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Journal of Arid Land  2023, Vol. 15 Issue (9): 1107-1128    DOI: 10.1007/s40333-023-0025-4
Research article     
Leguminosae plants play a key role in affecting soil physical-chemical and biological properties during grassland succession after farmland abandonment in the Loess Plateau, China
SUN Lin, YU Zhouchang, TIAN Xingfang, ZHANG Ying, SHI Jiayi, FU Rong, LIANG Yujie, ZHANG Wei*()
College of Grassland Agriculture, Northwest A&F University, Yangling 712100, China
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Leguminosae are an important part of terrestrial ecosystems and play a key role in promoting soil nutrient cycling and improving soil properties. However, plant composition and species diversity change rapidly during the process of succession, the effect of leguminosae on soil physical-chemical and biological properties is still unclear. This study investigated the changes in the composition of plant community, vegetation characteristics, soil physical-chemical properties, and soil biological properties on five former farmlands in China, which had been abandoned for 0, 5, 10, 18, and 30 a. Results showed that, with successional time, plant community developed from annual plants to perennial plants, the importance of Leguminosae and Asteraceae significantly increased and decreased, respectively, and the importance of grass increased and then decreased, having a maximum value after 5 a of abandonment. Plant diversity indices increased with successional time, and vegetation coverage and above- and below-ground biomass increased significantly with successional time after 5 a of abandonment. Compared with farmland, 30 a of abandonment significantly increased soil nutrient content, but total and available phosphorus decreased with successional time. Changes in plant community composition and vegetation characteristics not only change soil properties and improve soil physical-chemical properties, but also regulate soil biological activity, thus affecting soil nutrient cycling. Among these, Leguminosae have the greatest influence on soil properties, and their importance values and community composition are significantly correlated with soil properties. Therefore, this research provides more scientific guidance for selecting plant species to stabilize soil ecosystem of farmland to grassland in the Loess Plateau, China.

Key wordssecondary succession      leguminosae      plant diversity      plant community composition      soil physical-chemical properties      soil biological properties     
Received: 21 March 2023      Published: 30 September 2023
Corresponding Authors: * ZHANG Wei (E-mail:
About author: First author contact:The first and second authors contributed equally to this work.
Cite this article:

SUN Lin, YU Zhouchang, TIAN Xingfang, ZHANG Ying, SHI Jiayi, FU Rong, LIANG Yujie, ZHANG Wei. Leguminosae plants play a key role in affecting soil physical-chemical and biological properties during grassland succession after farmland abandonment in the Loess Plateau, China. Journal of Arid Land, 2023, 15(9): 1107-1128.

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Fig. 1 Sampling process of the Zhifanggou watershed in the Loess Plateau, China
Index Years of farmland abandonment F P
5 a 10 a 18 a 30 a
Coverage (%) 58.87±3.82a 40.44±2.47c 51.59±2.61b 58.94±1.70a 30.04 <0.001
Above-ground biomass (g/m2) 145.99±5.97c 116.12±8.04d 228.27±16.27b 275.06±8.53a 147.06 <0.001
Below-ground biomass (g/m2) 52.46±4.53c 44.58±4.08c 91.87±7.15b 117.54±8.21a 90.34 <0.001
Margalef richness index 1.62±0.08c 2.67±0.27b 2.92±0.27b 3.51±0.09a 48.17 <0.001
Shannon-Wiener diversity index 1.13±0.07d 2.50±0.13c 2.89±0.29b 3.65±0.12a 114.38 <0.001
Pielou evenness index 0.47±0.03d 0.65±0.02c 0.80±0.03b 0.95±0.02a 227.99 <0.001
Table 1 Plant community characteristics over different successional time
Plant species Years of farmland abandonment
5 a 10 a 18 a 30 a
Patrinia heterophylla Bunge - 4.56±0.49a 1.89±0.26b 0.58±0.09c
Dracocephalum moldavica L. 4.03±0.48a - - -
Lespedeza dahurica Schindler 9.86±0.11d 16.36±0.66c 21.20±0.53a 17.79±0.40b
Medicago sativa L. - - 3.56±0.44b 6.53±0.57a
Vicia sepium L. - - 2.33±0.51a -
Thermopsis lanceolata R. Br. - - - 3.07±0.64a
Glycyrrhiza uralensis Fisch. - - 1.22±0.39a -
Astragalus melilotoides Pall. - 3.89±0.28b 2.98±0.62b 6.11±0.46a
Gueldenstaedtia verna Boriss. - - - 2.99±0.34a
Setaria viridis (L.) Beauv. 10.29±0.36a 5.36±0.46b 0.68±0.06c -
Poa pratensis L. 6.41±0.27a 2.09±0.15c 3.30±0.33b -
Stipa bungeana Trin. - 19.41±0.77a 13.45±0.99b 8.60±0.79c
Cleistogenes chinensis (Maxim.) Keng - - 2.50±0.58a 0.55±0.36b
Roegneria kamoji Ohwi - - 4.54±0.50a 0.30±0.11b
Phragmites australis (Cav.) Trin.ex Steud - - - 0.75±0.18a
Leymus secalinus (Georgi) Tzvel. - - - 1.41±0.13a
Bothriochloa ischaemum (Linnaeus) Keng - - 4.96±0.65b 12.74±0.64a
Viola philippica Cav. - 1.21±0.27a - -
Artemisia capillaris Thunb. 35.02±0.44a 18.31±0.78b 3.68±0.18c 2.07±0.76d
Heteropappus altaicus (Willd.) Novopokr 17.37±0.80a 8.29±0.74b 2.17±0.44c -
Cirsium setosum (Willd.) MB. 1.70±0.14a - - -
Artemisia sacrorum Ledeb. - 7.83±0.72c 16.49±0.90a 10.11±0.62b
Ixeris polycephala Cass. - 2.30±0.38a - 1.84±0.32a
Saussurea japonica (Thunb.) DC. - - 2.59±0.25a 1.23±0.25b
Artemisia giraldii Pamp. - 2.24±0.21c 5.12±0.55b 18.88±0.93a
Ixeridium sonchifolium (Maxim.) Shih - 1.59±0.13a - 0.50±0.12b
Bidens parviflora Willd. - - 1.05±0.09a -
Salsola collina Pall. 4.87±0.06a 1.24±0.26b - 0.85±0.11c
Chenopodium glaucum L. - 2.79±0.78a - -
Potentilla discolor Bge. 6.99±0.61a 2.53±0.47b - -
Duchesnea indica (Andr.) Focke - - 3.37±0.43a -
Polygala tenuifolia Willd. 1.17±0.33b - 0.57±0.06c 1.86±0.10a
Incarvillea sinensis Lam. 2.29±0.20a - 2.34±0.38a 1.26±0.06b
Compositae 54.09±0.66a 40.57±0.21b 31.10±0.38d 34.63±0.22c
Gramineae 16.70±0.44d 26.86±0.17b 29.43±1.22a 24.35±0.26c
Leguminosae 9.86±0.11d 20.25±0.93c 31.30±1.62b 36.49±0.08a
Sum of Compositae, Gramineae, and Leguminosae 80.65±0.70d 87.67±1.27c 91.82±0.63b 95.48±0.59a
Table 2 Importance value of plant species over different successional time
Fig. 2 Changes of soil nutrient contents over different successional time. (a), soil organic carbon (SOC); (b), total nitrogen (TN); (c), total phosphorus (TP); (d), available phosphorus (AP); (e), nitrate nitrogen (NO3--N); (f), ammonium nitrogen (NH4+-N). Different lowercase letters indicate significant difference among different successional time at P<0.05 level. Bars are standard errors.
Parameter Farmland Years of farmland abandonment F P
5 a 10 a 18 a 30 a
Soil water content (%) 7.24±0.18e 6.71±0.16d 7.82±0.13c 9.28±0.26b 9.86±0.10a 177.30 <0.001
pH 8.49±0.02c 8.54±0.03bc 8.62±0.02a 8.49±0.03c 8.56±0.03b 10.72 0.001
Soil bulk density (g/cm3) 1.29±0.02a 1.26±0.02a 1.21±0.03b 1.18±0.02bc 1.15±0.02c 23.30 <0.001
Soil temperature (°C) 18.71±0.24a 18.14±0.13b 17.52±0.30c 16.44±0.26d 17.56±0.32c 32.16 <0.001
Clay (%) 18.86±0.81c 19.22±0.28c 19.52±0.26bc 20.19±0.33ab 20.81±0.29a 9.21 0.002
SOC (g/kg) 3.53±0.31d 3.94±0.22cd 4.31±0.23c 4.98±0.28b 5.70±0.20a 35.28 <0.001
TN (g/kg) 0.41±0.02cd 0.36±0.05d 0.45±0.02c 0.51±0.02b 0.59±0.03a 32.47 <0.001
TP (g/kg) 0.60±0.01a 0.58±0.01ab 0.57±0.01bc 0.53±0.01d 0.55±0.01c 14.43 <0.001
NH4+-N (mg/kg) 9.45±0.44b 7.56±0.38d 8.66±0.42c 9.82±0.30b 10.68±0.56a 23.13 <0.001
NO3--N (mg/kg) 3.12±0.23e 2.47±0.12d 3.59±0.29c 4.47±0.21b 5.15±0.26a 63.54 <0.001
AP (mg/kg) 1.95±0.17a 1.64±0.07b 1.20±0.16c 0.78±0.07d 0.61±0.07d 69.98 <0.001
Table 3 Soil physical-chemical properties over different successional time
Fig. 3 Redundancy analysis (RDA) for the relationships among soil physical-chemical properties, vegetation characteristics, and plant community composition. (a), relationships between soil physical-chemical properties (red arrows) and vegetation characteristics (grey arrows); (b), relationships between soil physical-chemical properties (red arrows) and plant community composition (grey arrows). SOC, soil organic carbon; TN, total nitrogen; TP, total phosphorus; AP, available phosphorus; SWC, soil water content; BD, bulk density; T, soil temperature; M, Margalef richness index; H, Shannon-Wiener diversity index; E, Pielou evenness index; AB, above-ground biomass; BB, below-ground biomass; sum of C, G, and L, sum of Compositae, Gramineae, and Leguminosae.
Fig. 4 Changes of soil biological properties over different successional time. Different lowercase letters indicate significant difference among different successional time at P<0.05 level. Bars are standard errors. C, carbon; N, nitrogen. (a), saccharase; (b), urease; (c), alkiline phosphatase; (d), catalase; (e), microbial biomass C (MBC); (f), microbial biomass N; (g), soil microbial respiration; (h), metabolic quotient.
Fig. 5 Redundancy analysis (RDA) for relationship among soil biological properties, vegetation characteristics, and plant community composition. (a), relationships between soil biological properties (red arrows) and vegetation characteristics (grey arrows); (b), relationships between soil biological properties (red arrows) and plant community composition (grey arrows); MBC, soil microbial biomass carbon; MBN, soil microbial biomass nitrogen; ALP, alkaline phosphatase; URE, urease; CAT, catalase; SAC, saccharase; MR, microbial respiration; qCO2, metabolic quotient; M, Margalef richness index; H, Shannon-Wiener diversity index; E, Pielou evenness index; AB, above-ground biomass; BB, below-ground biomass; sum of C, G, and L, sum of importance values of Compositae, Gramineae, and Leguminosae plants.
Parameter Contribution (%) F P
Leguminosae 75.90 27.4 0.002
Compositae 66.10 17.6 0.002
Gramineae 37.90 5.8 0.016
Above-ground biomass 59.40 13.4 0.004
Below-ground biomass 59.10 13.3 0.002
Margalef richness index 53.30 10.6 0.002
Shannon-Wiener diversity index 58.30 12.9 0.002
Pielou evenness index 56.20 11.8 0.002
Table 4 Effects of vegetation characteristics on soil physical-chemical properties over successional time
Parameter Contribution (%) F P
Leguminosae 81.60 31.6 0.002
Compositae 59.00 13.6 0.006
Gramineae 27.30 3.6 0.052
Above-ground biomass 73.70 25.8 0.004
Below-ground biomass 77.70 31.8 0.002
Margalef richness index 45.10 7.9 0.020
Shannon-Wiener diversity index 47.60 8.4 0.004
Pielou evenness index 46.60 8.4 0.010
Table 5 Effects of vegetation characteristics on soil biological properties over successional time
Fig. 6 Variance partitioning analysis of dominant species of Leguminosae on soil physical-chemical properties (a) and soil biological properties (b)
Plot Slope degree (°) Aspect (°) Elevation (m) Location Dominant species
Farmland 26 North by east 45 1187.3 36°46′N, 109°15′E -
5 a 28 North by west 40 1152.4 36°45′N, 109°15′E A. capillaris, H. altaicus
10 a 30 North by east 55 1239.1 36°44′N, 109°16′E S. bungeana, A. capillaris
18 a 27 North by west 38 1195.6 36°45′N, 109°16′E L. dahurica, A. sacrorum
30 a 32 North by east 42 1251.2 36°44′N, 109°16′E A. giraldii, L. dahurica, B. ischaemum
Table S1 Geographical features and dominant species at different plots
Type of enzyme Detailed measurement method
Soil catalase activity Soil catalase activity was determined by addition of 40 mL distilled water and 5 mL 0.3% H2O2 to 2 g fresh soil. The mixture was shaken for 20 min (at 150 r/m) and filtered (Whatman 2V) immediately. Then the filtrate was titrated with 0.1 mol/L KMnO4 under the conditions of sulfuric acid. Finally, the results were expressed as 0.1 mol KMnO4/(g·20 min).
Soil saccharase activity Soil saccharase activity was determined using 8% glucose solution as substrates. About 5 g fresh soil was incubated with 15 mL substrates, 5 mL 0.2 M phosphate buffer (pH 5.5), and 5 drops of toluene for 24 h at 37.8°C. After incubation, the mixture was filtered (Whatman 2V) immediately and 1-mL aliquot was reacted with 3 mL 3, 5-dinitrylsalicylate in a volumetric flask, and then heated for 5 min. Soil solution in the flask was quantified in an ultraviolet spectrometer subsystem (UVS) at 508 nm when it reached room temperature. Finally, results were also expressed as mg glucose/(g·24 h).
Soil urease activity Soil urease activities was routinely determined using 10% urea solution as substrates. About 5 g fresh soil was incubated for 24 h at 37.8°C with 5 mL citrate solution at pH 6.7 and 5 mL substrates. The reaction mixture was then diluted to 50 mL with distilled water. After incubation, the mixture was immediately filtered and 1 mL supernatant was treated with 4 mL sodium phenol solution and 3 mL 0.9% sodium hypochlorite solution. The released ammonium released from urea hydrolysis was quantified in an ultraviolet spectrometer subsystem (UVS) at 578 nm. Results were expressed as mg NH4+-N/(g·24 h).
Soil alkaline phosphatase activity Soil alkaline phosphatase activity was determined by addition of 10 g fresh soil, 2 mL toluene, 10 mL disodium phenyl phosphate solution, and 10 mL 0.05 M borate buffer. The reaction mixture was incubated for 2 h at 37.8°C. After incubation, the mixture was immediately filtered, then the filtrate was treated with 0.5 mL of 2% 4-aminoantipyrine and 8% potassium ferrocyanide; the phenol released was determined in an ultraviolet spectrometer subsystem (UVS) at 510 nm. Results were expressed as mg phenol/(g·24 h).
Table S2 Methods for determination of soil enzymatic activities
Fig. S1 Nonmetric multidimensional scaling (NMDS) analysis of plant community composition over successional time
Fig. S2 Nonmetric multidimensional scaling (NMDS) analysis of soil physical-chemical properties over successional time
Fig. S3 Nonmetric multidimensional scaling (NMDS) analysis of soil biological properties over successional time
[1]   An S, Zheng F, Zhang F, et al. 2008. Soil quality degradation processes along a deforestation chronosequence in the Ziwuling area, China. CATENA, 75(3): 248-256.
doi: 10.1016/j.catena.2008.07.003
[2]   Baličević R, Ravlić M, Kleflin J, et al. 2016. Allelopathic activity of plant species from Asteraceae and Polygonaceae family on lettuce. Herbologia, 16(1): 23-30.
[3]   Bao S. 2000. Soil and Agricultural Chemistry Analysis. Beijing: China Agriculture Press, 11-12. (in Chinese)
[4]   Bao Y, Gao M, Luo D, et al. 2022. The influence of plant community characteristics in urban parks on the microclimate. Forests, 13(9): 1342, doi: 10.3390/f13091342.
doi: 10.3390/f13091342
[5]   Berg B. 2014. Decomposition patterns for foliar litter-A theory for influencing factors. Soil Biology and Biochemistry, 78: 222-232.
doi: 10.1016/j.soilbio.2014.08.005
[6]   Chabrerie O, Laval K, Puget P, et al. 2003. Relationship between plant and soil microbial communities along a successional gradient in a chalk grassland in north-western France. Applied Soil Ecology, 24(1): 43-56.
doi: 10.1016/S0929-1393(03)00062-3
[7]   Chen C R, Condron L M, Davis M R, et al. 2000. Effects of afforestation on phosphorus dynamics and biological properties in a New Zealand grassland soil. Plant and Soil, 220(1-2): 151-163.
doi: 10.1023/A:1004712401721
[8]   Chen Y C, Li W P, You Y, et al. 2022. Soil properties and substrate quality determine the priming of soil organic carbon during vegetation succession. Plant and Soil, 471(1-2): 559-575.
doi: 10.1007/s11104-021-05241-z
[9]   Chen Y X, Wei T X, Sha G L, et al. 2022. Soil enzyme activities of typical plant communities after vegetation restoration on the Loess Plateau, China. Applied Soil Ecology, 170: 104292, doi: 10.1016/j.apsoil.2021.104292.
doi: 10.1016/j.apsoil.2021.104292
[10]   Chen Z F, Xiong P F, Zhou J J, et al. 2021. Effects of plant diversity on semiarid grassland stability depends on functional group composition and dynamics under N and P addition. Science of the Total Environment, 799: 149482, doi: 10.1016/j.scitotenv.2021.149482.
doi: 10.1016/j.scitotenv.2021.149482
[11]   Chou C B, Hedin L O, Pacala S W. 2018. Functional groups, species and light interact with nutrient limitation during tropical rainforest sapling bottleneck. Journal of Ecology, 106(1): 157-167.
doi: 10.1111/jec.2018.106.issue-1
[12]   Cotrufo M F, Haddix M L, Kroeger M E, et al. 2022. The role of plant input physical-chemical properties, and microbial and soil chemical diversity on the formation of particulate and mineral-associated organic matter. Soil Biology and Biochemistry, 168: 108648, doi: 10.1016/j.soilbio.2022.108648.
doi: 10.1016/j.soilbio.2022.108648
[13]   Criquet S, Farnet A M, Tagger S, et al. 2000. Annual variations of phenoloxidase activities in an evergreen oak litter: Influence of certain biotic and abiotic factors. Soil Biology and Biochemistry, 32(11-12): 1505-1513.
doi: 10.1016/S0038-0717(00)00027-4
[14]   DeBerry J W, Atkinson R B. 2014. Aboveground forest biomass and litter production patterns in Atlantic white cedar swamps of differing hydroperiods. Southeastern Naturalist, 13(4): 673-690.
doi: 10.1656/058.013.0414
[15]   Fan H B, Wu J P, Liu W F, et al. 2015. Linkages of plant and soil C:N:P stoichiometry and their relationships to forest growth in subtropical plantations. Plant and Soil, 392(1-2): 127-138.
doi: 10.1007/s11104-015-2444-2
[16]   Fanin N, Bertrand I. 2016. Aboveground litter quality is a better predictor than belowground microbial communities when estimating carbon mineralization along a land-use gradient. Soil Biology and Biochemistry, 94: 48-60.
doi: 10.1016/j.soilbio.2015.11.007
[17]   Fterich A, Mahdhi M, Mars M. 2014. The effects of Acacia tortilis subsp. raddiana, soil texture and soil depth on soil microbial and biochemical characteristics in arid zones of Tunisia. Land Degradation and Development, 25(2): 143-152.
doi: 10.1002/ldr.v25.2
[18]   Fu B J, Chen L D, Ma K M, et al. 2000. The relationships between land use and soil conditions in the hilly area of the loess plateau in northern Shaanxi, China. CATENA, 39(1): 69-78.
doi: 10.1016/S0341-8162(99)00084-3
[19]   Gu Q, Yu Q, Grogan P. 2023. Cryptogam plant community stability: Warming weakens influences of species richness but enhances effects of evenness. Ecology, 104(1): 3842, doi: 10.1002/ecy.3842.
doi: 10.1002/ecy.3842
[20]   Guan H L, Fan J W, Lu X K. 2022. Soil specific enzyme stoichiometry reflects nitrogen limitation of microorganisms under different types of vegetation restoration in the Karst areas. Applied Soil Ecology, 169: 104253, doi: 10.1016/j.apsoil.2021.104253.
doi: 10.1016/j.apsoil.2021.104253
[21]   Háněl L. 2003. Recovery of soil nematode populations from cropping stress by natural secondary succession to meadow land. Applied Soil Ecology, 22(3): 255-270.
doi: 10.1016/S0929-1393(02)00152-X
[22]   Hao J W, Chu L M. 2021. Vegetation types attributed to deforestation and secondary succession drive the elevational changes in diversity and distribution of terrestrial mosses in a tropical mountain forest in Southern China. Forests, 12(8): 961, doi: 10.3390/f12080961.
doi: 10.3390/f12080961
[23]   HilleRisLambers J, Harpole W S, Tilman D, et al. 2004. Mechanisms responsible for the positive diversity-productivity relationship in Minnesota grasslands. Ecology Letters, 7(8): 661-668.
doi: 10.1111/ele.2004.7.issue-8
[24]   Horodecki P, Jagodzinski A M. 2017. Tree species effects on litter decomposition in pure stands on afforested post-mining sites. Forest Ecology and Management, 406: 1-11.
doi: 10.1016/j.foreco.2017.09.059
[25]   Hu G Z, Liu H Y, Yin Y, et al. 2016. The role of legumes in plant community succession of degraded grasslands in northern China. Land Degradation and Development, 27(2): 366-372.
doi: 10.1002/ldr.v27.2
[26]   Hu S, van Bruggen A H C. 1997. Microbial dynamics associated with multiphasic decomposition of 14C-labeled cellulose in soil. Microbial Ecology, 33(2): 134-143.
pmid: 9052647
[27]   Huang W, Liu J, Wang Y P, et al. 2013. Increasing phosphorus limitation along three successional forests in southern China. Plant and Soil, 364(1-2): 181-191.
doi: 10.1007/s11104-012-1355-8
[28]   Ile O J, Aguilos M, Morkoc S, et al. 2021. Root biomass distribution and soil physical properties of short-rotation coppice American sycamore (Platanus occidentalis L.) grown at different planting densities. Forests, 12(12): 1806, doi: 10.3390/f12121806.
doi: 10.3390/f12121806
[29]   Jiao F, Wen Z M, An S S. 2011. Changes in soil properties across a chronosequence of vegetation restoration on the Loess Plateau of China. CATENA, 86(2): 110-116.
doi: 10.1016/j.catena.2011.03.001
[30]   Jing Z, Cheng J, Jin J, et al. 2014. Revegetation as an efficient means of improving the diversity and abundance of soil eukaryotes in the Loess Plateau of China. Ecological Engineering, 70: 169-174.
doi: 10.1016/j.ecoleng.2014.05.011
[31]   Kardol P, Wardle D A. 2010. How understanding aboveground-belowground linkages can assist restoration ecology. Trends in Ecology & Evolution, 25(11): 670-679.
doi: 10.1016/j.tree.2010.09.001
[32]   Kong Y S, Zhang H K, Tian L L, et al. 2023. Relationships between denitrification rates and functional gene abundance in a wetland: The roles of single- and multiple-species plant communities. Science of the Total Environment, 863: 160913, doi: 10.1016/j.scitotenv.2022.160913.
doi: 10.1016/j.scitotenv.2022.160913
[33]   Li J, Liu Y, Hai X, et al. 2019. Dynamics of soil microbial C:N:P stoichiometry and its driving mechanisms following natural vegetation restoration after farmland abandonment. Science of the Total Environment, 693: 133613, doi: 10.1016/j.scitotenv.2019.133613.
doi: 10.1016/j.scitotenv.2019.133613
[34]   Li J H, Jiao S M, Gao R Q, et al. 2012. Differential effects of legume species on the recovery of soil microbial communities, and carbon and nitrogen contents, in abandoned fields of the Loess Plateau, China. Environmental Management, 50(6): 1193-1203.
doi: 10.1007/s00267-012-9958-7 pmid: 23064665
[35]   Li J W, Liu Y L, Hai X Y, et al. 2019. Dynamics of soil microbial C:N:P stoichiometry and its driving mechanisms following natural vegetation restoration after farmland abandonment. Science of the Total Environment, 693: 133613, doi: 10.1016/j.scitotenv.2019.133613.
doi: 10.1016/j.scitotenv.2019.133613
[36]   Li Q, Song Y T, Li G D, et al. 2015. Grass-legume mixtures impact soil N, species recruitment, and productivity in temperate steppe grassland. Plant and Soil, 394(1-2): 271-285.
doi: 10.1007/s11104-015-2525-2
[37]   Li W, Zhang R, Liu S, et al. 2018. Effect of loss of plant functional group and simulated nitrogen deposition on subalpine ecosystem properties on the Tibetan Plateau. Science of the Total Environment, 631-632: 289-297.
doi: 10.1016/j.scitotenv.2018.02.287
[38]   Li X Y, Jin H J, He R X, et al. 2023. Impact of wildfire on soil carbon and nitrogen storage and vegetation succession in the Nanweng? The National Natural Wetlands Reserve, Northeast China. CATENA, 221: 106797, doi: 10.1016/j.catena.2022.106797.
doi: 10.1016/j.catena.2022.106797
[39]   Liao J J, Dou Y X, An S S. 2023. Plant community productivity is associated with multiple ecological stoichiometry in restoration grasslands. Ecological Engineering, 187: 106845, doi: 10.1016/j.ecoleng.2022.106845.
doi: 10.1016/j.ecoleng.2022.106845
[40]   Liu H F, Liang C T, Ai Z M, et al. 2019. Plant-mycorrhizae association affects plant diversity, biomass, and soil nutrients along temporal gradients of natural restoration after farmland abandonment in the Loess Plateau, China. Land Degradation & Development, 30(14): 1677-1690.
doi: 10.1002/ldr.v30.14
[41]   Liu M Y, Li P, Liu M M, et al. 2021. The trend of soil organic carbon fractions related to the successions of different vegetation types on the tableland of the Loess Plateau of China. Journal of Soils and Sediments, 21(1): 203-214.
doi: 10.1007/s11368-020-02710-3
[42]   Liu W X, Xu W H, Hong J P, et al. 2010. Interannual variability of soil microbial biomass and respiration in responses to topography, annual burning and N addition in a semiarid temperate steppe. Geoderma, 158(3-4): 259-267.
doi: 10.1016/j.geoderma.2010.05.004
[43]   Liu Y F, Meng L C, Huang Z, et al. 2022. Contribution of fine roots mechanical property of Poaceae grasses to soil erosion resistance on the Loess Plateau. Geoderma, 426: 116122, doi: 10.1016/j.geoderma.2022.116122.
doi: 10.1016/j.geoderma.2022.116122
[44]   Liu Y L, Song X Z, Wang K B, et al. 2022. Changes in soil microbial metabolic activity following long-term forest succession on the central Loess Plateau, China. Land Degradation & Development, 34(3): 723-735.
doi: 10.1002/ldr.v34.3
[45]   Liu Z Y, Michalet R, Wang C Y, et al. 2023. Contrasting effects of two phenotypes of an alpine cushion plant on understory species drive community assembly. Science of the Total Environment, 859: 160154, doi: 10.1016/j.scitotenv.2022.160154.
doi: 10.1016/j.scitotenv.2022.160154
[46]   Lozano Y M, Hortal S, Armas C, et al. 2014. Interactions among soil, plants, and microorganisms drive secondary succession in a dry environment. Soil Biology and Biochemistry, 78: 298-306.
doi: 10.1016/j.soilbio.2014.08.007
[47]   Lucas-Borja M E, Hedo J, Cerda A, et al. 2016. Unravelling the importance of forest age stand and forest structure driving microbiological soil properties, enzymatic activities and soil nutrients content in Mediterranean Spanish black pine (Pinus nigra Ar. ssp salzmannii) Forest. Science of the Total Environment, 562: 145-154.
doi: 10.1016/j.scitotenv.2016.03.160
[48]   Lucas-Borja M E, Delgado-Baquerizo M. 2019. Plant diversity and soil stoichiometry regulates the changes in multifunctionality during pine temperate forest secondary succession. Science of the Total Environment, 697: 134204, doi: 10.1016/j.scitotenv.2019.134204.
doi: 10.1016/j.scitotenv.2019.134204
[49]   Luo Y, Xiao H B, Wu X, et al. 2022. Mitigation of nutrient runoff loss using reduced nitrogen application and green manure planting in citrus orchard in Hubei, China. Journal of Soils and Sediments, 23(2): 582-595.
doi: 10.1007/s11368-022-03356-z
[50]   Marriott C A, Hudson G, Hamilton D, et al. 1997. Spatial variability of soil total C and N and their stable isotopes in an upland Scottish grassland. Plant and Soil, 196(1): 151-162.
doi: 10.1023/A:1004288610550
[51]   O'dea J K, Jones C A, Zabinski C A, et al. 2015. Legume, cropping intensity, and N-fertilization effects on soil attributes and processes from an eight-year-old semiarid wheat system. Nutrient Cycling in Agroecosystems, 102(2): 179-194.
doi: 10.1007/s10705-015-9687-4
[52]   O'Halloran L R, Borer E T, Seabloom E W, et al. 2013. Regional contingencies in the relationship between aboveground biomass and litter in the world's grasslands. PloS ONE, 8(2): e054988, doi: 10.1371/journal.pone.0054988.
doi: 10.1371/journal.pone.0054988
[53]   Ohtsuka T, Shizu Y, Nishiwaki A, et al. 2010. Carbon cycling and net ecosystem production at an early stage of secondary succession in an abandoned coppice forest. Journal of Plant Research, 123(4): 393-401.
doi: 10.1007/s10265-009-0274-0 pmid: 20033468
[54]   Olsen S R, Sommers L E, Page A L. 1982. Methods of Soil Analysis. Part 2. Chemical and Microbiological Properties of Phosphorus. Madison: American Society of Agronomy, 403-430.
[55]   Peichl M, Leava N A, Kiely G. 2012. Above- and belowground ecosystem biomass, carbon and nitrogen allocation in recently afforested grassland and adjacent intensively managed grassland. Plant and Soil, 350(1-2): 281-296.
doi: 10.1007/s11104-011-0905-9
[56]   Plaza C, Hernandez D, Garcia-Gil J C, et al. 2004. Microbial activity in pig slurry-amended soils under semi-arid conditions. Soil Biology and Biochemistry, 36(10): 1577-1585.
doi: 10.1016/j.soilbio.2004.07.017
[57]   Qiu D X, Xu R R, Wu C X, et al. 2022. Vegetation restoration improves soil hydrological properties by regulating soil physicochemical properties in the Loess Plateau, China. Journal of Hydrology, 609: 127730, doi: 10.1016/j.jhydrol.2022.127730.
doi: 10.1016/j.jhydrol.2022.127730
[58]   Reich P B. 2005. Global biogeography of plant chemistry: Filling in the blanks. New Phytologist, 168(2): 263-266.
pmid: 16219064
[59]   Ren C J, Kang D, Wu J P, et al. 2016a. Temporal variation in soil enzyme activities after afforestation in the Loess Plateau, China. Geoderma, 282: 103-111.
doi: 10.1016/j.geoderma.2016.07.018
[60]   Ren C J, Zhao F, Kang D, et al. 2016b. Linkages of C:N:P stoichiometry and bacterial community in soil following afforestation of former farmland. Forest Ecology and Management, 376: 59-66.
doi: 10.1016/j.foreco.2016.06.004
[61]   Ren C J, Chen J, Deng J, et al. 2017. Response of microbial diversity to C:N:P stoichiometry in fine root and microbial biomass following afforestation. Biology and Fertility of Soils, 53(4): 457-468.
doi: 10.1007/s00374-017-1197-x
[62]   Ren C J, Wang T, Xu Y D, et al. 2018. Differential soil microbial community responses to the linkage of soil organic carbon fractions with respiration across land-use changes. Forest Ecology and Management, 409: 170-178.
doi: 10.1016/j.foreco.2017.11.011
[63]   Sidlauskaite G, Kemesyte V, Toleikiene M, et al. 2022. Plant diversity, functional group composition and legumes effects versus fertilisation on the yield and forage quality. Sustainability, 14(3): 1182, doi: 10.3390/su14031182.
doi: 10.3390/su14031182
[64]   Sivaram A K, Logeshwaran P, Subashchandrabose S R, et al. 2018. Comparison of plants with C3 and C4 carbon fixation pathways for remediation of polycyclic aromatic hydrocarbon contaminated soils. Scientific Reports, 8: 2100, doi: 10.1038/s41598-018-20317-0.
doi: 10.1038/s41598-018-20317-0
[65]   Song G, Hui R, Yang H T, et al. 2022. Biocrusts mediate the plant community composition of dryland restoration ecosystems. Science of the Total Environment, 844: 157315, doi: 10.1016/j.scitotenv.2022.157135.
doi: 10.1016/j.scitotenv.2022.157135
[66]   Sun C, Chai Z, Liu G, et al. 2017. Changes in species diversity patterns and spatial heterogeneity during the secondary succession of grassland vegetation on the Loess Plateau, China. Frontiers in Plant Science, 8: 1465, doi: 10.3389/fpls.2017.01465.
doi: 10.3389/fpls.2017.01465 pmid: 28900433
[67]   Sun Y, Chen L, Zhang S Y, et al. 2022. Plant interaction patterns shape the soil microbial community and nutrient cycling in different intercropping scenarios of aromatic plant species. Frontiers in Microbiology, 13: 888789, doi: 10.3389/fmicb.2022.888789.
doi: 10.3389/fmicb.2022.888789
[68]   Ter Braak C J, Smilauer P. 2002. CANOCO reference manual and CanoDraw for Windows user's guide: software for canonical community ordination (version 4.5). [2022-10-12].
[69]   Tessema Z K, Belay E F. 2017. Effect of tree species on understory vegetation, herbaceous biomass and soil nutrients in a semi-arid savanna of Ethiopia. Journal of Arid Environments, 139: 76-84.
doi: 10.1016/j.jaridenv.2016.12.007
[70]   Tian F P, Zhang Z N, Chang X F, et al. 2016. Effects of biotic and abiotic factors on soil organic carbon in semi-arid grassland. Journal of Soil Science and Plant Nutrition, 16(4): 1087-1096.
[71]   van der Heijden M G A, Bardgett R D, van Straalen N M. 2008. The unseen majority: soil microbes as drivers of plant diversity and productivity in terrestrial ecosystems. Ecology Letters, 11(3): 296-310.
doi: 10.1111/j.1461-0248.2007.01139.x pmid: 18047587
[72]   van der Putten W H, Bardgett R D, Bever J D, et al. 2013. Plant-soil feedbacks: The past, the present and future challenges. Journal of Ecology, 101(2): 265-276.
doi: 10.1111/jec.2013.101.issue-2
[73]   Walker L R, Wardle D A, Bardgett R D, et al. 2010. The use of chronosequences in studies of ecological succession and soil development. Journal of Ecology, 98(4): 725-736.
doi: 10.1111/j.1365-2745.2010.01664.x
[74]   Wang G L, Liu G B, Xu M X. 2009. Above- and belowground dynamics of plant community succession following abandonment of farmland on the Loess Plateau, China. Plant and Soil, 316(1-2): 227-239.
doi: 10.1007/s11104-008-9773-3
[75]   Wang L L, Zhang G H, Zhu P Z, et al. 2022. Soil C, N and P contents and their stoichiometry as affected by typical plant communities on steep gully slopes of the Loess Plateau, China. CATENA, 208: 105740, doi: 10.1016/j.catena.2021.105740.
doi: 10.1016/j.catena.2021.105740
[76]   Wardle D A, Ghani A. 1995. A critique of the microbial metabolic quotient (qCO2) as a bio-indicator of disturbance and ecosystem development. Soil Biology & Biochemistry, 27(12): 1601-1610.
doi: 10.1016/0038-0717(95)00093-T
[77]   Wickander N J, Rasmussen P U, Marteinsdottir B, et al. 2021. Ecological and evolutionary responses of an arctic plant to variation in microclimate and soil. Oikos, 130(2): 211-218.
doi: 10.1111/oik.2021.v130.i2
[78]   Williams M A, Jangid K, Shanmugam S G, et al. 2013. Bacterial communities in soil mimic patterns of vegetative succession and ecosystem climax but are resilient to change between seasons. Soil Biology and Biochemistry, 57: 749-757.
doi: 10.1016/j.soilbio.2012.08.023
[79]   Wu G L, Liu Y, Tian F P, et al. 2017. Legumes functional group promotes soil organic carbon and nitrogen storage by increasing plant diversity. Land Degradation & Development, 28(4): 1336-1344.
doi: 10.1002/ldr.v28.4
[80]   Xiao C W, Zhou Y, Su J Q, et al. 2017. Effects of plant functional group loss on soil microbial community and litter decomposition in a steppe vegetation. Frontiers in Plant Science, 8: 2040, doi: 10.3389/fpls.2017.02040.
doi: 10.3389/fpls.2017.02040 pmid: 29234343
[81]   Xiao J, Wang B L, Qiu X L, et al. 2021. Interaction between carbon cycling and phytoplankton community succession in hydropower reservoirs: Evidence from stable carbon isotope analysis. Science of the Total Environment, 774: 145141, doi: 10.1016/j.scitotenv.2021.145141.
doi: 10.1016/j.scitotenv.2021.145141
[82]   Yahdjian L, Tognetti P M, Chaneton E J. 2017. Plant functional composition affects soil processes in novel successional grasslands. Functional Ecology, 31(9): 1813-1823.
doi: 10.1111/fec.2017.31.issue-9
[83]   Yang T, Song L Y, Lin H Y, et al. 2023. Within-species plant phylogeny drives ectomycorrhizal fungal community composition in tree roots along a timberline. Soil Biology and Biochemistry, 176: 108880, doi: 10.1016/j.soilbio.2022.108880.
doi: 10.1016/j.soilbio.2022.108880
[84]   Yang Y, Liang C, Wang Y Q, et al. 2020. Soil extracellular enzyme stoichiometry reflects the shift from P- to N-limitation of microorganisms with grassland restoration. Soil Biology and Biochemistry, 149: 107928, doi: 10.1016/j.soilbio.2020.107928.
doi: 10.1016/j.soilbio.2020.107928
[85]   Yao Z Y, Xu Q, Chen Y P, et al. 2021. Leguminous green manure enhances the soil organic nitrogen pool of cropland via disproportionate increase of nitrogen in particulate organic matter fractions. CATENA, 207: 105574, doi: 10.1016/j.catena.2021.105574.
doi: 10.1016/j.catena.2021.105574
[86]   Zeng Q, An S, Liu Y. 2017. Soil bacterial community response to vegetation succession after fencing in the grassland of China. Science of the Total Environment, 609: 2-10.
doi: 10.1016/j.scitotenv.2017.07.102
[87]   Zhang C, Liu G B, Xue S, et al. 2011. Rhizosphere soil microbial activity under different vegetation types on the Loess Plateau, China. Geoderma, 161(3-4): 115-125.
doi: 10.1016/j.geoderma.2010.12.003
[88]   Zhang C, Liu G B, Xue S, et al. 2016. Soil bacterial community dynamics reflect changes in plant community and soil properties during the secondary succession of abandoned farmland in the Loess Plateau. Soil Biology and Biochemistry, 97: 40-49.
doi: 10.1016/j.soilbio.2016.02.013
[89]   Zhang D B, Zhang C, Ren H L, et al. 2021. Trade-offs between winter wheat production and soil water consumption via leguminous green manures in the Loess Plateau of China. Field Crops Research, 272: 108278, doi: 10.1016/j.fcr.2021.108278.
doi: 10.1016/j.fcr.2021.108278
[90]   Zhang J T. 2005. Succession analysis of plant communities in abandoned croplands in the eastern Loess Plateau of China. Journal of Arid Environments, 63(2): 458-474.
doi: 10.1016/j.jaridenv.2005.03.027
[91]   Zhang W, Gao D, Chen Z, et al. 2018a. Substrate quality and soil environmental conditions predict litter decomposition and drive soil nutrient dynamics following afforestation on the Loess Plateau of China. Geoderma, 325: 152-161.
doi: 10.1016/j.geoderma.2018.03.027
[92]   Zhang W, Qiao W, Gao D, et al. 2018b. Relationship between soil nutrient properties and biological activities along a restoration chronosequence of Pinus tabulaeformis plantation forests in the Ziwuling Mountains, China. CATENA, 161(Suppl. C): 85-95.
doi: 10.1016/j.catena.2017.10.021
[93]   Zhang X Y, Zhao W Q, Liu Y J, et al. 2022. Dominant plant species and soil properties drive differential responses of fungal communities and functions in the soils and roots during secondary forest succession in the subalpine region. Rhizosphere, 21: 100483, doi: 10.1016/j.rhisph.2022.100483.
doi: 10.1016/j.rhisph.2022.100483
[94]   Zhang Y H, Xu X L, Li Z W, et al. 2021. Improvements in soil quality with vegetation succession in subtropical China Karst. Science of the Total Environment, 775: 145876, doi: 10.1016/j.scitotenv.2021.145876.
doi: 10.1016/j.scitotenv.2021.145876
[95]   Zhao F Z, Kang D, Han X, et al. 2015. Soil stoichiometry and carbon storage in long-term afforestation soil affected by understory vegetation diversity. Ecological Engineering, 74: 415-422.
doi: 10.1016/j.ecoleng.2014.11.010
[96]   Zhao F Z, Ren C J, Han X H, et al. 2018. Changes of soil microbial and enzyme activities are linked to soil C, N and P stoichiometry in afforested ecosystems. Forest Ecology and Management, 427: 289-295.
doi: 10.1016/j.foreco.2018.06.011
[97]   Zhao F Z, Bai L, Wang J Y, et al. 2019. Change in soil bacterial community during secondary succession depend on plant and soil characteristics. CATENA, 173: 246-252.
doi: 10.1016/j.catena.2018.10.024
[98]   Zheng F L. 2006. Effect of vegetation changes on soil erosion on the Loess Plateau. Pedosphere, 16(4): 420-427.
doi: 10.1016/S1002-0160(06)60071-4
[99]   Zheng Y, Hu Z K, Pan X, et al. 2021. Carbon and nitrogen transfer from litter to soil is higher in slow than rapid decomposing plant litter: A synthesis of stable isotope studies. Soil Biology and Biochemistry, 156: 108196, doi: 10.1016/j.soilbio.2021.108196.
doi: 10.1016/j.soilbio.2021.108196
[100]   Zhou Z H, Wang C K, Jiang L F, et al. 2017. Trends in soil microbial communities during secondary succession. Soil Biology and Biochemistry, 115: 92-99.
doi: 10.1016/j.soilbio.2017.08.014
[101]   Zhu H H, He X Y, Wang K L, et al. 2012. Interactions of vegetation succession, soil bio-chemical properties and microbial communities in a Karst ecosystem. European Journal of Soil Biology, 51: 1-7.
doi: 10.1016/j.ejsobi.2012.03.003
[102]   Zhu P Z, Zhang G H, Wang C S, et al. 2021a. Variation in land surface roughness under typical plant communities on steep gully slopes on the Loess Plateau of China. CATENA, 206: 105549, doi: 10.1016/j.catena.2021.105549.
doi: 10.1016/j.catena.2021.105549
[103]   Zhu P Z, Zhang G H, Wang H X, et al. 2021b. Effectiveness of typical plant communities in controlling runoff and soil erosion on steep gully slopes on the Loess Plateau of China. Journal of Hydrology, 602: 126714, doi: 10.1016/j.jhydrol.2021.126714.
doi: 10.1016/j.jhydrol.2021.126714
[104]   Zou H, Gao G Y, Yuan C, et al. 2022. Interactions between soil water and plant community during vegetation succession in the restored grasslands on the Loess Plateau of China. Land Degradation & Development, 34(5): 1582-1592.
doi: 10.1002/ldr.v34.5
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