Short-term nitrogen pulse affects nitrogen fixation and ammonia oxidation in biocrusts: New insights into dryland nitrogen dynamics

Wei Hang , Waseem Shoukat , Zihan Kan , Shihang Zhang , Jungang Yang , Rongliang Jia , Ye Tao , Bengfeng Yin , Huiliang Liu , Yongxing Lu , Yuanming Zhang , Xiaobing Zhou

Soil Ecology Letters ›› 2026, Vol. 8 ›› Issue (4) : 260429

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Soil Ecology Letters ›› 2026, Vol. 8 ›› Issue (4) :260429 DOI: 10.1007/s42832-026-0429-0
RESEARCH ARTICLE
Short-term nitrogen pulse affects nitrogen fixation and ammonia oxidation in biocrusts: New insights into dryland nitrogen dynamics
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Abstract

In drylands, biocrusts function as essential components of the nitrogen cycle and display pronounced sensitivity to external nitrogen inputs. Episodic rainfall events can mobilize dry-deposited nitrogen into short-term pulse that influences nitrogen retention and transformation. However, the effects of short-term nitrogen pulse, commonly encountered in drylands, on biocrust nitrogen dynamics remain poorly understood. This study simulates rainfall-driven short-term nitrogen pulse to examine how varying pulse concentrations impact biocrusts nitrogen fixation, ammonia oxidation, and overall nitrogen balance under conditions of intensified nitrogen deposition after a 13-year nitrogen addition experiment in Gurbantunggut Desert. The nitrogen pulse sharply disrupted biocrusts' nitrogen cycling. Both nitrogen fixation and ammonia oxidation rates declined precipitously immediately after the pulse. However, within 14–21 days, these rates rebounded to or even surpassed pre-pulse levels. This pattern reflects the biocrusts' acute sensitivity to nitrogen perturbations, as well as their ecological resilience. Over 21 days, cumulative nitrogen fixation decreased by 47%–72% relative to the control, indicating suppression by nitrogen addition; in contrast, cumulative ammonia oxidation increased by 39%–91% but stabilized at moderate to high nitrogen inputs (1.0 g N m–2 yr–1, N1.0), revealing a saturation threshold. Moreover, the risk of nitrogen loss increased with higher nitrogen concentrations and plateaued at approximately an 88% elevation under N1.0, indicating that high nitrogen inputs exacerbate biocrusts nitrogen-loss vulnerabi-lity. These findings elucidate the impacts of acute nitrogen pulse on nitrogen dynamics in biocrusts, highlighting that amid escalating global nitrogen deposition, precise evaluation of nitrogen balance in drylands must integrate pulse effects, biocrusts nitrogen-carrying capacity, input modes, concentrations, durations, and interactions with environmental conditions.

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Keywords

biocrusts / nitrogen pulse / nitrogen fixation / ammonia oxidation

Highlight

● Short-term nitrogen pulses significantly alter nitrogen cycling dynamics in biocrusts.

● The mechanism of short-term sensitivity and recovery in biocrusts nitrogen dynamics to nitrogen disturbance was elucidated.

● Nitrogen pulses exacerbate nitrogen loss risks, providing new insights for dryland nitrogen deposition research.

● Nitrogen loss risks increase with nitrogen concentration, exposing the ecological vulnerability of drylands.

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Wei Hang, Waseem Shoukat, Zihan Kan, Shihang Zhang, Jungang Yang, Rongliang Jia, Ye Tao, Bengfeng Yin, Huiliang Liu, Yongxing Lu, Yuanming Zhang, Xiaobing Zhou. Short-term nitrogen pulse affects nitrogen fixation and ammonia oxidation in biocrusts: New insights into dryland nitrogen dynamics. Soil Ecology Letters, 2026, 8(4): 260429 DOI:10.1007/s42832-026-0429-0

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References

[1]

Antoninka, A., Chuckran, P.F., Mau, R.L., Slate, M.L., Mishler, B.D., Oliver, M.J., Coe, K.K., Stark, L.R., Fisher, K.M., Bowker, M.A., 2022. Responses of biocrust and associated soil bacteria to novel climates are not tightly coupled. Frontiers in Microbiology13, 821860.

[2]

Barger, N.N., Castle, S.C., Dean, G.N., 2013. Denitrification from nitrogen-fixing biologically crusted soils in a cool desert environment, southeast Utah, USA. Ecological Processes2, 16.

[3]

Belnap, J., 2002. Nitrogen fixation in biological soil crusts from southeast Utah, USA. Biology and Fertility of Soils35, 128–135.

[4]

Benvenutto-Vargas, V.P., Ochoa-Hueso, R., 2020. Effects of nitrogen deposition on the spatial pattern of biocrusts and soil microbial activity in a semi-arid Mediterranean shrubland. Functional Ecology34, 923–937.

[5]

Bremer, E., Krämer, R., 2019. Responses of microorganisms to osmotic stress. Annual Review of Microbiology73, 313–334.

[6]

Chang, C.N., Kwok, O.M., 2022. Partitioning variance for a within-level predictor in multilevel models. Structural Equation Modeling: A Multidisciplinary Journal29, 716–730.

[7]

Chen, R.L., Tan, X.Y., Zhang, Y.M., Chen, H., Yin, B.F., Zhu, X.L., Chen, J., 2023. Monitoring rainfall events in desert areas using the spectral response of biological soil crusts to hydration: evidence from the Gurbantunggut Desert, China. Remote Sensing of Environment286, 113448.

[8]

Chen, Y.N., Wang, Q., Li, W.H., Ruan, X., 2007. Microbiotic crusts and their interrelations with environmental factors in the Gurbantonggut desert, western China. Environmental Geology52, 691–700.

[9]

Cronin, J.P., Schoolmaster, D.R., 2018. A causal partition of trait correlations: using graphical models to derive statistical models from theoretical language. Ecosphere9, e02422.

[10]

Delgado-Baquerizo, M., Morillas, L., Maestre, F.T., Gallardo, A., 2013. Biocrusts control the nitrogen dynamics and microbial functional diversity of semi-arid soils in response to nutrient additions. Plant and Soil372, 643–654.

[11]

Dixon, P., 2003. VEGAN, a package of R functions for community ecology. Journal of Vegetation Science14, 927–930.

[12]

Evans, S.E., Wallenstein, M.D., 2012. Soil microbial community response to drying and rewetting stress: does historical precipitation regime matter. Biogeochemistry109, 101–116.

[13]

Fan, K.K., Delgado-Baquerizo, M., Guo, X.S., Wang, D.Z., Wu, Y.Y., Zhu, M., Yu, W., Yao, H.Y., Zhu, Y.G., Chu, H.Y., 2019. Suppressed N fixation and diazotrophs after four decades of fertilization. Microbiome7, 143.

[14]

Fernandes, V.M.C., Rudgers, J.A., Collins, S.L., Garcia-Pichel, F., 2022. Rainfall pulse regime drives biomass and community composition in biological soil crusts. Ecology103, e3744.

[15]

García, L.V., 2004. Escaping the bonferroni iron claw in ecological studies. Oikos105, 657–663.

[16]

Göransson, H., Godbold, D.L., Jones, D.L., Rousk, J., 2013. Bacterial growth and respiration responses upon rewetting dry forest soils: impact of drought-legacy. Soil Biology and Biochemistry57, 477–486.

[17]

Hang, W., Shoukat, W., Yang, J.G., Lu, Y.X., Qin, S.P., Liu, H.L., Rong, X.Y., Zhou, X.B., Zhang, Y.M., 2025. Effects of simulated nitrogen deposition on microbial dynamics: altered nitrogen fixation and ammonia oxidation in biological soil crusts. Journal of Environmental Management380, 125141.

[18]

James, J.J., Richards, J.H., 2006. Plant nitrogen capture in pulse-driven systems: interactions between root responses and soil processes. Journal of Ecology94, 765–777.

[19]

Jenerette, G.D., Chatterjee, A., 2012. Soil metabolic pulses: water, substrate, and biological regulation. Ecology93, 959–966.

[20]

Lefcheck, J.S., 2016. PIECEWISESEM: piecewise structural equation modelling in R for ecology, evolution, and systematics. Methods in Ecology and Evolution7, 573–579.

[21]

Liang, C., Schimel, J.P., Jastrow, J.D., 2017. The importance of anabolism in microbial control over soil carbon storage. Nature Microbiology2, 17105.

[22]

Liu, C., Liu, J.J., Wang, J., Ding, X.Y., 2024. Effects of short-term nitrogen additions on biomass and soil phytochemical cycling in alpine grasslands of Tianshan, China. Plants13, 1103.

[23]

Lu, M.Z., Cheng, S.L., Fang, H.J., Xu, M., Yang, Y., Li, Y.N., Zhang, J.B., Müller, C., 2021. Organic nitrogen addition causes decoupling of microbial nitrogen cycles by stimulating gross nitrogen transformation in a temperate forest soil. Geoderma385, 114886.

[24]

Luo, G.W., Ling, N., Xue, C., Dippold, M.A., Firbank, L.G., Guo, S.W., Kuzyakov, Y., Shen, Q.R., 2019. Nitrogen-inputs regulate microbial functional and genetic resistance and resilience to drying–rewetting cycles, with implications for crop yields. Plant and Soil441, 301–315.

[25]

Maier, S., Kratz, A.M., Weber, J., Prass, M., Liu, F., Clark, A.T., Abed, R.M.M., Su, H., Cheng, Y., Eickhorst, T., Fiedler, S., Pöschl, U., Weber, B., 2022. Water-driven microbial nitrogen transformations in biological soil crusts causing atmospheric nitrous acid and nitric oxide emissions. The ISME Journal16, 1012–1024.

[26]

Manzoni, S., Schaeffer, S.M., Katul, G., Porporato, A., Schimel, J.P., 2014. A theoretical analysis of microbial eco-physiological and diffusion limitations to carbon cycling in drying soils. Soil Biology and Biochemistry73, 69–83.

[27]

Mao, C., Kou, D., Peng, Y.F., Qin, S.Q., Zhang, Q.W., Yang, Y.H., 2021. Soil nitrogen transformations respond diversely to multiple levels of nitrogen addition in a Tibetan alpine steppe. Journal of Geophysical Research: Biogeosciences126, e2020JG006211.

[28]

Meisner, A., Rousk, J., Bååth, E., 2015. Prolonged drought changes the bacterial growth response to rewetting. Soil Biology and Biochemistry88, 314–322.

[29]

Micks, P., Aber, J.D., Boone, R.D., Davidson, E.A., 2004. Short-term soil respiration and nitrogen immobilization response to nitrogen applications in control and nitrogen-enriched temperate forests. Forest Ecology and Management196, 57–70.

[30]

Miller, A.E., Schimel, J.P., Meixner, T., Sickman, J., Melack, J., 2005. Episodic rewetting enhances carbon and nitrogen release from chaparral soils. Soil Biology and Biochemistry37, 2195–2204.

[31]

Moreau, D., Bardgett, R.D., Finlay, R.D., Jones, D.L., Philippot, L., 2019. A plant perspective on nitrogen cycling in the rhizosphere. Functional Ecology33, 540–552.

[32]

Niu, Y.Y., Duan, Y.L., Li, Y.Q., Wang, X.Y., Chen, Y., Wang, L.L., 2021. Soil microbial community responses to short-term nitrogen addition in China’s Horqin Sandy Land. PLoS One16, e0242643.

[33]

Ramond, J.B., Jordaan, K., Díez, B., Heinzelmann, S.M., Cowan, D.A., 2022. Microbial biogeochemical cycling of nitrogen in arid ecosystems. Microbiology and Molecular Biology Reviews86, e00109–21.

[34]

Rodriguez-Caballero, E., Belnap, J., Büdel, B., Crutzen, P.J., Andreae, M.O., Pöschl, U., Weber, B., 2018. Dryland photoautotrophic soil surface communities endangered by global change. Nature Geoscience11, 185–189.

[35]

Rong, X.Y., Zhou, X.B., Li, X.Z., Yao, M.J., Lu, Y.X., Xu, P., Yin, B.F., Li, Y.G., Aanderud, Z.T., Zhang, Y.M., 2022. Biocrust diazotrophs and bacteria rather than fungi are sensitive to chronic low N deposition. Environmental Microbiology24, 5450–5466.

[36]

Schwinning, S., Sala, O.E., 2004. Hierarchy of responses to resource pulses in arid and semi-arid ecosystems. Oecologia141, 211–220.

[37]

Sinsabaugh, R.L., Manzoni, S., Moorhead, D.L., Richter, A., 2013. Carbon use efficiency of microbial communities: stoichiometry, methodology and modelling. Ecology Letters16, 930–939.

[38]

Strauss, S.L., Day, T.A., Garcia-Pichel, F., 2012. Nitrogen cycling in desert biological soil crusts across biogeographic regions in the Southwestern United States. Biogeochemistry108, 171–182.

[39]

Sun, F.H., Xiao, B., Kidron, G.J., Heitman, J., 2024. Biocrusts critical regulation of soil water vapor transport (diffusion, sorption, and late-stage evaporation) in drylands. Water Resources Research60, e2023WR036520.

[40]

Tian, D.S., Niu, S.L., 2015. A global analysis of soil acidification caused by nitrogen addition. Environmental Research Letters10, 024019.

[41]

Vourlitis, G.L., Fernandez, J.S., 2015. Carbon and nitrogen mineralization of semi-arid shrubland soils exposed to chronic nitrogen inputs and pulses of labile carbon and nitrogen. Journal of Arid Environments122, 37–45.

[42]

Weber, B., Wu, D.M., Tamm, A., Ruckteschler, N., Rodríguez-Caballero, E., Steinkamp, J., Meusel, H., Elbert, W., Behrendt, T., Sörgel, M., Cheng, Y.F., Crutzen, P.J., Su, H., Pöschl, U., 2015. Biological soil crusts accelerate the nitrogen cycle through large NO and HONO emissions in drylands. Proceedings of the National Academy of Sciences of the United States of America112, 15384–15389.

[43]

Wei, L., Ge, T.D., Zhu, Z.K., Ye, R.Z., Peñuelas, J., Li, Y.H., Lynn, T.M., Jones, D.L., Wu, J.S., Kuzyakov, Y., 2022. Paddy soils have a much higher microbial biomass content than upland soils: a review of the origin, mechanisms, and drivers. Agriculture, Ecosystems & Environment326, 107798.

[44]

Wei, T., Simko, V., 2024. R Package “corrplot”: Visualization of A Correlation Matrix [Online]. .

[45]

Xu, W., Luo, X.S., Pan, Y.P., Zhang, L., Tang, A.H., Shen, J.L., Zhang, Y., Li, K.H., Wu, Q.H., Yang, D.W., Zhang, Y.Y., Xue, J., Li, W.Q., Li, Q.Q., Tang, L., Lu, S.H., Liang, T., Tong, Y.A., Liu, P., Zhang, Q., Xiong, Z.Q., Shi, X.J., Wu, L.H., Shi, W.Q., Tian, K., Zhong, X.H., Shi, K., Tang, Q.Y., Zhang, L.J., Huang, J.L., He, C.E., Kuang, F.H., Zhu, B., Liu, H., Jin, X., Xin, Y.J., Shi, X.K., Du, E.Z., Dore, A.J., Tang, S., Collett, J.L.J., Goulding, K., Sun, Y.X., Ren, J., Zhang, F.S., Liu, X.J., 2015. Quantifying atmospheric nitrogen deposition through a nationwide monitoring network across China. Atmospheric Chemistry and Physics15, 12345–12360.

[46]

Zar, J.H., 2013. Biostatistical Analysis. 5th ed. Harlow: Pearson Education Limited.

[47]

Zhang, J., Zhang, Y.M., 2020. Ecophysiological responses of the biocrust moss Syntrichia caninervis to experimental snow cover manipulations in a temperate desert of central Asia. Ecological Research35, 198–207.

[48]

Zhang, T., Chen, H.Y.H., Ruan, H.H., 2018. Global negative effects of nitrogen deposition on soil microbes. The ISME Journal12, 1817–1825.

[49]

Zhang, Y., Zhang, F., Abalos, D., Luo, Y.Q., Hui, D.F., Hungate, B.A., García-Palacios, P., Kuzyakov, Y., Olesen, J.E., Jørgensen, U., Chen, J., 2022. Stimulation of ammonia oxidizer and denitrifier abundances by nitrogen loading: poor predictability for increased soil N2O emission. Global Change Biology28, 2158–2168.

[50]

Zheng, M.H., Xu, M.C., Li, D.J., Deng, Q., Mo, J.M., 2023. Negative responses of terrestrial nitrogen fixation to nitrogen addition weaken across increased soil organic carbon levels. Science of the Total Environment877, 162965.

[51]

Zhou, G.Y., Zhou, L.Y., Shao, J.J., Zhou, X.H., 2020. Effects of extreme drought on terrestrial ecosystems: review and prospects. Chinese Journal of Plant Ecology44, 515–525.

[52]

Zhou, X.B., Smith, H., Silva, A.G., Belnap, J., Garcia-Pichel, F., 2016a. Differential responses of dinitrogen fixation, diazotrophic cyanobacteria and ammonia oxidation reveal a potential warming-induced imbalance of the N-cycle in biological soil crusts. PLoS One11, e0164932.

[53]

Zhou, X.B., Zhang, Y.M., Yin, B.F., 2016b. Divergence in physiological responses between cyanobacterial and lichen crusts to a gradient of simulated nitrogen deposition. Plant and Soil399, 121–134.

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