Abstract
Nitrogen (N)-fixing species are widely used in forestation and agriculture. The effects of planting N-fixing species on soil organic carbon (SOC) stock, however, remain uncertain, limiting policy development and their application towards a possible climate change mitigation strategy. Here we conduct a global meta-analysis of 385 datapoints from 136 studies comparing SOC stock with planting N-fixing versus non-N-fixing species. Planting N-fixing species increases SOC stock by 16% compared with non-N-fixing species. This SOC increase is closely accompanied by soil N increases, with an average accumulation of 7.8 g of SOC per gram of soil N increase. Climate mediates SOC responses, with greater SOC sequestration observed in drier and warmer regions, particularly in the tropics. We estimate that an additional increase of 0.29–0.75 PgC yr−1 in global SOC stock could be achieved by adopting N-fixing species for forestation, agriculture and regeneration of marginal lands, highlighting their potential for climate change mitigation.
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Data availability
The data used in the meta-analysis are available via figshare at https://doi.org/10.6084/m9.figshare.25458238 (ref. 194). Global maps of planted forests and cropland were obtained from SDPT v.1.0 (https://www.wri.org/research/spatial-database-planted-trees-sdpt-version-10) and GLAD (https://glad.umd.edu/dataset/croplands), respectively. Global maps of SOC and soil N stocks at 0–30 cm were obtained from ISRIC Data Hub (https://data.isric.org). Global maps of MAT and MAP were obtained from WorldClim v.2 (https://worldclim.org). Source data are provided with this paper.
Code availability
The code used in this study is available via figshare at https://doi.org/10.6084/m9.figshare.25458238 (ref. 194).
References
Hulvey, K. B. et al. Benefits of tree mixes in carbon plantings. Nat. Clim. Change 3, 869–874 (2013).
Zhao, J. et al. Global systematic review with meta-analysis reveals yield advantage of legume-based rotations and its drivers. Nat. Commun. 13, 4926 (2022).
Reed, S. C., Cleveland, C. C. & Townsend, A. R. Functional ecology of free-living nitrogen fixation: a contemporary perspective. Annu. Rev. Ecol. Evol. Syst. 42, 489–512 (2011).
Levy-Varon, J. H. et al. Tropical carbon sink accelerated by symbiotic dinitrogen fixation. Nat. Commun. 10, 5637 (2019).
Resh, S. C., Binkley, D. & Parrotta, J. A. Greater soil carbon sequestration under nitrogen-fixing trees compared with Eucalyptus species. Ecosystems 5, 217–231 (2002).
Forrester, D. I., Pares, A., O’Hara, C., Khanna, P. K. & Bauhus, J. Soil organic carbon is increased in mixed-species plantations of Eucalyptus and nitrogen-fixing. Acacia Ecosyst. 16, 123–132 (2013).
Batterman, S. A. et al. Key role of symbiotic dinitrogen fixation in tropical forest secondary succession. Nature 502, 224–227 (2013).
Crews, T. E. & Peoples, M. B. Legume versus fertilizer sources of nitrogen: ecological tradeoffs and human needs. Agric. Ecosyst. Environ. 102, 279–297 (2004).
Erisman, J. W. et al. Consequences of human modification of the global nitrogen cycle. Philos. Trans. R. Soc. B 368, 20130116 (2013).
Niu, S., Song, L., Wang, J., Luo, Y. & Yu, G. Dynamic carbon–nitrogen coupling under global change. Sci. China Life Sci. 66, 771–782 (2023).
Wang, Y. P., Law, R. M. & Pak, B. A global model of carbon, nitrogen and phosphorus cycles for the terrestrial biosphere. Biogeosciences 7, 2261–2282 (2010).
Heimann, M. & Reichstein, M. Terrestrial ecosystem carbon dynamics and climate feedbacks. Nature 451, 289–292 (2008).
Gama-Rodrigues, E. F. et al. Carbon storage in soil size fractions under two cacao agroforestry systems in Bahia, Brazil. Environ. Manag. 45, 274–283 (2010).
Mao, R. et al. Soil microbiological and chemical effects of a nitrogen-fixing shrub in poplar plantations in semi-arid region of Northeast China. Eur. J. Soil Biol. 46, 325–329 (2010).
Wang, Q., Wang, S. & Zhang, J. Assessing the effects of vegetation types on carbon storage fifteen years after reforestation on a Chinese fir site. For. Ecol. Manag. 258, 1437–1441 (2009).
Parenti, A., Zegada-Lizarazu, W., Pagani, E. & Monti, A. Soil organic carbon dynamics in multipurpose cropping systems. Ind. Crops Prod. 187, 115315 (2022).
Wolf, A. A., Funk, J. L. & Menge, D. N. L. The symbionts made me do it: legumes are not hardwired for high nitrogen concentrations but incorporate more nitrogen when inoculated. New Phytol. 213, 690–699 (2017).
Li, Q., Zhou, D., Denton, M. D. & Cong, S. Alfalfa monocultures promote soil organic carbon accumulation to a greater extent than perennial grass monocultures or grass–Alfalfa mixtures. Ecol. Eng. 131, 53–62 (2019).
Thilakarathna, M. S., McElroy, M. S., Chapagain, T., Papadopoulos, Y. A. & Raizada, M. N. Belowground nitrogen transfer from legumes to non-legumes under managed herbaceous cropping systems. A review. Agron. Sustain. Dev. 36, 58 (2016).
Wang, J. et al. Interplanting leguminous shrubs boosts the trophic interactions of soil micro-food web in a karst grassland. Soil Biol. Biochem. 188, 109224 (2024).
Ye, X. et al. Tree species richness and N-fixing tree species enhance the chemical stability of soil organic carbon in subtropical plantations. Soil Biol. Biochem. 174, 108828 (2022).
Kuzyakov, Y., Friedel, J. K. & Stahr, K. Review of mechanisms and quantification of priming effects. Soil Biol. Biochem. 32, 1485–1498 (2000).
Liu, M. et al. C:N stoichiometry of stable and labile organic compounds determine priming patterns. Geoderma 362, 114122 (2020).
Ball, K. R. et al. Soil organic carbon and nitrogen pools are increased by mixed grass and legume cover crops in vineyard agroecosystems: detecting short-term management effects using infrared spectroscopy. Geoderma 379, 114619 (2020).
Menge, D. N. L., Wolf, A. A. & Funk, J. L. Diversity of nitrogen fixation strategies in Mediterranean legumes. Nat. Plants 1, 15064 (2015).
Forrester, D. I., Bauhus, J., Cowie, A. L. & Vanclay, J. K. Mixed-species plantations of Eucalyptus with nitrogen-fixing trees: a review. For. Ecol. Manag. 233, 211–230 (2006).
Michaletz, S. T., Cheng, D., Kerkhoff, A. J. & Enquist, B. J. Convergence of terrestrial plant production across global climate gradients. Nature 512, 39–43 (2014).
Carvalhais, N. et al. Global covariation of carbon turnover times with climate in terrestrial ecosystems. Nature 514, 213–217 (2014).
Binkley, D. in Tree Species Effects on Soils: Implications for Global Change (eds Binkley, D. & Menyailo, O.) 155–164 (Springer, 2005).
McGroddy, M. E., Daufresne, T. & Hedin, L. O. Scaling of C:N:P stoichiometry in forests worldwide: implications of terrestrial redfield-type ratios. Ecology 85, 2390–2401 (2004).
Bytnerowicz, T. A., Akana, P. R., Griffin, K. L. & Menge, D. N. L. Temperature sensitivity of woody nitrogen fixation across species and growing temperatures. Nat. Plants 8, 209–216 (2022).
Cheng, Y., Wang, J., Wang, S., Zhang, J. & Cai, Z. Effects of soil moisture on gross N transformations and N2O emission in acid subtropical forest soils. Biol. Fertil. Soils 50, 1099–1108 (2014).
Dijkstra, F. A. & Cheng, W. Increased soil moisture content increases plant N uptake and the abundance of 15N in plant biomass. Plant Soil 302, 263–271 (2008).
Campo, J. & Merino, A. Variations in soil carbon sequestration and their determinants along a precipitation gradient in seasonally dry tropical forest ecosystems. Glob. Change Biol. 22, 1942–1956 (2016).
Balting, D. F., AghaKouchak, A., Lohmann, G. & Ionita, M. Northern Hemisphere drought risk in a warming climate. npj Clim. Atmos. Sci. 4, 61 (2021).
Jackson, R. B. et al. A global analysis of root distributions for terrestrial biomes. Oecologia 108, 389–411 (1996).
Jackson, R. B. et al. The ecology of soil carbon: pools, vulnerabilities, and biotic and abiotic controls. Annu. Rev. Ecol. Evol. Syst. 48, 419–445 (2017).
Berhongaray, G., Cotrufo, F. M., Janssens, I. A. & Ceulemans, R. Below-ground carbon inputs contribute more than above-ground inputs to soil carbon accrual in a bioenergy poplar plantation. Plant Soil 434, 363–378 (2019).
Zhou, X., Wu, H., Koetz, E., Xu, Z. & Chen, C. Soil labile carbon and nitrogen pools and microbial metabolic diversity under winter crops in an arid environment. Appl. Soil Ecol. 53, 49–55 (2012).
Six, J., Elliott, E. T. & Paustian, K. Soil macroaggregate turnover and microaggregate formation: a mechanism for C sequestration under no-tillage agriculture. Soil Biol. Biochem. 32, 2099–2103 (2000).
Zomer, R. J., Bossio, D. A., Sommer, R. & Verchot, L. V. Global sequestration potential of increased organic carbon in cropland soils. Sci. Rep. 7, 15554 (2017).
Shi, L., Feng, W., Xu, J. & Kuzyakov, Y. Agroforestry systems: meta-analysis of soil carbon stocks, sequestration processes, and future potentials. Land Degrad. Dev. 29, 3886–3897 (2018).
Bastin, J. F. et al. The global tree restoration potential. Science 365, 76–79 (2019).
Friedlingstein, P. et al. Global carbon budget 2024. Earth Syst. Sci. Data 17, 965–1039 (2025).
Parazoo, N. C. et al. Terrestrial gross primary production inferred from satellite fluorescence and vegetation models. Glob. Change Biol. 20, 3103–3121 (2014).
Vitousek, P. M., Porder, S., Houlton, B. Z. & Chadwick, O. A. Terrestrial phosphorus limitation: mechanisms, implications, and nitrogen–phosphorus interactions. Ecol. Appl. 20, 5–15 (2010).
Vitousek, P. M. & Howarth, R. W. Nitrogen limitation on land and in the sea: how can it occur? Biogeochemistry 13, 87–115 (1991).
Liao, W., Menge, D. N. L., Lichstein, J. W. & Ángeles-Pérez, G. Global climate change will increase the abundance of symbiotic nitrogen-fixing trees in much of North America. Glob. Change Biol. 23, 4777–4787 (2017).
Global Forest Resources Assessment 2015 (FAO, 2015).
Ferreira, G. W. D. & Aubrey, D. P. A functional trait framework for integrating nitrogen-fixing cover crops into short-rotation woody crop systems. GCB Bioenergy 15, 663–679 (2023).
Dynarski, K. A., Pett-Ridge, J. C. & Perakis, S. S. Decadal-scale decoupling of soil phosphorus and molybdenum cycles by temperate nitrogen-fixing trees. Biogeochemistry 149, 355–371 (2020).
Li, D., Niu, S. & Luo, Y. Global patterns of the dynamics of soil carbon and nitrogen stocks following afforestation: a meta-analysis. New Phytol. 195, 172–181 (2012).
Feng, J. & Zhu, B. A global meta-analysis of soil respiration and its components in response to phosphorus addition. Soil Biol. Biochem. 135, 38–47 (2019).
Romero-Olivares, A. L., Allison, S. D. & Treseder, K. K. Soil microbes and their response to experimental warming over time: a meta-analysis of field studies. Soil Biol. Biochem. 107, 32–40 (2017).
An, R. et al. Analysis of bacterial community structure and diversity characteristics of mixed forest of Robinia pseudoacacia and Ailanthus altissima and there pure forest in the Yellow River Delta. Acta Ecol. Sin. 39, 7960–7967 (2019).
Andy, O. N. Smallholder agriculture land use impact on soil organic carbon stock in Federal Capital Territory of Nigeria. J. Agric. Environ. Int. Dev. 112, 109–119 (2018).
Arshad, M. A., Soon, Y. K. & Ripmeester, J. A. Quality of soil organic matter and C storage as influenced by cropping systems in northwestern Alberta, Canada. Nutr. Cycl. Agroecosyst. 89, 71–79 (2011).
Aschi, A. et al. Introduction of faba bean in crop rotation: impacts on soil chemical and biological characteristics. Appl. Soil Ecol. 120, 219–228 (2017).
Balota, E. L. & Chaves, J. C. Enzymatic activity and mineralization of carbon and nitrogen in soil cultivated with coffee and green manures. Rev. Bras. Cienc. Solo 34, 1573–1583 (2010).
Balota, E. L., Colozzi, A., Andrade, D. S. & Dick, R. P. Long-term tillage and crop rotation effects on microbial biomass and C and N mineralization in a Brazilian Oxisol. Soil Tillage Res. 77, 137–145 (2004).
Beedy, T. L., Snapp, S. S., Akinnifesi, F. K. & Sileshi, G. W. Impact of Gliricidia sepium intercropping on soil organic matter fractions in a maize-based cropping system. Agric. Ecosyst. Environ. 138, 139–146 (2010).
Benintende, S. M., Benintende, M. C., Sterren, M. A. & De Battista, J. J. Soil microbiological indicators of soil quality in four rice rotations systems. Ecol. Indic. 8, 704–708 (2008).
Bernhard-Reversat, F. Soil nitrogen mineralization under a Eucalyptus plantation and a natural Acacia forest in Senegal. For. Ecol. Manag. 23, 233–244 (1988).
Binkley, D., Sollins, P., Bell, R., Sachs, D. & Myrold, D. Biogeochemistry of adjacent conifer and alder–conifer stands. Ecology 73, 2022–2033 (1992).
Borase, D. N. et al. Long-term impact of grain legumes and nutrient management practices on soil microbial activity and biochemical properties. Arch. Agron. Soil Sci. 67, 2015–2032 (2021).
Broersma, K., Juma, N. G. & Robertson, J. A. Net nitrogen mineralization from a Gray Luvisol under diverse cropping systems in the Peace River region of Alberta. Can. J. Soil Sci. 76, 117–123 (1996).
Cao, B. & Wu, L. Studies on soil enzyme activity and soil nutrient content of mixed stands with Robinia pseudoacacia and Frascinus velutina in coastal saline soil. J. Soil Water Conserv. 22, 128–133 (2008).
Casals, P., Romero, J., Rusch, G. M. & Ibrahim, M. Soil organic C and nutrient contents under trees with different functional characteristics in seasonally dry tropical silvopastures. Plant Soil 374, 643–659 (2014).
Chen, J., Shen, W., Xu, H., Li, Y. & Luo, T. The composition of nitrogen-fixing microorganisms correlates with soil nitrogen content during reforestation: a comparison between legume and non-legume plantations. Front. Microbiol. 10, 508 (2019).
Chen, X., Liu, Q. & Zhang, G. Effects of different crop rotation modes on soil fertility and rice yield in Taihu Region. Jiangsu J. Agric. Sci. 37, 874–883 (2021).
Collins, H. P., Rasmussen, P. E. & Douglas, C. L. Jr Crop rotation and residue management effects on soil carbon and microbial dynamics. Soil Sci. Soc. Am. J. 56, 783–788 (1992).
Conceicao, P. C., Dieckow, J. & Bayer, C. Combined role of no-tillage and cropping systems in soil carbon stocks and stabilization. Soil Tillage Res. 129, 40–47 (2013).
Conrad, K. A., Dalal, R. C., Dalzell, S. A., Allen, D. E. & Menzies, N. W. The sequestration and turnover of soil organic carbon in subtropical leucaena-grass pastures. Agric. Ecosyst. Environ. 248, 38–47 (2017).
Dad, J. M., Dand, S. A. & Pala, N. A. The effect of bi-culture cover crops on soil quality, carbon sequestration, and growth characteristics in apple orchards of North Western Himalayas. Agrofor. Syst. 95, 1745–1758 (2021).
Dai, H., Hu, X., Cao, M., Yang, M. & Wang, J. Effects of intercropping with leguminous crops on tomato yield, soil nutrients and enzyme activity. Acta Pedol. Sin. 52, 911–918 (2015).
Deng, L. et al. Soil physical and chemical properties of six fast-growing tree species plantations after afforestation for six years. Guangxi For. Sci. 48, 371–376 (2019).
Deng, S. & Tabatabai, M. A. Effect of cropping systems on nitrogen mineralization in soils. Biol. Fertil. Soils 31, 211–218 (2000).
Dou, X., Li, F., Cheng, X. & Zhu, P. Soil organic carbon and nitrogen dynamics induced by continuous maize cropping compared to maize–soya bean rotation. Eur. J. Soil Sci. 69, 535–544 (2018).
Feng, H. et al. Soil quality indicators as influenced by 5-year diversified and monoculture cropping systems. J. Agric. Sci. 158, 594–605 (2020).
Gao, D. et al. Ecological stoichiometry characteristics of soil and leaves during the recovery process of typical vegetation on the Loess Plateau. Acta Ecol. Sin. 39, 3622–3630 (2019).
Garcia-Montiel, D. C. & Binkley, D. Effect of Eucalyptus saligna and Albizia falcataria on soil processes and nitrogen supply in Hawaii. Oecologia 113, 547–556 (1998).
Gei, M. & Powers, J. S. Do legumes and non-legumes tree species affect soil properties in unmanaged forests and plantations in Costa Rican dry forests? Soil Biol. Biochem. 57, 264–272 (2013).
Ghimire, R., Bista, P. & Machado, S. Long-term management effects and temperature sensitivity of soil organic carbon in grassland and agricultural soils. Sci. Rep. 9, 12151 (2019).
Ghosh, P. K. et al. Increasing soil organic carbon through crop diversification in cereal–cereal rotations of Indo-Gangetic plain. Proc. Indian Natl Sci. Acad. B 89, 429–440 (2019).
Gijsman, A. J., Oberson, A., Friesen, D. K., Sanz, J. I. & Thomas, R. J. Nutrient cycling through microbial biomass under rice–pasture rotations replacing native savanna. Soil Biol. Biochem. 29, 1433–1441 (1997).
Gitari, H. I. et al. Potato–legume intercropping on a sloping terrain and its effects on soil physico-chemical properties. Plant Soil 438, 447–460 (2019).
Gregorich, E. G., Drury, C. F. & Baldock, J. A. Changes in soil carbon under long-term maize in monoculture and legume-based rotation. Can. J. Soil Sci. 81, 21–31 (2001).
Guan, A., Zhang, Y., Liu, Y., Luo, S. & Wang, J. Effects of reduced nitrogen application and sugarcane-soybean intercropping on carbon balance in sugarcane fields. Chin. J. Eco-Agric. 24, 478–488 (2016).
Gurlevik, N. & Karatepe, Y. Long-term effects of afforestation on soil characteristics and net nitrogen mineralization in sandy soils. Austrian J. For. Sci. 133, 187–202 (2016).
Han, J., Yao, Q., Hong, C. & Lin, W. Study on the growth and ecological benefit of mixed stand of Eucalyptus grandis × E. urophylla and Acacia mangium × A. auriculiformis. Eucalypt. Sci. Technol. 72, 15–18 (2008).
He, B., Jia, L., Jin, D. & Qin, W. Studies on soil fertility change in Acacia mangium plantation in Nanning, Guangxi. Sci. Silvae Sin. 43, 10–16 (2007).
He, H. et al. The structure and diversity of nitrogen functional groups from different cropping systems in Yellow River Delta. Microorganisms 8, 424 (2020).
Higashi, T. et al. Tillage and cover crop species affect soil organic carbon in Andosol, Kanto, Japan. Soil Tillage Res. 138, 64–72 (2014).
Hoogmoed, M., Cunningham, S. C., Baker, P., Beringer, J. & Cavagnaro, T. R. N-fixing trees in restoration plantings: effects on nitrogen supply and soil microbial communities. Soil Biol. Biochem. 77, 203–212 (2014).
Hoogmoed, M., Cunningham, S. C., Baker, P. J., Beringer, J. & Cavagnaro, T. R. Is there more soil carbon under nitrogen-fixing trees than under non-nitrogen-fixing trees in mixed-species restoration plantings? Agric. Ecosyst. Environ. 188, 80–84 (2014).
Hu, B. et al. Comparison of nitrogen nutrition and soil carbon status of afforested stands established in degraded soil of the Loess Plateau, China. For. Ecol. Manag. 389, 46–58 (2017).
Huang, X. et al. Changes of soil microbial biomass carbon and community composition through mixing nitrogen-fixing species with Eucalyptus urophylla in subtropical China. Soil Biol. Biochem. 73, 42–48 (2014).
Huang, X. et al. Microbial community and associated enzymes activity influence soil carbon chemical composition in Eucalyptus urophylla plantation with mixing N2-fixing species in subtropical China. Plant Soil 414, 199–212 (2017).
Jeddi, K., Cortina, J. & Chaieb, M. Acacia salicina, Pinus halepensis and Eucalyptus occidentalis improve soil surface conditions in arid southern Tunisia. J. Arid. Environ. 73, 1005–1013 (2009).
Jia, B., Jia, L., Zhang, Y., Mou, X. & Li, X. Leguminous Caragana korshinskii evidently enhances microbial necromass carbon accumulation in dryland soils. Catena 215, 106342 (2022).
Jing, Y. et al. Effects of N-fixing tree species (Alnus sibirica) on amino sugars in soil aggregates of Larix kaempferi plantation in eastern Liaoning Province, China. Chin. J. Appl. Ecol. 29, 1753–1758 (2018).
Johnson, D. W. Soil properties beneath ceanothus and pine stands in the eastern Sierra Nevada. Soil Sci. Soc. Am. J. 59, 918–924 (1995).
Juhi et al. Crop yields and soil organic matter pools in zero-till direct-seeded rice-based cropping systems as influenced by fertigation levels in the Indo-Gangetic plains in India. Carbon Manag. 13, 78–89 (2022).
Kaonga, M. L. & Bayliss-Smith, T. P. Carbon pools in tree biomass and the soil in improved fallows in eastern Zambia. Agrofor. Syst. 76, 37–51 (2009).
Kaye, J. P., Binkley, D., Zou, X. & Parrotta, J. A. Non-labile soil 15nitrogen retention beneath three tree species in a tropical plantation. Soil Sci. Soc. Am. J. 66, 612–619 (2002).
Kaye, J. P., Resh, S. C., Kaye, M. W. & Chimner, R. A. Nutrient and carbon dynamics in a replacement series of Eucalyptus and Albizia trees. Ecology 81, 3267–3273 (2000).
Kumar, M., Kundu, D. K., Ghorai, A. K., Mitra, S. & Singh, S. R. Carbon and nitrogen mineralization kinetics as influenced by diversified cropping systems and residue incorporation in inceptisols of eastern Indo-Gangetic plain. Soil Tillage Res. 178, 108–117 (2018).
Li, H. et al. Changes in carbon, nutrients and stoichiometric relations under different soil depths, plant tissues and ages in black locust plantations. Acta Physiol. Plant. 35, 2951–2964 (2013).
Li, J., Jiao, S., Gao, R. & Bardgett, R. D. 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. Environ. Manag. 50, 1193–1203 (2012).
Li, Q. et al. Changes in soil organic carbon and total nitrogen stocks along a chronosequence of Caragana intermedia plantations in alpine sandy land. Ecol. Eng. 133, 53–59 (2019).
Li, Z., Wen, H. & Yu, Z. The impact of human activities on the soil nitrogen mineralization in artificial forests. Chin. J. Bot. 12, 142–148 (1995).
Liu, C., Nie, Y., Zhang, Y., Tang, J. & Siddique, K. H. M. Introduction of a leguminous shrub to a rubber plantation changed the soil carbon and nitrogen fractions and ameliorated soil environments. Sci. Rep. 8, 17324 (2018).
Liu, L. et al. Effect of monospecific and mixed Cunninghamia lanceolata plantations on microbial community and two functional genes involved in nitrogen cycling. Plant Soil 327, 413–428 (2010).
Lopez-Bellido, L., Lopez-Garrido, F. J., Fuentes, M., Castillo, J. E. & Fernandez, E. J. Influence of tillage, crop rotation and nitrogen fertilization on soil organic matter and nitrogen under rain-fed Mediterranean conditions. Soil Tillage Res. 43, 277–293 (1997).
Ma, H. et al. Intercropping improves soil ecosystem multifunctionality through enhanced available nutrients but depends on regional factors. Plant Soil 480, 71–84 (2022).
Ma, J. & Li, K. Effects of plantation on soil nutrient and evaluation in Yuanmou dry-hot valley. For. Res. 19, 467–471 (2006).
Macedo, M. O. et al. Changes in soil C and N stocks and nutrient dynamics 13 years after recovery of degraded land using leguminous nitrogen-fixing trees. For. Ecol. Manag. 255, 1516–1524 (2008).
Matusso, J. M. M., Mugwe, J. N. & Mucheru-Muna, M. Effects of different maize (Zea mays L.)–soybean (Glycine max (L.) Merrill) intercropping patterns on soil mineral-N, N-uptake and soil properties. Afr. J. Agric. Res. 9, 42–55 (2014).
McDaniel, M. D. & Grandy, A. S. Soil microbial biomass and function are altered by 12 years of crop rotation. Soil 2, 583–599 (2016).
McDaniel, M. D., Grandy, A. S., Tiemann, L. K. & Weintraub, M. N. Crop rotation complexity regulates the decomposition of high and low quality residues. Soil Biol. Biochem. 78, 243–254 (2014).
Mo, Q. et al. Reforestation in southern China: revisiting soil N mineralization and nitrification after 8 years restoration. Sci. Rep. 6, 19770 (2016).
Montagnini, F. & Sancho, F. Net nitrogen mineralization in soils under six indigenous tree species, an abandoned pasture and a secondary forest in the Atlantic lowlands of Costa Rica. Plant Soil 162, 117–124 (1994).
Nath, C. P. et al. Including grain legume in rice–wheat cropping system improves soil organic carbon pools over time. Ecol. Eng. 129, 144–153 (2019).
Novara, A. et al. Cover crop impact on soil organic carbon, nitrogen dynamics and microbial diversity in a Mediterranean semiarid vineyard. Sustainability 12, 3256 (2020).
Omay, A. B., Rice, C. W., Maddux, L. D. & Gordon, W. B. Changes in soil microbial and chemical properties under long-term crop rotation and fertilization. Soil Sci. Soc. Am. J. 61, 1672–1678 (1997).
Pereira, E. L., Santos, S. A. P., Arrobas, M. & Patricio, M. S. Microbial biomass and N mineralization in mixed plantations of broadleaves and nitrogen-fixing species. For. Syst. 20, 516–524 (2011).
Qiu, L., Zhang, X., Cheng, J. & Yin, X. Effects of black locust (Robinia pseudoacacia) on soil properties in the loessial gully region of the Loess Plateau, China. Plant Soil 332, 207–217 (2010).
Radrizzani, A., Shelton, H. M., Dalzell, S. A. & Kirchhof, G. Soil organic carbon and total nitrogen under Leucaena leucocephala pastures in Queensland. Crop Pasture Sci. 62, 337–345 (2011).
Regehr, A., Oelbermann, M., Videla, C. & Echarte, L. Gross nitrogen mineralization and immobilization in temperate maize–soybean intercrops. Plant Soil 391, 353–365 (2015).
Robles, M. D. & Burke, I. C. Legume, grass, and conservation reserve program effects on soil organic matter recovery. Ecol. Appl. 7, 345–357 (1997).
Rothe, A., Cromack, K., Resh, S. C., Makineci, E. & Son, Y. Soil carbon and nitrogen changes under Douglas-fir with and without red alder. Soil Sci. Soc. Am. J. 66, 1988–1995 (2002).
Rui, Y. et al. Persistent soil carbon enhanced in mollisols by well-managed grasslands but not annual grain or dairy forage cropping systems. Proc. Natl Acad. Sci. USA 119, e2118931119 (2022).
Russell, A. E., Raich, J. W., Valverde-Barrantes, O. J. & Fisher, R. F. Tree species effects on soil properties in experimental plantations in tropical moist forest. Soil Sci. Soc. Am. J. 71, 1389–1397 (2007).
Sainju, U. M. et al. Cover crop effect on soil carbon fractions under conservation tillage cotton. Soil Tillage Res. 96, 205–218 (2007).
Santos, E. R. S. et al. Particulate soil organic matter in bahiagrass–rhizoma peanut mixtures and their monocultures. Soil Sci. Soc. Am. J. 83, 658–665 (2019).
Scalise, A. et al. Legume–barley intercropping stimulates soil N supply and crop yield in the succeeding durum wheat in a rotation under rainfed conditions. Soil Biol. Biochem. 89, 150–161 (2015).
Singh, A. N., Raghubanshi, A. S. & Singh, J. S. Impact of native tree plantations on mine spoil in a dry tropical environment. For. Ecol. Manag. 187, 49–60 (2004).
Tao, J. et al. Impact of intercropping and arbuscular mycorrhizal fungi on soil fertility and corn yield in a newly cultivated mountain land. Acta Agr. Zhejiang. 32, 115–123 (2020).
Tayir, M. et al. Distinct leaf functional traits of Tamarix chinensis at different habitats in the hinterland of the Taklimakan desert. Front. Plant Sci. 13, 1094049 (2023).
TerAvest, D. et al. Soil carbon pools, nitrogen supply, and tree performance under several groundcovers and compost rates in a newly planted apple orchard. Hortscience 46, 1687–1694 (2011).
Tian, Y., Liu, J., Zhang, X. & Gao, L. Effects of summer catch crop, residue management, soil temperature and water on the succeeding cucumber rhizosphere nitrogen mineralization in intensive production systems. Nutr. Cycl. Agroecosyst. 88, 429–446 (2010).
Tiemann, L. K., Grandy, A. S., Atkinson, E. E., Marin-Spiotta, E. & McDaniel, M. D. Crop rotational diversity enhances belowground communities and functions in an agroecosystem. Ecol. Lett. 18, 761–771 (2015).
van der Pol, L. K. et al. Addressing the soil carbon dilemma: legumes in intensified rotations regenerate soil carbon while maintaining yields in semi-arid dryland wheat farms. Agric. Ecosyst. Environ. 330, 107906 (2022).
Veloso, M. G. et al. High carbon storage in a previously degraded subtropical soil under no-tillage with legume cover crops. Agric. Ecosyst. Environ. 268, 15–23 (2018).
Venkatesh, M. S., Hazra, K. K., Ghosh, P. K., Praharaj, C. S. & Kumar, N. Long-term effect of pulses and nutrient management on soil carbon sequestration in Indo-Gangetic plains of India. Can. J. Soil Sci. 93, 127–136 (2013).
Voigtlaender, M. et al. Introducing Acacia mangium trees in Eucalyptus grandis plantations: consequences for soil organic matter stocks and nitrogen mineralization. Plant Soil 352, 99–111 (2012).
Wang, B. et al. Changes in soil nutrient and enzyme activities under different vegetations in the Loess Plateau area, Northwest China. Catena 92, 186–195 (2012).
Wang, F. et al. Effects of nitrogen-fixing and non-nitrogen-fixing tree species on soil properties and nitrogen transformation during forest restoration in southern China. Soil Sci. Plant Nutr. 56, 297–306 (2010).
Wang, F. & Wu, S. Effects of afforestation model on soil and microorganism stoichiometric characteristics. Shaanxi For. Sci. Technol. 48, 9–13 (2020).
Wang, F., Zhu, W., Xia, H., Fu, S. & Li, Z. Nitrogen mineralization and leaching in the early stages of a subtropical reforestation in southern China. Restor. Ecol. 18, 313–322 (2010).
Wang, P., Wang, Y. & Wu, Q. S. Effects of soil tillage and planting grass on arbuscular mycorrhizal fungal propagules and soil properties in citrus orchards in southeast China. Soil Tillage Res. 155, 54–61 (2016).
Wang, Q., Xia, J., Zhang, J., Zhang, J. & Liu, J. Soil enzyme activities and nutrient characteristics of different improving patterns in the Yellow River Delta Area. J. Soil Water Conserv. 26, 133–137 (2012).
Wang, S., Zhang, W. & Sanchez, F. Relating net primary productivity to soil organic matter decomposition rates in pure and mixed Chinese fir plantations. Plant Soil 334, 501–510 (2010).
Wang, W. et al. Effects of labile carbon and phosphorus addition on N transformations with N- vs. non-N-fixing tree plantations. Ecosphere 9, e02165 (2018).
Wang, X., Tang, C., Severi, J., Butterly, C. R. & Baldock, J. A. Rhizosphere priming effect on soil organic carbon decomposition under plant species differing in soil acidification and root exudation. New Phytol. 211, 864–873 (2016).
Wani, S. P., Rego, T. J., Rajeswari, S. & Lee, K. K. Effect of legume-based cropping systems on nitrogen mineralization potential of vertisol. Plant Soil 175, 265–274 (1995).
Wei, Z., Fan, J., Pan, W., Chen, G. & Liu, H. Soil amelioration effects of mixed plantations of Robinia Pseudoacacia and Populus Liaoningensis in saline-alkali soils along muddy seacoast. Bull. Soil Water Conserv. 32, 106–110,170 (2012).
Wu, P. & Xue, J. Effects of three different plantations on soil physicochemical and microbial characteristics in Krast region. J. Nanjing For. Univ. 39, 67–72 (2015).
Wu, Z., Wu, Y., Song, X., Peng, X. & Yang, L. Research on the mixed forest of Pinus tabulaeformis and Robinia pseudoacacia in the dry valley of the upper Minjiang river. J. Sichuan For. Sci. Technol. 35, 17–21 (2014).
Xiao, W. et al. Study of biomass and soil impact of Eucalyptus with Acacia crassicarpa mixed forest. Guangdong For. Sci. Technol. 15, 8–15 (1999).
Xue, L. et al. Effect of different mixed proportion on soil quantity in mixed Robinia pseudoacacia and Pinus tabulaeformis plantations. Sci. Soil Water Conserv. 17, 87–93 (2019).
Xue, S., Liu, G., Dai, Q., Dang, X. & Zhou, P. Effect of different vegetation restoration models on soil microbial biomass in eroded hilly loess plateau. J. Nat. Resour. 22, 20–27 (2007).
Yan, Z. et al. Diversified cropping systems benefit soil carbon and nitrogen stocks by increasing aggregate stability: results of three fractionation methods. Sci. Total Environ. 824, 153878 (2022).
Yao, Y., Zhao, Z., Wei, X. & Shao, M. Effects of shrub species on soil nitrogen mineralization in the desert–loess transition zone. Catena 173, 330–338 (2019).
Yao, Z. et al. Dynamics and sequestration potential of soil organic carbon and total nitrogen stocks of leguminous green manure-based cropping systems on the Loess Plateau of China. Soil Tillage Res. 191, 108–116 (2019).
Yuan, Z. et al. Effects of legume species introduction on vegetation and soil nutrient development on abandoned croplands in a semi-arid environment on the Loess Plateau, China. Sci. Total Environ. 541, 692–700 (2016).
Yuan, Z. et al. Medicago sativa improves soil carbon sequestration following revegetation of degraded arable land in a semi-arid environment on the Loess Plateau, China. Agric. Ecosyst. Environ. 232, 93–100 (2016).
Zavyalova, N. E., Vasbieva, M. T. & Fomin, D. S. Microbial biomass, respiratory activity and nitrogen fixation in soddy-podzolic soils of the pre-urals area under various agricultural uses. Eurasia. Soil Sci. 53, 383–388 (2020).
Zeng, Q., Li, X., Dong, Y., Li, Y. & An, S. Soil microbial biomass nitrogen and carbon, water soluble nitrogen and carbon under different arbors forests on the Loess Plateau. Acta Ecol. Sin. 35, 3598–3605 (2015).
Zhang, D. et al. Building up the soil carbon pool via the cultivation of green manure crops in the Loess Plateau of China. Geoderma 337, 425–433 (2019).
Zhang, J. et al. Understory management and fertilization affected soil greenhouse gas emissions and labile organic carbon pools in a Chinese chestnut plantation. For. Ecol. Manag. 337, 126–134 (2015).
Zhang, J. et al. Short-term effects on the heterogeneity of soil nutrients in mixed poplar–alder plantation. Ecol. Environ. Sci. 27, 216–223 (2018).
Zhang, Y., Dong, X., Xue, L., Xiao, L. & Zhu, S. Effect of young Tephrosia candida and Leucaena glauca plantations on soil fertility. Hunan For. Sci. Technol. 42, 6–9 (2015).
Zhang, Y. et al. Effect of long-term tillage and cropping system on portion of fungal and bacterial necromass carbon in soil organic carbon. Soil Tillage Res. 218, 105307 (2022).
Zhao, S., Shi, S., Chen, J., Chen, X. & He, J. Effects of different rotation patterns on soil carbon and nitrogen contents and enzyme activities in the arid region of central Gansu. Acta Agrestia Sin. 27, 817–824 (2019).
Zhao, W. et al. Effects of tree species transformation on soil carbon, nitrogen, phosphorus and stoichiometry in mountainous areas of southern Fujian. Acta Agric. Univ. Jiangxiensis 44, 299–310 (2022).
Zhao, Y., Ji, T., Zhang, D. & Zhang, J. Soil physi-chemical properties, biomass and nutrient contents of forest litter in mixed plantations of Eucalyptus grandis and three tree species. Chin. J. Appl. Environ. Biol. 21, 948–953 (2015).
Zhou, L. et al. Characteristics of soil nutrient and enzyme activities in plantations of Eucalyptus urophylla × E. grandis and five Acacia species. J. Trop. Subtrop. Bot. 29, 483–493 (2021).
Zhou, X. et al. Symbiotic nitrogen fixation and soil N availability under legume crops in an arid environment. J. Soils Sediment. 11, 762–770 (2011).
Zhu, X. et al. Effects of nitrogen addition on litter decomposition and nutrient release in two tropical plantations with N2-fixing vs. non-N2-fixing tree species. Plant Soil 399, 61–74 (2016).
Zou, L. Study on productivity and soil physical and chemical property of mixed stand of Pinus massoniana with Acacia dealbata link. Prot. For. Sci. Technol. 5, 5–7 (2006).
Muggeo, V. M. R. Segmented: an R package to fit regression models with broken-line relationships. R News 8, 20–25 (2008).
Lin, Y. S. et al. Optimal stomatal behaviour around the world. Nat. Clim. Change 5, 459–464 (2015).
Tedersoo, L. et al. Global database of plants with root-symbiotic nitrogen fixation: NodDB. J. Veg. Sci. 29, 560–568 (2018).
R Core Team. A Language and Environment for Statistical Computing (R Foundation for Statistical Computing, 2023).
Hedges, L. V., Gurevitch, J. & Curtis, P. S. The meta-analysis of response ratios in experimental ecology. Ecology 80, 1150–1156 (1999).
Viechtbauer, W. Conducting meta-analyses in R with the metafor package. J. Stat. Softw. 36, 1–48 (2010).
Egger, M., Smith, G. D., Schneider, M., & Minder, C. Bias in meta-analysis detected by a simple, graphical test. Br. Med. J. 315, 629 (1997).
Chen, J. et al. A keystone microbial enzyme for nitrogen control of soil carbon storage. Sci. Adv. 4, eaaq1689 (2018).
Calcagno, V. & De Mazancourt, C. glmulti: an R package for easy automated model selection with (generalized) linear models. J. Stat. Softw. 34, 1–29 (2010).
Terrer, C. et al. Nitrogen and phosphorus constrain the CO2 fertilization of global plant biomass. Nat. Clim. Change 9, 684–689 (2019).
Hijmans, R. J. et al. raster: Geographic data analysis and modeling. R package version 3.6-26 (2023).
Global Forest Resources Assessment 2020—Key Findings (FAO, 2020).
Sun, X. & Chen, H. Meta-analysis shows that planting nitrogen-fixing species increases soil organic carbon stock. figshare https://doi.org/10.6084/m9.figshare.25458238 (2025).
Acknowledgements
We sincerely thank X. Chen and Z. Ma for their suggestions on data analysis. This work was supported by Guangdong Basic and Applied Basic Research Foundation (2025A1515011004), Shenzhen Science and Technology Program (JCYJ20220530150015035), National Natural Science Foundation of China (42367035, 32471685, 42361144886) and the Shaanxi Province Natural Science Foundation for Distinguished Young Scholar (2024JC-JCQN-32).
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X. S. and H. C. designed research and compiled and analysed data. X. S. wrote the initial draft with significant contribution from H. C. and J. C. Y.K., Y.Y., G.W.D.F., R.O.-H., C.W.M., Z.W., Y.H. and D.L. contributed to the revision of the paper.
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A list of identified invasive N-fixing species in different climate zones.
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Sun, X., Chen, J., Kuzyakov, Y. et al. Meta-analysis shows that planting nitrogen-fixing species increases soil organic carbon stock. Nat Ecol Evol (2025). https://doi.org/10.1038/s41559-025-02861-x
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DOI: https://doi.org/10.1038/s41559-025-02861-x