Scaling and stoichiometry of seed nitrogen and phosphorus across Chinese grasslands

doi:10.1016/j.pld.2026.03.006

Plant Diversity ›› 2026, Vol. 48 ›› Issue (04): 782-790.DOI: 10.1016/j.pld.2026.03.006

• Articles • Previous Articles Next Articles

Scaling and stoichiometry of seed nitrogen and phosphorus across Chinese grasslands

Nan Hu^a, Li Zhang^b, Sean T. Michaletz^c,d, Milos Simovic^c,d, Josep Peñuelas^e,f, Jordi Sardans^e,f, Haiyang Gong^a, Haiyan Bu^g, Zhiqiang Wang^a

a. Sichuan Zoige Alpine Wetland Ecosystem National Observation and Research Station, College of Grassland Resources, Southwest Minzu University, Chengdu 610041, China;
b. Co-Innovation Center for Sustainable Forestry in Southern China, Bamboo Research Institute, Nanjing Forestry University, Nanjing 210037, China;
c. Department of Botany, The University of British Columbia, Vancouver, British Columbia, V6T 1Z4, Canada;
d. Biodiversity Research Centre, The University of British Columbia, Vancouver, British Columbia, V6T 1Z4, Canada;
e. CSIC, Global Ecology Unit, CREAF-CSIC-UAB, Bellaterra, Catalonia 08193, Spain;
f. CREAF, Cerdanyola del Vallès, Catalonia 08193, Spain;
g. State Key Laboratory of Herbage Improvement and Grassland Agro-ecosystems, College of Ecology, Lanzhou University, Lanzhou 730000, China

Received:2025-08-31 Revised:2026-03-04 Accepted:2026-03-06 Online:2026-03-12 Published:2026-07-25
Contact: Haiyan Bu,E-mail:bohy@lzu.edu.cn;Zhiqiang Wang,E-mail:wangzq@swun.edu.cn
Supported by:
We thank Xiang Liu for assistance in the field sampling. The determination of nitrogen and phosphorus contents in seed were performed by BaiHui Biotechnology Co., Ltd at Chengdu, China. This study was financially supported by the Southwest Minzu University Research Startup Funds (RQD2024017), the National Key Research and Development Program of China (Grant No. 2023YFF1304404), the National Natural Science Foundation of China (32171518), and the Sichuan Science and Technology Program (2024NSFSC0106).

Scaling and stoichiometry of seed nitrogen and phosphorus across Chinese grasslands

Nan Hu^a, Li Zhang^b, Sean T. Michaletz^c,d, Milos Simovic^c,d, Josep Peñuelas^e,f, Jordi Sardans^e,f, Haiyang Gong^a, Haiyan Bu^g, Zhiqiang Wang^a

a. Sichuan Zoige Alpine Wetland Ecosystem National Observation and Research Station, College of Grassland Resources, Southwest Minzu University, Chengdu 610041, China;
b. Co-Innovation Center for Sustainable Forestry in Southern China, Bamboo Research Institute, Nanjing Forestry University, Nanjing 210037, China;
c. Department of Botany, The University of British Columbia, Vancouver, British Columbia, V6T 1Z4, Canada;
d. Biodiversity Research Centre, The University of British Columbia, Vancouver, British Columbia, V6T 1Z4, Canada;
e. CSIC, Global Ecology Unit, CREAF-CSIC-UAB, Bellaterra, Catalonia 08193, Spain;
f. CREAF, Cerdanyola del Vallès, Catalonia 08193, Spain;
g. State Key Laboratory of Herbage Improvement and Grassland Agro-ecosystems, College of Ecology, Lanzhou University, Lanzhou 730000, China

通讯作者: Haiyan Bu,E-mail:bohy@lzu.edu.cn;Zhiqiang Wang,E-mail:wangzq@swun.edu.cn
基金资助:
We thank Xiang Liu for assistance in the field sampling. The determination of nitrogen and phosphorus contents in seed were performed by BaiHui Biotechnology Co., Ltd at Chengdu, China. This study was financially supported by the Southwest Minzu University Research Startup Funds (RQD2024017), the National Key Research and Development Program of China (Grant No. 2023YFF1304404), the National Natural Science Foundation of China (32171518), and the Sichuan Science and Technology Program (2024NSFSC0106).

Abstract

Abstract: Nitrogen (N) and phosphorus (P) play crucial roles in early seedling establishment, influencing germination, survival, and overall performance. However, large-scale multispecies seed element survey data are extremely scarce, and studies on the N, P concentrations and N:P ratios as well as scaling relationships of N and P in seeds remain limited. To examine the N versus P scaling relationship and N:P stoichiometry in seeds, we conducted a large-scale survey and collected 1652 seed samples from 446 species in 51 families spanning 236 sites across the Chinese grasslands, including typical steppe, desert steppe, and alpine meadow. We found that the arithmetic means of seed N and P concentrations and N:P ratios were 28.95 ± 0.32 mg/g, 4.26 ± 0.05 mg/g, and 7.24 ± 0.07, respectively. Seed N, P concentrations and N:P ratios varied across different plant groups and grassland types. For example, N-fixers had the highest N, P concentrations and N:P ratios across plant groups. Among the grassland types, alpine meadow displayed the highest N and P concentrations but the lowest N:P ratios. Using a recent bootstrapping approach to resolve data imbalance, the overall seed N vs. P scaling exponent was 0.75, supporting the growth rate hypothesis. We also detected strong phylogenetic signals in these traits. After accounting for phylogeny, the scaling exponent in independently evolving lineages was 0.66. These findings highlight how both environment and evolutionary history regulate seed nutrient allocation, advancing our understanding of plant reproductive strategies and providing key parameters for modelling early plant growth.

Key words: Grasslands, Nitrogen, Phosphorus, Scaling exponent, Seed

摘要： Nitrogen (N) and phosphorus (P) play crucial roles in early seedling establishment, influencing germination, survival, and overall performance. However, large-scale multispecies seed element survey data are extremely scarce, and studies on the N, P concentrations and N:P ratios as well as scaling relationships of N and P in seeds remain limited. To examine the N versus P scaling relationship and N:P stoichiometry in seeds, we conducted a large-scale survey and collected 1652 seed samples from 446 species in 51 families spanning 236 sites across the Chinese grasslands, including typical steppe, desert steppe, and alpine meadow. We found that the arithmetic means of seed N and P concentrations and N:P ratios were 28.95 ± 0.32 mg/g, 4.26 ± 0.05 mg/g, and 7.24 ± 0.07, respectively. Seed N, P concentrations and N:P ratios varied across different plant groups and grassland types. For example, N-fixers had the highest N, P concentrations and N:P ratios across plant groups. Among the grassland types, alpine meadow displayed the highest N and P concentrations but the lowest N:P ratios. Using a recent bootstrapping approach to resolve data imbalance, the overall seed N vs. P scaling exponent was 0.75, supporting the growth rate hypothesis. We also detected strong phylogenetic signals in these traits. After accounting for phylogeny, the scaling exponent in independently evolving lineages was 0.66. These findings highlight how both environment and evolutionary history regulate seed nutrient allocation, advancing our understanding of plant reproductive strategies and providing key parameters for modelling early plant growth.

关键词: Grasslands, Nitrogen, Phosphorus, Scaling exponent, Seed

Nan Hu, Li Zhang, Sean T. Michaletz, Milos Simovic, Josep Peñuelas, Jordi Sardans, Haiyang Gong, Haiyan Bu, Zhiqiang Wang. Scaling and stoichiometry of seed nitrogen and phosphorus across Chinese grasslands[J]. Plant Diversity, 2026, 48(04): 782-790.

References

[1] Adler, P.B., Salguero-Gomez, R., Compagnoni, A., et al., 2014. Functional traits explain variation in plant life history strategies. Proc. Natl. Acad. Sci. U.S.A., 111, 740-745. https://doi.org/10.1073/pnas.1315179111.
[2] Aerts, R., Berendse, F., 1989. Above-ground nutrient turnover and net primary production of an evergreen and a deciduous species in a heathland ecosystem. J. Ecol. 77, 343-356. https://doi.org/10.2307/2260754.
[3] Aerts, R., 1996. Nutrient resorption from senescing leaves of perennials: are there general patterns? J. Ecol. 84, 597-608. https://doi.org/10.2307/2261481.
[4] Aerts, R., Chapin, F.S. III, 2000. The mineral nutrition of wild plants revisited: a re-evaluation of processes and patterns. Adv. Ecol. Res. 30, 1-67. https://doi.org/10.1016/S0065-2504(08)60016-1.
[5] Agren, G.I., 2004. The C: N : P stoichiometry of autotrophs-theory and observations. Ecol. Lett. 7, 185-191. https://doi.org/10.1111/j.1461-0248.2004.00567.x.
[6] Bai, Y., Wu, J., Clark, C.M., et al., 2010. Tradeoffs and thresholds in the effects of nitrogen addition on biodiversity and ecosystem functioning: Evidence from inner Mongolia Grasslands. Glob. Change Biol. 16, 358-372. https://doi.org/10.1111/j.1365-2486.2009.01950.x.
[7] Bewley, J.D., Bradford, K.J., Hilhorst, H.W.M., et al., 2013. Seeds: Physiology of Development, Germination and Dormancy, 3rd Edn. New York, NY: Springer.
[8] Blomberg, S.P., Garland, T.J., Ives, A.R., 2003. Testing for phylogenetic signal in comparative data: behavioral traits are more labile. Evolution. 57, 717-745. https://doi.org/10.1111/j.0014-3820.2003.tb00285.x.
[9] Bonfil, C., 1998. The effects of seed size, cotyledon reserves, and herbivory on seedling survival and growth in Quercus rugosa and Q. laurina (Fagaceae). Am. J. Bot. 85, 79-87. https://doi.org/10.2307/2446557.
[10] Broadley, M.R., Bowen, H.C., Cotterill, H.L., et al., 2004. Phylogenetic variation in the shoot mineral concentration of angiosperms. J. Exp. Bot. 55, 321-336. https://doi.org/10.1093/jxb/erh002.
[11] Bu, H.Y., Du, G.Z., Chen, X.L., et al., 2008. Community-wide germination strategies in an alpine meadow on the eastern Qinghai-Tibetan Plateau: phylogenetic and life-history correlates. Plant Ecol. 195, 87-98. https://doi.org/10.1007/s11258-007-9301-1.
[12] Bu, H.Y., Zhang, Y.M., Zhao, D., et al., 2019. The evolutionary correlation associated with seed mass and altitude on nutrient allocation of seeds. Seed Sci. Res. 29, 38-43. https://doi.org/10.1017/S0960258518000387.
[13] Chen, S., Poschlod, P., Antonelli, A., et al., 2020. Trade-off between seed dispersal in space and time. Ecol. Lett. 23, 1635-1642. https://doi.org/10.1111/ele.13595.
[14] Cheng, J., Cheng, X., Wang, L., et al., 2015. Physiological characteristics of seed reserve utilization during the early seedling growth in rice. Braz. J. Bot. 38, 751-759. https://doi.org/10.1007/s40415-015-0213-4.
[15] Cochrane, A., Yates, C.J., Hoyle, G.L., et al., 2015. Will among-population variation in seed traits improve the chance of species persistence under climate change? Glob. Ecol. Biogeogr. 24, 12-24. https://doi.org/10.1111/geb.12234.
[16] De Frenne, P., Kolb, A., Graae, B.J., et al., 2011. A latitudinal gradient in seed nutrients of the forest herb Anemone nemorosa. Plant Biol. 13, 493-501. https://doi.org/10.1111/j.1438-8677.2010.00404.x.
[17] De Menezes, L.C.C.R., Klein, J., Kestring, D., et al., 2010. Bottom-up and top-down effects in a pre-dispersal seed predation system: Are non-predated seeds damaged? Basic Appl. Ecol. 11, 126-134. https://doi.org/10.1016/j.baae.2009.11.004.
[18] Delatte, E., Chabrerie, O., 2008. Performances des plantes herbacees forestieres dans la dispersion de leurs graines par la fourmi Myrmica ruginodis. C. R. Biol. 331, 309-320. https://doi.org/10.1016/j.crvi.2008.02.002.
[19] Elser, J.J., Sterner, R.W., Gorokhova, E., et al., 2000. Biological stoichiometry from genes to ecosystems. Ecol. Lett. 3, 540-550. https://doi.org/10.1111/j.1461-0248.2000.00185.x.
[20] Elser, J.J., Acharya, K., Kyle, M., et al., 2003. Growth rate-stoichiometry couplings in diverse biota. Ecol. Lett. 6, 936-943. https://doi.org/10.1046/j.1461-0248.2003.00518.x.
[21] Elser, J.J., Fagan, W.F., Kerkhoff, A.J., et al., 2010. Biological stoichiometry of plant production: Metabolism, scaling and ecological response to global change. New Phytol. 186, 593-608. https://doi.org/10.1111/j.1469-8137.2010.03214.x.
[22] Felsenstein, J., 1985. Phylogenies and the comparative method. Am. Nat. 125, 1-15. https://doi.org/10.1086/284325.
[23] Geng, Y., Wang, L., Jin, D., et al., 2014. Alpine climate alters the relationships between leaf and root morphological traits but not chemical traits. Oecologia. 175, 445-455. https://doi.org/10.1007/s00442-014-2919-5.
[24] Ghimire, B., Riley, W.J., Koven, C.D., et al., 2017. A global trait-based approach to estimate leaf nitrogen functional allocation from observations. Ecol. Appl. 27, 1421-1434. https://doi.org/10.1002/eap.1542.
[25] Gong, H.Y., Niu, Y., Niklas, K.J., et al., 2024a. Nitrogen and phosphorus allocation in bark across diverse tree species. Sci. Total Environ. 908, 168327. https://doi.org/10.1016/j.scitotenv.2023.168327.
[26] Gong, H.Y., Sardans, J., Huang, H., et al., 2024b. Global patterns and controlling factors of tree bark C:N:P stoichiometry in forest ecosystems consistent with biogeochemical niche hypothesis. New Phytol. 244, 1303-1314. https://doi.org/10.1111/nph.20119.
[27] Gusewell, S., 2004. N:P ratios in terrestrial plants: variation and functional significance. New Phytol. 164, 243-266. https://doi.org/10.1111/j.1469-8137.2004.01192.x.
[28] Han, W.X., Fang, J.Y., Guo, D.L., et al., 2005. Leaf nitrogen and phosphorus stoichiometry across 753 terrestrial plant species in China. New Phytol. 168, 377-385. https://doi.org/10.1111/j.1469-8137.2005.01530.x.
[29] He, J.S., Wang, L., Flynn, D.F.B., et al., 2008. Leaf nitrogen:phosphorus stoichiometry across Chinese grassland biomes. Oecologia. 155, 301-310. https://doi.org/10.1007/s00442-007-0912-y.
[30] He, J.S., Wang, Z.H., Wang, X.P., et al., 2006. A test of the generality of leaf trait relationships on the Tibetan Plateau. New Phytol. 170, 835-848. https://doi.org/10.1111/j.1469-8137.2006.01704.x.
[31] Huang, H., Ran, J.Z., Ji, M.F., et al., 2020a. Water content quantitatively affects metabolic rates over the course of plant ontogeny. New Phytol. 228, 1524-1534. https://doi.org/10.1111/nph.16808.
[32] Huang, H., Ran, J.Z., Li, X.W., et al., 2020b. A general model for seed and seedling respiratory metabolism. Am. Nat. 195, 534-546. https://doi.org/10.1086/707072.
[33] Jimenez-Alfaro, B., Silveira, F.A.O., Fidelis, A., et al., 2016. Seed germination traits can contribute better to plant community ecology. J. Veg. Sci. 27, 637-645. https://doi.org/10.1111/jvs.12375.
[34] Jin, Y., Qian, H., 2019. V.PhyloMaker: an R package that can generate very large phylogenies for vascular plants. Ecography. 42, 1353-1359. https://doi.org/10.1111/ecog.04434.
[35] Kang, L., Han, X., Zhang, Z., et al., 2007. Grassland ecosystems in China: review of current knowledge and research advancement. Philos. Trans. R. Soc. B. 362, 997-1008. https://doi.org/10.1098/rstb.2007.2029.
[36] Kerkhoff, A.J., Fagan, W.F., Elser, J.J., et al., 2006. Phylogenetic and Growth Form Variation in the Scaling of Nitrogen and Phosphorus in the Seed Plants. Am. Nat. 168, E103-E122. https://doi.org/10.1086/507879.
[37] Koerselman, W., Meuleman, A.F.M., 1996. The vegetation N:P ratio: a new tool to detect the nature of nutrient limitation. J. Appl. Ecol. 33, 1441-1450. https://doi.org/10.2307/2404783.
[38] Korner, C., 1989. The nutritional status of plants from high altitudes: A worldwide comparison. Oecologia. 81, 379-391. https://doi.org/10.1007/BF00377088.
[39] Lambers, H., Poorter, H., 1992. Inherent variation in growth rate between higher plants: A search for physiological causes and ecological consequences. Adv. Ecol. Res. 34, 283-362. https://doi.org/10.1016/S0065-2504(03)34004-8.
[40] Legendre, P., 2018. lmodel2: Model II Regression (R Package Version 1.7-3). http://CRAN.R-project.org/package=lmodel2.
[41] Liang, X.S., Ning, Y., Wang, H.Y., et al., 2026. Why leaf N:P is not reliable for diagnosing nutrient limitation of productivity: answer from leaf phosphorus fractions. New Phytol. 249, 2800-2807. https://doi.org/10.1111/nph.70893.
[42] Lord, J., Westoby, M., Leishman, M., 1995. Seed size and phylogeny in six temperate floras: Constraints, niche conservatism, and adaptation. Am. Nat. 146, 349-364. https://doi.org/10.1086/285804.
[43] Lott, J.N., Ockenden, I., Raboy, V., et al., 2000. Phytic acid and phosphorus in crop seeds and fruits: a global estimate. Seed Sci. Res. 10, 11-33. https://doi.org/10.1017/S0960258500000039.
[44] Marschner, P., 2012. Marschner's mineral nutrition of higher plants. Amsterdam, Netherlands: The Netherland Academic Press.
[45] Muthukumar, T., Udaiyan, K., 2000. The role of seed reserves in arbuscular mycorrhizal formation and growth of Leucaena leucocephala (Lam.) de Wit. and Zea mays L. Mycorrhiza. 9, 323-330. https://doi.org/10.1007/s005720050276.
[46] Naegle, E.R., Burton, J.W., Carter, T.E., et al., 2005. Influence of seed nitrogen content on seedling growth and recovery from nitrogen stress. Plant Soil. 271, 329-340. https://doi.org/10.1007/s11104-004-3242-4.
[47] Niklas, K.J., 2006. Plant allometry, leaf nitrogen and phosphorus stoichiometry, and interspecific trends in annual growth rates. Ann. Bot. 97, 155-163. https://doi.org/10.1093/aob/mcj021.
[48] Niklas, K.J., Cobb, E.D., 2005. N, P, and C stoichiometry of Eranthis hyemalis (Ranunculaceae) and the allometry of plant growth. Am. J. Bot. 92, 1256-1263. https://doi.org/10.3732/ajb.92.8.1256.
[49] Niklas, K.J., Owens, T., Reich, P.B., et al., 2005. Nitrogen/phosphorus leaf stoichiometry and the scaling of plant growth. Ecol. Lett. 8, 636-642. https://doi.org/10.1111/j.1461-0248.2005.00759.x.
[50] Pagel, M., 1999. The maximum likelihood approach to reconstructing ancestral character states of discrete characters on phylogenies. Syst. Biol. 48, 612-622. https://doi.org/10.1080/106351599260184.
[51] Peoples, M.B., Craswell, E.T., 1992. Biological nitrogen fixation: Investments, expectations and actual contributions to agriculture. Plant Soil. 141, 13-39. https://doi.org/10.1007/BF00011308.
[52] R Core Team, 2024. R: A language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing.
[53] Reich, P.B., Oleksyn, J., 2004. Global patterns of plant leaf N and P in relation to temperature and latitude. Proc. Natl. Acad. Sci. U.S.A. 101, 11001-11006. https://doi.org/10.1073/pnas.0403588101.
[54] Reich, P.B., Oleksyn, J., Wright, I.J., et al., 2010. Evidence of a general 2/3-power law of scaling leaf nitrogen to phosphorus among major plant groups and biomes. Proc. R. Soc. B. 277, 877-883. https://doi.org/10.1098/rspb.2009.1818.
[55] Sardans, J., Janssens, I.A., Alonso, R., et al., 2015. Foliar elemental composition of E uropean forest tree species associated with evolutionary traits and present environmental and competitive conditions. Glob. Ecol. Biogeogr. 24, 240-255. https://doi.org/10.1111/geb.12253.
[56] Simovic, M., Michaletz, S.T., 2025a. Harnessing the full power of data to characterise biological scaling relationships. Glob. Ecol. Biogeogr. 34, e70019. https://doi.org/10.1111/geb.70019.
[57] Simovic, M., Michaletz, S.T., 2025b. Hydraulics and structural mechanics jointly shape root-to-leaf scaling of xylem conduit traits. Plant Cell Environ. https://doi.org/10.1111/pce.15660.
[58] Slot, M., Palow, D.T., Kitajima, K., 2013. Seed reserve dependency of Leucaena leucocephala seedling growth for nitrogen and phosphorus. Funct. Plant Biol. 40, 244-250. https://doi.org/10.1071/fp12255.
[59] Soltani, A., Gholipoor, M., Zeinali, E., 2006. Seed reserve utilization and seedling growth of wheat as affected by drought and salinity. Environ. Exp. Bot. 55, 195-200. https://doi.org/10.1016/j.envexpbot.2004.10.012.
[60] Soriano, D., Orozco-Segovia, A., Marquez-Guzman, J., et al., 2011. Seed reserve composition in 19 tree species of a tropical deciduous forest in Mexico and its relationship to seed germination and seedling growth. Ann. Bot. 107, 939-951. https://doi.org/10.1093/aob/mcr041.
[61] Sterner, R.W., Elser, J.J., 2002. Ecological stoichiometry: The biology of elements from molecules to the biosphere. Princeton, NJ: Princeton University Press.
[62] Stock, W.D., Verboom, G.A., 2012. Phylogenetic ecology of foliar N and P concentrations and N:P ratios across mediterranean-type ecosystems. Glob. Ecol. Biogeogr. 21, 1147-1156. https://doi.org/10.1111/j.1466-8238.2011.00752.x.
[63] Tang, Z.Y., Xu, W.T., Zhou, G.Y., et al., 2018. Patterns of plant carbon, nitrogen and phosphorus concentrations in relation to productivity in the terrestrial ecosystems of China. Proc. Natl. Acad. Sci. USA 115, 4033-4038.
[64] Te, N.W., Luo, W.T., Torok, P., et al., 2025. The context dependency of nitrogen deposition impacts on the compositional coupling between aboveground vegetation and soil seed bank. Plant Diversity. In press. https://doi.org/10.1016/j.pld.2025.12.009.
[65] Tian, D., Yan, Z.B., Niklas, K.J., et al., 2018. Global leaf nitrogen and phosphorus stoichiometry and their scaling exponent. Natl. Sci. Rev. 5, 728-739. https://doi.org/10.1093/nsr/nwx142.
[66] Turnbull, L.A., Philipson, C.D., Purves, D.W., et al., 2012. Plant growth rates and seed size: a re-evaluation. Ecology. 93, 1283-1289. https://doi.org/10.1890/11-0261.1.
[67] Valverde-Barrantes, O.J., Freschet, G.T., Roumet, C., et al., 2017. A worldview of root traits: The influence of ancestry, growth form, climate and mycorrhizal association on the functional trait variation of fine-root tissues in seed plants. New Phytol. 215, 1562-1573. https://doi.org/10.1111/nph.14571.
[68] Wang, X.T., Ji, M.F., Zhang, Y.H., et al., 2023. Plant trait networks reveal adaptation strategies in the drylands of China. BMC Plant Biol. 23, 266. https://doi.org/10.1186/s12870-023-04273-0.
[69] Wang, Z.Q., Bu, H.Y., Wang, M.C., et al., 2020. Allocation strategies for seed nitrogen and phosphorus in an alpine meadow along an altitudinal gradient on the Tibetan Plateau. Front. Plant Sci. 11, 614644. https://doi.org/10.3389/fpls.2020.614644.
[70] Wang, Z.Q., Gong, H.Y., Sardans, J., et al., 2022. Divergent nitrogen and phosphorus allocation strategies in terrestrial plant leaves and fine roots: A global meta-analysis. J. Ecol. 110, 2745-2758. https://doi.org/10.1111/1365-2745.13985.
[71] Wang, Z.Q., Huang, H., Yao, B.Q., et al., 2021. Divergent scaling of fine-root nitrogen and phosphorus in different root diameters, orders and functional categories: A meta-analysis. For. Ecol. Manag. 495, 119384. https://doi.org/10.1016/j.foreco.2021.119384.
[72] Wang, Z.Q., Ji, M.F., Deng, J.M., et al., 2015. A theoretical framework for whole-plant carbon assimilation efficiency based on metabolic scaling theory: a test case using Picea seedling. Tree Physiol. 35, 599-607. https://doi.org/10.1093/treephys/tpv030.
[73] Wang, Z.Q., Niklas, K.J., Ma, Z.Q., et al., 2022. The 2/3 scaling of twig nitrogen to phosphorus in woody plants. For. Ecosyst. 9, 100049. https://doi.org/10.1016/j.fecs.2022.100049.
[74] Wang, Z.Q., Yu, K.L., Lv, S.Q., et al., 2019. The scaling of fine root nitrogen versus phosphorus in terrestrial plants: A global synthesis. Funct. Ecol. 33, 2081-2094. https://doi.org/10.1111/1365-2435.13434.
[75] Warton, D.I., Wright, I.J., Falster, D.S., et al., 2006. Bivariate line-fitting methods for allometry. Biol. Rev. 81, 259-291. https://doi.org/10.1017/s1464793106007007.
[76] Wright, I.J., Reich, P.B., Westoby, M., et al., 2004. The worldwide leaf economics spectrum. Nature. 428, 821-827. https://doi.org/10.1038/nature02403.
[77] Xiong, J.L., Dong, L.W., Lu, J.L., et al., 2022. Variation in plant carbon, nitrogen and phosphorus contents across the drylands of China. Funct. Ecol. 36, 174-186. https://doi.org/10.1111/1365-2435.13937.
[78] Xiong, J.L., Hu, W.G., Degan, A., et al., 2025. Morphological and physiological responses to aridity in narrowly- and widely-distributed plant species in drylands. Sci. China Life Sci. 68, 2784-2795. https://doi.org/10.1007/s11427-024-2969-7.
[79] Yao, S.R., Hu, W.G., Ji, M.F., et al., 2025. Distribution, species richness, and relative importance of different plant life forms across drylands in China. Plant Diversity. 47, 273-281. https://doi.org/10.1016/j.pld.2024.09.007.
[80] Yuan, Z.Y., Chen, H.Y.H., Reich, P.B., 2011. Global-scale latitudinal patterns of plant fine-root nitrogen and phosphorus. Nat. Commun. 2, 344. https://doi.org/10.1038/ncomms1346.
[81] Zhao, M.Y., Luo, Y.K., Chen, Y.H., et al., 2021. Varied nitrogen versus phosphorus scaling exponents among shrub organs across eastern China. Ecol. Indic. 121, 107024. https://doi.org/10.1016/j.ecolind.2020.107024.

Scaling and stoichiometry of seed nitrogen and phosphorus across Chinese grasslands

Scaling and stoichiometry of seed nitrogen and phosphorus across Chinese grasslands

PDF (PC)

Knowledge

Abstract

Cite this article

share this article

References

Related Articles 15

Recommended Articles

Metrics

Comments

[1]	Ke Xia, Matthew I. Daws, Zi-Qi Zhu. Climate drives patterns of response of recalcitrant embryonic axes of Quercus species to cryopreservation [J]. Plant Diversity, 2026, 48(04): 670-677.
[2]	Fui Ying Tsan, Louise Colville, Hugh W. Pritchard. Regeneration from seeds with physical defects: Contrasts in survival strategies in recalcitrant and orthodox seeds [J]. Plant Diversity, 2026, 48(04): 641-654.
[3]	Li-Qin Zhu, Xiao-Dong Yao, Xiao-Hong Wang, David Robinson, Wei-Le Chen, Ting-Ting Chen, Qi Jiang, Lin-Qiao Jia, Ai-Lian Fan, Guang-Shui Chen. Responses of root exudates to phosphorus-rich patches in a subtropical evergreen broad-leaved forest and their module affiliation in the trait network [J]. Plant Diversity, 2026, 48(03): 608-617.
[4]	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.
[5]	Yu Chen, Xingjiang Song, Gang Liu, Jia Wang, Chunling Zhang, Xiaojian Chang, Jiabin Zou, Zhihong Zhu. Unraveling the invasion patterns of Galinsoga quadriradiata in mountain ranges: Insights from human activities, phenotypic and genetic variations [J]. Plant Diversity, 2026, 48(01): 192-203.
[6]	Renyu Zhou, Pengfei Yang, Xufang Chen, Minshu Song, Hang Sun, Jianguo Chen. Simulated climate warming strongly constrains the seedling establishment of alpine cushion Arenaria oreophila [J]. Plant Diversity, 2025, 47(06): 889-898.
[7]	Xiaoyan Hu, Jinyu Zhang, Bo Wang. Seed size distributions of fleshy-fruited species are more right-skewed compared to that of dry-fruited species [J]. Plant Diversity, 2025, 47(03): 522-525.
[8]	Yufeng Xu, Chenyang Xue, Xuezhi Wang, Lin Meng, Ying Gao, Mengyang Yu, Lin Geng, Meini Shao, Bo Qu. Morphological adaptation strategy of invasive plant Ambrosia trifida seed distributed along rivers is closely related to soil nutrients [J]. Plant Diversity, 2025, 47(03): 499-510.
[9]	Fu-Qiang Huang, Josep Peñuelas, Jordi Sardans, Scott L. Collins, Kai-Liang Yu, Man-Qiong Liu, Jiu-Ying Pei, Wen-Bin Ke, Jian-Sheng Ye. Plant use of water across soil depths regulates species dominance under nitrogen addition [J]. Plant Diversity, 2025, 47(03): 479-488.
[10]	Qinfeng Guo, Hong Qian, Shenhua Qian. Climate space, traits, and the spread of nonnative plants in North America [J]. Plant Diversity, 2025, 47(02): 255-263.
[11]	Min-Fei Jin, Xiang-Hai Cai, Gao Chen. Seed dispersal by deception: A game between mimetic seeds and their bird dispersers [J]. Plant Diversity, 2025, 47(02): 169-177.
[12]	Jianghua Duan, Liu Yang, Ting Tang, Jiesheng Rao, Wencong Liu, Xi Chen, Rong Li, Zehao Shen. Environment and management jointly shape the spatial patterns of plant species diversity of moist grasslands in the mountains of northeastern Yunnan [J]. Plant Diversity, 2024, 46(06): 744-754.
[13]	Libing Pan, Xiaoyang Song, Wenfu Zhang, Jie Yang, Min Cao. Seedling dynamics differ between canopy species and understory species in a tropical seasonal rainforest, SW China [J]. Plant Diversity, 2024, 46(05): 671-677.
[14]	Tana Wuyun, Lu Zhang, Tiina Tosens, Bin Liu, Kristiina Mark, José ángel Morales-Sánchez, Jesamine Jöneva Rikisahedew, Vivian Kuusk, ülo Niinemets. Extremely thin but very robust: Surprising cryptogam trait combinations at the end of the leaf economics spectrum [J]. Plant Diversity, 2024, 46(05): 621-629.
[15]	Meixu Han, Haiyang Zhang, Mingchao Liu, Jinqi Tang, Xiaocheng Guo, Weizheng Ren, Yong Zhao, Qingpei Yang, Binglin Guo, Qinwen Han, Yulong Feng, Zhipei Feng, Honghui Wu, Xitian Yang, Deliang Kong. Increased dependence on nitrogen-fixation of a native legume in competition with an invasive plant [J]. Plant Diversity, 2024, 46(04): 510-518.