Plant diversity is key for microbial necromass carbon accrual in alpine grasslands

1. Bai, Y. & Cotrufo, M. F. Grassland soil carbon sequestration: current understanding, challenges, and solutions. Science 377, 603–608 (2022).

   Google Scholar 

2. Wang, Y. et al. Grassland changes and adaptive management on the Qinghai-Tibetan Plateau. Nat. Rev. Earth Environ. 3, 668–683 (2022).

   Google Scholar 

3. Bardgett, R. D. et al. Combatting global grassland degradation. Nat. Rev. Earth Environ. 2, 720–735 (2021).

   Google Scholar 

4. Chen, H. et al. The impacts of climate change and human activities on biogeochemical cycles on the Qinghai-Tibetan Plateau. Glob. Change Biol. 19, 2940–2955 (2013).

   Google Scholar 

5. Tilman, D., Isbell F. & Cowles J. M. Biodiversity and ecosystem functioning. In: Annual Review of Ecology, Evolution & Systematics, Vol 45 (ed Futuyma D. J.) (2014).

6. Tilman, D., Reich, P. B. & Isbell, F. Biodiversity impacts ecosystem productivity as much as resources, disturbance, or herbivory. Proc. Natl. Acad. Sci. USA 109, 10394–10397 (2012).

   Google Scholar 

7. Chen, L. et al. Soil carbon persistence governed by plant input and mineral protection at regional and global scales. Ecol. Lett. 24, 1018–1028 (2021).

   Google Scholar 

8. Chen, X. et al. Effects of plant diversity on soil carbon in diverse ecosystems: a global meta-analysis. Biol. Rev. 95, 167–183 (2020).

   Google Scholar 

9. Yang, Y., Tilman, D., Furey, G. & Lehman, C. Soil carbon sequestration accelerated by restoration of grassland biodiversity. Nat. Commun. 10, 718 (2019).

   Google Scholar 

10. Spohn, M. et al. The positive effect of plant diversity on soil carbon depends on climate. Nat. Commun. 14, 6624 (2023).

    Google Scholar 

11. Chen, H. et al. Carbon and nitrogen cycling on the Qinghai-Tibetan Plateau. Nat. Rev. Earth Environ. 3, 701–716 (2022).

    Google Scholar 

12. Angst, Š, Angst, G., Mueller, K. E., Lange, M. & Eisenhauer, N. Un (der) explored links between plant diversity and particulate and mineral-associated organic matter in soil. Nat. Commun. 16, 5548 (2025).

    Google Scholar 

13. Wang, B., An, S., Liang, C., Liu, Y. & Kuzyakov, Y. Microbial necromass as the source of soil organic carbon in global ecosystems. Soil Biol. Biochem. 162, 108422 (2021).

    Google Scholar 

14. Hu, H. et al. Reducing the uncertainty in estimating soil microbial-derived carbon storage. Proc. Natl. Acad. Sci. USA 121, e2401916121 (2024).

    Google Scholar 

15. Liang, C., Schimel, J. P. & Jastrow, J. D. The importance of anabolism in microbial control over soil carbon storage. Nat. Microbiol. 2, 17105 (2017).

    Google Scholar 

16. Buckeridge, K. M., Creamer, C. & Whitaker, J. Deconstructing the microbial necromass continuum to inform soil carbon sequestration. Funct. Ecol. 36, 1396–1410 (2022).

    Google Scholar 

17. Manzoni, S., Taylor, P., Richter, A., Porporato, A. & Ågren, G. I. Environmental and stoichiometric controls on microbial carbon-use efficiency in soils. N. Phytol. 196, 79–91 (2012).

    Google Scholar 

18. Lange, M. et al. Plant diversity increases soil microbial activity and soil carbon storage. Nat. Commun. 6, 6707 (2015).

    Google Scholar 

19. Prommer, J. et al. Increased microbial growth, biomass, and turnover drive soil organic carbon accumulation at higher plant diversity. Glob. Change Biol. 26, 669–681 (2020).

    Google Scholar 

20. Mou, X. et al. Plant species richness mediates the responses of microbial necromass carbon accumulation to climate aridity in alpine meadows. J. Ecol. 113, 883–895 (2025).

    Google Scholar 

21. Domeignoz-Horta, L. A. et al. Plant diversity drives positive microbial associations in the rhizosphere enhancing carbon use efficiency in agricultural soils. Nat. Commun. 15, 8065 (2024).

    Google Scholar 

22. He, M. et al. Depth-dependent drivers of soil microbial necromass carbon across Tibetan alpine grasslands. Glob. Change Biol. 28, 936–949 (2022).

    Google Scholar 

23. Zhang, X. et al. Links between microbial biomass and necromass components in the top- and subsoils of temperate grasslands along an aridity gradient. Geoderma 379, 114623 (2020).

    Google Scholar 

24. Fierer, N., Schimel, J. P. & Holden, P. A. Variations in microbial community composition through two soil depth profiles. Soil Biol. Biochem. 35, 167–176 (2003).

    Google Scholar 

25. Tilman, D. et al. Diversity and productivity in a long-term grassland experiment. Science 294, 843–845 (2001).

    Google Scholar 

26. Loreau, M. & Hector, A. Partitioning selection and complementarity in biodiversity experiments. Nature 412, 72–76 (2001).

    Google Scholar 

27. Chen, C., Xiao, W. & Chen, H. Y. Meta-analysis reveals global variations in plant diversity effects on productivity. Nature 638, 435–440 (2025).

    Google Scholar 

28. Chen, X. & Chen, H. Y. H. Plant diversity loss reduces soil respiration across terrestrial ecosystems. Glob. Change Biol. 25, 1482–1492 (2019).

    Google Scholar 

29. Tilman, D., Wedin, D. & Knops, J. Productivity and sustainability influenced by biodiversity in grassland ecosystems. Nature 379, 718–720 (1996).

    Google Scholar 

30. Duan, P. et al. Tree species diversity increases soil microbial carbon use efficiency in a subtropical forest. Glob. Change Biol. 29, 7131–7144 (2023).

    Google Scholar 

31. Chen, S. et al. Plant diversity enhances productivity and soil carbon storage. Proc. Natl. Acad. Sci. USA 115, 4027–4032 (2018).

    Google Scholar 

32. Chen, X. et al. Tree diversity increases decadal forest soil carbon and nitrogen accrual. Nature 618, 94–101 (2023).

    Google Scholar 

33. Delgado-Baquerizo, M. et al. It is elemental: soil nutrient stoichiometry drives bacterial diversity. Environ. Microbiol. 19, 1176–1188 (2017).

    Google Scholar 

34. Grosso, F., Baath, E. & De Nicola, F. Bacterial and fungal growth on different plant litter in Mediterranean soils: effects of C/N ratio and soil pH. Appl. Soil Ecol. 108, 1–7 (2016).

    Google Scholar 

35. Hu, Y., Zheng, Q., Noll, L., Zhang, S. & Wanek, W. Direct measurement of the in situ decomposition of microbial-derived soil organic matter. Soil Biol. Biochem. 141, 107660 (2020).

    Google Scholar 

36. Liang, C., Amelung, W., Lehmann, J. & Kaestner, M. Quantitative assessment of microbial necromass contribution to soil organic matter. Glob. Change Biol. 25, 3578–3590 (2019).

    Google Scholar 

37. Zak, D. R., Holmes, W. E., White, D. C., Peacock, A. D. & Tilman, D. Plant diversity, soil microbial communities, and ecosystem function: Are there any links? Ecology 84, 2042–2050 (2003).

    Google Scholar 

38. Thakur, M. P. et al. Plant diversity drives soil microbial biomass carbon in grasslands irrespective of global environmental change factors. Glob. Change Biol. 21, 4076–4085 (2015).

    Google Scholar 

39. Wang, J., Zhao, W., Wang, G. & Pereira, P. Afforestation changes the trade-off between soil moisture and plant species diversity in different vegetation zones on the Loess Plateau. Catena 219, 106583 (2022).

    Google Scholar 

40. Chen, Y., Ma, S., Jiang, H., Hu, Y. & Lu, X. Influences of litter diversity and soil moisture on soil microbial communities in decomposing mixed litter of alpine steppe species. Geoderma 377, 114577 (2020).

    Google Scholar 

41. Vogel, A., Eisenhauer, N., Weigelt, A. & Scherer-Lorenzen, M. Plant diversity does not buffer drought effects on early-stage litter mass loss rates and microbial properties. Glob. Change Biol. 19, 2795–2803 (2013).

    Google Scholar 

42. Borer, E. T. et al. Herbivores and nutrients control grassland plant diversity via light limitation. Nature 508, 517–520 (2014).

    Google Scholar 

43. Hautier, Y., Niklaus, P. A. & Hector, A. Competition for light causes plant biodiversity loss after eutrophication. Science 324, 636–638 (2009).

    Google Scholar 

44. Eskelinen, A., Harpole, W. S., Jessen, M.-T., Virtanen, R. & Hautier, Y. Light competition drives herbivore and nutrient effects on plant diversity. Nature 611, 301–305 (2022).

    Google Scholar 

45. Yang, Z., Hautier, Y., Borer, E. T., Zhang, C. & Du, G. Abundance- and functional-based mechanisms of plant diversity loss with fertilization in the presence and absence of herbivores. Oecologia 179, 261–270 (2015).

    Google Scholar 

46. Schmidtke, A., Rottstock, T., Gaedke, U. & Fischer, M. Plant community diversity and composition affect individual plant performance. Oecologia 164, 665–677 (2010).

    Google Scholar 

47. Mao, C. et al. Permafrost nitrogen status and its determinants on the Tibetan Plateau. Glob. Change Biol. 26, 5290–5302 (2020).

    Google Scholar 

48. Kou, D. et al. Progressive nitrogen limitation across the Tibetan alpine permafrost region. Nat. Commun. 11, 3331 (2020).

    Google Scholar 

49. He, H., Zhang, W., Zhang, X., Xie, H. & Zhuang, J. Temporal responses of soil microorganisms to substrate addition as indicated by amino sugar differentiation. Soil Biol. Biochem. 43, 1155–1161 (2011).

    Google Scholar 

50. Ding, X. et al. Warming increases microbial residue contribution to soil organic carbon in an alpine meadow. Soil Biol. Biochem. 135, 13–19 (2019).

    Google Scholar 

51. Zhang, R. et al. Dryness weakens the positive effects of plant and fungal beta diversities on above- and belowground biomass. Glob. Change Biol. 28, 6629–6639 (2022).

    Google Scholar 

52. Pan, J. et al. Biotic factors dominantly determine soil inorganic carbon stock across Tibetan alpine grasslands. Soil 8, 687–698 (2022).

    Google Scholar 

53. Gale, M. R. & Grigal, D. F. Vertical root distributions of northern tree species in relation to successional status. Can. J. For. Res. 17, 829–834 (1987).

    Google Scholar 

54. Yang, Y., Fang, J., Ji, C. & Han, W. Above- and belowground biomass allocation in Tibetan grasslands. J. Veg. Sci. 20, 177–184 (2009).

    Google Scholar 

55. Zhang, X. D. & Amelung, W. Gas chromatographic determination of muramic acid, glucosamine, mannosamine, and galactosamine in soils. Soil Biol. Biochem. 28, 1201–1206 (1996).

    Google Scholar 

56. Yuan, Y. et al. Phosphorus addition decreases microbial residual contribution to soil organic carbon pool in a tropical coastal forest. Glob. Change Biol. 27, 454–466 (2021).

    Google Scholar 

57. Appuhn, A. & Joergensen, R. G. Microbial colonisation of roots as a function of plant species. Soil Biol. Biochem. 38, 1040–1051 (2006).

    Google Scholar 

58. Shao, S. et al. Linkage of microbial residue dynamics with soil organic carbon accumulation during subtropical forest succession. Soil Biol. Biochem. 114, 114–120 (2017).

    Google Scholar 

59. Engelking, B., Flessa, H. & Joergensen, R. G. Shifts in amino sugar and ergosterol contents after addition of sucrose and cellulose to soil. Soil Biol. Biochem. 39, 2111–2118 (2007).

    Google Scholar 

60. Brookes, P. C., Landman, A., Pruden, G. & Jenkinson, D. S. Chloroform fumigation and the release of soil nitrogen-a rapid direct extraction method to measure microbial biomass nitrogen in soil. Soil Biol. Biochem. 17, 837–842 (1985).

    Google Scholar 

61. Lovell, R. D., Jarvis, S. C. & Bardgett, R. D. Soil microbial biomass and activity in long-term grassland-effects of management changes. Soil Biol. Biochem. 27, 969–975 (1995).

    Google Scholar 

62. Tatti, E., McKew, B. A., Whitby, C. & Smith, C. J. Simultaneous DNA-RNA Extraction from Coastal Sediments and Quantification of 16S rRNA Genes and Transcripts by Real-time PCR. J. Vis. Exp. 112, e54067 (2016).

    Google Scholar 

63. Rousk, J. et al. Soil bacterial and fungal communities across a pH gradient in an arable soil. ISME J. 4, 1340–1351 (2010).

    Google Scholar 

64. Bokulich, N. A. et al. Quality-filtering vastly improves diversity estimates from Illumina amplicon sequencing. Nat. Methods 10, 57–59 (2013).

    Google Scholar 

65. Paz-Ferreiro, J., Gasco, G., Gutierrez, B. & Mendez, A. Soil biochemical activities and the geometric mean of enzyme activities after application of sewage sludge and sewage sludge biochar to soil. Biol. Fertil. Soils 48, 511–517 (2012).

    Google Scholar 

66. German, D. P. et al. Optimization of hydrolytic and oxidative enzyme methods for ecosystem studies. Soil Biol. Biochem. 43, 1387–1397 (2011).

    Google Scholar 

67. Ye, C. et al. Reconciling multiple impacts of nitrogen enrichment on soil carbon: plant, microbial and geochemical controls. Ecol. Lett. 21, 1162–1173 (2018).

    Google Scholar 

68. Pinheiro, J. Bates, D., DebRoy, S.S., Sarkar, D. Nlme: Linear and Nonlinear Mixed Effects Models. R package version 3, 1–113 (2013). 31–110.

69. Rosseel, Y. lavaan: an R Package for Structural Equation Modeling. J. Stat. Softw. 48, 1–36 (2012).

    Google Scholar 

70. R Core Team. R: A language and environment for statistical computing. (2021).

71. Yan, Y. et al. Plant diversity is key for microbial necromass carbon accrual in alpine grasslands. figshare. https://doi.org/10.6084/m9.figshare.31708387 (2026).

Download references