The context dependency of nitrogen deposition impacts on the compositional coupling between aboveground vegetation and soil seed bank

doi:10.1016/j.pld.2025.12.009

Plant Diversity ›› 2026, Vol. 48 ›› Issue (02): 373-380.DOI: 10.1016/j.pld.2025.12.009

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The context dependency of nitrogen deposition impacts on the compositional coupling between aboveground vegetation and soil seed bank

Niwu Te^a,b,c, Wen-Tao Luo^a, Péter Török^d,e,f, Xiao-Ru Zhang^a,b, Xiao-Sa Liang^a, Yuan-Xiu Wu^a,b, Xiao-Jing Zhang^a,b, Anke Jentsch^c, Xiao-Tao Lü^a

a. Erguna Forest-Steppe Ecotone Research Station, CAS Key Laboratory of Forest Ecology and Silviculture, Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang 110016, China;
b. College of Resources and Environment, University of Chinese Academy of Sciences, Beijing 100081, China;
c. Disturbance Ecology and Vegetation Dynamics, University of Bayreuth, Bayreuth 95447, Germany;
d. Department of Ecology, University of Debrecen, Debrecen 4032, Hungary;
e. HUN-REN-UD Functional and Restoration Ecology Research Group, Debrecen 4032, Hungary;
f. Polish Academy of Sciences, Botanical Garden - Centre for Biological Diversity Conservation in Powsin, Warszawa 02-973, Poland

Received:2025-09-17 Revised:2025-12-13 Online:2026-05-19 Published:2026-03-25
Contact: Xiao-Tao Lü,E-mail:lvxiaotao@iae.ac.cn
Supported by:
We appreciate the logistical support from the National Field Observation and Research Station of Shenyang Agro-ecosystems and the constructive comments and suggestions from three anonymous reviewers. This study was supported by the National Key Research and Development Program of China (2023YFD1301702), Natural Science Foundation of China (32171543 and 32422057), and the China Scholarship Council joint Ph.D. program. PT was supported by the National Research, Development and Innovation Office [KKP144068 and K137573] during the manuscript preparation.

The context dependency of nitrogen deposition impacts on the compositional coupling between aboveground vegetation and soil seed bank

Niwu Te^a,b,c, Wen-Tao Luo^a, Péter Török^d,e,f, Xiao-Ru Zhang^a,b, Xiao-Sa Liang^a, Yuan-Xiu Wu^a,b, Xiao-Jing Zhang^a,b, Anke Jentsch^c, Xiao-Tao Lü^a

a. Erguna Forest-Steppe Ecotone Research Station, CAS Key Laboratory of Forest Ecology and Silviculture, Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang 110016, China;
b. College of Resources and Environment, University of Chinese Academy of Sciences, Beijing 100081, China;
c. Disturbance Ecology and Vegetation Dynamics, University of Bayreuth, Bayreuth 95447, Germany;
d. Department of Ecology, University of Debrecen, Debrecen 4032, Hungary;
e. HUN-REN-UD Functional and Restoration Ecology Research Group, Debrecen 4032, Hungary;
f. Polish Academy of Sciences, Botanical Garden - Centre for Biological Diversity Conservation in Powsin, Warszawa 02-973, Poland

通讯作者: Xiao-Tao Lü,E-mail:lvxiaotao@iae.ac.cn
基金资助:
We appreciate the logistical support from the National Field Observation and Research Station of Shenyang Agro-ecosystems and the constructive comments and suggestions from three anonymous reviewers. This study was supported by the National Key Research and Development Program of China (2023YFD1301702), Natural Science Foundation of China (32171543 and 32422057), and the China Scholarship Council joint Ph.D. program. PT was supported by the National Research, Development and Innovation Office [KKP144068 and K137573] during the manuscript preparation.

Abstract

Abstract: The compositional coupling between aboveground vegetation and soil seed bank is critical for community stability and successional trajectories, but such relationships are highly sensitive to environmental changes. Although nitrogen (N) deposition has been reported to decouple the associations between aboveground vegetation and the soil seed bank in grasslands, it remains unclear whether and how grassland management practices, such as mowing, mediate N deposition effects. We evaluated the impacts of N input and mowing on the diversity and composition of the aboveground vegetation and soil seed bank, as well as their compositional coupling in a decadal grassland experiment. N addition increased the compositional dissimilarity between the soil seed bank and aboveground vegetation by increasing abundance gradient, but only in unmown plots. N addition did not affect the compositional dissimilarity between aboveground vegetation and the soil seed bank in mown plots. Mowing increased the graminoid abundance in the aboveground vegetation, but increased the forb abundance in the soil seed bank, and thus increased the compositional dissimilarity between aboveground vegetation and the soil seed bank by promoting balanced variation in abundance. Our results reveal the importance of context dependence in the consequences of N inputs on the compositional coupling between aboveground vegetation and the soil seed bank in grasslands. The land-use context should be fully considered when evaluating the impacts of global change drivers on grassland community dynamics and succession.

Key words: Compositional coupling, Grassland management, Hay harvest, Nitrogen enrichment, Soil seed bank, Temperate steppe

摘要： The compositional coupling between aboveground vegetation and soil seed bank is critical for community stability and successional trajectories, but such relationships are highly sensitive to environmental changes. Although nitrogen (N) deposition has been reported to decouple the associations between aboveground vegetation and the soil seed bank in grasslands, it remains unclear whether and how grassland management practices, such as mowing, mediate N deposition effects. We evaluated the impacts of N input and mowing on the diversity and composition of the aboveground vegetation and soil seed bank, as well as their compositional coupling in a decadal grassland experiment. N addition increased the compositional dissimilarity between the soil seed bank and aboveground vegetation by increasing abundance gradient, but only in unmown plots. N addition did not affect the compositional dissimilarity between aboveground vegetation and the soil seed bank in mown plots. Mowing increased the graminoid abundance in the aboveground vegetation, but increased the forb abundance in the soil seed bank, and thus increased the compositional dissimilarity between aboveground vegetation and the soil seed bank by promoting balanced variation in abundance. Our results reveal the importance of context dependence in the consequences of N inputs on the compositional coupling between aboveground vegetation and the soil seed bank in grasslands. The land-use context should be fully considered when evaluating the impacts of global change drivers on grassland community dynamics and succession.

关键词: Compositional coupling, Grassland management, Hay harvest, Nitrogen enrichment, Soil seed bank, Temperate steppe

Niwu Te, Wen-Tao Luo, Péter Török, Xiao-Ru Zhang, Xiao-Sa Liang, Yuan-Xiu Wu, Xiao-Jing Zhang, Anke Jentsch, Xiao-Tao Lü. The context dependency of nitrogen deposition impacts on the compositional coupling between aboveground vegetation and soil seed bank[J]. Plant Diversity, 2026, 48(02): 373-380.

References

[1] Angert, A.L., Huxman, T.E., Chesson, P., et al., 2009. Functional tradeoffs determine species coexistence via the storage effect. Proc. Natl. Acad. Sci. U.S.A. 106 (28), 11641-11645. https://doi.org/10.1073/pnas.0904512106.
[2] Bahn, M., Ingrisch, J., Jentsch, A., 2022. Grasslands. In: Wohlgemuth, T., Jentsch, A., Seidl, R. (Eds.), Disturbance Ecology. Landscape Series, vol. 32. Springer, Cham, pp. 349-376. https://doi.org/10.1007/978-3-030-98756-5.
[3] Bardgett, R.D., Bullock, J.M., Lavorel, S., et al., 2021. Combatting global grassland degradation. Nat. Rev. Earth Environ. 2, 720-735. https://doi.org/10.1038/s43017-021-00207-2.
[4] Baselga, A., 2013. Separating the two components of abundance-based dissimilarity: balanced changes in abundance vs. abundance gradients. Methods Ecol. Evol. 4, 552-557. https://doi.org/10.1111/2041-210X.12029.
[5] Bengtsson, J., Bullock, J.M., Egoh, B., et al., 2019. Grasslands-More important for ecosystem services than you might think. Ecosphere 10 (2), e02582. https://doi.org/10.1002/ecs2.2582.
[6] Borer, E.T., Seabloom, E.W., Gruner, D.S., et al., 2014. Herbivores and nutrients control grassland plant diversity via light limitation. Nature 508, 517-520. https://doi.org/10.1038/nature13144.
[7] Chen, Q., Blowes, S.A., Ladouceur, E., et al., 2025. Local nutrient addition drives plant biodiversity losses but not biotic homogenization in global grasslands. Nat. Commun. 16, 4903. https://doi.org/10.1038/s41467-025-59166-7.
[8] Clark, C.M., Tilman, D., 2008. Loss of plant species after chronic low-level nitrogen deposition to prairie grasslands. Nature 451, 712-715. https://doi.org/10.1038/nature06503.
[9] DeMalach, N., Kigel, J., Sternberg, M., 2021. The soil seed bank can buffer long-term compositional changes in annual plant communities. J. Ecol. 109, 1275-1283. https://doi.org/10.1111/1365-2745.13555.
[10] Eskelinen, A., Jessen, M., Bahamonde, H.A., et al., 2023. Herbivory and nutrients shape grassland soil seed banks. Nat. Commun. 14, 3949. https://doi.org/10.1038/s41467-023-39677-x.
[11] Habenczyus, A.A., Weiterova, I., Foremnik, K., et al., 2024. Long-term effects of meadow management on seed bank diversity and composition. J. Veg. Sci. 35, e13282. https://doi.org/10.1111/jvs.13282.
[12] Hautier, Y., Niklaus, P.A., Hector, A., 2009. Competition for light causes plant biodiversity loss after eutrophication. Science 324, 636-638. https://doi.org/10.1126/science.1169640.
[13] Hopfensperger, K.N., 2007. A review of similarity between seed bank and standing vegetation across ecosystems. Oikos 116, 1438-1448. https://doi.org/10.1111/j.0030-1299.2007.15818.x.
[14] Kiss, R., Deak, B., Torok, P., et al., 2018. Grassland seed bank and community resilience in a changing climate. Restor. Ecol. 26, 141-150. https://doi.org/10.1111/rec.12694.
[15] Klaus, V.H., Schafer, D., Prati, D., et al., 2018. Effects of mowing, grazing, and fertilization on soil seed banks in temperate grasslands in Central Europe. Agric. Ecosyst. Environ. 256, 211-217. https://doi.org/10.1016/j.agee.2017.11.008.
[16] Klimesova, J., Martinkova, J., Bartuskova, A., et al., 2023. Belowground plant traits and their ecosystem functions along aridity gradients in grasslands. Plant Soil 487, 39-48. https://doi.org/10.1007/s11104-023-05964-1.
[17] Ma, M., Collins, S.L., Ratajczak, Z., et al., 2021. Soil seed banks, alternative stable state theory, and ecosystem resilience. Bioscience 71(7), 697-707. https://doi.org/10.1093/biosci/biab011.
[18] Ma, M., Eskelinen, A., Zhao, Y., et al., 2025. Multiple mechanisms associated with loss of seed bank diversity under nitrogen enrichment. J. Ecol. 00, 1-13. https://doi.org/10.1111/1365-2745.14486.
[19] Miao, R., Guo, M., Ma, J., et al., 2020. Shifts in soil seed bank and plant community under nitrogen addition and mowing in an Inner Mongolian steppe. Ecol. Eng. 153, 105900. https://doi.org/10.1016/j.ecoleng.2020.105900.
[20] Molina, C.D., Tognetti, P.M., Graff, P., et al., 2021. Mowing does not redress the negative effect of nutrient addition on alpha and beta diversity in a temperate grassland. J. Ecol. 109, 1501-1510. https://doi.org/10.1111/1365-2745.13573.
[21] Ott, J.P., Klimesova, J., Hartnett, D.C., 2019. The ecology and significance of below-ground bud banks in plants. Ann. Bot. 123(7), 1099-1118. https://doi.org/10.1093/aob/mcz051.
[22] R Core Team, 2022. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria.
[23] Simkin, S.M., Allen, E.B., Bowman, W.D., et al., 2016. Conditional vulnerability of plant diversity to atmospheric nitrogen deposition across the United States. Proc. Natl. Acad. Sci. U.S.A. 113(15), 4086-4091. https://doi.org/10.1073/pnas.1515241113.
[24] Stevens, C.J., Dise, N.B., Mountford, J.O., et al., 2004. Impact of nitrogen deposition on the species richness of grasslands. Science 303, 1876-1879. https://doi.org/10.1126/science.1094678.
[25] Suding, K.N., Collins, S.L., Gough, L., et al., 2005. Functional- and abundance-based mechanisms explain diversity loss due to N fertilization. Proc. Natl. Acad. Sci. U.S.A. 102(12), 4387-4392. https://doi.org/10.1073/pnas.0408648102.
[26] Talle, M., Deak, B., Poschlod, P., et al., 2016. Grazing vs. mowing: A meta-analysis of biodiversity benefits for grassland management. Agric. Ecosyst. Environ. 222, 200-212. https://doi.org/10.1016/j.agee.2016.02.008.
[27] Te, N., Griffin-Nolan, R.J., Slette, I.J., et al., 2025. Grassland bud and shoot demographic responses to single and recurrent droughts vary across an aridity gradient. Ecology 106(2), e70002. https://doi.org/10.1002/ecy.70002.
[28] Tognetti, P.M., Prober, S.M., Baez, S., et al., 2021. Negative effects of nitrogen override positive effects of phosphorus on grassland legumes worldwide. Proc. Natl. Acad. Sci. U.S.A. 118(28), e2023718118. https://doi.org/10.1073/pnas.2023718118.
[29] Torok, P., Kelemen, A., Valko, O., et al., 2018. Succession in soil seed banks and its implications for restoration of calcareous sand grasslands. Restor. Ecol. 26, S134-S140. https://doi.org/10.1111/rec.12611.
[30] Vitousek, P.M., Aber, J.D., Howarth, R.W., et al., 1997. Human alteration of the global nitrogen cycle: Sources and consequences. Ecol. Appl. 7(3), 737-750. https://doi.org/10.1890/1051-0761(1997)007[0737:HAOTGN]2.0.CO;2.
[31] Wang, X., Ge, W., Zhang, M., et al., 2024. Large and non-spherical seeds are less likely to form a persistent soil seed bank. Proc. R. Soc. B. 291, 20232764. https://doi.org/10.1098/rspb.2023.2764.
[32] Yang, G.J., Lu, X.T., Stevens, C.J., et al., 2019. Mowing mitigates the negative impacts of N addition on plant species diversity. Oecologia 189, 769-779. https://doi.org/10.1007/s00442-019-04353-9.
[33] Yang, Z., Hautier, Y., Borer, E.T., et al., 2015. Abundance- and functional-based mechanisms of plant diversity loss with fertilization in the presence and absence of herbivores. Oecologia 179, 261-270. https://doi.org/10.1007/s00442-015-3313-7.
[34] Zhang, C., Willis, C.G., Ma, Z., et al., 2019. Direct and indirect effects of long-term fertilization on the stability of the persistent seed bank. Plant Soil 438, 239-250. https://doi.org/10.1007/s11104-019-04024-x.
[35] Zhang, X., Yang, G., Ning, Y., et al., 2025. Dominant species drove the balance between biodiversity and productivity in mown grasslands under nitrogen fertilization. Ecol. Appl. 35(1), e70009. https://doi.org/10.1002/eap.70009.
[36] Zhang, Y., Loreau, M., He, N., et al., 2017. Mowing exacerbates the loss of ecosystem stability under nitrogen enrichment in a temperate grassland. Funct. Ecol. 31, 1637-1646. https://doi.org/10.1111/1365-2435.12850.
[37] Zhao, M., Loreau, M., Ochoa-Hueso, R., et al., 2023. Decoupled responses of above- and below-ground beta-diversity to nitrogen enrichment in a typical steppe. Ecol. Lett. 27, e14339. https://doi.org/10.1111/ele.14339.

The context dependency of nitrogen deposition impacts on the compositional coupling between aboveground vegetation and soil seed bank

The context dependency of nitrogen deposition impacts on the compositional coupling between aboveground vegetation and soil seed bank

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