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Arbuscular mycorrhizal association regulates global root–seed coordination

Abstract

Terrestrial plants exhibit immense variation in their form and function among species. Coordination between resource acquisition by roots and reproduction through seeds could promote the fitness of plant populations. How root and seed traits covary has remained unclear until our analysis of the largest-ever compiled joint global dataset of root traits and seed mass. Here we demonstrate that seed mass and seed phosphorus mass scale positively with root diameter in arbuscular mycorrhizal (AM) plants, depending on variation in root cortical thickness instead of root vessel size. These findings suggest a dual role of AM association in phosphorus uptake and pathogen resistance which drives the global root–seed coordination, instead of initially expected resource transport via root vessels as the main driver. In contrast, we found no relationship between root traits and seed mass in ectomycorrhizal plants. Overall, our study reveals coordination between roots and seeds in AM plants, which is probably regulated by root–mycorrhizal symbiosis, and may be crucial in shaping global plant diversity and species distributions.

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Fig. 1: Three hypotheses regarding the relationships between roots and seeds.
Fig. 2: Integration of seed mass within the above- and belowground plant trait space.
Fig. 3: Global relationship between root diameter and seed mass.
Fig. 4: Relationships between root anatomical traits and seed mass.
Fig. 5: Relationships between soil pathogen richness and root traits.
Fig. 6: Evolutionary trends of absorptive root diameter and seed mass.

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Data availability

The raw data in this study are available in figshare (https://doi.org/10.6084/m9.figshare.28300658)74. Literature data were extracted from the Global Root Trait database (https://groot-database.github.io/GRooT/)40.

Code availability

The code utilized for this study is publicly available and is hosted in figshare (https://doi.org/10.6084/m9.figshare.28300658)74.

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Acknowledgements

We thank S. Chen for comments on an early draft of the manuscript; J. Bergmann for discussion of the idea regarding hypothesis 2 in the early draft; J. Chen, J. Cao, M. Han, H. Wang, Y. Hou, Y. Tian, Y. Dong, M. Liu, D. Mo, C. Xiang, Y. Jiang, Y. Liang, S. Dong, Z. Meng, L. Zhao and C. Hu for assistance in field sampling. We also thank the following field research stations and government agencies for their kind support: Xishuangbanna Station for Tropical Forest Studies of Xishuangbanna Tropical Botanical Garden; Dinghushan Station for Subtropical Forest Studies of South China Botany Garden; Gutianshan Biodiversity Science Research Station; Shennongjia National Park Administration; Forest Ecosystem Research Station, Institute of Botany, Chinese Academy of Sciences; Beijing Forest Ecosystem Research Station, Chinese Academy of Sciences; Liangshui Experimental Forest Farm, Northeast Forestry University; Changbai Mountain Forest Ecosystem Positioning Station, Chinese Academy of Sciences. This study was supported by the National Natural Science Foundation of China (32471824, 32171746 and 31870522) to D.K., the leading talents of basic research in Henan Province (24XM0375) to D.K., Excellent Youth Creative Research Group Project in Henan Province (252300421002) to D.K., Foreign Scientists Studio in Henan Province (GZS2025011) to D.K., the Scientific Research Foundation of Henan Agricultural University (30500854) to D.K., the National Natural Science Foundation of China (42122054 and 42477227) to J.W., the Guangdong Provincial Key Laboratory of Soil and Groundwater Pollution Control (2023B1212060002) to J.W., the High-level University Special Fund (G030290001) to J.W., the National Natural Science Foundation of China (42077450) to Y. Z. and the Estonian Research Council (PRG2142) to C.P.C.

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Authors and Affiliations

Authors

Contributions

Q.Y. and D.K. conceived of the idea. Q.Y. and B.G. completed the creation of figures. Q.Y., D.K., M.L. and P.B.R. conducted the statistical analyses. Q.Y., D.K., J.W., G.L., H.W. and Y.L. discussed and contributed to the final framework of this study. Q.Y and D.K. wrote the first draft of the paper with significant help from P.K., R.D.B., J.H.C.C., S.D., I.J.W. and J.A.H. Q.Y., B.G., M.L., Y.L., P.K., P.B.R., R.D.B., J.H.C.C., N.J.B.K., S.D., I.J.W., N.H., J.A.H., Y.P., Q.H., Z.L., Z.W., W.Y., J.D., Z.Y., H.W., C.P.C., O.J.V.-B., D.L., J.C., H.Z., Y. Zhang, W.R., Y. Zhao, X.Y., G.F., J.W., G.L. and D.K. contributed to the completion and revision of the paper.

Corresponding authors

Correspondence to Junjian Wang, Guoyong Li or Deliang Kong.

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Nature Plants thanks Wenming Bai, Christina Birnbaum and Thomas Kuyper for their contribution to the peer review of this work.

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Extended data

Extended Data Fig. 1 Phylogenetic tree of 626 taxa in the study.

The internal purple section of the phylogeny corresponds to the Cornales, the red section to the Ranunculales, the green section to the Magnoliales, the coffee-colored section to the Gymnosperms, the yellow section to the Rosales, and the blue section to the Saxifragales. The various color bars on the outside of the phylogeny’s inner circle correspond to the mycorrhizal group, growth form, and climatic zone (in that order from inside to outside) of the respective plant species (the tips of the phylogeny). AM = arbuscular mycorrhizal; ECM = ectomycorrhizal; ERM = ericoid mycorrhizal; Non = unknown / non-mycorrhizal. Due to the limited number of ERM species, we only use AM and ECM species in our analysis.

Extended Data Fig. 2 Relationships between root diameter and seed mass.

Analyses were performed using our field-measured trait data of this study (a, c) and existing data from the literature (b, d) using the standardized major axis (SMA) regression. For arbuscular mycorrhizal (AM) plant species, a significant positive correlation is shown: Field-measured data (a): Green filled circles; regression equation: log10(y) = 5.80 log10(x) + 4.46, r = 0.43, P = 2.2 × 10−16, n = 489. Literature data: (b): Tan filled circles; regression equation: log10(y) = 6.10 log10(x) + 4.41, r = 0.43, P = 2.2 × 10−16, n = 506. The scaling exponent for the field-measured data (a) is 5.80 (95% confidence interval (CI) = 5.35−6.28), equivalent to the exponent derived from the literature data (b) (6.10; 95% CI = 5.61−6.57) (P = 2.22 × 10−16; see the Method section). In contrast, no significant correlation was found for ectomycorrhizal (ECM) plant species: Field-measured data (c): Tan filled circles; r = 0.01, P = 0.91, n = 78. Literature data (d): Tan filled circles; r = 0.24, P = 0.07, n = 55. Significance was tested using a two-sided t-test. All data are plotted on logarithmic scales (log10) for both axes, with each point representing a single plant species.

Extended Data Fig. 3 Relationships between root anatomical traits and seed mass for different plant growth forms and mycorrhizal types.

Analyses were performed using field-measured data using the standardized major axis (SMA) regression. There is a significant positive relationship between seed mass and root diameter in arbuscular mycorrhizal (AM) tree and shrub species (green filled circles; a, log10(y) = 5.75 log10(x) + 4.51, r = 0.40, P = 2.24 × 10−13, 95% CI = 5.19−6.38, n = 303; tan filled circles, d, log10(y) = 5.55 log10(x) + 4.21, r = 0.38, P = 6.70 × 10−6, 95% CI = 4.85−6.34, n = 186). The cortical thickness and seed mass of AM plant species show a positive relationship across two growth forms (green filled circles, b, log10(y) = 4.73 log10(x) - 7.34, r = 0.33, P = 1.26 × 10−9, 95% CI = 4.25−5.26, n = 303; tan filled circles; e, log10(y) = 4.63 log10(x) - 7.40, r = 0.26, P = 0.0003, 95% CI = 4.02−5.33, n = 185). The vessel diameter and seed mass is positively correlated only in tree species (green filled circles, c, log10(y) = 7.82 log10(x) - 4.37, r = 0.20, P = 0.0006, 95% CI = 6.98−8.78, n = 283), but not in shrub species (tan filled circles, f, r = 0.05, P = 0.51, n = 175). Conditional correlations by considering the significant relationship between cortical thickness and vessel diameter (r = 0.52, P < 0.01) shows no correlation between vessel diameter and seed mass (r = 0.03, P = 0.49) but a positive correlation between cortical thickness and seed mass (r = 0.22, P < 0.01). In contrast, no correlation is observed for ectomycorrhizal (ECM) species (green open circles, a, r = 0.05, P = 0.73, n = 62; b, r = 0.18, P = 0.15, n = 62; c, r = 0.01, P = 0.92, n = 57; tan open circles, d, r = 0.09, P = 0.73, n = 16; e, r = 0.44, P = 0.09, n = 16; f, r = 0.13, P = 0.67, n = 14). Significance was tested using a two-sided t-test. Data are plotted on a logarithmic scale (log10) for both axes, with each point representing one species.

Extended Data Fig. 4 Relationships between root traits and seed mass of different mycorrhizal types across different climatic zones.

Analyses were performed using field-measured data using the standardized major axis (SMA) regression. Seed mass with both root diameter and cortical thickness of arbuscular mycorrhiza (AM) plant species are positively correlated in tropics and subtropics (green filled circles, a, log10(y) = 5.72 log10(x) + 4.42, r = 0.38, P = 1.83 × 10−8, 95% CI = 5.04−6.50, n = 206; d, log10(y) = 5.80 log10(x) + 4.50, r = 0.44, P = 5.88 × 10−10, 95% CI = 5.08−6.63, n = 177; tan filled circles, b, log10(y) = 4.40 log10(x) −6.61, r = 0.31, P = 4.58 × 10−6, 95% CI = 3.86−5.01, n = 206; e, log10(y) = 4.76 log10(x) −7.56, r = 0.38, P = 1.92 × 10−7, 95% CI = 4.15−5.46, n = 177), but not in temperate (green filled circles, g, r = 0.17, P = 0.09, n = 106. tan filled circles, h, r = 0.1, P = 0.31, n = 105). In contrast, no correlation is observed for ectomycorrhizal (ECM) plant species (green open circles, a, R2 = 0.27, P = 0.45, n = 10; d, r = 0.12, P = 0.57, n = 25; g, r = 0.07, P = 0.65, n = 43; tan open circles, b, r = 0.48, P = 0.16, n = 10; e, r = 0.08, P = 0.71, n = 25; h, r = 0.29, P = 0.06, n = 43). The vessel diameter and seed mass of AM and ECM plant species show no correlation across three climatic zones (blue filled circles, c, r = 0.13, P = 0.07, n = 192; f, r = 0.15, P = 0.052, n = 170; i, r = 0.002, P = 0.95, n = 96; blue open circles, c, r = 0.62, P = 0.10, n = 8; f, r = 0.06, P = 0.79, n = 24; i, r = 0.06, P = 0.68, n = 39). Significance was tested using a two-sided t-test. Data are plotted on a logarithmic scale (log10) for both axes, with each point representing one species.

Extended Data Fig. 5 Relationships between root diameter and seed mass for global woody and non-woody species.

Analyses were performed using field-measured data of this study and literature data using the standardized major axis (SMA) regression. There is a positive correlation between root diameter and seed mass in global woody arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) plant species (AM + ECM) plant species (Total, a, log10(y) = 5.93 log10(x) + 4.52, r = 0.26, P = 2.2 × 10−16, CI = 5.56−6.33, n = 854). The positive correlation between root diameter and seed mass persists for both global woody and non-woody AM plant species (green filled circles, a, log10(y) = 5.55 log10(x) + 4.27, r = 0.31, P = 2.2 × 10−16, n = 729; tan filled circles, b, log10(y) = 4.76 log10(x) + 4.26, r = 0.41, P = 2.79 × 10−12, n = 266). For global woody AM plant species, the scaling exponent is 5.55 (95% (CI) = 5.18−5.95), which is significantly different from that of global non-woody AM plant species (exponent = 4.76, 95% CI = 4.26−5.31) (P = 2.22 × 10−16, see the Methods section). In contrast, no correlation is observed for ECM woody plant species (green open circles, a, r = 0.14, P = 0.11, n = 137). Non-woody ECM plant species are not analyzed due to the limited number of species. Significance was tested using a two-sided t-test. Data are presented on logarithmic scales (log10) for both axes, with each point representing one species.

Extended Data Fig. 6 Pairwise correlations for all plant traits used in this study.

Analyses were performed using field-measured data of this study. In each scatter plot, the blue scatter is represented by the raw data, and the red scatter is represented by PGLS data (removal of the phylogenetic signal). SM - seed mass, RD - root diameter, SRL - specific root length, RTD - root tissue density, RN- root nitrogen concentration, CT - root cortical thickness, VD – root vessel diameter, VDen – root vessel density, LA - single leaf area, SLA – specific leaf area, LN - leaf nitrogen concentration, and H - height of mature plants. Regression lines represent raw data correlations (blue) and phylogenetically-corrected bivariate relationships calculated by fitting Phylogenetic Generalized Least Square models (red). Correlation coefficients are obtained using Pearson correlation, with the raw data shown in blue and the phylogenetic correction data in red (* 0.05 > P > 0.01; ** 0.01 > P > 0.001; *** P < 0.001) Note that the two regression lines and scatter points are too close to be distinguished by the naked eye.

Extended Data Fig. 7 Relationships between mature plant root diameter and seedling root diameter of a same plant species.

Pearson correlation is conducted using our field-measured data and literature data (Siqueira and Saggin-Júnior28). There is a positive relationship between root diameter of mature plants and root diameter of seedlings (y = 1.04x + 0.02, r = 0.89, P = 2.14 × 10−12, n = 39). Shaded areas indicates 95% confidence intervals of the regression lines. Significance was tested using a two-sided t-test. Each point represents one species.

Extended Data Fig. 8 Indirect evidence supporting the Pathogen Resistance Hypothesis.

Analyses were performed using field-measured data of this study and literature data. a, there are differences in arbuscular mycorrhizal (AM) colonization rates across different climatic zones, with colonization rates in the tropics (blue) and subtropics (red) being higher than colonization rates in the temperate zone (green) (n = 237; *** P < 0.001). Pearson correlation indicates a positive correlation between soil pathogen richness and both root diameter and cortical thickness in arbuscular mycorrhizal (AM) plant species (b, y = 208.43x−22.41, r = 0.67, P = 6.59 × 10−9; d, y = 1.43x−70.55, r = 0.85, P = 2.2 × 10−16), but not in ectomycorrhizal (ECM) plant species (c, r = 0.14, P = 0.25; e, r = 0.17, P = 0.20). Shaded areas indicates 95% confidence intervals of the regression lines. The soil pathogen richness here were obtained through sequencing of soil samples collected from XSBN, DHS, SNJ, JGS, CBS, HZ sites (see the Methods section for details of these sites). The arbuscular mycorrhizal colonization data were collected from the GRooT database. Violin plots indicate the median value (solid line), the 25th and 75th percentiles (box), and the density of the data (width of the violin).

Extended Data Fig. 9 Relationships between arbuscular mycorrhizal colonization rates and root diameter in woody and non-woody plants.

Analyses were performed using field-measured data (a), as well as data from GRooT and literature (b, c). a, root diameter is positively related with seed mass in temperate non-woody species using the standardized major axis (SMA) regression (non-woody; a, log10(y) = 3.94 log10(x) + 1.01, r = 0.46, P = 2.55 × 10-11, 95% CI = 3.46−4.50, n = 285), but not in temperate woody species (woody, a, r = 0.06, P = 0.32, n = 284). b, Pearson correlation indicates a significantly positive relationship between arbuscular mycorrhizal (AM) colonization and root diameter in both woody and non-woody plants (woody, y = 55.88x + 33.25, r = 0.34, P = 9.66 × 10-7, n = 202; non-woody, y = 57.24x + 1.44, r = 0.53, P = 0.001, n = 32). Shaded areas indicates 95% confidence intervals of the regression lines. Mixed linear model showed no effect of plant growth form (woody versus non-woody) on the linear regression between arbuscular mycorrhizal colonization and root diameter (P = 0.51). c, boxplots of AM colonization rates by plant growth form; the red points represent woody plants, and the blue points represent non-woody plants. Horizontal lines in the middle of the box plots are the median AM colonization rates. There is a difference in AM colonization rate between woody and non-woody plants (n = 234, P = 0.003, F = 50.57****). Each point representing one species. Box plots indicate the median value (solid line), 25th and 75th percentiles (box), and the data range (whiskers).

Extended Data Table 1 Summary of sampling information for all Chinese field sites where root traits were measured

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Yang, Q., Guo, B., Lu, M. et al. Arbuscular mycorrhizal association regulates global root–seed coordination. Nat. Plants 11, 1759–1768 (2025). https://doi.org/10.1038/s41477-025-02089-4

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