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Journal of Arid Land  2021, Vol. 13 Issue (3): 257-273    DOI: 10.1007/s40333-021-0004-6
Research article     
Soil bacterial characteristics between surface and subsurface soils along a precipitation gradient in the Alxa Desert, China
TENG Zeyu1,2,*(), XIAO Shengchun1, CHEN Xiaohong3, HAN Chao1,2
1Key Laboratory of Ecohydrology of Inland River Basin, Northwest Institute of Eco-Environmental Resources, Chinese Academy of Sciences, Lanzhou 730000, China
2University of Chinese Academy of Sciences, Beijing 100049, China
3Key Laboratory of Desert and Desertification, Northwest Institute of Eco-Environment and Resources, Chinese Academy of Sciences, Lanzhou 730000, China
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Bacteria in desert soil have unique phylogeny and important ecological functions, and their responses to changes in precipitation need further attention. However, relevant studies have mainly focused on the surface soil, and studies on the responses of bacteria at different soil depths to variations in precipitation are rare. Thus, we used 16S rDNA high-throughput sequencing to investigate the changes in soil bacterial distribution along a mean annual precipitation gradient (50-150 mm) in the Alxa Desert, China, and compared the variation characteristics in the surface soil layer (0-10 cm) and subsurface soil layer (10-20 cm). Results showed that soil bacterial communities significantly changed along the precipitation gradient in both soil layers. However, the subsurface soil layer could support bacterial communities with higher diversity and closer internal relationships but more internal competition than the surface soil layer. Additionally, compared with the surface soil layer, variations in diversity and co-occurrence patterns in the subsurface soil layer were more in line with the changes in the mean annual precipitation, while bacterial community structure was less variable in the subsurface soil layer. Compared with the mean annual precipitation, soil moisture had little influence on the structure and diversity of soil bacterial community but had a high correlation with intercommunity connectivity. Therefore, soil moisture might play a complex role in mediating environmental conditions and soil bacterial community characteristics. Due to the different responses of surface and subsurface soil bacteria to the changes in precipitation, it is necessary to distinguish different soil layers when predicting the trends in desert soil bacterial conditions associated with precipitation, and prediction of subsurface soil bacteria may be more accurate.

Key wordsprecipitation gradient      soil layer      soil bacterial community structure      diversity      co-occurrence pattern     
Received: 22 September 2020      Published: 10 March 2021
Corresponding Authors: TENG Zeyu     E-mail:
About author: * TENG Zeyu (E-mail:
Cite this article:

TENG Zeyu, XIAO Shengchun, CHEN Xiaohong, HAN Chao. Soil bacterial characteristics between surface and subsurface soils along a precipitation gradient in the Alxa Desert, China. Journal of Arid Land, 2021, 13(3): 257-273.

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Type Sampling site Latitude and longitude Elevation
Dominant species Coverage
Soil type
Typical desert
(MAP=50 mm)
Td1 40°55′22″N, 100°52′40″E 1031 Nitraria tangutorum, Haloxylon
Reaumuria songarica, Calligonum mongolicum
7.01 Gray-brown calcic soil
Td2 41°23′48″N, 102°21′26″E 904 2.95
Td3 41°17′53″N, 104°08′45″E 794 3.18
Steppe desert
(MAP=100 mm)
Sd1 40°21′21″N, 104°44′19″E 1278
Nitraria tangutorum, Zygophyllum xanthoxylon,
Reaumuria songarica, Caragana korshinskii,
Pearl russianthistle
17.93 Gray-brown desert soil
Sd2 39°28′13″N, 105°36′16″E 1089 15.45
Sd3 40°03′12″N, 103°55′08″E 1471 26.87
Sd4 39°37′16″N, 103°07′54″E 1232 12.29
Desertified steppe
(MAP=150 mm)
Ds1 38°16′34″N, 103°56′35″E 1394 Nitraria tangutorum, Caragana korshinskii,
Oxytropis aciphylla, Reaumuria songarica, Ammopiptanthus mongolicus,
Zygophyllum xanthoxylon
35.66 Gray desert soil
Ds2 38°54′33″N, 105°30′15″E 1345 52.76
Ds3 37°54′02″N, 105°12′31″E 1416 28.44
Table 1 Location and environmental characteristics of sampling sites
Fig. 1 Relative abundance of soil bacterial at phylum level in different MAP (mean annual precipitation)-related subregions and soil layers. Td, typical desert (50 mm precipitation); Sd, steppe desert (100 mm precipitation); Ds, desertified steppe (150 mm precipitation); A, 0-10 cm soil layer; B, 10-20 cm soil layer.
Fig. 2 Alpha diversity index in different MAP-related subregions and soil layers. Td, typical desert (50 mm precipitation); Sd, steppe desert (100 mm precipitation); Ds, desertified steppe (150 mm precipitation); A, 0-10 cm soil layer; B, 10-20 cm soil layer; PD, phylogenetic diversity. The lowercase letters indicate significant differences among different treatments at P<0.05 level.
Group F value P value Group F value P value
Ds-Sd 3.021 0.008 Td.A-Sd.A 16.727 <0.001
Sd-Td 12.343 <0.001 Td.A-Ds.A 15.883 <0.001
Ds-Td 12.203 <0.001 Sd.A-Ds.A 4.081 0.004
Td.A-Td.B 5.069 0.007 Td.B-Sd.B 4.615 <0.001
Sd.A-Sd.B 7.222 <0.001 Td.B-Ds.B 7.022 <0.001
Ds.A-Ds.B 17.155 <0.001 Sd.B-Ds.B 2.925 0.014
A-B 13.404 <0.001
Table 2 Result of analysis of molecular variance (AMOVA) in different MAP-related subregions and soil layers
Fig. 3 Cladograms generated according to LEfSe analysis for the precipitation divisions in layer A (a) and layer B (b). Biomarkers were statistically significant at LDA ≥4. Cladograms indicating the phylogenetic distribution of bacterial lineages associated with the three subregions. Phylum, class, order, family, genus and species levels are listed in order from inside to outside of the cladogram. Red, green and blue circles represent the bacteria enriched in Ds, Sd and Td subregions, respectively, whereas yellow circle represents the taxa showing no significant difference among the three subregions. Labels for biomarkers for the class, order and family levels are abbreviated with a single letter. For the genus and species levels, please see Figures S1 and S2 for details.
Fig. 4 Cladogram generated according to LEfSe analysis for the soil layers and biomarkers showing statistical significance at LDA ≥4. Red and green circles represent the bacteria enriched in the A and B soil layers, respectively.
Index Td.A Td.B Sd.A Sd.B Ds.A Ds.B
SOC (g/kg) 1.11±0.71a 0.89±0.35a 1.09±0.53a 0.84±0.21a 1.62±1.15a 1.33±0.86a
TN (g/kg) 0.16±0.06b 0.21±0.05ab 0.26±0.10ab 0.21±0.10ab 0.31±0.11a 0.20±0.10ab
TC (g/kg) 3.85±0.66a 4.76±0.53a 5.87±4.16a 4.10±1.99a 4.61±4.02a 4.07±3.78a
TP (g/kg) 0.28±0.07a 0.30±0.11a 0.25±0.03a 0.21±0.05b 0.16±0.04c 0.15±0.02c
AN (mg/kg) 15.10±9.82b 16.28±12.12b 32.39±9.59a 33.95±3.69a 27.54±10.5ab 25.27±9.11ab
AP (mg/kg) 1.58±0.20b 1.42±0.54b 2.15±0.12ab 2.25±1.91ab 3.10±1.03a 1.58±0.63b
AK (mg/kg) 99.60±31.33b 113.43±16.60b 155.63±40.83a 112.05±39.01b 127.27±47.86ab 135.57±46.21ab
SMC (%) 1.33±0.78b 2.47±0.01a 1.30±0.60b 2.66±1.00a 3.07±0.76a 3.92±1.55a
SO2- 4 (cmol/kg) 4.77±4.75b 10.31±5.83a 1.95±3.31b 1.92±3.31b 0.12±0.10c 0.09±0.06c
Cl- (cmol/kg) 17.55±28.02a 5.76±6.24a 0.50±0.10b 0.15±0.10bc 0.07±0.01c 0.07±0.01c
HCO3- (cmol/kg) 0.27±0.06b 0.23±0.04c 0.36±0.08ab 0.38±0.08a 0.38±0.06a 0.39±0.06a
Ca2+ (cmol/kg) 2.97±2.09b 7.71±4.36a 1.90±3.07bc 1.82±3.07bc 0.32±0.08c 0.27±0.05c
Mg2+ (cmol/kg) 0.63±0.80a 0.44±0.29a 0.18±0.13b 0.17±0.13b 0.11±0.08b 0.12±0.04b
K+ (cmol/kg) 0.13±0.11a 0.12±0.06a 0.04±0.04b 0.04±0.04b 0.04±0.02b 0.05±0.01b
Na+ (cmol/kg) 15.78±25.01a 6.10±6.29a 0.34±0.21b 0.32±0.21b 0.02±0.03c 0.05±0.02c
EC (dS/m) 8.04±11.31a 5.06±3.86a 1.36±6.14a 4.34±6.15a 0.31±0.03b 0.39±0.05b
pH 7.44±0.16b 7.38±0.05b 7.67±0.16a 7.80±0.16a 7.72±0.08a 7.70±0.12a
VCS (%) 5.91±2.71a 3.22±3.63b 0.22±0.44c 0.03±0.07d 0.72±0.66c 0.33±0.29c
CS (%) 14.30±5.99a 8.89±6.82ab 3.84±1.02b 5.95±5.24b 21.07±18.57a 13.05±11.79a
MS (%) 21.58±9.17a 21.28±8.50a 26.69±8.40a 25.57±15.04a 27.67±17.71a 27.31±16.48a
FS (%) 43.32±7.39a 48.24±7.74a 47.26±10.84a 50.13±21.77a 32.89±18.77a 40.77±12.22a
VFS (%) 11.13±5.53ab 12.66±3.23a 8.78±3.91ab 7.90±5.14b 11.43±9.37ab 11.47±6.58ab
Silt (%) 3.20±1.61a 4.82±2.31a 11.20±11.19a 9.10±9.37a 5.33±7.42a 6.20±7.99a
Clay (%) 0.57±0.50a 0.88±0.67a 2.03±2.27a 1.32±1.67a 0.89±1.55a 0.87±1.51a
TCn (g/kg) 384.71±6.85a 377.66 ± 4.24b 377.23±7.59b
TNn (g/kg) 18.39±1.92a 20.45 ± 7.31a 21.32±6.94a
TPn (g/kg) 1.23±0.08ab 1.52 ± 0.78a 1.15±0.09b
COV (%) 4.38±1.98c 18.14 ± 5.67b 38.95±10.82a
Table 3 Differences in environmental factors (soil nutrients, moisture, salinity, pH, particle size distribution, N. tangutorum stoichiometric characteristic and vegetation coverage) in different MAP-related subregions and soil layers
Fig. 5 Canonical correlation analysis (CCA) ordination of bacterial community structure at the genus level in relation to major environmental factors in layers A (a) and B (b)
Layer A r value P value Layer B r value P value
VCS 0.7811 <0.001 Ca2+ 0.7993 <0.001
Na+ 0.6744 <0.001 SO2- 4 0.7242 <0.001
pH 0.6366 <0.001 Cl- 0.6586 <0.001
MAP 0.6125 <0.001 Na+ 0.6384 <0.001
Cl- 0.5879 <0.001 HCO3- 0.5660 <0.001
SO2- 4 0.5877 <0.001 K+ 0.4798 <0.001
EC 0.5766 <0.001 TP 0.4403 <0.001
Ca2+ 0.5717 <0.001 Mg2+ 0.4013 0.002
Mg2+ 0.4498 <0.001 pH 0.3987 <0.001
K+ 0.4090 <0.001 EC 0.3601 <0.001
AN 0.3987 <0.001 VCS 0.3558 <0.006
COV 0.3099 0.003 MAP 0.3397 <0.001
TCn 0.2961 <0.001 AN 0.2644 <0.001
AP 0.2855 0.004 COV 0.2077 0.009
VFS 0.2020 0.008
Table 4 Mantel test comparing the correlation between environmental factors and soil bacterial community structure in layer A and layer B
Td.A 0.6502 0.4980 0.0242 4.443 24.41
Sd.A 0.5014 0.5539 0.0230 4.272 39.15
Ds.A 0.3332 0.5299 0.0354 3.691 25.14
Td.B 0.5216 0.5267 0.0332 4.072 28.19
Sd.B 0.5512 0.5407 0.0489 3.576 36.01
Ds.B 0.3704 0.5721 0.0527 3.303 40.14
Table 5 Parameters of network topology analysis (genus level) in different MAP-related subregions and soil layers
Fig. S1 Histogram of linear discriminant analysis (LDA) score distribution shows the categories with LDA score greater than the set value (≥ 4) in layer A (0-10 cm) along the precipitation gradient. The length of the histogram indicates the magnitude of the influence of biomarkers.
Fig. S2 Histogram of linear discriminant analysis (LDA) score distribution shows the categories with LDA score greater than the set value (≥ 4) in layer B (10-20 cm) along the precipitation gradient. The length of the histogram indicates the magnitude of the influence of biomarkers.
Fig. S3 Major environmental factors associated with alpha diversity in layers A (a) and B (b). Environmental factors are significantly correlated with at least one alpha diversity index. Asterisks indicate significance of the Spearman's r correlation. *, P<0.05 level; **, P<0.01 level.
Fig. S4 Major environmental factors associated with parameters of network topology analysis. Environmental factors are significantly correlated with at least one parameter. Asterisks indicate significance of the Spearman's r correlation. *, P<0.05 level; **, P<0.01 level.
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