Root structural remodeling under soil compaction for herbaceous plants

doi:10.1016/j.pld.2025.12.008

Plant Diversity ›› 2026, Vol. 48 ›› Issue (01): 128-139.DOI: 10.1016/j.pld.2025.12.008

Root structural remodeling under soil compaction for herbaceous plants

Qinwen Han^a, Qingpei Yang^a, Binglin Guo^a, Tino Colombi^b, Junjian Wang^c,d, Huifang Wu^a, Zhipei Feng^a, Zhi Zheng^e, Zhenjiang Li^a, Yue Zhang^a, Meixu Han^a, Qiang Li^a, Junxiang Ding^f, Xitian Yang^a, Hannah M. Schneider^g, Ying Zhao^h, Deliang Kong^a,i

a College of Forestry, Henan Agricultural University, Zhengzhou 450002, China;
b School of Biosciences, University of Nottingham, Sutton Bonington, LE12 5RD, United Kingdom;
c State Key Laboratory of Soil Pollution Control and Safety, Southern University of Science and Technology, Shenzhen 518055, China;
d Guangdong Provincial Key Laboratory of Soil and Groundwater Pollution Control, School of Environmental Science and Engineering, Southern University of Science and Technology, Shenzhen 518055, China;
e College of Life Science, Northwest Normal University, Lanzhou, China;
f College of Ecology and Environment, Zhengzhou University, Zhengzhou 450001, China;
g Department of Physiology and Cell Biology, Leibniz Institute for Plant Genetics and Crop Plant Research (IPK), OT Gatersleben, Seeland 06466, Germany;
h. College of Resources and Environmental Engineering, Ludong University, Yantai 264025, China;
i. Henan Province Engineering Research Center of Crop Synthetic Biology, Henan Agricultural University, Zhengzhou 450046, China

收稿日期:2025-08-02 修回日期:2025-12-14 出版日期:2026-01-25 发布日期:2026-03-05
通讯作者: Qinwen Han,E-mail:hanqw0111@163.com;Qingpei Yang,E-mail:yangqingpei1992@126.com;Binglin Guo,E-mail:3113142671@qq.com;Tino Colombi,E-mail:Tino.Colombi@nottingham.ac.uk;Junjian Wang,E-mail:wangjj@sustech.edu.cn;Huifang Wu,E-mail:huifangwu1999@126.com;Zhipei Feng,E-mail:fzp@henau.edu.cn;Zhi Zheng,E-mail:zhengzhi1982@126.com;Zhenjiang Li,E-mail:18336457322@163.com;Yue Zhang,E-mail:zhangyue5724@163.com;Meixu Han,E-mail:h20219898@126.com;Qiang Li,E-mail:liqiang@henau.edu.cn;Junxiang Ding,E-mail:dingjunxiang@zzu.edu.cn;Xitian Yang,E-mail:yangxt@henau.edu.cn;Hannah M. Schneider,E-mail:schneiderh@ipk-gatersleben.de;Ying Zhao,E-mail:yzhaosoils@gmail.com;Deliang Kong,E-mail:deliangkong1999@126.com
基金资助:
This study was funded by the National Natural Science Foundation of China (32471824, 32171746, 31870522, 42477227, and 32560282), the leading talents of basic research in Henan Province (24XM0375), Excellent Youth Creative Research Group Project in Henan Province (252300421002), Foreign Scientists Studio in Henan Province (GZS2025011), MOHRSS National Foreign Expert Individual Projectsand (110000264820258001) and Natural Science Foundation of Henan (242300420604). Tino Colombi acknowledges the University of Nottingham for funding (Nottingham Research Fellowship). Junjian Wang was supported by the Guangdong Provincial Key Laboratory of Soil and Groundwater Pollution Control (2023B1212060002) and the High-level University Special Fund (G03050K001). Zhipei Feng acknowledges the China Postdoctoral Science Foundation (No. 2021M690922).

Root structural remodeling under soil compaction for herbaceous plants

a College of Forestry, Henan Agricultural University, Zhengzhou 450002, China;
b School of Biosciences, University of Nottingham, Sutton Bonington, LE12 5RD, United Kingdom;
c State Key Laboratory of Soil Pollution Control and Safety, Southern University of Science and Technology, Shenzhen 518055, China;
d Guangdong Provincial Key Laboratory of Soil and Groundwater Pollution Control, School of Environmental Science and Engineering, Southern University of Science and Technology, Shenzhen 518055, China;
e College of Life Science, Northwest Normal University, Lanzhou, China;
f College of Ecology and Environment, Zhengzhou University, Zhengzhou 450001, China;
g Department of Physiology and Cell Biology, Leibniz Institute for Plant Genetics and Crop Plant Research (IPK), OT Gatersleben, Seeland 06466, Germany;
h. College of Resources and Environmental Engineering, Ludong University, Yantai 264025, China;
i. Henan Province Engineering Research Center of Crop Synthetic Biology, Henan Agricultural University, Zhengzhou 450046, China

Received:2025-08-02 Revised:2025-12-14 Online:2026-01-25 Published:2026-03-05
Contact: Qinwen Han,E-mail:hanqw0111@163.com;Qingpei Yang,E-mail:yangqingpei1992@126.com;Binglin Guo,E-mail:3113142671@qq.com;Tino Colombi,E-mail:Tino.Colombi@nottingham.ac.uk;Junjian Wang,E-mail:wangjj@sustech.edu.cn;Huifang Wu,E-mail:huifangwu1999@126.com;Zhipei Feng,E-mail:fzp@henau.edu.cn;Zhi Zheng,E-mail:zhengzhi1982@126.com;Zhenjiang Li,E-mail:18336457322@163.com;Yue Zhang,E-mail:zhangyue5724@163.com;Meixu Han,E-mail:h20219898@126.com;Qiang Li,E-mail:liqiang@henau.edu.cn;Junxiang Ding,E-mail:dingjunxiang@zzu.edu.cn;Xitian Yang,E-mail:yangxt@henau.edu.cn;Hannah M. Schneider,E-mail:schneiderh@ipk-gatersleben.de;Ying Zhao,E-mail:yzhaosoils@gmail.com;Deliang Kong,E-mail:deliangkong1999@126.com
Supported by:
This study was funded by the National Natural Science Foundation of China (32471824, 32171746, 31870522, 42477227, and 32560282), the leading talents of basic research in Henan Province (24XM0375), Excellent Youth Creative Research Group Project in Henan Province (252300421002), Foreign Scientists Studio in Henan Province (GZS2025011), MOHRSS National Foreign Expert Individual Projectsand (110000264820258001) and Natural Science Foundation of Henan (242300420604). Tino Colombi acknowledges the University of Nottingham for funding (Nottingham Research Fellowship). Junjian Wang was supported by the Guangdong Provincial Key Laboratory of Soil and Groundwater Pollution Control (2023B1212060002) and the High-level University Special Fund (G03050K001). Zhipei Feng acknowledges the China Postdoctoral Science Foundation (No. 2021M690922).

摘要/Abstract

摘要： Soil compaction often imposes stress on root development and plant survival. However, root anatomical responses that enable persistent root growth and functioning under soil compaction remain unclear. We grew 10 herbaceous species differing substantially in lateral root diameter, in soils with low (1.0 g cm^-3) and high (1.4 g cm^-3) bulk density, and assessed root traits including root biomass, anatomical structures, and respiration rates. Greater root thickening upon soil compaction was found in species with thicker first-order lateral roots, mainly due to larger cortical cell size. Both xylem vessel diameter and wall thickness increased more in compacted soils in these species. Despite these anatomical shifts, root respiration rate responded little to soil compaction across most species, likely due to the opposite investment in cortical cells and xylem vessels. Notably, root biomass, independent of root respiration rate and anatomical structures, determined whole-plant growth under soil compaction. Our study reveals two independent strategies of root response to soil compaction: anatomical remodeling for mechanical and metabolic maintenance, and root biomass investment for resource acquisition. These findings offer new insights for breeding and selecting species tolerant to soil compaction and highlight multidimensional strategies of plant adaptation to physical stress.

关键词: Root anatomy, Root respiration rate, Soil compaction, Cortex, Xylem vessel, Root biomass

Abstract: Soil compaction often imposes stress on root development and plant survival. However, root anatomical responses that enable persistent root growth and functioning under soil compaction remain unclear. We grew 10 herbaceous species differing substantially in lateral root diameter, in soils with low (1.0 g cm^-3) and high (1.4 g cm^-3) bulk density, and assessed root traits including root biomass, anatomical structures, and respiration rates. Greater root thickening upon soil compaction was found in species with thicker first-order lateral roots, mainly due to larger cortical cell size. Both xylem vessel diameter and wall thickness increased more in compacted soils in these species. Despite these anatomical shifts, root respiration rate responded little to soil compaction across most species, likely due to the opposite investment in cortical cells and xylem vessels. Notably, root biomass, independent of root respiration rate and anatomical structures, determined whole-plant growth under soil compaction. Our study reveals two independent strategies of root response to soil compaction: anatomical remodeling for mechanical and metabolic maintenance, and root biomass investment for resource acquisition. These findings offer new insights for breeding and selecting species tolerant to soil compaction and highlight multidimensional strategies of plant adaptation to physical stress.

Key words: Root anatomy, Root respiration rate, Soil compaction, Cortex, Xylem vessel, Root biomass

Qinwen Han, Qingpei Yang, Binglin Guo, Tino Colombi, Junjian Wang, Huifang Wu, Zhipei Feng, Zhi Zheng, Zhenjiang Li, Yue Zhang, Meixu Han, Qiang Li, Junxiang Ding, Xitian Yang, Hannah M. Schneider, Ying Zhao, Deliang Kong. Root structural remodeling under soil compaction for herbaceous plants[J]. Plant Diversity, 2026, 48(01): 128-139.

参考文献

Ahmed, MA., Zarebanadkouki, M., Kaestner, A., et al., 2016. Measurements of water uptake of maize roots: the key function of lateral roots. Plant Soil 398, 59-77.
Ahmed, MA., Zarebanadkouki, M., Meunier, F., et al., 2018. Root type matters: measurement of water uptake by seminal, crown, and lateral roots in maize. Journal of Experimental Botany 69, 1199-1206.
Bello-Bello, E., Lopez-Arredondo, D., Rico-Chambron, T.Y., et al., 2022. Conquering compacted soils: uncovering the molecular components of root soil penetration. Trends in Plant Science 27, 814-827.
Berisso, F.E., Schjoenning, P., Keller, T., et al., 2012. Persistent effects of subsoil compaction on pore size distribution and gas transport in a loamy soil. Soil and Tillage Research 122, 42-51.
Cao, J-J., Chen, J., Yang Q., et al., 2023. Leaf hydraulics coordinated with leaf economics and leaf size in mangrove species along a salinity gradient. Plant Diversity 45, 309-314.
Cao, J., Wang, J., Yang, Q., et al., 2025. Root anatomy governs bi-directional resource transfer in mycorrhizal symbiosis. Nat Commun 16, 8731.
Chen, J., Cao, J., Guo, B., et al., 2025. Increased dependence on mycorrhizal fungi for nutrient acquisition under carbon limitation by tree girdling. Plant Diversity 47, 466-478.
Chimungu, J.G., Loades, K.W., Lynch, J.P., 2015. Root anatomical phenes predict root penetration ability and biomechanical properties in maize (Zea Mays). Journal of Experimental Botany 66, 3151-3162.
Chimungu, J., Brown, K., Lynch, J., 2014. Large Root Cortical Cell Size Improves Drought Tolerance in Maize. Plant Physiology 166, 2166-2178.
Colombi, T., Herrmann, A.M., Vallenback, P., et al., 2019. Cortical Cell Diameter Is Key To Energy Costs of Root Growth in Wheat. Plant Physiology 180, 2049-2060.
Colombi, T., Keller, T., 2019. Developing strategies to recover crop productivity after soil compaction-A plant eco-physiological perspective. Soil and Tillage Research 191, 156-161.
Corner, E. J. H., 1949. The Durian theory or the origin of the modern tree. Ann. Bot 13, 367-414.
Cui, Y.N., Man, Y., Song, C.W., et al., 2020. Chemical components, physiological functions and regulation mechanism of plant Casparian strips (in Chinese). Sci Sin-Vitae 50, 102-110.
Frene, J.P., Pandey, B.K., Castrillo, G., et al., 2024. Under pressure: elucidating soil compaction and its effect on soil functions. Plant and Soil 502, 267-278.
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, 1247-1260.
Freschet, G.T., Pages, L., Iversen, C.M., et al., 2021. A starting guide to root ecology: strengthening ecological concepts and standardising root classification, sampling, processing and trait measurements. New Phytologist 232, 973-1122.
Fujikawa, T., Miyazaki, T., 2005. EFFECTS OF BULK DENSITY AND SOIL TYPE ON THE GAS DIFFUSION COEFFICIENT IN REPACKED AND UNDISTURBED SOILS. Soil Science 170.
Gao, Y.Q., Su, Y., Chao, D.Y., 2024. Exploring the function of plant root diffusion barriers in sealing and shielding for environmental adaptation. Nature Plants 10, 1865-1874.
Gleason, S., Westoby, M., Jansen, S., et al., 2015. Weak tradeoff between xylem safety and xylem-specific hydraulic efficiency across the world's woody plant species. New Phytologist 209, 123-136.
Goldberg-Yehuda, N., Nachshon, U., Assouline, S., et al., 2024. The effect of mechanical compaction on the soil water retention curve: Insights from a rapid image analysis of micro-CT scanning. CATENA 242, 108068.
Guet, J., Fichot, R., Ledee, C., et al., 2015. Stem xylem resistance to cavitation is related to xylem structure but not to growth and water-use efficiency at the within-population level in Populus nigra L. Journal of Experimental Botany 66, 4643-4652.
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, 673-683.
Hamza, M.A., Anderson, W.K., 2005. Soil compaction in cropping systems: A review of the nature, causes and possible solutions. Soil and Tillage Research 82, 121-145.
Han, M, Zhang, H, Liu, M, et al., 2024. Increased dependence on nitrogen-fixation of a native legume in competition with an invasive plant. Plant Diversity 46, 510-518.
Han, Q., Yang, Q., Guo, B., et al., 2024. Linking root cell wall width with plant functioning under drought conditions. Journal of Experimental Botany 75, 5463-5466.
Han, M., Zhu, B., 2021. Linking root respiration to chemistry and morphology across species. Global Change Biology 27(1):190-201.
Herbin, T., Hennessy, D., Richards, K.G., et al., 2011. The effects of dairy cow weight on selected soil physical properties indicative of compaction. Soil Use and Management 27, 36-44.
Hou, J., McCormack, M.L., Reich, P.B., et al., 2024. Linking fine root lifespan to root chemical and morphological traits-A global analysis. Proceedings of the National Academy of Sciences 121, e2320623121.
Huang, G., Kilic, A., Karady, M., et al., 2022. Ethylene inhibits rice root elongation in compacted soil via ABA- and auxin-mediated mechanisms. Proceedings of the National Academy of Sciences 119, e2201072119.
Kalmbach, L., Hematy, K., De, Bellis, D., et al., 2017. Transient cell-specific EXO70A1 activity in the CASP domain and Casparian strip localization. Nature Plants 3, 17058.
Keller, T., Or, D., 2022. Farm vehicles approaching weights of sauropods exceed safe mechanical limits for soil functioning. Proceedings of the National Academy of Sciences 119, e2117699119.
Kirfel, K., Leuschner, C., Hertel, D., et al., 2017. Influence of Root Diameter and Soil Depth on the Xylem Anatomy of Fine- to Medium-Sized Roots of Mature Beech Trees in the Top- and Subsoil. Frontiers in Plant Science 8, 1194.
Kong, D., Ma, C., Zhang, Q., et al., 2014. Leading dimensions in absorptive root trait variation across 96 subtropical forest species. New Phytologist 203, 863-872.
Kong, D., Wang, J., Zeng, H., et al., 2017. The nutrient absorption transportation hypothesis: optimizing structural traits in absorptive roots. New Phytologist 213, 1569-1572.
Kong, D., Wang, J., Wu, H., et al., 2019. Nonlinearity of root trait relationships and the root economics spectrum. Nature Communications 10, 2203.
Kong, D., Wang, J., Valverde-Barrantes, O.J., et al., 2021. A framework to assess the carbon supply–consumption balance in plant roots. New Phytologist 229, 659-664.

10.1079/9780851994321.0031 Leishman, M., Wright, I., Moles, A., et al., 2000. in Seeds: The Ecology of Regeneration in Plant Communities (ed. Fenner, M.). CABI, 31-57.
Liang, S., Guo, H., McCormack, M.L., et al., 2023. Positioning absorptive root respiration in the root economics space across woody and herbaceous species. Journal of Ecology 111, 2710-2720.
Li, H., Liu, B., McCormack, M.L., et al., 2017. Diverse belowground resource strategies underlie plant species coexistence and spatial distribution in three grasslands along a precipitation gradient. New Phytologist 216, 1140-1150.
Li, L., McCormack, M.L., Ma, C., et al., 2015. Leaf economics and hydraulic traits are decoupled in five species-rich tropical-subtropical forests. Ecology Letters 18, 899-906.
Long, Z., Wang, Y., Sun, B., et al., 2024. Impact of mechanical compaction on crop growth and sustainable agriculture. Front. Agr. Sci. Eng 11, 243-252.
Lynch, J.P., Chimungu, J.G., Brown, K.M., 2014. Root anatomical phenes associated with water acquisition from drying soil: targets for crop improvement. Journal of Experimental Botany 65, 6155-6166.
Ma, Z., Guo, D., Xu, X., et al., 2018. Evolutionary history resolves global organization of root functional traits. Nature 555, 94-97.
Ma, Z., Buckley, T.N., Sack, L., et al., 2025. The determination of leaf size on the basis of developmental traits. New Phytol 246, 461-480.
Makita, N., Hirano, Y., Dannoura, M., et al., 2009. Fine root morphological traits determine variation in root respiration of Quercus serrata. Tree Physiology 29, 579-585.
Mao, Z., Roumet, C., Rossi, L., et al., 2023. Intra- and inter-specific variation in root mechanical traits for twelve herbaceous plants and their link with the root economics space. Oikos 2023, e09032.
McCormack, M.L., Guo, D., Iversen, C.M., et al., 2017. Building a better foundation: improving root-trait measurements to understand and model plant and ecosystem processes. New Phytologist 215, 27-37.
Meijer, G.J., Lynch, J.P., Chimungu, J.G., et al., 2024. Root anatomy and biomechanical properties: improving predictions through root cortical and stele properties. Plant Soil 500, 587-603.
Mo, J., Song, Z., Che, Y., et al., 2024. Effects of aeolian deposition on soil properties and microbial carbon metabolism function in farmland of Songnen Plain, China. Scientific Reports 14, 14791.
Nawaz, M.F., Bourrie, G., Trolard, F., 2013. Soil compaction impact and modelling. A review. Agronomy for Sustainable Development 33, 291-309.
Nazari, M., Arthur, E., Lamandé, M., et al., 2023. A Meta-analysis of Soil Susceptibility to Machinery-Induced Compaction in Forest Ecosystems Across Global Climatic Zones. Curr. For. Rep. 9, 370–381.
Olson, M., Rosell, J., 2012. Vessel diameter-stem diameter scaling across woody angiosperms and the ecological causes of xylem vessel diameter variation. New Phytologist 197, 1204-1213.
Pandey, B.K., Bennett, M.J., 2024. Uncovering root compaction response mechanisms: new insights and opportunities. Journal of Experimental Botany 75, 578-583.
Pandey, B.K., Huang, G., Bhosale, R., et al., 2021. Plant roots sense soil compaction through restricted ethylene diffusion. Science 371, 276-280.
Poorter, L., McDonald, I., Alarcon, A., et al., 2010. The importance of wood traits and hydraulic conductance for the performance and life history strategies of 42 rainforest tree species. New Phytologist 185, 481-492.
Pregitzer, K.S., DeForest, J.L., Burton, A.J., et al., 2002. FINE ROOT ARCHITECTURE OF NINE NORTH AMERICAN TREES. Ecological Monographs 72, 293-309.
Schneider, H.M., Strock, C.F., Hanlon, M.T., et al., 2021. Multiseriate cortical sclerenchyma enhance root penetration in compacted soils. Proceedings of the National Academy of Sciences 118, e2012087118.
Shaheb, M.R., Venkatesh, R., Shearer, S.A., 2021. A Review on the Effect of Soil Compaction and its Management for Sustainable Crop Production. Journal of Biosystems Engineering 46, 417-439.
Sidhu, J.S., Lopez-Valdivia, I., Strock, C.F., et al., 2024. Cortical parenchyma wall width regulates root metabolic cost and maize performance under suboptimal water availability. Journal of Experimental Botany 75, 5750-5767.
Soane, B.D., Van, Ouwerkerk, C., 1994. Soil compaction problems in world agriculture. Developments in Agricultural Engineering 11, 1-21.
Song, T., Tian, Y.Q., Liu, C.B., et al., 2023. A new family of proteins is required for tethering of Casparian strip membrane domain and nutrient homoeostasis in rice. Nature Plants 9, 1749-1759.
Tjoelker, M.G., Oleksyn, J., Reich, P.B., 2001. Modelling respiration of vegetation: evidence for a general temperature-dependent Q10. Global Change Biology 7, 223-230.
Tracy, S.R., Black, C.R., Roberts, J.A., et al., 2011. Soil compaction: a review of past and present techniques for investigating effects on root growth. Journal of the Science of Food and Agriculture 91, 1528-1537.
Vanhees, D.J., Schneider, H.M., Sidhu, J.S., et al., 2022. Soil penetration by maize roots is negatively related to ethylene-induced thickening. Plant, Cell & Environment 45, 789-804.
Wen, Z., Li, H., Shen, Q., et al., 2019. Tradeoffs among root morphology, exudation and mycorrhizal symbioses for phosphorus-acquisition strategies of 16 crop species. New Phytologist 223, 882-895.
Wang, M., Kong, D., Mo, X., et al.,2024. Molecular-level carbon traits underlie the multidimensional fine root economics space. Nature Plants 10, 901-909.
Wang, X., Liu, X., Chen, S., et al., 2024. Elevational variation in anatomical traits of the first-order roots and their adaptation mechanisms. Plant Diversity 47, 291-299.
Weigelt, A., Mommer, L., Andraczek, K., et al., 2023. The importance of trait selection in ecology. Nature 618, E29-E30.
Wu, H., Yang, Q., Chen, J., et al., 2025. Divergent leaf-root coordination between mangroves and non-mangroves. Journal of Plant Ecology rtaf023.
Yang, Q., Guo, B., Lu, M., et al., 2025. Arbuscular mycorrhizal association regulates global root-seed coordination. Nat. Plants 11, 1759-1768.
Zarebanadkouki, M., Kim, Y.X., Carminati, A., 2013. Where do roots take up water? Neutron radiography of water flow into the roots of transpiring plants growing in soil. New Phytologist 199, 1034-1044.
Zhang, Y., Cao, J-J., Yang, Q-P., et al., 2023. The worldwide allometric relationship in anatomical structures for plant roots. Plant Diversity 45, 621-629.
Zhang, Y., Cao, J., Lu, M., et al., 2024. The origin of bi-dimensionality in plant root traits. Trends in Ecology & Evolution 39, 78-88.
Zhao, J., Guo, B., Hou, Y., et al., 2024. Multi-dimensionality in plant root traits: progress and challenges. Journal of Plant Ecology 17, rtae043.
Zheng, Z., Dong, F., Li, Z., et al., 2025. The unique root form and function on the Tibetan Plateau. New Phytol 248, 2280-2294.
Zhou, M., Bai, W., Li, Q., et al., 2021. Root anatomical traits determined leaf-level physiology and responses to precipitation change of herbaceous species in a temperate steppe. New Phytologist 229, 1481-1491.
Zhou, M., Guo, Y., Sheng, J., et al., 2022. Using anatomical traits to understand root functions across root orders of herbaceous species in a temperate steppe. New Phytologist 234, 422-434.

Root structural remodeling under soil compaction for herbaceous plants

Root structural remodeling under soil compaction for herbaceous plants

PDF (PC)

可视化

摘要/Abstract

引用本文

使用本文

参考文献

相关文章 1

编辑推荐

Metrics

本文评价