%–88% of the total erosion rate on
the 19% and 26% steep slopes, which are hereafter
referred to as TES-I and TES-II, respectively (Table
I). The details of the calculation of the soil erosion rates
on the four slopes have been reported previously (Nie
et al., 2013b). Therefore, in the present study, the two
gentle slopes and two steep slopes were considered as
two replicates of water and tillage erosion-dominated
landscapes, respectively. Soil sampling was conducted
using a soil corer (6.8 cm in diameter) along a downs-
lope transect at 10-m intervals on the two gentle slopes
and at 5-m intervals on the two steep slopes. The coor-
dinates of each sampling point as well as the elevation
TABLE I
Soil erosion of the selected eroded slopes in the Sichuan Basin of the southwestern part of China (Nie et al., 2013b)
Erosion Water erosion-dominated slope (WES)a)
WES-I
Total erosion rate (t ha−1 year−1 ) 24.41
Tillage erosion
Rate (t ha−1 year−1 ) 6.96
Percentage of total erosion (%) 29
Water erosion
Rate (t ha−1 year−1 ) 17.45
Percentage of total erosion (%) 71
a) WES-I = 4% gentle slope; WES-II = 7% gentle slope.
b) TES-I = 19% steep slope; TES-II = 26% steep slope.
491
were measured using a survey-grade differential global
positioning system (DGPS). Overall, 11 samples were
collected on the water erosion-dominated landscapes,
including 6 samples from WES-I (2 from the upper
part, 2 from the middle part and 2 from the lower
part) and 5 samples from WES-II (2 from the upper
part, 1 from the middle part and 2 from the lower
part). On the tillage erosion-dominated landscapes, 9
samples were collected, including 5 samples from TES-I
(2 from the upper part, 1 from the middle part and 2
from the lower part) and 4 samples from TES-II (2
from the upper part and 2 from the lower part).
Five soil cores were collected at each sampling
point. Two cores were taken from depths of 20 to 40
cm, depending on the total thickness of the soil lay-
ers at different slope positions, and then combined to
make a composite sample for determining the 137 Cs in-
ventory extending down to the bedrock. At the points
sampled in this study, the soil reached a maximum
depth of 40 cm; therefore, sampling to this depth
incorporated all of the 137 Cs in the soil profile into
the samples. Three additional cores for determining
physical-chemical and biochemical properties were col-
lected from the till layer (0–20 cm). One of the major
considerations here was that soil redistribution by wa-
ter and tillage occurs mainly in the till layer. Another
was that soil microbes are most active in the till layer,
where a large portion of organic matter, such as crop
residues, is concentrated and therefore the effects of
soil erosion on enzyme activities are expected to domi-
nate. The three soil cores were mixed into a composite
sample and were immediately stored at 4 ◦ C in plastic
bags. To measure soil biochemical properties, including
enzyme activities and microbial biomass carbon, the
fresh soil samples were passed through a 2-mm mesh
sieve and analyzed within one week. One portion of
the soil was air-dried for the analysis of soil physical-
chemical properties.
Tillage erosion-dominated slope (TES)b)
WES-II TES-I TES-II
28.01 39.77 44.83
9.29 26.16 39.67
33 66 88
18.72 13.61 5.16
67 34 12
492
Laboratory analysis
The soil samples to be used for 137 Cs determination
were air-dried, crushed and passed through a 2-mm
mesh sieve to remove gravel. The samples sieved to
< 2 mm were packed into plastic (PVC) beakers with
a volume of 320 cm3 , and 137 Cs activity was measured
using a hyperpure lithium-drifted germanium detec-
tor (HpC, 40% efficiency) coupled to a Nuclear Da-
ta 6700 multichannel γ-ray spectrophotometer (Ortec,
USA). The detection of 137 Cs was performed at 662
KeV, and the count time for each sample ranged from
40 000 to 60 000 s, providing a measurement precision
of ±5%. The content of 137 Cs was initially expressed
on a unit mass basis (Bq kg−1 ) and then converted
into an area basis (Bq m−2 ) using the total weight of
the bulk core sample and the cross-sectional area of
the sampling device. The successful application of this
technique has been widely reported in other study a-
reas (Chappell et al., 2012; Mekuria et al., 2012; Biro
et al., 2013; Dungait et al., 2013; Li et al., 2014).
The SOC content was determined through wet oxi-
dation with a mixture of potassium dichromate and
concentrated sulfuric acid, and total N was measured
following the classic Kjeldahl digestion method (Liu,
1996). The determination of the total P was carried
out using the NaOH digestion method (Liu, 1996). An
automatic acid-base titrator (Metrohm 702, Metrohm,
Switzerland) was employed to determine the soil pH
value in a 1:2.5 soil/water suspension. The soil bulk
density was determined based on the oven-dried weight
and sample volume. Soil particle size fractions were de-
termined in a 1 000 mL soil solution using the pipette
method, following H2 O2 treatment to destroy organic
matter and dispersion of the soil suspensions with Na-
hexametaphosphate (Liu, 1996).
Enzyme activities were determined according to
the methods described by Guan (1986). The activity
of invertase was measured through 3,5-dinitrosalicylic
acid colorimetry using sucrose as a substrate. Briefly, 5
g of soil, 15 mL of 8% (weight/volume) sucrose, 5 mL
of phosphate buffer (pH 5.5) and 5 drops of methyl-
benzene were mixed and incubated at 37 ◦ C for 24
h. After filtration, 1 mL of the filtrate was reacted with
3 mL of 3,5-dinitrosalicylic acid in a 50-mL volumet-
ric flask, followed by incubation in a boiling bath for 5
min. The incubated solution was diluted to 50 mL. The
absorbance was determined at 508 nm using a spec-
trophotometer. The invertase activity was expressed
in mg glucose g−1 dry soil. The activity of urease was
measured through indophenol colorimetry using urea
as a substrate. Briefly, 5 g of soil and 1 mL of methyl-
X. J. NIE et al.
benzene were combined in a 150-mL conical flask. Af-
ter 15 min, 10 mL of 10% (weight/volume) urea and 20
mL of citrate buffer (pH 6.7) were mixed with the soil
sample. The mixture was incubated at 37 ◦ C for 24
h and then diluted and filtered. Following filtration, 1
mL of the filtrate was transferred to a 50-mL volumet-
ric flask and mixed with 4 mL of sodium phenoxide and
3 mL of sodium hypochlorite. After the mixture was
diluted to 50 mL, the absorbance was determined at
578 nm using a spectrophotometer. The urease activity
was expressed in mg NH3 -N g−1 dry soil. The activity
of alkaline phosphatase was measured through disodi-
um phenyl phosphate colorimetry using p-nitrophenyl
phosphate (p-NPP) as a substrate. Briefly, 10 g of soil,
1.5 mL of methylbenzene, 10 mL of borate buffer (pH
10) and 10 mL of disodium phenyl phosphate were
added to a 100-mL volumetric flask. The mixture was
incubated at 37 ◦ C for 24 h. After incubation, 100 mL
of 0.3% (weight/volume) Al2 (SO4 )3 was added to ter-
minate the reaction, followed by filtration. After filtra-
tion, 3 mL of the filtrate was transferred to a 50-mL
volumetric flask and then diluted to 50 mL. The ab-
sorbance of the supernatant was determined at 660 nm
using a spectrophotometer. The alkaline phosphatase
activity was expressed in mg phenol g−1 dry soil. Soil
microbial biomass carbon was determined through fu-
migation with ethanol-free CHCl3 and extraction with
K2 SO4 (Vance et al., 1987). All of the enzyme activi-
ty and microbial biomass carbon determinations were
performed in triplicate.
Statistical analysis
Statistical analyses were conducted using SPSS
11.0 for Windows (SPSS Inc., USA). Pearson corre-
lation coefficients were calculated to test the rela-
tionships between 137 Cs, soil physical-chemical pro-
perties and enzyme activities on both water and tillage
erosion-dominated landscapes. A general linear model
(GLM) was employed to evaluate the effects of erosion
type (i.e., water erosion and tillage erosion), slope posi-
tion (i.e., upper, middle and lower parts of the slope)
and their interactions on soil enzyme activity. Diffe-
rences at P < 0.05 were considered statistically signifi-
cant.
RESULTS
Soil redistribution
Similar 137 Cs distributions were observed on all
eroded slopes: the 137 Cs inventory increased downs-
lope (Fig. 1). For the water erosion-dominated land-
scapes, the mean 137 Cs inventory increased from 786
SOIL ENZYME ACTIVITIES ON ERODED SLOPES
(WES-I) and 861 (WES-II) Bq m−2 at upper slope
parts to 1 296 (WES-I) and 1 335 (WES-II) Bq m−2 at
lower slope parts (Fig. 1a). From the upper to lower s-
lope parts of the tillage erosion-dominated landscapes,
the mean 137 Cs inventory increased from a low of 402
(TES-I) and 668 (TES-II) Bq m−2 to a high of 1 117
(TES-I) and 1 002 (TES-II) Bq m−2 (Fig. 1b). The re-
sults suggest that soil loss occurred in the upper sec-
tions, whereas soil accumulation occurred in the lower
sections of the eroded slopes.
Fig. 1 Distribution of 137 Cs inventories on the water erosion-
dominated slopes (WES) (a) and tillage erosion-dominated slo-
pes (TES) (b) studied. WES-I = 4% gentle slope; WES-II = 7%
gentle slope; TES-I = 19% steep slope; TES-II = 26% steep
slope.
Soil physical-chemical properties
On both WES-I and WES-II, the contents of clay,
SOC and total N gradually increased downslope
(Fig. 2a, d, e). The total P content roughly increased
downslope (Fig. 2f), whereas the soil bulk density and
pH decreased downslope (Fig. 2b, c). In addition, the
patterns of clay, SOC and total N were similar to
the redistribution patterns of the 137 Cs inventory on
the two water erosion-dominated slopes. However, the
bulk density showed a distribution opposite to that of
the 137 Cs inventory. On both TES-I and TES-II, the
SOC content increased downslope (Fig. 2d) and exhibi-
493
ted a similar distribution pattern to the 137 Cs inven-
tory. Unlike the SOC content, the other five physical-
chemical properties did not display an obvious spa-
tial pattern on the two tillage erosion-dominated slopes
(Fig. 2a–c, e, f). These results indicate that soil redis-
tribution had an apparent impact on the spatial pat-
terns of clay, bulk density, SOC and total N on the
water erosion-dominated landscapes, whereas only the
SOC distribution pattern was related to soil redistri-
bution on the tillage erosion-dominated landscapes.
Soil enzyme activity
On both WES-I and WES-II, the invertase and
urease activities increased downslope, with the inver-
tase activity showing a greater increase than the urea-
se activity (Fig. 3a, b). These patterns were consistent
with the distribution pattern of the 137 Cs invento-
ry. However, the alkaline phosphatase activity varied
irregularly on the two water erosion-dominated slo-
pes (Fig. 3c), exhibiting a distinctly dissimilar pattern
from that of the 137 Cs inventory. Consequently, it is
clear that the spatial patterns of the invertase and
urease activities were closely associated with soil re-
distribution, whereas the spatial pattern of the alkaline
phosphatase activity was independent of soil redistri-
bution on the water erosion-dominated slopes.
On both TES-I and TES-II, the most noticeable
changes in the soil enzyme parameters included in-
creased activities of invertase (Fig. 3a) and alkaline
phosphatase (Fig. 3c) along the downslope transect.
Furthermore, the invertase and alkaline phosphatase
activity patterns were consistent with the 137 Cs dis-
tribution on these two tillage erosion-dominated slo-
pes. However, the changes in urease activity were irreg-
ular (Fig. 3b), showing a clearly dissimilar pattern from
the 137 Cs distribution. This finding suggests that the
spatial patterns of invertase and alkaline phosphatase
activities were markedly impacted by soil redistribu-
tion on sloped landscapes dominated by tillage erosion,
whereas the pattern of urease activity was independent
of soil redistribution.
The impacts of erosion type and slope position
and their interactions on soil enzyme activity (on a
dry soil basis) were as presented in Tables II and
III. Overall, the activities of invertase and alkaline
phosphatase were significantly different between the
water and tillage erosion-dominated slopes, whereas
no difference in urease activity was observed (Table
II). The soil invertase activity on the water erosion-
dominated was significantly higher than that on the
tillage erosion-dominated slopes (Table III), whereas
the alkaline phosphatase activity on the water erosion-
494 X. J. NIE et al.

Fig. 2 Spatial patterns of the soil physical-chemical properties on the water erosion-dominated slopes (WES) and tillage erosion-
dominated slopes (TES) studied: clay (a), bulk density (b), pH (c), soil organic C (SOC) (d), total N (e) and total P (f). WES-I =
4% gentle slope; WES-II = 7% gentle slope; TES-I = 19% steep slope; TES-II = 26% steep slope.

TABLE II
Analysis of variance on soil enzyme activity affected by erosion types and slope positions on the water erosion-dominated slopes (WES)
and tillage erosion-dominated slopes (TES) studied

Item Erosion type (ET) Slope position (SP) ET × SP

F value P value F value P value F value P value

Activity on a dry soil basis
Invertase 26.86 < 0.001 8.55 0.004 1.90 0.187
Urease 0.03 0.870 1.64 0.230 0.04 0.963
Alkaline phosphatase 31.72 < 0.001 1.90 0.186 0.50 0.619
Soil organic C (SOC) 4.02 0.065 12.91 0.001 0.20 0.822
Activity on an organic matter basisa)
Invertase 9.93 0.007 0.70 0.512 0.49 0.623
Urease 4.80 0.046 6.93 0.008 0.00 0.999
Alkaline phosphatase 41.17 < 0.001 0.42 0.663 0.27 0.770
Microbial biomass C (Cmic )b) 25.83 < 0.001 2.57 0.112 2.29 0.137
Activity on a microbial biomass basisc)
Invertase 98.43 < 0.001 3.84 0.047 0.15 0.863
Urease 53.87 < 0.001 3.05 0.080 5.40 0.019
Alkaline phosphatase 5.01 0.042 0.91 0.424 0.59 0.570
a) Calculated as the ratio of enzyme activity to SOC.
b) From Nie et al., 2013b.
c) Calculated as the ratio of enzyme activity to C
mic .

dominated slopes was significantly lower than that on fected by slope position (Table II), with significantly
the tillage erosion-dominated slopes (Table III). In ad- lower values at the upper and middle slope parts com-
dition, only the invertase activity was significantly af- pared with the lower slope parts of the eroded landsca-
SOIL ENZYME ACTIVITIES ON ERODED SLOPES
Fig. 3 Distribution patterns of soil enzyme activities on the
water erosion-dominated slopes (WES) and tillage erosion-
dominated slopes (TES) studied: invertase (a), urease (b) and
alkaline phosphatase (c). WES-I = 4% gentle slope; WES-II =
7% gentle slope; TES-I = 19% steep slope; TES-II = 26% steep
slope.
pes (Table III). No interaction between erosion type
and slope position on each enzyme activity was de-
tected (Table II).
In this study, the ratios of enzyme activity to SOC
and microbial biomass carbon (Cmic ) were employed
to express soil enzyme activity on an organic mat-
ter basis and a microbial biomass basis, respective-
ly (Tables II and III). All of the enzyme activity-to-
SOC and -Cmic ratios were significantly different be-
tween the water and tillage erosion-dominated land-
scapes (Table II). Both the invertase activity-to-SOC
(INV/SOC) and -Cmic (INV/Cmic ) ratios were signifi-
495
cantly higher on the water erosion-dominated slopes
than the tillage erosion-dominated slopes (Table III).
However, both the alkaline phosphatase activity-to-
SOC (ALP/SOC) and -Cmic (ALP/Cmic ) ratios were
significantly lower on the water erosion-dominated slo-
pes than the tillage erosion-dominated slopes (Table
III). The changes in INV/SOC and INV/Cmic ratios
between the two contrasting landscape types were con-
sistent with the change in invertase activity, and those
in ALP/SOC and ALP/Cmic ratios were also consistent
with alkaline phosphatase activity. Unlike urease ac-
tivity, which did not present a difference between the
two contrasting landscape types, the urease activity-
to-SOC ratio (UR/SOC) was significantly lower and
urease activity-to-Cmic ratio (UR/Cmic ) was higher on
the water erosion-dominated slopes than the tillage
erosion-dominated slopes (Table III).
The UR/SOC and INV/Cmic ratios were signifi-
cantly impacted by the slope position (Table II). The
values of the UR/SOC ratio significantly decreased at
upper and middle slope parts compared with the lower
slope parts of the eroded landscapes (Table III). Si-
milar to the invertase activity, which varied with the
slope position, the values of the INV/Cmic ratio were
also significantly lower at the upper and middle slope
parts relative to the lower slope parts (Table III). In
addition, the changes in the UR/Cmic ratio with slope
position were significantly affected by erosion type (Ta-
ble III).
Relationships between soil enzyme activities and phy-
sical-chemical properties
The Pearson correlation analysis was performed to
quantify the relationships between soil enzyme activi-
ty and physical-chemical properties, as shown in Ta-
ble IV. The activities of invertase and urease were
significantly (P < 0.01) correlated with the SOC and
total N contents (r = 0.84 and 0.82) on the wa-
ter erosion-dominated slopes, whereas on the tillage
erosion-dominated slopes, the only detected signifi-
cant (P < 0.05) correlation was between the inver-
tase activity and SOC content (r = 0.76). Further-
more, the activities of invertase and urease exhibited a
stronger significant (P < 0.01) correlation with the
clay content on the water erosion-dominated slopes
(r = 0.84 and 0.81) than that (P < 0.05) on the tillage
erosion-dominated slopes (r = 0.78 and 0.69). The al-
kaline phosphatase activity did not show any signifi-
cant correlation with the total P content on either the
water or tillage erosion-dominated slopes. Likewise, the
three enzyme parameters were not significantly corre-
lated with the soil bulk density and pH on the two
contrasting landscape types.
496 X. J. NIE et al.

TABLE III
Comparison of soil enzyme activities between erosion types and slope positions on the water erosion-dominated slopes (WES) and
tillage erosion-dominated slopes (TES) studied

Item Erosion type Slope position

WES TES Upper slope part Middle slope part Lower slope part
Activity on a dry soil basis
Invertase (mg glucose g−1 soil) 8.94aa) 6.46b 7.06b 7.16b 8.89a
Urease (mg NH3 -N g−1 soil) 0.35a 0.36a 0.34a 0.35a 0.37a
Alkaline phosphatase (mg phenol g−1 soil) 0.44b 0.80a 0.57a 0.60a 0.69a
Soil organic C (SOC) (g kg−1 ) 10.56a 9.27a 8.80b 9.09b 11.84a
Activity on an organic matter basisb)
Invertase (mg glucose g−1 SOC) 849a 708b 803a 782a 751a
Urease (mg NH3 -N g−1 SOC) 34.18b 39.23a 39.67a 38.83a 31.62b
Alkaline phosphatase (mg phenol g−1 SOC) 41.60b 86.91a 65.63a 66.99a 60.15a
Microbial biomass C (Cmic ) (mg kg−1 )c) 196.78b 270.74a 212.61a 251.93a 236.74a
Activity on a microbial biomass basisd)
Invertase (mg glucose mg−1 Cmic ) 45.32a 24.37b 35.12b 30.94b 38.49a
Urease (mg NH3 -N mg−1 Cmic ) 1.82a 1.34b 1.68a 1.47b 1.58ab
Alkaline phosphatase (mg phenol mg−1 Cmic ) 2.21b 2.98a 2.63a 2.29a 2.88a
a) Mean values followed by the same letter(s) in a row within the erosion type or slope position are not significantly different at
P < 0.05.
b) Calculated as the ratio of enzyme activity to SOC.
c) From Nie et al., 2013b.
d) Calculated as the ratio of enzyme activity to C
mic .

TABLE IV
Correlation coefficients (r) between soil enzyme activities and physical-chemical properties including soil organic carbon (SOC), total
N, total P, pH, clay and bulk density on the water and tillage erosion-dominated slopes studied

Slopes Enzyme Clay Bulk density pH SOC Total N Total P

Water erosion-dominated Invertase 0.84** −0.33 −0.44 0.84** – –
Urease 0.81** 0.19 −0.32 – 0.82** –
Alkaline phosphatase 0.30 −0.03 −0.37 – – 0.28
Tillage erosion-dominated Invertase 0.78* 0.33 −0.13 0.76* – –
Urease 0.69* −0.54 −0.21 – 0.55 –
Alkaline phosphatase 0.52 −0.31 −0.24 – – 0.59

*, **Significant at the 0.05 and 0.01 probability levels, respectively.

DISCUSSION lective soil transport by water) and tillage erosion (i.e.,

In this study, the 137 Cs data revealed a similar
soil redistribution pattern on both water and tillage
erosion-dominated slopes, with soil loss occurring at
the upper slope parts and soil accumulation at the lo-
wer slope parts. The results are consistent with those
of Zhang et al. (2013) and Dungait et al. (2013), in-
dicating that soil erosion varies with slope position.
However, it is also worth noting that without high-
intensity thunderstorms, the slope position does not
affect soil erosion (Cerdà, 1998). The clay content was
observed to increase downslope on the water erosion-
dominated slopes but changed irregularly on the tillage
erosion-dominated slopes, which is consistent with pre-
vious studies (Li and Lindstrom, 2001; Zhang et al.,
2006) that have suggested a difference in the dominant
transport mechanism between water erosion (i.e., se-
entire transport).
Soil redistribution was also found to dominate the
spatial patterns of invertase activity on both water and
tillage erosion-dominated landscapes. This could ex-
plain the significant impact of slope position on inver-
tase activity, with lower invertase activity observed in
the upper sections of the eroded slopes because of soil
loss and higher invertase activity observed in the lower
sections because of soil accumulation. However, sig-
nificant differences in urease and alkaline phosphatase
activities were not observed between the upper and
lower sections of the eroded slopes, which was most
likely a result of differential impacts of soil redistribu-
tion on the two enzyme parameters between the two
contrasting landscape types, including the domination
of spatial patterns of urease and alkaline phosphatase
activities by soil redistribution on the water and tillage
SOIL ENZYME ACTIVITIES ON ERODED SLOPES
erosion-dominated slopes, respectively.
In this study, the activities of invertase and alka-
line phosphatase were affected by the erosion type.
The higher invertase activity on the water erosion-
dominated slopes relative to that of the tillage erosion-
dominated slopes could be explained by the different
soil transport mechanisms that occur with water and
tillage. Selective transport of water erosion may easi-
ly disrupt soil aggregates relative to the complete
transport that occurs in tillage erosion (Zhang et al.,
2006). Based on the physical protection of organic mat-
ter inside soil aggregates (Berhe et al., 2012), water
erosion would liberate more organic substrates from
aggregates to increase the catalysis of invertase rela-
tive to tillage erosion. The higher alkaline phosphatase
activity on the tillage erosion-dominated slopes may
have a significantly (P < 0.001) greater dependency
on a higher soil pH (a mean of 8.51) compared with
that of the water erosion-dominated slopes (a mean of
7.65) (data not shown). Alkaline phosphatase reactions
are known to be most active under soil pH values of
8.0–10.0; under other conditions, the reactions are sub-
stantially restricted (Frankenberger et al., 1983).
Microbial biomass determines the quantity of en-
zymes, and organic matter provides a substrate for en-
zyme reactions. Therefore, the levels of these two bio-
chemical parameters in soils may be considered when
evaluating enzyme activities. Barriuso et al. (1988) su-
ggested that the enzyme activity per unit of organic
matter is a more useful measure and can avoid the
misuses of both organic matter and enzyme activity
in soil quality assessment. Landi et al. (2000) indi-
cated that the ratio of enzyme activity to microbial
biomass C represents a combination of two different
measurements into a single criterion that can provide
an indication of the changes that occur during micro-
bial activity. Several researchers have clearly identi-
fied the impacts of vegetation changes on soil microbi-
ological activity using these ratio parameters (Basti-
da et al., 2006; Wang et al., 2012). In the present
study, consistent results were obtained for INV/SOC,
INV/Cmic and invertase activity impacted by different
erosion types and slope positions and for ALP/SOC,
ALP/Cmic and alkaline phosphatase activity impacted
by different erosion types. Interestingly, different re-
sults were detected for UR/SOC, UR/Cmic and urease
activity, including 1) evident impact of erosion type
on both UR/SOC and UR/Cmic and no impact of ero-
sion type on urease activity and 2) significant effect
of slope position on UR/SOC and UR/Cmic . More-
over, UR/Cmic was found to be higher on the water
erosion-dominated slopes relative to the tillage erosion-
497
dominated slopes, whereas the opposite result was ob-
served for UR/SOC. In terms of the more sensitive
response of microbial biomass to soil disturbance com-
pared with organic matter (Powlson and Jenkinson,
1981), we believe that the results reflected by the ra-
tio of enzyme activity to Cmic in our study should be
preferable. Overall, our results suggest that express-
ing soil enzyme activity on organic matter and micro-
bial biomass basis may provide insights into the im-
pact of soil erosion on microbial activity. It is antici-
pated that the lower activities of invertase and urease
would result in decreased soil biological fertility on the
tillage erosion-dominated slopes, relative to the water
erosion-dominated slopes in the Sichuan Basin. Effe-
ctive bioremediation of tillage erosion-dominated soils
should be taken. Previous studies have indicated that
soil microbial activity is related to land management
practices such as fertilization, organic amendment ap-
plications, cropping and tillage systems and pesti-
cide applications (Garcı́a-Orenes et al., 2010; Álvaro-
Fuentes et al., 2013; Wu et al., 2013; Balota et al.,
2014; Tejada and Benı́tez, 2014). Based on the results
of our study, the impact of soil erosion on soil micro-
bial activity should also be evaluated.
Generally speaking, extracellular enzymes, such as
invertase, urease and phosphatase, in soils may be
tightly adsorbed onto soil inorganic colloids such as
clay; therefore, their activities are associated with the
clay content (Paul, 2007). In our study, the corre-
lation analysis between enzyme activities and clay con-
tent suggested that both invertase and urease activi-
ties were closely associated with the clay content in
the two contrasting landscape types; however, a poor
relationship between alkaline phosphatase activity and
clay content was observed, which could be related to
P fixation in soil (Li et al., 1982). A portion of the
bioavailable P substrates would be inaccessible to soil
microbes, potentially disturbing the catalysis of alka-
line phosphatase adsorbed onto clay. The soil bulk den-
sity and pH, which reflect soil compactability/aeration
and acidity-alkalinity, respectively, are considered to
be two key factors influencing enzyme activity. Seve-
ral studies have demonstrated that there is a negative
or no correlation between enzyme activity and bulk
density in agricultural soils (Dick et al., 1988; Li et al.,
2002). Positive, negative or null correlations between
enzyme activity and pH have also been reported (Kang
and Freeman, 1999; Acosta-Martı́nez and Tabatabai,
2000; Gianfreda et al., 2005). In the present study,
no correlation was detected between bulk density and
pH and the activities of invertase, urease and alka-
line phosphatase on either the water or tillage erosion-
498
dominated slopes.
Enzyme activity is essential for the soil nutrient cy-
cle. Numerous studies have demonstrated that soil in-
vertase, urease and alkaline phosphatase activities are
generally correlated with the SOC, total N and total P
contents, respectively (Gianfreda et al., 2005; Melero
et al., 2006; Yu et al., 2006; Cheng et al., 2013). In the
present study, soil redistribution dominated the dis-
tribution patterns of SOC and invertase activity on
the two contrasting landscape types, with a positive
correlation between them (r = 0.84 and P < 0.01
for the water erosion-dominated slopes; r = 0.76 and
P < 0.05 for the tillage erosion-dominated slopes). The
results confirmed that invertase maintained an influ-
ential role in the C cycle under water erosion and
tillage erosion, especially water erosion. There was a
significantly positive correlation (r = 0.82, P < 0.01)
between the urease activity and total N content on
the water erosion-dominated slopes, but no correlation
(r = 0.55, P > 0.05) was found on the tillage erosion-
dominated slopes. This finding suggested that involve-
ment of urease in the N cycle might be stronger under
water erosion than tillage erosion, which was ascribed
to the dependence of urease dynamics on soil redistri-
bution by water. However, there was a lack of corre-
lation between the alkaline phosphatase activity and
total P content on the two contrasting landscape types
(r = 0.28 and 0.59, P > 0.05), indicating that both wa-
ter erosion and tillage erosion constrained the involve-
ment of alkaline phosphatase in the P cycle. On the
tillage erosion-dominated slopes, the lack of a corre-
lation between the alkaline phosphatase activity and
total P content was attributed to the dependence on
alkaline phosphatase dynamics but independence of
the dynamics of total P from soil redistribution. Ne-
vertheless, the absence of a correlation between these
two parameters on the water erosion-dominated slopes
could not be explained by soil redistribution.
CONCLUSIONS
Soil redistribution had different impacts on soil en-
zyme activity on the two contrasting eroded landscape
types selected in the Sichuan Basin of China. On the s-
lopes dominated by water erosion, the distribution pat-
terns of the invertase and urease activities were associ-
ated with soil redistribution. On the slopes dominated
by tillage erosion, the distribution patterns of the in-
vertase and alkaline phosphatase activities were linked
to soil redistribution. The erosion type had an obvious
effect on the activities of invertase, urease and alkaline
phosphatase. The urease activity may be affected by
X. J. NIE et al.
the interaction between erosion type and slope posi-
tion. The ratios of enzyme activities to Cmic could be
used to better evaluate soil microbiological activity on
eroded slopes. Our results generally supported the hy-
pothesis that the activities of invertase and urease were
higher on water erosion-dominated slopes than tillage
erosion-dominated slopes. In addition, soil redistribu-
tion might influence the involvement of urease in the
N cycle and alkaline phosphatase in the P cycle.
ACKNOWLEDGEMENT
The authors wish to acknowledge the financial sup-
port for this study provided by the National Natu-
ral Science Foundation of China (No. 41001157), the
135 Strategic Program of the Institute of Mountain
Hazards and the Environment, Chinese Academy of
Sciences (No. SDS-135-1206) and the Young Teacher
Foundation of Henan Polytechnic University, China.
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