黄土高原生物结皮形成过程中土壤胞外酶活性及其化学计量变化特征
收稿日期: 2021-08-24
修回日期: 2021-11-24
网络出版日期: 2022-03-30
基金资助
国家自然科学基金面上项目(41671280);国家自然科学基金面上项目(41877074);国家自然科学基金面上项目(42077072)
Variation in soil extracellular enzyme activities stoichiometry during biological soil crust formation in the Loess Plateau
Received date: 2021-08-24
Revised date: 2021-11-24
Online published: 2022-03-30
生物土壤结皮在增强土壤抵抗水蚀风蚀能力、改善土壤养分等方面发挥着重要作用,土壤胞外酶活性可作为土壤生化反应强度的微生物指标,对理解荒漠生态系统微生物参与的养分循环过程具有重要意义。选取黄土高原神木县六道沟小流域5个生物结皮发育阶段(裸沙地、全藻结皮、藻-藓混生结皮、藓-藻混生结皮、全藓结皮)的土壤作为研究对象,研究生物结皮发育过程中土壤胞外酶活性(β-1,4-葡萄糖苷酶、β-1,4-N-乙酰基氨基葡萄糖苷酶、亮氨酸氨基多肽酶和碱性磷酸酶)及其化学计量的变化特征。结果表明:(1) 土壤4种胞外酶活性均随生物结皮发育序列变化显著增加,全藓结皮土壤显著大于全藻结皮土壤(P<0.05)。(2) 生物土壤结皮层胞外酶活性显著高于下层土壤,且胞外酶活性随着土层深度的加深不断降低。(3) 相关性分析表明,土壤C、N、P、土壤C:P、土壤N:P均与土壤胞外酶活性呈极显著正相关关系(P<0.05)。(4) 有结皮覆盖土壤的养分含量及土壤胞外酶活性显著高于裸沙地,全藓结皮土壤的养分含量及土壤胞外酶活性显著高于全藻结皮土壤。(5) 标准化主轴估计表明,土壤胞外酶活性随着生物结皮的发育均有明显增强并呈现稳态特征,N-获取酶和P-获取酶相对于C-获取酶之间的斜率呈现等容关系,这一结果表明土壤微生物通过稳态调控胞外酶化学计量特征,在干旱半干旱生态系统养分循环过程中具有重要作用。
姚宏佳,王宝荣,安韶山,杨娥女,黄懿梅 . 黄土高原生物结皮形成过程中土壤胞外酶活性及其化学计量变化特征[J]. 干旱区研究, 2022 , 39(2) : 456 -468 . DOI: 10.13866/j.azr.2022.02.13
Biological soil crusts (BSCs) play an important role in enhancing soil resistance to water and wind erosion, and improving soil nutrients. Soil extracellular enzyme activity can be used as a microbial indicator of the intensity of soil biochemical reactions, which is important for understanding the nutrient cycling processes involving microorganisms in desert ecosystems. However, there is a lack of understanding of the variation in soil extracellular enzymes and their stoichiometry during the formation of BSCs on the Loess Plateau. Soils from five biological crust formation stages (CK, A, AM, MA, and M) in Liudaogou watershed, Shenmu County, on the Loess Plateau were selected for the study. Soil extracellular enzyme activities [β-1,4-glucosidase (BG), β-1,4-N-acetylaminoglucosidase (NAG), leucine aminopeptidase (LAP), and alkaline phosphatase (AP)] and their stoichiometric changes during the formation of biocrusts were characterized. The activities of soil BG, NAG, LAP, and AP increased significantly with the sequence of biological crust development, and activity was significantly greater in Mo soil than that in Al soil (P<0.05). Extracellular enzyme activities were significantly higher in the BSCs layer than those in the lower soil layer, and decreased continuously with the depth of the soil. Soil C, N, P, C:P, and N:P all showed highly significant positive correlations with soil extracellular enzyme activities (P<0.05). The nutrient content and soil extracellular enzyme activity of soil with crust cover were significantly higher than those of bare land, and those of Mo soil were significantly higher than those of Al soil. Standardized major axis estimates indicated that soil extracellular enzyme activities were significantly enhanced and showed homeostasis with formation of biological crusts, and the slopes between N- and P-acquiring enzymes relative to C-acquiring enzymes showed isotropic relationships. Soil microorganisms play an important role in nutrient cycling in arid and semi-arid ecosystems through homeostatic regulation of extracellular enzyme stoichiometry.
[1] | Li X R, Jia R L, Chen Y W, et al. Association of ant nests with successional stages of biological soil crusts in the Tengger Desert, Northern China[J]. Applied Soil Ecology, 2011, 47(1):59-66. |
[2] | Rodriguez-Caballero E, Belnap J, Büdel B, et al. Dryland photoautotrophic soil surface communities endangered by global change[J]. Nature Geoscience, 2018, 11(3):185-189. |
[3] | Liu Y M, Yang H Y, Li X R, et al. Effects of biological soil crusts on soil enzyme activities in revegetated areas of the Tengger Desert, China[J]. Applied Soil Ecology, 2014, 80:6-14. |
[4] | Spohn M, Schleuss P. Addition of inorganic phosphorus to soil leads to desorption of organic compounds and thus to increased soil respiration[J]. Soil Biology and Biochemistry, 2019, 130:220-226. |
[5] | Liang C, Schimel J P, Jastrow J D. The importance of anabolism in microbial control over soil carbon storage[J]. Nature Microbiology, 2017, 2, 17105. |
[6] | Zi H B, Hu L, Wang C T, et al. Responses of soil bacterial community and enzyme activity to experimental warming of an alpine meadow[J]. European Journal of Soil Science, 2018, 69(3):429-438. |
[7] | Yang Y, Liang C, Wang Y Q, et al. Soil extracellular enzyme stoichiometry reflects the shift from P-to N-limitation of microorganisms with grassland restoration[J]. Soil Biology and Biochemistry, 2020, 149:107928. |
[8] | 勒佳佳, 苏原, 彭庆文, 等. 氮添加对天山高寒草原土壤酶活性和酶化学计量特征的影响[J]. 干旱区研究, 2020, 37(2):382-389. |
[8] | [ Le Jiajia, Su Yuan, Peng Qingwen, et al. Effects of nitrogen addition on soil enzyme activities and ecoenzymatic stoichiometry in alpine grassland of the Tianshan Mountains[J]. Arid Zone Research, 2020, 37(2):382-389. ] |
[9] | Xu Z W, Yu G R, Zhang X Y, et al. Soil enzyme activity and stoichiometry in forest ecosystems along the North-South transect in eastern China (NSTEC)[J]. Soil Biology and Biochemistry, 2017, 104:152-163. |
[10] | Cui Y X, Fang L C, Guo X B, et al. Ecoenzymatic stoichiometry and microbial nutrient limitation in rhizosphere soil in the arid area of the northern Loess Plateau, China[J]. Soil Biology and Biochemistry, 2018, 116:11-21. |
[11] | 吴秀芝, 阎欣, 王波, 等. 荒漠草地沙漠化对土壤-微生物-胞外酶化学计量特征的影响[J]. 植物生态学报, 2018, 42(10) : 1022-1032. |
[11] | [ Wu Xiuzhi, Yan Xin, Wang Bo, et al. Effects of desertification on the C : N : P stoichiometry of soil, microbes, and extracellular enzymes in a desert grassland[J]. Chinese Journal of Plant Ecology, 2018, 42(10):1022-1032. ] |
[12] | 郭天斗, 于露, 孙忠超, 等. 宁夏东部荒漠草原向灌丛地人为转变过程中土壤胞外酶活性响应[J]. 应用生态学报, 2020, 31(8) : 2541-2548. |
[12] | [ Guo Tiandou, Yu Lu, Sun Zhongchao, et al. Responses of soil extracellular enzyme activities to the anthropogenic transition from desert grassland to shrubland in eastern Ningxia, China[J]. Chinese Journal of Applied Ecology, 2020, 31(8):2541-2548. ] |
[13] | 左宜平, 张馨月, 曾辉, 等. 大兴安岭森林土壤胞外酶活力的时空动态及其对潜在碳矿化的影响[J]. 北京大学学报(自然科学版), 2018, 54(6):1311-1324. |
[13] | [ Zuo Yiping, Zhang Xinyue, Zeng Hui, et al. Spatiotemporal dynamics of soil extracellular enzyme activity and its influence on potential mineralization rate of soil organic carbon in forests of Daxing’an Mountain range[J]. Acta Scientiarum Naturalium Universitatis Pekinensis, 2018, 54(6):1311-1324. ] |
[14] | Xiao L, Liu G B, Li P, et al. Ecoenzymatic stoichiometry and microbial nutrient limitation during secondary formation of natural grassland on the Loess Plateau, China[J]. Soil & Tillage Research, 2020, 200:104605. |
[15] | 王嘉维. 黄土高原风蚀水蚀交错区典型植被对土壤酶活性的影响[J]. 山西农业科学, 2018, 46(6):976-980, 997. |
[15] | [ Wang Jiawei. Effects of typical vegetation on soil enzyme activities in wind erosion and water erosion crisscross region in Loess Plateau[J]. Journal of Shanxi Agricultural Sciences, 2018, 46(6):976-980, 997. ] |
[16] | 王彦峰, 肖波, 王兵, 等. 黄土高原水蚀风蚀交错区藓结皮对土壤酶活性的影响[J]. 应用生态学报, 2017, 28(11):3553-3561. |
[16] | [ Wang Yanfeng, Xiao Bo, Wang Bing, et al. Effects of moss-dominated biological soil crusts on soil enzyme activities in water-wind erosion crisscross region on the Loess Plateau of China[J]. Chinese Journal of Applied Ecology, 2017, 28(11):3553-3561. ] |
[17] | 李渊博, 李胜龙, 肖波, 等. 黄土高原水蚀风蚀交错区藓结皮覆盖土壤的蒸发特征[J]. 水土保持学报, 2020, 34(5):208-215. |
[17] | [ Li Yuanbo, Li Shenglong, Xiao Bo, et al. Evaporation characteristics of soil covered with moss crust in the wind-water erosion crisscross region of the Loess Plateau[J]. Journal of Soil and Water Conservation, 2020, 34(5):208-215. ] |
[18] | 张健, 徐明, 邹晓, 等. 不同土壤和植被生境下生物结皮对土壤性质的影响[J]. 水土保持学报, 2019, 33(5):323-328. |
[18] | [ Zhang Jian, Xu Ming, Zou Xiao, et al. Effects of biological crusts on soil properties under different soil and vegetation habitats[J]. Journal of Soil and Water Conservation, 2019, 33(5):323-328. ] |
[19] | 王彦峰, 王兵, 肖波, 等. 陕北黄土高原藓结皮的7种土壤酶活性及其剖面分布特征[J]. 西北农林科技大学学报(自然科学版), 2017, 45(3):161-169. |
[19] | [ Wang Yanfeng, Wang Bing, Xiao Bo, et al. Soil enzyme activities and vertical distribution of moss-dominated biological soil crusts on the Loess Plateau in Northern Shaanxi[J]. Journal of Northwest A & F University(Natural Science Edition), 2017, 45(3):161-169. ] |
[20] | 黄婷婷, 史扬子, 曹琦, 等. 黄土高原六道沟小流域近30年来土壤侵蚀变化评价[J]. 中国水土保持科学, 2020, 18(1):8-17. |
[20] | [ Huang Tingting, Shi Yangzi, Cao Qi, et al. Soil erosion evaluation of Liudaogou catchment in the Loess Plateau during the past 30 years[J]. Science of Soil and Water Conservation, 2020, 18(1):8-17. ] |
[21] | 王宝荣, 杨佳佳, 安韶山, 等. 黄土丘陵区植被与地形特征对土壤和土壤微生物生物量生态化学计量特征的影响[J]. 应用生态学报, 2018, 29(1):247-259. |
[21] | [ Wang Baorong, Yang Jiajia, An Shaoshan, et al. Effects of vegetation and topography features on ecological stoichiometry of soil and soil microbial biomass in the hilly-gully region of the Loess Plateau, China[J]. Chinese Journal of Applied Ecology, 2018, 29(1):247-259. ] |
[22] | Fanin N, Fromin N, Buatois B, et al. An experimental test of the hypojournal of non-homeostatic consumer stoichiometry in a plant litter-microbe system[J]. Ecology Letters, 2013, 16:764-772. |
[23] | Li X R, Song G, Hui R, et al. Precipitation and topsoil attributes determine the species diversity and distribution patterns of crustal communities in desert ecosystems[J]. Plant Soil, 2017, 420(1-2):163-175. |
[24] | 李云飞, 马晓俊, 李小军. 固沙植被演替过程中藓类结皮及其表层土壤理化性质变化[J]. 兰州大学学报(自然科学版), 2020, 56(4):463-470. |
[24] | [ Li Yunfei, Ma Xiaojun, Li Xiaojun. Changes in physicochemical properties of moss crusts and the underlying soil during the succession of sand-binding-vegetation[J]. Journal of Lanzhou University (Natural Sciences Edition), 2020, 56(4):463-470. ] |
[25] | 杨巧云, 赵允格, 包天莉, 等. 黄土丘陵区不同类型生物结皮下的土壤生态化学计量特征[J]. 应用生态学报, 2019, 30(8) : 2699-2706. |
[25] | [ Yang Qiaoyun, Zhao Yunge, Bao Tianli, et al. Soil ecological stoichiometry characteristics under different types of biological soil crusts in the hilly Loess Plateau region, China[J]. Chinese Journal of Applied Ecology, 2019, 30(8):2699-2706. ] |
[26] | 曾全超, 李鑫, 董扬红, 等. 黄土高原延河流域不同植被类型下土壤生态化学计量学特征[J]. 自然资源学报, 2016, 31(11):1881-1891. |
[26] | [ Zeng Quanchao, Li Xin, Dong Yanghong, et al. Ecological stoichiometry of soils in the Yanhe Watershed in the Loess Plateau: The influence of different vegetation zones[J]. Journal of Natural Resources, 2016, 31(11):1881-1891. ] |
[27] | 高丽倩, 赵允格, 许明祥, 等. 生物土壤结皮演替对土壤生态化学计量特征的影响[J]. 生态学报, 2018, 38(2):678-688. |
[27] | [ Gao Liqian, Zhao Yunge, Xu Mingxiang, et al. The effects of biological soil crust succession on soil ecological stoichiometry characteristics[J]. Acta Ecologica Sinica, 2018, 38(2):678-688. ] |
[28] | Zhao Y, Xu M, Belnap J. Potential nitrogen fixation activity of different aged biological soil crusts from rehabilitated grasslands of the hilly Loess Plateau, China[J]. Journal of Arid Environments, 2010, 74(10):1186-1191. |
[29] | Chen J, Sinsabaugh R L. Linking microbial functional gene abundance and soil extracellular enzyme activity: Implications for soil carbon dynamics[J]. Global Change Biology, 2021, 27:1322-1325. |
[30] | Miralles I, Domingo F, Cantón Y, et al. Hydrolase enzyme activities in a formation gradient of biological soil crusts in arid and semi-arid zones[J]. Soil Biology and Biochemistry, 2012, 53:124-132. |
[31] | 王荣女. 毛乌素沙地不同类型生物土壤结皮-土壤呼吸的季节变化特征[J]. 干旱区研究, 2021, 38(4):961-972. |
[31] | [ Wang Rongnv. Seasonal variation characteristics of different types of biological soil crust-soil system respiration in Mu Us Sandy Land[J]. Arid Zone Research, 2021, 38(4):961-972. ] |
[32] | 李彬, 武志芳, 陶冶, 等. 古尔班通古特沙漠不同类型生物结皮对草本植物多样性影响[J]. 干旱区研究, 2021, 38(2):438-449. |
[32] | [ Li Bin, Wu Zhifang, Tao Ye, et al. Effects of biological soil crust type on herbaceous diversity in the Gurbantunggut Desert[J]. Arid Zone Research, 2021, 38(2):438-449. ] |
[33] | Maxwell T L, Augusto L, Bon L, et al. Effect of a tree mixture and water availability on soil nutrients and extracellular enzyme activities along the soil profile in an experimental forest[J]. Soil Biology and Biochemistry, 2020, 148:107864. |
[34] | Liu J G, Gou X H, Zhang F, et al. Spatial patterns in the C:N:P stoichiometry in Qinghai spruce and the soil across the Qilian Mountains, China[J]. Catena, 2021, 196:104814. |
[35] | 程才, 李玉杰, 张远东, 等. 石漠化地区苔藓结皮对土壤养分及生态化学计量特征的影响[J]. 生态学报, 2020, 40(24):9234-9244. |
[35] | [ Cheng Cai, Li Yujie, Zhang Yuandong, et al. Effects of moss crusts on soil nutrients and ecological stoichiometry characteristics in karst rocky desertification region[J]. Acta Ecologica Sinica, 2020, 40(24):9234-9244. ] |
[36] | 娄泊远, 王永东, 闫晋升, 等. 亚寒带荒漠草原不同树种人工林土壤生态化学计量特征[J]. 干旱区研究, 2021, 38(5):1385-1392. |
[36] | [ Lou Boyuan, Wang Yongdong, Yan Jinsheng, et al. Characteristics of soil ecological stoichiometry of different tree spcies in sub-frigid desert steppe[J]. Arid Zone Research, 2021, 38(5):1385-1392. ] |
[37] | Wang H, Hu G Q, Xu W H, et al. Effects of nitrogen addition on soil organic carbon mineralization after maize stalk addition[J]. European Journal of Soil Biology, 2018, 89:33-38. |
[38] | Guan P T, Yang J J, Yang Y R, et al. Land conversion from cropland to grassland alleviates climate warming effects on nutrient limitation: Evidence from soil enzymatic activity and stoichiometry[J]. Global Ecology and Conservation, 2020, 24, e01328. |
[39] | Peng X Q, Wang W. Stoichiometry of soil extracellular enzyme activity along a climatic transect in temperate grasslands of northern China[J]. Soil Biology and Biochemistry, 2016, 98:74-84. |
[40] | Wang B, Xue S, Liu G B, et al. Changes in soil nutrient and enzyme activities under different vegetations in the Loess Plateau area, Northwest China[J]. Catena, 2012, 92:186-195. |
[41] | Jiao F, Wu J J, Qian Z, et al. Stimulation of nitrogen-hydrolyzing enzymes in soil aggregates mitigates nitrogen constraint for carbon sequestration following afforestation in subtropical China[J]. Soil Biology and Biochemistry, 2018, 123:136-144. |
[42] | 于德良, 雷泽勇, 张岩松, 等. 沙地樟子松人工林土壤酶活性及其影响因子[J]. 干旱区研究, 2019, 36(3):621-629. |
[42] | [ Yu Deliang, Lei Zeyong, Zhang Yansong, et al. Soil enzyme activity and its affecting factors under Pinus sylvestris var. mongolica plantation in sandy land[J]. Arid Zone Research, 2019, 36(3):621-629. ] |
[43] | 贾培龙, 安韶山, 李程程, 等. 黄土高原森林带土壤养分和微生物量及其生态化学计量变化特征[J]. 水土保持学报, 2020, 34(1):315-321. |
[43] | [ Jia Peilong, An Shaoshan, Li Chengcheng, et al. Dynamics of soil nutrients and their ecological stoichiometry characteristics under different longitudes in the East-West forest belt of the Loess Plateau[J]. Journal of Soil and Water Conservation, 2020, 34(1):315-321. ] |
[44] | Kiikkilä O, Kanerva S, Kitunen V, et al. Soil microbial activity in relation to dissolved organic matter properties under different tree species[J]. Plant Soil, 2014, 377(1-2):169-177. |
/
〈 | 〉 |