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\begin{document}

\title{Slope gradient but not slope position affecting microbial community
composition in the cropland of typical black soil region}



\author[1]{lei sun}%
\author[1]{Abdul Hakim Jamshidi}%
\author[2]{Yong Niu}%
\author[1]{Xia Liu}%
\author[3]{Zhaofei Fan}%
\author[1]{Wenying Yi}%
\author[1]{Shouhao Zhang}%
\affil[1]{Co-Innovation Center for Sustainable Forestry in Southern China Jiangsu Province Key Laboratory of Soil and Water Conservation and Ecological Restoration Nanjing Forestry University Nanjing Jiangsu 210037 China}%
\affil[2]{College of Forestry Shandong Agricultural University Shandong 271018 China}%
\affil[3]{Auburn University School of Forestry and Wildlife Sciences}%


\vspace{-1em}



  
  \date{May 26, 2024}


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\selectlanguage{english}
\begin{abstract}
Soil microorganisms are the main driving force in soil ecological and
biogeochemical processes, such as carbon mineralization, humus
formation, and nutrient cycles. Soil erosion may dramatically alter
microbial properties. However, it is still unclear how soil microbial
characteristics affected by soil erosion. In hilly areas, soil erosion
is highly attributed to topographic characteristics such as slope
position and slope gradient. Herein, we compared the soil microbial
characteristics between different slope gradients and positions in the
farmland in the typical black soil area of Northeast China. The relative
abundances of major microbial phyla are illustrated. Proteobacteria,
Acidobacteria, Gemmatimonadetes, Planctomycetes, and Actinobacteria
accounted for 83\% of the total bacterial sequences, and Ascomycota,
Basidiomycota, and chytridiomycosis accounted for 78\% of total fungal
sequences in the studied area. Microbial alpha-diversity (Bacteria and
Fungi) was similar among the slope gradients or slope positions. The
bacterial and fungal community composition was only affected by slope
gradient, which was significantly lower in the highest slope gradient
level than in others. This study suggests that slope gradient but not
slope gradient affecting microbial community composition in the cropland
of black soil region.%
\end{abstract}%



\sloppy


\textbf{Slope gradient but not slope position affecting microbial
community composition in the cropland of typical black soil region}

Abdul Hakim Jamshidi\textsuperscript{a}, Lei Sun\textsuperscript{a},
Yong Niu\textsuperscript{b}, Xia Liu\textsuperscript{a*}, Zhaofei
Fan\textsuperscript{c}, Wenying Yi\textsuperscript{a}, Shouhao
Zhang\textsuperscript{a}

\emph{a. Co-Innovation Center for Sustainable Forestry in Southern
China, Jiangsu Province Key Laboratory of Soil and Water Conservation
and Ecological Restoration, Nanjing Forestry University, Nanjing,
Jiangsu 210037, China;}

\emph{b. College of Forestry, Shandong Agricultural University, Shandong
271018, China;}

\emph{c. School of Forestry and Wildlife Sciences, Auburn University,
Auburn, AL 36849;}

\textsuperscript{\emph{*}} \emph{Corresponding author. E-mail address:
liuxia@njfu.edu.cn}

Text pages: 19 Tables: 5 Figures: 3

Submitted to: Land Degradation and Development

\textbf{Abstract}

Soil microorganisms are the main driving force in soil ecological and
biogeochemical processes, such as carbon mineralization, humus
formation, and nutrient cycles. Soil erosion may dramatically alter
microbial properties. However, it is still unclear how soil microbial
characteristics affected by soil erosion. In hilly areas, soil erosion
is highly attributed to topographic characteristics such as slope
position and slope gradient. Herein, we compared the soil microbial
characteristics between different slope gradients and positions in the
farmland in the typical black soil area of Northeast China. The relative
abundances of major microbial phyla are illustrated. Proteobacteria,
Acidobacteria, Gemmatimonadetes, Planctomycetes, and Actinobacteria
accounted for 83\% of the total bacterial sequences, and Ascomycota,
Basidiomycota, and chytridiomycosis accounted for 78\% of total fungal
sequences in the studied area. Microbial alpha-diversity (Bacteria and
Fungi) was similar among the slope gradients or slope positions. The
bacterial and fungal community composition was only affected by slope
gradient, which was significantly lower in the highest slope gradient
level than in others. This study suggests that slope gradient but not
slope gradient affecting microbial community composition in the cropland
of black soil region.

Keywords: Black soil; soil microbial community; bacterial; fungal;
slope.

\textbf{Introduction}

Land degradation refers to deterioration of soil properties related to
crop production, infrastructure maintenance, and natural resource
quality (Lal, 2001). More than 20\% of cultivated areas in the world
experience degradation, and its severity extends to many parts (Bai et
al., 2008). The NRC (1994) highlighted that land degradation is complex
and implicates transformations in the soil and vegetation's physical,
chemical, and biological properties. Soil erosion, one of the
considerable critical environmental problems, is the primary process
leading to soil degradation worldwide (Wilkinson and Mcelory, 2007),
particularly in hilly areas, highly attributed to topographic effects
such as slope position, and slope gradient (Khormali et al., 2009).
Topographic properties determine the receipt and redistribution of
light, heat, water, and sediments over the land surface, affecting the
variations in soil properties and leading to reduced productivity (FAO,
2015).

Soil microorganisms are the main driving force in soil ecological and
biogeochemical processes, such as carbon mineralization, humus
formation, and nutrient cycles. Soil erosion may dramatically alter
microbial properties and enzyme activities. Research has indicated that
microbial counts, biomass, diversity, and enzyme activity decline
drastically during soil erosion (Salam et al., 2001; Moreno-De Las
Heras, 2009; Mabuhay et al., 2004; An et al., 2008). Erosion directs to
the loss of fertile topsoil and nutrients, which sustain microbial
habitats and the ecosystem services function, resulting in reduced
productivity (Van Oost and Bakker, 2012). The composition, structure,
and functionality of soil microorganisms are sensitive to environmental
change and disturbance (Chang et al., 2007). Soil carbon and nutrient
availability to microbes would be modified through rain splash and
surface wash, which is more pronounced in sloped land. The available
substrates would move with fine particles along the slope.

Wang et al. (2017) found that the bottom slope position is rich in soil
moisture, clay, and organic carbon content compared with the top slope
position, leading to a larger microbial community and greater microbial
biomass and enzyme activity in the bottom slope position. The varying
soil water and substrate conditions can further lead to distinctive
vegetation coverage, above- and below-ground plant biomass on different
topographic positions (Wang et al., 2015), which would generate a
variety of substrates for different microbial communities through fresh
C supply (Li et al., 2017). Soil loss would increase with the increase
of slope gradient because of the more extensive surface runoff and lower
infiltration rate under a higher slope gradient (Zhang and Hosoyamada,
1996). Du et al. (2020) observed that relative abundances of bacterial
phyla at eroding slopes (10\selectlanguage{ngerman}° and 20°) and depositional zones differed
from reference slopes (5°). The changes in soil microbiota due to
erosion would profoundly impact the environment, such as nutrient
cycling (Li et al., 2019). Due to the significance of soil microbiota
for ecosystem service function, they should be considered when exploring
the mechanisms behind the response of soil to erosion (Qiu et al.,
2021).

Hence, few studies have thoroughly examined the soil microbial
characteristics, extracellular enzyme, and soil physicochemical process,
which represent the main understanding gap given that soils are a
complex system. The main objectives of our study are to find out the
effect of slope gradient and slope position on soil physicochemical
properties, microbial community composition and diversity, and enzyme
activity. Our hypothesis is higher slope gradient and position would
have lower microbial community composition and diversity, and enzyme
activity due to the deterioration of soil properties than lower slope
gradient and position.

\textbf{2. Materials and methods}

\textbf{2.1 Study site and soil sampling}

Soil samples were taken from sloped cropland in Baiquan and Keshan
Counties, Qiqihar, Heilongjiang province (125°30\selectlanguage{english}'E--126deg31\selectlanguage{english}'E,
47deg20\selectlanguage{english}'N--47deg55\selectlanguage{english}'N) (Fig. 1), which are both located in the typical black
soil region (Luvic Phaeozem, FAO; Liu et al., 2021). The region has a
temperate continental monsoon climate with an annual range from - 4 to 4
oC. Strong winds are frequent in spring, winters are long and frigid,
and summers are short and cool. Annual rainfall averages 500-600mm,
concentrated primarily in the summer. We used a Global Positioning
System (GPS) to acquire each sampling site's longitude, latitude, and
elevation. We used remotely sensed images to calculate the ridge-slope
angle for ridge interpretation and a digital elevation model (DEM;
Resolution of 30 m) for generating the slope raster image layer of the
sampling site using the ArcGIS 10.2 raster calculator. We interviewed
farmers to obtain crop yield and management at each soil sampling site.
Therefore, data and information on farmers' knowledge about the
management practices of their fields are composed in the area. A total
of 108 soil samples were collected randomly from study sites with
similar soil types (black soil), vegetation types (soybean), and
cultivation systems (Crop Rotation). This study selected four slope
gradients (0o-1.5o, 1.5o-3o, 3o-5o, \textgreater{}5o) with the slope
ridge angle between 10o-80o. In each slope gradient, soil samples were
collected at three positions (top, middle, and bottom) from the surface
layer (0-10 cm). To analyze microbial community characteristics, we
consider three slope gradient levels: low (0o-1.5o), medium (3o-5o), and
high (\textgreater{}5o). Totally 54 disturbed (mixed of five samples
collected in an ``S'' shape) soil samples (for soil physicochemical and
biological properties) and 54 undisturbed soil samples (for soil bulk
density and water stable aggregate) were collected. The soil samples
were transferred to the laboratory for air-drying and preparation for
measurements of soil physicochemical and biological properties.

\textbf{2.2 Soil analysis}

Samples were chosen randomly and analyzed in the laboratory after
passing through a 2-mm sieve and air dried. For measuring microbial
characteristics and soil enzyme activity, samples were incubated at 25
oC and 60\% water holding capacity (WHC). Soil variables were measured
using standard methods, bulk density (BD), water stable aggregate
content (WSA), soil organic matter (SOM), available nitrogen (AN),
available phosphorus (AP), available potassium (AK), and cation exchange
capacity (CEC) (Bao, 2000). To determine the clay content, we used a
Microtrac S3500 laser particle size analyzer (Microtrac Inc., USA) to
obtain the volume ratio, which was converted to mass ratio using an
empirical equation (Li et al., 2015). Soil pH was measured in MIlli-Q
water (soil: solution ratio = 1: 2.5(w: v)). TOC/TN analyzer (Shimaduz
Corp, Kyoto, Japan) used to measure soil dissolved organic carbon (DOC),
and total dissolved nitrogen (TDN) (extracts 1M KCL (1:10 (w: v) for 60
min)) (Jones et al., 2005). Chloroform fumigation-extraction method
(48h) determined Soil microbial biomass carbon (MBC) and microbial
biomass nitrogen (MBN) (Vance et al., 1987). MBC and MBN amounts were
calculated by subtraction concentration of the fumigated sample from the
non-fumigated sample, and the result was multiplied by a conversion
factor of 2.22 and 1.85, respectively (Brookes et al., 1985). Invertase
(INV), Catalases (CAT), Phosphates (ACP), and Urease (U.R. were measured
by colorimetric enzyme assay in soil suspensions (Sinsabaugha, et al,
2000; Hou Q, et al, 2020).

\textbf{2.3 DNA extraction and PCR (Polymerase chain reaction)
amplification}

Microbial DNA was extracted using the HiPure Soil DNA Kits (or HiPure
Stool DNA Kits) (Magen, Guangzhou, China). The 16S rDNA V3-V4 region of
the ribosomal RNA gene was amplified for bacteria by PCR\selectlanguage{english}(95degC for 2min,
followed by 27 cycles at 98degC for 10s, 62degC for the 30s, and 68degC for
the 30s, and a final extension at 68degC for 10min using primers 341F:
CCTACGGGNGGCWGCAG; 806R: GGACTACHVGGGTATCTAAT, and the ITS region of the
Eukaryotic ribosomal RNA gene were amplified for fungi, using primers
ITS3\_KYO2F (5'- GATGAAGAACGYAGYRAA -3') and ITS4R
(5'TCCTCCGCTTATTGATATGC -3'), where the barcode is an eight-base
sequence individual to each sample. PCR reactions were executed in a
triplicate 50 \selectlanguage{greek}μ\selectlanguage{english}L mixture containing 5 \selectlanguage{greek}μ\selectlanguage{english}L of 10 \selectlanguage{ngerman}× KOD Buffer, 5 \selectlanguage{greek}μ\selectlanguage{english}L of 2.5
mM dNTPs, 1.5 \selectlanguage{greek}μ\selectlanguage{english}L of each primer (5 \selectlanguage{greek}μ\selectlanguage{english}M), 1\selectlanguage{greek}μ\selectlanguage{english}L of KOD Polymerase, and 100ng
of template DNA. Sequencing was performed using the Illumina HiSeq2500
(PE250) platform. Amplicons were extracted from 2\% agarose gels and
filtered using the AxyPrep DNA Gel Extraction Kit (Axygen Biosciences,
Union City, CA, USA) and quantified using ABI StepOnePlus Real-Time PCR
System (Life Technologies, Foster City, USA). Purified amplicons were
pooled in equimolar and paired-end sequenced (2 \selectlanguage{ngerman}×250) on an Illumina
platform. The raw reads were deposited into the NCBI Sequence Read
Archive (SRA) database.

\textbf{2.4 Statistical analysis}

Statistical analyses were conducted using SPSS 25.0 (SPSS Inc., USA).
Two-way ANOVA was employed to evaluate the effects of slope gradients
(i.e., 0º-1.5º, 1.5º-3º, 3º-5º, \textgreater{}5º), slope positions
(i.e., top, middle, and bottom parts of the slope) on soil
physicochemical and biological properties. Differences at the
significance level of p \textless{} 0.05 were considered statistically
significant. Alpha diversity indexes, including Shannon index, Chao1,
Simpson, and abundance-based coverage estimate (ACE), were calculated in
QIIME1. Tukey's HSD test computed alpha index comparison in the R
Project (version 3.4.1). The expected sequences were classified into
organisms by a naive Bayesian model employing the RDP classifier
(version 2.2) based on the SILVA database (version 132) and ITS2
database (version update\_2015), with the confidence threshold values
ranging from 0.8 to 1 (Nilsson et al. 2019). The abundance statistics of
each taxonomy were visualized using Krona (version 2.6). Nonmetric
multidimensional scaling (NMDS) base of Bray-Curtis distances analysis
was performed using the `vegan' package to visualize microbial community
composition. A permutational multivariate ANOVA (PERMANOVA) test was
calculated (plotted in R ggplot2) and used to test the effect of slope
gradient and slope position on microbial community composition.
Redundancy analysis (RDA) performed was adopted to explore the
relationships between the composition of bacterial and fungal
communities and soil environmental factors.

\textbf{3. Results}

\textbf{3.1 Basic soil properties and enzyme activities}

Soil BD, ranging from 1.14 to 1.30 g cm\textsuperscript{-3}, was
significantly increased with slope gradient (p\textless{}0.05; Table 1).
Slope position did not affect soil BD. Soil CC
(18.64\%\textasciitilde{}27.23\%) was significantly higher at 0º-1.5º
slope gradient than other slope gradients (p\textless{}0.05) but did not
change significantly among slope positions. WSA content
(66.67\%\textasciitilde{}82.01\%) was significantly lower at 3º-5º and
\textgreater{}5º than other slope gradients but did not change with
slope positions significantly. Soil organic matter increased with the
decrease of slope gradient while there was no difference among slope
positions. Soil pH, CEC, AN, AP, AK, DOC, and TDN were not different
among slope gradients or positions. Either slope gradient or slope
position, but the interaction affected soil MBC and MBN (Table 1). Soil
INV activity decreased with increasing slope gradient. Slope position
did not affect soil INV activity. Soil UR, PHS, and CAT activities were
not different either among slope gradients or among slope positions.
There is no interaction between slope gradient and slope position on
enzyme activities (Table 2).

\textbf{3.2 Soil microbial community structure}

The relative abundances of major microbial phyla are illustrated
(Fig.2a, b). Proteobacteria, Acidobacteria, Gemmatimonadetes,
Planctomycetes, and Actinobacteria accounted for 83\% of the total
bacterial sequences, and Ascomycota, Basidiomycota, and chytridiomycota
accounted for 78\% of fungal. Bacteria alpha-diversity Chao1 and ACE
indices did not significantly differ among the slope gradients. Simpson
indices significantly differed among slope gradients and declined with
increasing slope gradient (Table 3). These alpha-diversity indices had
no significant differences between slope positions. Fungi
alpha-diversity indices were not significantly affected by slope
gradient or position. NMDS analysis based on the Bray-Curtis distance
clustered bacterial and fungal communities, and distinct differences
were observed in the communities' composition (Fig. 2c, d). PERMANOVA
revealed significant differences in bacterial and fungal community
composition, which is significantly lower in the highest slope gradient
level than in the other two levels. However, differences were not
significant among slope positions (Table 4).

\textbf{3.3 Relationships between soil properties, microbial community
composition, slope gradient and position}

Soil bacterial composition was positively correlated with WSA, CC, SOM,
CEC, AN, AP, AK, MBC, UR, and PHS but negatively correlated with BD, pH,
DOC, TDN, MBN, CAT, INV, slope gradient and slope position
(p\textless{}0.05) (Table 5). The fungi community was positively
correlated with BD, WSA, AP, TDN, MBC, phosphatase, catalases, and slope
position but negatively correlated with CC, OM, CEC, PH, AN, AK, DOC,
MBN, UR, INV, slope gradient and position (Table 5). RDA indicated a
clear difference in the composition of bacterial and fungal communities
at different slope gradients (Fig. 3). The first two axes explained
96.9\% of the overall variability for the bacterial community
composition (Fig. 3a). The first two axes explained 94.7\% of the
overall variability for the fungal community (Fig. 3b). Soil BD, CAT,
pH, INV, UR, CC, OC, and AN were significantly correlated with soil
microbial community.

\textbf{4. Discussion}

The effects of slope gradient and slope position on soil properties and
vegetation have been studied extensively (Bennie et al., 2006), but
their effects on soil microbial communities have remained largely
unexamined. Soil bacterial and fungal communities are ecologically
important groups, as soil bacteria have the largest biomass among soil
microorganisms to maintain soil structure. Soil fungi are essential in
soil aggregation and improving plant tolerance to drought and heavy
metal contamination (Fitter, 2005; Rillig and Mummey, 2006; van der
Heijden, 2008). In this study, the lower slope gradient may have suited
habitat for soil bacterial communities due to significantly higher
bacterial alpha-diversity and species richness. The community was
dominated by Proteobacteria, which is similar to the observations from
other soils collected from the Arctic (Chu et al., 2010), subalpine
(Shen et al., 2013), and agricultural soil environments (Sun et al.,
2015). The subsequent analysis of soil bacterial community composition
according to different groups in each slope gradient revealed increasing
differences among slope gradients. These results clarified soil
bacterial community distribution with increasingly apparent effects of
topography. Unlike bacterial alpha-diversity, the fungal alpha-diversity
was untouched among slope gradients (Table 3), suggesting its resistance
to slope gradients-induced interpretation in moisture, root biomass, and
soil C, N fractions (Sun et al., 2018). However, fungal community
composition showed strong sensitivity to slope gradients (Table 4),
indicating that the phyla-type composition of the fungal communities
could adapt accordingly to environmental changes without a general loss
in diversity (Yuste et al., 2014).

Soil physicochemical characteristics influence the soil microbial
community's composition, activity, and microbial mass level (Dick, 1994;
Xue et al., 2022). Clay content were higher in the lower slope gradient
which might be the reason of high presence of microbial communities.
Previous studies clarified that different soil textures influence the
structure of the microbial community, especially bacterial populations
(Fang et al., 2005). A possible explanation for the higher number of
bacteria in soil with high clay contents was documented by Carney and
Matson (2005). They mentioned that fine-textured soils support more
microbial biomass than coarse-textured soils. The distribution of
microorganisms in varied soil textures might be related to soil moisture
and nutrient contents, as Herioge et al. (2003) explained. RDA analysis
shows that pH highly correlated with microbial community (Fig. 3). The
importance of soil pH in shaping the bacterial population by vertical
soil profile has long been recognized (Yun et al., 2016). The close
relationship of soil pH with the bacterial community is owning to the
dependency of most of the bacterial community under narrow pH
(Charokopos et al., 2010), and the slight change in pH significantly
affects the bacterial community (Fernández-Calviño et al., 2010). Wei et
al. (2019) revealed that N, P, K, and pH are essential sources of
variation in soil fungal richness. Fungi play crucial roles in
decomposing organic matter and forming an association with the roots for
belowground carbon transport and respiration in the boreal forest
(Clemmensen et al., 2013). Soil nutrients released by microbes are
available for aboveground communities, supporting essential ecosystem
services such as food and fiber production (Wardle et al., 2004).

\textbf{5. Conclusion}

The effect of slope gradient and slope position on soil bacterial and
fungal diversities and community compositions in black soil were
analyzed using the 16S and ITS rRNA Illumina sequencing. The dominant
phyla were Proteobacteria, Acidobacteria, Gemmatimonadetes,
Planctomycetes, and Actinobacteria, representing the most dominant phyla
of bacteria. The most abundant phyla for the fungal community were
Ascomycota, Anthophyta, Basidiomycota, Chlorophyte, and Chytridimycota.
Microbial alpha-diversity (Bacteria and Fungi) was similar among the
slope gradients or slope positions. The bacterial and fungal community
composition was only affected by slope gradient, which was significantly
lower in the highest slope gradient level than others. This study
suggests that slope gradient but not slope position affecting microbial
community composition in the sloped cropland area.

\textbf{Acknowledgement} We wish to thank the National Key Research and
Development Project (Grant No. 2018YFC0507005), National Natural Science
Foundation of China (Grant No. 32071840) and National Natural Science
Foundation of China (Grant No. 31070627) for funding this research.
Abdul Hakim Jamshidi and Lei Sun contributed equally to this work. We
further thanks to Yannan Xu, Xiang Li, Xiangwei Yang, Wentao Di, Wenjuan
Huo and many others for assisting with soil sample collection and
analyses. We are extremely appreciative of the help given by Gene Denovo
Corp. during microbial community analysis.

Table 1 Effect of slope gradient and slope position on physico-chemical
and biological properties.\selectlanguage{english}
\begin{longtable}[]{@{}lllllllllllllll@{}}
\toprule
\begin{minipage}[b]{0.04\columnwidth}\raggedright\strut
Factor\strut
\end{minipage} & \begin{minipage}[b]{0.04\columnwidth}\raggedright\strut
Levels\strut
\end{minipage} & \begin{minipage}[b]{0.04\columnwidth}\raggedright\strut
BD\strut
\end{minipage} & \begin{minipage}[b]{0.04\columnwidth}\raggedright\strut
CC\strut
\end{minipage} & \begin{minipage}[b]{0.04\columnwidth}\raggedright\strut
WSA\strut
\end{minipage} & \begin{minipage}[b]{0.04\columnwidth}\raggedright\strut
pH\strut
\end{minipage} & \begin{minipage}[b]{0.04\columnwidth}\raggedright\strut
CEC\strut
\end{minipage} & \begin{minipage}[b]{0.04\columnwidth}\raggedright\strut
OM\strut
\end{minipage} & \begin{minipage}[b]{0.04\columnwidth}\raggedright\strut
AN\strut
\end{minipage} & \begin{minipage}[b]{0.04\columnwidth}\raggedright\strut
AP\strut
\end{minipage} & \begin{minipage}[b]{0.04\columnwidth}\raggedright\strut
AK\strut
\end{minipage} & \begin{minipage}[b]{0.04\columnwidth}\raggedright\strut
DOC\strut
\end{minipage} & \begin{minipage}[b]{0.04\columnwidth}\raggedright\strut
TDN\strut
\end{minipage} & \begin{minipage}[b]{0.04\columnwidth}\raggedright\strut
MBC\strut
\end{minipage} & \begin{minipage}[b]{0.04\columnwidth}\raggedright\strut
MBN\strut
\end{minipage}\tabularnewline
\midrule
\endhead
\begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
SG\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
0\selectlanguage{ngerman}º-1.5º\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
1.22 \selectlanguage{ngerman}±0.01a\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
27.23 \selectlanguage{ngerman}±0.81a\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
81.53 \selectlanguage{ngerman}±1.35a\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
5.38 \selectlanguage{ngerman}±0.78\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
32.76 \selectlanguage{ngerman}±1.21\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
55.26 \selectlanguage{ngerman}±0.80a\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
212.56 \selectlanguage{ngerman}±9.32\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
61.69 \selectlanguage{ngerman}±4.99\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
294.61 \selectlanguage{ngerman}±37.83\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
24.05 \selectlanguage{ngerman}±1.34\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
4.84 \selectlanguage{ngerman}±0.64\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
39.93 \selectlanguage{ngerman}±7.94\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
3.88 \selectlanguage{ngerman}±0.94\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
1.5\selectlanguage{ngerman}º-3º\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
1.14 \selectlanguage{ngerman}±0.02b\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
18.64 \selectlanguage{ngerman}±1.15b\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
82.01 \selectlanguage{ngerman}±1.54a\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
5.20 \selectlanguage{ngerman}±0.22\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
37.03 \selectlanguage{ngerman}±1.09\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
43.12 \selectlanguage{ngerman}±3.01a\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
232.47 \selectlanguage{ngerman}±11.41\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
62.12 \selectlanguage{ngerman}±8.40\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
338.55 \selectlanguage{ngerman}±15.73\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
21.07 \selectlanguage{ngerman}±1.28\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
8.55 \selectlanguage{ngerman}±1.57\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
22.92 \selectlanguage{ngerman}±5.82\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
8.18 \selectlanguage{ngerman}±2.05\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
3\selectlanguage{ngerman}º-5º\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
1.28 \selectlanguage{ngerman}±0.03a\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
21.46 \selectlanguage{ngerman}±0.79b\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
78.92 \selectlanguage{ngerman}±2.26a\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
5.38 \selectlanguage{ngerman}±0.07\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
30.46 \selectlanguage{ngerman}±2.94\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
31.17 \selectlanguage{ngerman}±4.94b\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
191.48 \selectlanguage{ngerman}±18.69\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
56.02 \selectlanguage{ngerman}±11.61\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
296.11 \selectlanguage{ngerman}±22.28\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
22.21 \selectlanguage{ngerman}±1.65\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
5.59 \selectlanguage{ngerman}±1.97\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
18.70 \selectlanguage{ngerman}±6.06\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
6.85 \selectlanguage{ngerman}±1.03\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
\textgreater{}5\selectlanguage{ngerman}º\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
1.30 \selectlanguage{ngerman}±0.03a\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
18.86 \selectlanguage{ngerman}±0.89b\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
66.67 \selectlanguage{ngerman}±3.08b\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
5.64 \selectlanguage{ngerman}±0.07\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
32.53 \selectlanguage{ngerman}±3.44\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
38.18 \selectlanguage{ngerman}±3.44b\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
200.48 \selectlanguage{ngerman}±13.65\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
51.10 \selectlanguage{ngerman}±6.35\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
221.90 \selectlanguage{ngerman}±10.48\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
24.28 \selectlanguage{ngerman}±1.85\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
7.46 \selectlanguage{ngerman}±0.96\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
18.75 \selectlanguage{ngerman}±7.11\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
11.01 \selectlanguage{ngerman}±3.20\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
SP\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
Top\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
1.23 \selectlanguage{ngerman}±0.03\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
20.72 \selectlanguage{ngerman}±1.53\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
79.65 \selectlanguage{ngerman}±1.96\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
5.36 \selectlanguage{ngerman}±0.07\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
31.22 \selectlanguage{ngerman}±2.21\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
43.53 \selectlanguage{ngerman}±3.53\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
222.39 \selectlanguage{ngerman}±11.42\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
53.10 \selectlanguage{ngerman}±5.93\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
296.85 \selectlanguage{ngerman}±25.22\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
22.50 \selectlanguage{ngerman}±1.25\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
6.22 \selectlanguage{ngerman}±1.20\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
33.75 \selectlanguage{ngerman}±7.43\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
5.88 \selectlanguage{ngerman}±1.78\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
Middle\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
1.23 \selectlanguage{ngerman}±0.02\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
22.15 \selectlanguage{ngerman}±1.32\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
77.34 \selectlanguage{ngerman}±2.20\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
5.42 \selectlanguage{ngerman}±0.07\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
35.08 \selectlanguage{ngerman}±1.23\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
41.89 \selectlanguage{ngerman}±3.98\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
209.53 \selectlanguage{ngerman}±11.90\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
65.22 \selectlanguage{ngerman}±8.50\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
291.87 \selectlanguage{ngerman}±23.69\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
22.78 \selectlanguage{ngerman}±1.23\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
7.67 \selectlanguage{ngerman}±1.49\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
26.99 \selectlanguage{ngerman}±6.03\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
8.93 \selectlanguage{ngerman}±1.54\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
Bottom\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
1.21 \selectlanguage{ngerman}±0.02\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
22.66 \selectlanguage{ngerman}±1.25\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
79.09 \selectlanguage{ngerman}±2.85\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
5.32 \selectlanguage{ngerman}±0.20\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
33.17 \selectlanguage{ngerman}±2.43\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
41.89 \selectlanguage{ngerman}±4.92\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
199.29 \selectlanguage{ngerman}±14.74\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
56.03 \selectlanguage{ngerman}±6.69\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
300.20 \selectlanguage{ngerman}±28.05\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
22.90 \selectlanguage{ngerman}±1.57\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
5.38 \selectlanguage{ngerman}±1.19\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
16.63 \selectlanguage{ngerman}±4.65\strut
\end{minipage} & \begin{minipage}[t]{0.06\columnwidth}\raggedright\strut
5.92 \selectlanguage{ngerman}±1.34\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{P-value}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{SG}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
0.004\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
0.000\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
0.000\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.310}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.169}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
0.000\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.189}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.833}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.074}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.407}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.27}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.104}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.073}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{SP}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.880}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.610}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.763}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.878}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.38}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.951}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.469}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.477}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.973}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.978}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.47}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.180}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.287}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\selectlanguage{ngerman}\emph{SG×SP}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.122}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.593}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.922}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.618}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.445}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.073}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.592}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{1.059}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.466}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.944}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.888}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{1.495}\strut
\end{minipage} & \begin{minipage}[t]{0.04\columnwidth}\raggedright\strut
\emph{0.438}\strut
\end{minipage}\tabularnewline
\bottomrule
\end{longtable}

Note; \emph{BD} Bulk Density (g cm\textsuperscript{-3}), \emph{CC} Clay
Content (\%), \emph{WSA} Water Stable Aggregate (\%), \emph{CEC} Cation
Exchange Capacity (cmol (+) kg-1), \emph{OM} Organic
Matter(g/kg),\emph{AN} Available Nitrogen(mg/kg), \emph{AP} Available
Phosphorus(mg/kg), \emph{AK} Available Potassium(mg/kg),
\emph{DOC}Dissolved Organic Carbon(mg/L), \emph{TDN} Total Dissolved
Nitrogen(mg/L), \emph{MBC} Microbial Biomass Carbon(mg/kg),
\emph{MBN}Microbial Biomass Nitrogen(mg/kg), \emph{SG} slope gradient,
\emph{SP}slope position, se (\selectlanguage{ngerman}±) = Standard Error. Means observed by the
same letter within the same column for each site are not significantly
different at \selectlanguage{greek}α \selectlanguage{english}= 0.05 according to Tukey's test.

Table 2 Effect of slope gradient and slope positions on enzyme activity\selectlanguage{english}
\begin{longtable}[]{@{}llllll@{}}
\toprule
Factors & Levels & UR & ACP & CAT & SC\tabularnewline
\midrule
\endhead
SG & 0\selectlanguage{ngerman}º-1.5º & 23.23\selectlanguage{ngerman}±6.73 & 11.35\selectlanguage{ngerman}±0.63 & 95.50\selectlanguage{ngerman}±0.67 &
34.17\selectlanguage{ngerman}±3.07b\tabularnewline
& 1.5\selectlanguage{ngerman}º-3º & 13.27\selectlanguage{ngerman}±4.45 & 11.28\selectlanguage{ngerman}±0.50 & 95.50\selectlanguage{ngerman}±0.97 &
19.87\selectlanguage{ngerman}±3.76a\tabularnewline
& 3\selectlanguage{ngerman}º-5º & 21.12\selectlanguage{ngerman}±3.41 & 9.84\selectlanguage{ngerman}±0.53 & 97.80\selectlanguage{ngerman}±0.53 &
18.47\selectlanguage{ngerman}±3.14a\tabularnewline
& \textgreater{} 5\selectlanguage{ngerman}º & 18.82\selectlanguage{ngerman}±6.65 & 9.17\selectlanguage{ngerman}±0.46 & 97.61\selectlanguage{ngerman}±0.23 &
17.00\selectlanguage{ngerman}±2.62a\tabularnewline
SP & Top & 13.37\selectlanguage{ngerman}±3.07 & 10.42\selectlanguage{ngerman}±0.56 & 96.31\selectlanguage{ngerman}±0.82 &
19.21\selectlanguage{ngerman}±2.77\tabularnewline
& Middle & 21.47\selectlanguage{ngerman}±5.04 & 10.55\selectlanguage{ngerman}±0.64 & 96.49\selectlanguage{ngerman}±0.68 &
20.58\selectlanguage{ngerman}±3.36\tabularnewline
& Bottom & 22.14\selectlanguage{ngerman}±4.97 & 10.72\selectlanguage{ngerman}±0.36 & 96.62\selectlanguage{ngerman}±0.66 &
22.58\selectlanguage{ngerman}±3.61\tabularnewline
\emph{P-value} & \emph{SG} & \emph{0.541} & \emph{0.043} & \emph{0.048}
& \emph{0.012}\tabularnewline
& \emph{SP} & \emph{0.342} & \emph{0.935} & \emph{0.954} &
\emph{0.767}\tabularnewline
& \selectlanguage{ngerman}\emph{SG×SP} & \emph{0.572} & \emph{0.924} & \emph{0.306} &
\emph{0.549}\tabularnewline
\bottomrule
\end{longtable}

\textbf{Note;} \emph{UR} Urease (nmol NH4
g\textsuperscript{-1}h\textsuperscript{-1}), \emph{ACP} Phosphatase U/g
(1 nmol phenol released per gram of soil in 37\selectlanguage{ngerman}ºC is one enzyme active
unite per day),\emph{CAT} Catalases U/g (enzyme activity per g of
air-dried soil sample per day for catalytic degradation of 1mmol
H\textsubscript{2}O\textsubscript{2}.), \emph{SC} invertase U/g
(production of 1mg reducing sugars per g of soil samples per
day),\emph{SG} slope gradient, \emph{SP} slope position, se (±) =
Standard Error. Means observed by the same letter within the same column
for each site are not significantly different at \selectlanguage{greek}α \selectlanguage{english}= 0.05 according to
Tukey's test.

Table 3 Results of ANOVA for all data on Alpha diversity including
different slope gradients and slope positions\selectlanguage{english}
\begin{longtable}[]{@{}llll@{}}
\toprule
Alpha diversity & diff\_groups & p-value(B) & p-value(F)\tabularnewline
\midrule
\endhead
Simpson & LT-vs-LM-vs-LB & 0.837 & 0.133\tabularnewline
Simpson & MT-vs-MM-vs-MB & 0.708 & 0.474\tabularnewline
Simpson & HT-vs-HM-vs-HB & 0.534 & 0.697\tabularnewline
Simpson & LT-vs-MT-vs-HT & 0.076 & 0.426\tabularnewline
Simpson & LM-vs-MM-vs-HM & 0.088 & 0.673\tabularnewline
Simpson & LB-vs-MB-vs-HB & \textbf{0.014} & 0.51\tabularnewline
Chao1 & LT-vs-LM-vs-LB & 0.14 & 0.719\tabularnewline
Chao1 & MT-vs-MM-vs-MB & 0.786 & 0.873\tabularnewline
Chao1 & HT-vs-HM-vs-HB & 0.646 & 0.846\tabularnewline
Chao1 & LT-vs-MT-vs-HT & 0.434 & 0.51\tabularnewline
Chao1 & LM-vs-MM-vs-HM & 0.166 & 0.798\tabularnewline
Chao1 & LB-vs-MB-vs-HB & 0.497 & 0.998\tabularnewline
ACE & LT-vs-LM-vs-LB & 0.13 & 0.793\tabularnewline
ACE & MT-vs-MM-vs-MB & 0.753 & 0.779\tabularnewline
ACE & HT-vs-HM-vs-HB & 0.861 & 0.897\tabularnewline
ACE & LT-vs-MT-vs-HT & 0.429 & 0.604\tabularnewline
ACE & LM-vs-MM-vs-HM & 0.199 & 0.858\tabularnewline
ACE & LB-vs-MB-vs-HB & 0.574 & 0.965\tabularnewline
\bottomrule
\end{longtable}

Note; B; Bacteria, F; Fungi, Values in columns represent p-values based
on way ANOVA. Bold \emph{p} -values indicate significant differences
(\emph{p} \textless{} 0.05). L, low slope gradient, M medium slope
gradient, H high slope gradient, T top slope position, M middle slope
position, B bottom slope position.

Table 4 PERMANOVA analysis of soil bacterial and fungal community
composition among different slope gradients and slope positions in
sequencing data based on Bray-Curtis distances.\selectlanguage{english}
\begin{longtable}[]{@{}lllll@{}}
\toprule
Variables & \emph{Bacteria} & \emph{Bacteria} & \emph{Fungi} &
\emph{Fungi}\tabularnewline
\midrule
\endhead
& \emph{R2} & \emph{p-value} & \emph{R2} & \emph{p-value}\tabularnewline
LT-vs-LM-vs-LB & 0.1193 & 0.195 & 0.2181 & 0.157\tabularnewline
MT-vs-MM-vs-MB & -0.0617 & 0.585 & -0.284 & 0.886\tabularnewline
HT-vs-HM-vs-HB & -0.1852 & 0.828 & -0.1605 & 0.927\tabularnewline
LT-vs-MT-vs-HT & 0.9424 & \textbf{0.009} & 0.7695 &
\textbf{0.002}\tabularnewline
LM-vs-MM-vs-HM & 0.9342 & \textbf{0.001} & 0.7202 &
\textbf{0.006}\tabularnewline
LB-vs-MB-vs-HB & 0.6461 & \textbf{0.01} & 0.5967 &
\textbf{0.012}\tabularnewline
\bottomrule
\end{longtable}

\textbf{Note:} Significance analysis was based on 999 permutation tests.
Bold \emph{p} -values indicate significant differences
(\emph{p}\textless{} 0.05). L, low slope gradient, M medium slope
gradient, H high slope gradient, T top slope position, M middle slope
position, B bottom slope position.

Table 5 Correlation coefficients (Spearman) between soil
physico-chemical properties, enzyme activities, microbial community,
slope gradient and slope position.\selectlanguage{english}
\begin{longtable}[]{@{}lll@{}}
\toprule
& Bacterial OTU & Fungi OTU\tabularnewline
\midrule
\endhead
Bulk Density & -0.317 & 0.717*\tabularnewline
Water Stable Aggregate & 0.833** & 0.35\tabularnewline
Clay Content & 0.65 & -0.317\tabularnewline
Organic Matter & 0.333 & -0.45\tabularnewline
pH & -0.216 & -0.700*\tabularnewline
Cation Exchange Capacity & 0.483 & -0.117\tabularnewline
Available Nitrogen & 0.05 & -0.416\tabularnewline
Available Phosphorus & 0.133 & 0.116\tabularnewline
Available Potassium & 0.683* & -0.1\tabularnewline
Dissolved Organic Carbon & -0.1 & -0.483\tabularnewline
Total Dissolved Nitrogen & -0.383 & 0.016\tabularnewline
Microbial Biomass Carbon & 0.717* & 0.25\tabularnewline
Microbial Biomass Nitrogen & -0.883** & -0.2\tabularnewline
Urease & 0.05 & -.817**\tabularnewline
Phosphatase & 0.266 & 0.3\tabularnewline
Catalases & -0.283 & 0.416\tabularnewline
Invertase & -0.516 & -0.466\tabularnewline
Slope Gradient & -0.896** & -0.210\tabularnewline
Slope Position & -0.158 & 0.685*\tabularnewline
\bottomrule
\end{longtable}

Note; * and ** represent P \textless{} 0.05 and P \textless{} 0.01,
respectively.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image1/image1}

\end{center}
\end{figure}

Fig. 1 Geographical location of the study sites and distribution of
sampling points\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image2/image2}

\end{center}
\end{figure}

\textbf{Fig.2}Relative abundance of the bacterial and (a) and fungal (b)
communities at the phylum level based on Illumina sequencing of the 16S
and ITS2 region. Nonmetric multidimensional scaling (NMDS) based on
Bray-Curtis dissimilarity displaying effect of slope gradient and
position on soil bacterial (c) and fungal (d) communities' composition.
L, low slope gradient, M medium slope gradient, H high slope gradient, T
top slope position, M middle slope position, B bottom slope position.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image3/image3}

\end{center}
\end{figure}

\textbf{Fig. 3.} Redundancy analysis (RDA) of soil bacterial (a) and
fungal (b) community composition with soil physicochemical parameter. L,
low slope gradient, M medium slope gradient, H high slope gradient, T
top slope position, M middle slope position, B bottom slope
position.\emph{BD} Bulk Density (g cm\textsuperscript{-3}), \emph{CC}
Clay Content (\%), \emph{WSA} Water Stable Aggregate (\%), \emph{CEC}
Cation Exchange Capacity (cmol (+) kg-1), \emph{OM} Organic
Matter(g/kg),\emph{AN} Available Nitrogen(mg/kg), \emph{AP} Available
Phosphorus(mg/kg), \emph{AK} Available Potassium(mg/kg),
\emph{DOC}Dissolved Organic Carbon(mg/L), \emph{TDN} Total Dissolved
Nitrogen(mg/L), \emph{MBC} Microbial Biomass Carbon(mg/kg),
\emph{MBN}Microbial Biomass Nitrogen(mg/kg).\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/fig1/fig1}

\end{center}
\end{figure}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/fig-2/fig-2}

\end{center}
\end{figure}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/fig3/fig3}

\end{center}
\end{figure}

\selectlanguage{english}
\FloatBarrier
\end{document}

