Variation Patterns of Plant Root and Leaf Functional Traits, and Ecosystem Functioning: Research Progress and Prospect

Authors

  • Hailing Liao

    Department of Forest Science and Biodiversity, Faculty of Forestry and Environment, Universiti Putra Malaysia, Serdang 43400, Malaysia

    Nanchang Junhan Environmental Protection Consulting Co., Ltd., Nanchang 330224, China
  • Mohd Nazre

    Department of Forest Science and Biodiversity, Faculty of Forestry and Environment, Universiti Putra Malaysia, Serdang 43400, Malaysia
  • Beilei Yin

    Department of Forest Science and Biodiversity, Faculty of Forestry and Environment, Universiti Putra Malaysia, Serdang 43400, Malaysia

    College of Forestry Engineering, Guangxi Eco-engineering Vocational and Technical College, Liuzhou 545003, China
  • Johar Mohamed

    Department of Forest Science and Biodiversity, Faculty of Forestry and Environment, Universiti Putra Malaysia, Serdang 43400, Malaysia

DOI:

https://doi.org/10.30564/re.v8i2.12690
Received: 6 November 2025 | Revised: 25 December 2025 | Accepted: 12 January 2026 | Published Online: 20 April 2026

Abstract

Plant root and leaf functional traits link individual adaptation to ecosystem functioning, yet their coordination mechanisms remain insufficiently understood. We established 18 plots along an elevational gradient (1200–2700 m) in the Qinling Mountains, measuring 12 key traits of 12 tree species and assessing three ecosystem functions. Results showed: (1) Traits exhibited significant interspecific variation with environmental gradients explaining 42–58% of variance; (2) Root-leaf coordination was moderate globally (r = 0.58 for specific leaf area and specific root length) but strengthened significantly under high-elevation stress (r increased from 0.38 to 0.72), supporting the environmental stress hypothesis; (3) Principal component analysis identified three ecological strategies—competitive, stress-tolerant, and intermediate types—cumulatively explaining 78.6% of total trait variance, with environmental factors predicting strategy classification at 82.3% accuracy; (4) Structural equation modeling revealed environmental effects were predominantly transmitted through root traits (β = 0.62) to leaf traits (β = 0.43), with 61.3% of effects mediated by traits rather than direct pathways; (5) Specific leaf area and leaf nitrogen jointly explained 61.2% of productivity variation, while litter nitrogen content controlled decomposition rates (r = 0.78), and the lignin:nitrogen ratio explained 65.6% of decomposition variance. This research confirms functional traits as effective predictors of ecosystem functioning, providing theoretical foundations for trait-based ecosystem management under global change scenarios.

Keywords:

Plant Functional Traits; Root-Leaf Coordination; Trait Trade-Offs; Ecosystem Functioning; Elevational Gradient

References

[1] Cornelissen, J.H.C., Lavorel, S., Garnier, E., et al., 2003. A handbook of protocols for standardised and easy measurement of plant functional traits worldwide. Australian Journal of Botany. 51(4), 335–380. DOI: https://doi.org/10.1071/BT02124

[2] Reich, P.B., Tilman, D., Craine, J., et al., 2001. Do species and functional groups differ in acquisition and use of C, N and water under varying atmospheric CO2 and N availability regimes? A field test with 16 grassland species. New Phytologist. 150(2), 435–448. DOI: https://doi.org/10.1046/j.1469-8137.2001.00114.x

[3] McIntyre, S., Lavorel, S., Landsberg, J., et al., 1999. Disturbance response in vegetation – towards a global perspective on functional traits. Journal of Vegetation Science. 10(5), 621–630. DOI: https://doi.org/10.2307/3237077

[4] Shipley, B., Vile, D., Garnier, E., et al., 2005. Functional linkages between leaf traits and net photosynthetic rate: reconciling empirical and mechanistic models. Functional Ecology. 19(4), 602–615. DOI: https://doi.org/10.1111/j.1365-2435.2005.01008.x

[5] Wright, I.J., Reich, P.B., Westoby, M., et al., 2004. The worldwide leaf economics spectrum. Nature. 428(6985), 821–827. DOI: https://doi.org/10.1038/nature02403

[6] Eissenstat, D.M., Yanai, R.D., 1997. The Ecology of Root Lifespan. In Advances in Ecological Research, Vol. 27. Elsevier: New York, NY, USA. pp. 1–60. DOI: https://doi.org/10.1016/S0065-2504(08)60005-7

[7] McCormack, M.L., Guo, D., 2014. Impacts of environmental factors on fine root lifespan. Frontiers in Plant Science. 5. DOI: https://doi.org/10.3389/fpls.2014.00205

[8] Freschet, G.T., Swart, E.M., Cornelissen, J.H.C., 2015. Integrated plant phenotypic responses to contrasting above‐ and below‐ground resources: key roles of specific leaf area and root mass fraction. New Phytologist. 206(4), 1247–1260. DOI: https://doi.org/10.1111/nph.13352

[9] Comas, L.H., Eissenstat, D.M., 2009. Patterns in root trait variation among 25 co‐existing North American forest species. New Phytologist. 182(4), 919–928. DOI: https://doi.org/10.1111/j.1469-8137.2009.02799.x

[10] McCormack, M.L., Dickie, I.A., Eissenstat, D.M., et al., 2015. Redefining fine roots improves understanding of below‐ground contributions to terrestrial biosphere processes. New Phytologist. 207(3), 505–518. DOI: https://doi.org/10.1111/nph.13363

[11] Guo, D., Xia, M., Wei, X., et al., 2008. Anatomical traits associated with absorption and mycorrhizal colonization are linked to root branch order in twenty‐three Chinese temperate tree species. New Phytologist. 180(3), 673–683. DOI: https://doi.org/10.1111/j.1469-8137.2008.02573.x

[12] Grime, J.P., 2006. Trait convergence and trait divergence in herbaceous plant communities: Mechanisms and consequences. Journal of Vegetation Science. 17(2), 255–260. DOI: https://doi.org/10.1111/j.1654-1103.2006.tb02444.x

[13] Laughlin, D.C., Messier, J., 2015. Fitness of multidimensional phenotypes in dynamic adaptive landscapes. Trends in Ecology & Evolution. 30(8), 487–496. DOI: https://doi.org/10.1016/j.tree.2015.06.003

[14] Freschet, G.T., Valverde‐Barrantes, O.J., Tucker, C.M., et al., 2017. Climate, soil and plant functional types as drivers of global fine‐root trait variation. Journal of Ecology. 105(5), 1182–1196. DOI: https://doi.org/10.1111/1365-2745.12769

[15] Li, A., Guo, D., Wang, Z., et al., 2010. Nitrogen and phosphorus allocation in leaves, twigs, and fine roots across 49 temperate, subtropical and tropical tree species: A hierarchical pattern. Functional Ecology. 24(1), 224–232. DOI: https://doi.org/10.1111/j.1365-2435.2009.01603.x

[16] Meier, I.C., Leuschner, C., 2008. Below-ground drought response of European beech: Fine root biomass and carbon partitioning in 14 mature stands across a precipitation gradient. Global Change Biology. 14(9), 2081–2095. DOI: https://doi.org/10.1111/j.1365-2486.2008.01634.x

[17] Zimmermann, J., Link, R.M., Hauck, M., et al., 2021. 60-year record of stem xylem anatomy and related hydraulic modification under increased summer drought in ring- and diffuse-porous temperate broad-leaved tree species. Trees. 35(3), 919–937. DOI: https://doi.org/10.1007/s00468-021-02090-2

[18] Lloyd, J., Bloomfield, K., Domingues, T.F., et al., 2013. Photosynthetically relevant foliar traits correlating better on a mass vs an area basis: Of ecophysiological relevance or just a case of mathematical imperatives and statistical quicksand? New Phytologist. 199(2), 311–321. DOI: https://doi.org/10.1111/nph.12281

[19] Rao, C., Su, X., Xia, Y., et al., 2026. From Physical Architecture to Ecosystem Function: Tillage Exerts Indirect Control on Nitrogen Transformation by Restructuring Preferential Flow Paths. Environmental Science & Technology. 60(2), 1886–1899. DOI: https://doi.org/10.1021/acs.est.5c12351

[20] Liu, Y., Hu, M., Zeng, W., et al., 2026. Enhancing spatial representativeness and adaptive management of national parks by integrating biodiversity, ecosystem service multifunctionality, and trade-offs. Journal of Environmental Management. 398, 128472. DOI: https://doi.org/10.1016/j.jenvman.2025.128472

[21] Mao, W., Li, Y.L., Zhang, T.H., et al., 2012. Research advances of plant leaf traits at different ecology scales. Journal of Desert Research. 32, 33–41. (in Chinese)

[22] Akram, M.A., Zhang, Y., Wang, X., et al., 2022. Phylogenetic independence in the variations in leaf functional traits among different plant life forms in an arid environment. Journal of Plant Physiology. 272, 153671. DOI: https://doi.org/10.1016/j.jplph.2022.153671

[23] Meziane, D., Shipley, B., 1999. Interacting determinants of specific leaf area in 22 herbaceous species: Effects of irradiance and nutrient availability. Plant, Cell & Environment. 22(5), 447–459. DOI: https://doi.org/10.1046/j.1365-3040.1999.00423.x

[24] Suzuki, K.C., Kajino, H., Hirokawa, S., et al., 2025. The coordination between root and leaf functional traits across 33 woody plant species shifts between mycorrhizal types. Tree Physiology. 45, tpaf151. DOI: https://doi.org/10.1093/treephys/tpaf151

[25] Moles, A.T., Perkins, S.E., Laffan, S.W., et al., 2014. Which is a better predictor of plant traits: Temperature or precipitation? Journal of Vegetation Science. 25(5), 1167–1180. DOI: https://doi.org/10.1111/jvs.12190

[26] Rodríguez-Castilla, G., Carpio, A.J., Bastias, C.C., et al., 2025. Leaf functional traits as browsing indicators in understory woody species of Mediterranean forests. Forest Ecology and Management. 598, 123230. DOI: https://doi.org/10.1016/j.foreco.2025.123230

[27] Tan, L., Liang, J., Qin, Z., et al., 2025. Unveiling the sustained effects of plant root exudates on soil microbiome and resistome and the related functional traits. Journal of Environmental Management. 376, 124485. DOI: https://doi.org/10.1016/j.jenvman.2025.124485

[28] Niinemets, Ü., 2001. Global-Scale Climatic Controls of Leaf Dry Mass Per Area, Density, and Thickness in Trees and Shrubs. Ecology. 82(2), 453–469. DOI: https://doi.org/10.1890/0012-9658(2001)082%255B0453:GSCCOL%255D2.0.CO;2

[29] Zhu, C., Wang, Z., Luo, W., et al., 2022. Fungal phylogeny and plant functional traits structure plant–rhizosphere fungi networks in a subtropical forest. Oikos. 2022(8), e08992. DOI: https://doi.org/10.1111/oik.08992

[30] Ostonen, I., Püttsepp, Ü., Biel, C., et al., 2007. Specific root length as an indicator of environmental change. Plant Biosystems - An International Journal Dealing with all Aspects of Plant Biology. 141(3), 426–442. DOI: https://doi.org/10.1080/11263500701626069

[31] Ali, F., Jilani, G., Bai, L., et al., 2021. Effects of previous drying of sediment on root functional traits and rhizoperformance of emerged macrophytes. Frontiers of Environmental Science & Engineering. 15(6), 134. DOI: https://doi.org/10.1007/s11783-021-1427-1

[32] Poorter, H., Niinemets, Ü., Poorter, L., et al., 2009. Causes and consequences of variation in leaf mass per area (LMA): A meta‐analysis. New Phytologist. 182(3), 565–588. DOI: https://doi.org/10.1111/j.1469-8137.2009.02830.x

[33] Guzmán-Jacob, V., Weigelt, P., Craven, D., et al., 2021. Biovera-Epi: A new database on species diversity, community composition and leaf functional traits of vascular epiphytes along gradients of elevation and forest-use intensity in Mexico. Biodiversity Data Journal. 9, e71974. DOI: https://doi.org/10.3897/BDJ.9.e71974

[34] Pregitzer, K.S., DeForest, J.L., Burton, A.J., et al., 2002. Fine Root Architecture of Nine North American Trees. Ecological Monographs. 72(2), 293–309. DOI: https://doi.org/10.1890/0012-9615(2002)072%255B0293:FRAONN%255D2.0.CO;2

[35] Raich, J.W., Nadelhoffer, K.J., 1989. Below-ground Carbon Allocation in Forest Ecosystems: Global Trends. Ecology. 70(5), 1346–1354. DOI: https://doi.org/10.2307/1938194

[36] Poli, A., Bongiovanni, D., Stefanini, I., et al., 2026. Autochthonous microorganisms of a soil contaminated by polycyclic aromatic hydrocarbons: Allies or silent threats? Biodiversity and Conservation. 35(2), 42. DOI: https://doi.org/10.1007/s10531-025-03244-1

[37] Rodríguez-Gallego, C., Navarro, T., Meerts, P., 2015. A comparative study of leaf trait relationships in coastal dunes in southern Spain. Plant Ecology and Evolution. 148(1), 57–67. DOI: https://doi.org/10.5091/plecevo.2015.951

[38] Xiao, J., Olatunji, O.A., Wang, D., et al., 2026. Wildfire alters forest structure while belowground multifunctionality remains unchanged in a karst Pinus massoniana forest. Forest Ecology and Management. 605, 123515. DOI: https://doi.org/10.1016/j.foreco.2026.123515

[39] Kashif, M., Liang, Q., Meng, C., et al., 2026. Ecological and functional diversity of sulfur-oxidizing and sulfate-reducing bacteria in Chinese mangrove ecosystems: A comparative review with insights from Guangxi Beihai. Rhizosphere. 37, 101260. DOI: https://doi.org/10.1016/j.rhisph.2026.101260

[40] Shi, Y., Wen, Z.M., Gong, S.H., et al., 2012. Trait variations along a climatic gradient in hilly area of Loess Plateau. Research of Soil and Water Conservation. 19(1), 107–111. (in Chinese)

[41] Chaudhary, R.D., Gautam, K.R., Yousuf, B., et al., 2015. Nutrients, microbial community structure and functional gene abundance of rhizosphere and bulk soils of halophytes. Applied Soil Ecology. 91, 16–26. DOI: https://doi.org/10.1016/j.apsoil.2015.02.003

[42] Pan, F., Yu, X., Liang, Y., et al., 2026. Restoration boosts soil P-cycle multifunctionality in karst ecosystems by modulating soil properties and rare bacterial taxa. Journal of Integrative Agriculture. 25(2), 513–528. DOI: https://doi.org/10.1016/j.jia.2025.05.001

[43] Swenson, N.G., Enquist, B.J., Pither, J., et al., 2012. The biogeography and filtering of woody plant functional diversity in North and South America. Global Ecology and Biogeography. 21(8), 798–808. DOI: https://doi.org/10.1111/j.1466-8238.2011.00727.x

[44] Liu, Y., Zhang, R., Shangguan, Z., et al., 2026. Distributions and influencing factors of glomalin-related soil protein in aggregate following forest restoration. Ecological Engineering. 225, 107893. DOI: https://doi.org/10.1016/j.ecoleng.2026.107893

[45] Vendramini, F., Díaz, S., Gurvich, D.E., et al., 2002. Leaf traits as indicators of resource‐use strategy in floras with succulent species. New Phytologist. 154(1), 147–157. DOI: https://doi.org/10.1046/j.1469-8137.2002.00357.x

[46] Vogt, K.A., Vogt, D.J., Palmiotto, P.A., et al., 1995. Review of root dynamics in forest ecosystems grouped by climate, climatic forest type and species. Plant and Soil. 187(2), 159–219. DOI: https://doi.org/10.1007/BF00017088

[47] Raza, A., Ahmad, F., Asghar, A., et al., 2026. Structural and functional adaptations of Cenchrus ciliaris to tree-specific overstory microenvironments in a forest ecosystem. Agroforestry Systems. 100(1), 44. DOI: https://doi.org/10.1007/s10457-025-01422-2

[48] Wang, R.L., Yu, G.R., He, N.P., et al., 2015. Latitude pattern and influencing factors of functional traits of forest leaves in China. Acta Geographica Sinica. 70(11), 1735–1746. (in Chinese)

[49] Wang, Z.Q., Guo, D.L., 2008. Root ecology. Chinese Journal of Plant Ecology. (6), 6–9. (in Chinese)

[50] Liu, T., Kaviriri, D.K., Gontcharov, A.A., et al., 2026. Identifying broader distribution and permanent refugia of Korean pine (Pinus koraiensis) for adaptive management in a changing climate. European Journal of Forest Research. 145(1), 2. DOI: https://doi.org/10.1007/s10342-025-01850-w

[51] Withington, J.M., Reich, P.B., Oleksyn, J., et al., 2006. Comparisons of Structure and Life Span in Roots and Leaves Among Temperate Trees. Ecological Monographs. 76(3), 381–397. DOI: https://doi.org/10.1890/0012-9615(2006)076%255B0381:COSALS%255D2.0.CO;2

[52] Xia, M., Guo, D., Pregitzer, K.S., 2010. Ephemeral root modules in Fraxinus mandshurica. New Phytologist. 188(4), 1065–1074. DOI: https://doi.org/10.1111/j.1469-8137.2010.03423.x

[53] Li, K., Wu, W., Li, Z., et al., 2026. Ecosystem quality dynamics and attribution in protected areas: An integrated analysis framework based on the structure-function-resilience-pressure. Journal of Environmental Management. 398, 128600. DOI: https://doi.org/10.1016/j.jenvman.2026.128600

[54] Yin, X., Perry, J.A., 1991. Factors Affecting Nitrogen Concentration of Fine Roots in Forest Communities: Regression Analysis of Literature Data. Forest Science. 37(1), 374–382. DOI: https://doi.org/10.1093/forestscience/37.1.374

[55] Zhang, W., Wen, H.S., Zheng, M.J., et al., 2026. Pollution legacy effect alters the response of soil ecological functions to water limitation. Environmental Science. 47(1), 614–621. (in Chinese)

[56] de Conto, T., Armston, J., Dubayah, R., 2026. Scalable deep fusion of spaceborne lidar and synthetic aperture radar for global forest structural complexity mapping. Machine Learning: Earth. 2(1), 015002. DOI: https://doi.org/10.1088/3049-4753/ae2c01

[57] Shobade, S.O., Zabotina, O.A., Nilsen-Hamilton, M., 2024. Plant root associated chitinases: structures and functions. Frontiers in Plant Science. 15, 1344142. DOI: https://doi.org/10.3389/fpls.2024.1344142

[58] Nadeem, F., Liu, X., 2023. Editorial: Advanced insights into plant rhizosphere functionality from the perspective of declining soil fertility status in the era of climate change. Frontiers in Plant Science. 14, 1338575. DOI: https://doi.org/10.3389/fpls.2023.1338575

[59] Da Silva, J.R., Yule, T.S., Ribas, A.C.D.A., et al., 2023. Do root secondary xylem functional traits differ between growth forms in Fabaceae species in a seasonally dry Neotropical environment? Annals of Botany. 132(3), 401–412. DOI: https://doi.org/10.1093/aob/mcad131

[60] Zhan, Y.Y., Liu, Y.H., Xiong, W.J., 2010. Research status and prospect of endangered causes of Davidia involucrata. Hubei Forestry Science and Technology. 1, 41–56. (in Chinese)

[61] Feng, H., Fu, R., Luo, J., et al., 2023. Listening to plant’s Esperanto via root exudates: reprogramming the functional expression of plant growth‐promoting rhizobacteria. New Phytologist. 239(6), 2307–2319. DOI: https://doi.org/10.1111/nph.19086

[62] de Francisco Martínez, P., Morgante, V., González-Pastor, J.E., 2022. Isolation of novel cold-tolerance genes from rhizosphere microorganisms of Antarctic plants by functional metagenomics. Frontiers in Microbiology. 13, 1026463. DOI: https://doi.org/10.3389/fmicb.2022.1026463

[63] Zhu, H., Fu, B., Wang, S., et al., 2015. Reducing soil erosion by improving community functional diversity in semi‐arid grasslands. Journal of Applied Ecology. 52(4), 1063–1072. DOI: https://doi.org/10.1111/1365-2664.12442

Downloads

How to Cite

Liao, H., Nazre, M., Yin, B., & Mohamed, J. (2026). Variation Patterns of Plant Root and Leaf Functional Traits, and Ecosystem Functioning: Research Progress and Prospect. Research in Ecology, 8(2), 298–324. https://doi.org/10.30564/re.v8i2.12690

Issue

Vol.8, Iss.2 (April 2026)(in progress)

Article Type

Article

License

Copyright © 2026 Hailing Liao, Mohd Nazre, Beilei Yin, Johar Mohamed

Creative Commons License

This is an open access article under the Creative Commons Attribution-NonCommercial 4.0 International (CC BY-NC 4.0) License.

Crossref
Scopus
Google Scholar
Europe PMC