- Open Access
Soil Texture Regulates Bacterial Motility and Chemotactic Recruitment to Plant Roots
PRX Life 4, 013034 – Published 19 March, 2026
DOI: https://doi.org/10.1103/56qk-s5zw
Abstract
Soil microbial communities regulate critical ecological processes, including nutrient cycling, carbon sequestration, and plant growth. However, due to the opacity and structural complexity of soil, how physical constraints imposed by pore geometry influence bacterial motility and chemotactic recruitment to plant roots remains poorly understood. We use a transparent soil mimic composed of cryolite grains that replicates the structural characteristics of natural soils while enabling direct visualization of bacterial dynamics. Using Escherichia coli as a model bacterium, we combine macroscopic spreading assays with microscopic tracking of cellular trajectories to characterize how soil texture affects motility across pore scales. We find that bacterial motility shifts from run-and-tumble behavior in large, open pores to frequent trapping in smaller, more confined spaces. This transition is governed by the pore size distribution and leads to reduced effective diffusivity and slower population-scale spreading. Moreover, pore-scale confinement hinders the chemotactic recruitment of bacteria to Arabidopsis thaliana roots: recruitment is robust in sandy and loamy soils but negligible in highly confining textures. Our results establish soil texture as a critical factor regulating microbial dynamics and ecological interactions in the rhizosphere. This mechanistic understanding complements genomic surveys by identifying physical confinement as an ecological filter that shapes root-associated microbiomes. These findings highlight the essential and previously underappreciated role of soil texture, suggesting new strategies for managing microbial communities to promote plant health and sustainable agriculture.
Physics Subject Headings (PhySH)
Article Text
Supplemental Material
References (69)
- D. C. Coleman, C. P. P. Reid, and C. V. Cole, Biological strategies of nutrient cycling in soil systems, Adv. Ecol. Res. 13, 1 (1983).
- M. M. M. Kuypers, H. K. Marchant, and B. Kartal, The microbial nitrogen-cycling network, Nat. Rev. Microbiol. 16, 263 (2018).
- P. G. Falkowski, T. Fenchel, and E. F. DeLong, The microbial engines that drive earth's biogeochemical cycles, Science 320, 1034 (2008).
- J. Six, S. D. Frey, R. K. Thiet, and K. M. Batten, Bacterial and fungal contributions to carbon sequestration in agroecosystems, Soil Sci. Soc. Am. J. 70, 555 (2006).
- W. R. Wieder, G. B. Bonan, and S. D. Allison, Global soil carbon projections are improved by microbial processes, Nat. Clim. Change 3, 909 (2013).
- M. W. I. Schmidt, M. S. Torn, S. Abiven, T. Dittmar, G. Guggenberger, I. A. Janssens, M. Kleber, I. Kögel-Knabner, J. Lehmann, D. A. C. Manning, P. Nannipieri, D. P. Rasse, S. Weiner, and S. E. Trumbore, Persistence of soil organic matter as an ecosystem property, Nature (London) 478, 49 (2011).
- S. Compant, C. Clément, and A. Sessitsch, Plant growth-promoting bacteria in the rhizo- and endosphere of plants: Their role, colonization, mechanisms involved and prospects for utilization, Soil Biol. Biochem. 42, 669 (2010).
- R. L. Berendsen, C. M. J. Pieterse, and P. A. H. M. Bakker, The rhizosphere microbiome and plant health, Trends Plant Sci. 17, 478 (2012).
- R. Mendes, M. Kruijt, I. de Bruijn, E. Dekkers, M. van der Voort, J. H. M. Schneider, Y. M. Piceno, T. Z. DeSantis, G. L. Andersen, P. A. H. M. Bakker, and J. M. Raaijmakers, Deciphering the rhizosphere microbiome for disease-suppressive bacteria, Science 332, 1097 (2011).
- M. Hartmann and J. Six, Soil structure and microbiome functions in agroecosystems, Nat. Rev. Earth Environ. 4, 4 (2022).
- T. W. Crowther, J. van den Hoogen, J. Wan, M. A. Mayes, A. D. Keiser, L. Mo, C. Averill, and D. S. Maynard, The global soil community and its influence on biogeochemistry, Science 365, eaav0550 (2019).
- R. D. Bardgett and W. H. van der Putten, Belowground biodiversity and ecosystem functioning, Nature (London) 515, 505 (2014).
- M. G. A. van der Heijden, R. D. Bardgett, and N. M. van Straalen, The unseen majority: Soil microbes as drivers of plant diversity and productivity in terrestrial ecosystems, Ecol. Lett. 11, 296 (2008).
- C. Franche, K. Lindström, and C. Elmerich, Nitrogen-fixing bacteria associated with leguminous and non-leguminous plants, Plant Soil 321, 35 (2009).
- D. Haas and G. Défago, Biological control of soil-borne pathogens by fluorescent pseudomonads, Nat. Rev. Microbiol. 3, 307 (2005).
- O. M. Finkel, I. Salas-González, G. Castrillo, J. M. Conway, T. F. Law, P. J. P. L. Teixeira, E. D. Wilson, C. R. Fitzpatrick, C. D. Jones, and J. L. Dangl, A single bacterial genus maintains root growth in a complex microbiome, Nature (London) 587, 103 (2020).
- N. Zhang, D. Wang, Y. Liu, S. Li, Q. Shen, and R. Zhang, Effects of different plant root exudates and their organic acid components on chemotaxis, biofilm formation and colonization by beneficial rhizosphere-associated bacterial strains, Plant Soil 374, 689 (2014).
- H. P. Bais, T. L. Weir, L. G. Perry, S. Gilroy, and J. M. Vivanco, The role of root exudates in rhizosphere interactions with plants and other organisms, Annu. Rev. Plant Biol. 57, 233 (2006).
- J. Ramoneda, K. Fan, J. M. Lucas, H. Chu, A. Bissett, M. S. Strickland, and N. Fierer, Ecological relevance of flagellar motility in soil bacterial communities, ISME J. 18, wrae067 (2024).
- B. Karimi, S. Terrat, S. Dequiedt, N. P. A. Saby, W. Horrigue, M. Lelièvre, V. Nowak, C. Jolivet, D. Arrouays, P. Wincker, C. Cruaud, A. Bispo, P.-A. Maron, N. C. P. Bouré, and L. Ranjard, Biogeography of soil bacteria and archaea across France, Sci. Adv. 4, eaat1808 (2018).
- E. M. Bach, R. J. Williams, S. K. Hargreaves, F. Yang, and K. S. Hofmockel, Greatest soil microbial diversity found in micro-habitats, Soil Biol. Biochem. 118, 217 (2018).
- Q. Xia, T. Rufty, and W. Shi, Soil microbial diversity and composition: Links to soil texture and associated properties, Soil Biol. Biochem. 149, 107953 (2020).
- J. K. Jansson and K. S. Hofmockel, The soil microbiome—From metagenomics to metaphenomics, Curr. Opin. Microbiol. 43, 162 (2018).
- K. Zhalnina, K. B. Louie, Z. Hao, N. Mansoori, U. N. da Rocha, S. Shi, H. Cho, U. Karaoz, D. Loqué, B. P. Bowen, M. K. Firestone, T. R. Northen, and E. L. Brodie, Dynamic root exudate chemistry and microbial substrate preferences drive patterns in rhizosphere microbial community assembly, Nat. Microbiol. 3, 470 (2018).
- H. Toju, K. G. Peay, M. Yamamichi, K. Narisawa, K. Hiruma, K. Naito, S. Fukuda, M. Ushio, S. Nakaoka, Y. Onoda, K. Yoshida, K. Schlaeppi, Y. Bai, R. Sugiura, Y. Ichihashi, K. Minamisawa, and E. T. Kiers, Core microbiomes for sustainable agroecosystems, Nat. Plants 4, 247 (2018).
- I. Arif, M. Batool, and P. M. Schenk, Plant microbiome engineering: Expected benefits for improved crop growth and resilience, Trends Biotechnol. 38, 1385 (2020).
- C. Luo, Y. He, and Y. Chen, Rhizosphere microbiome regulation: Unlocking the potential for plant growth, Curr. Res. Microbiol. Sci. 8, 100322 (2025).
- A. Erktan, D. Or, and S. Scheu, The physical structure of soil: Determinant and consequence of trophic interactions, Soil Biol. Biochem. 148, 107876 (2020).
- M. Lucas, J. P. Santiago, J. Chen, A. Guber, and A. Kravchenko, The soil pore structure encountered by roots affects plant-derived carbon inputs and fate, New Phytol. 240, 515 (2023).
- N. Fierer and R. B. Jackson, The diversity and biogeography of soil bacterial communities, Proc. Natl. Acad. Sci. USA 103, 626 (2006).
- M. F. A. Leite, S. W. E. B. van den Broek, and E. E. Kuramae, Current challenges and pitfalls in soil metagenomics, Microorganisms 10, 1900 (2022).
- D. Garg, N. Patel, A. Rawat, and A. S. Rosado, Cutting edge tools in the field of soil microbiology, Curr. Res. Microbiol. Sci. 6, 100226 (2024).
- T. Bhattacharjee and S. S. Datta, Bacterial hopping and trapping in porous media, Nat. Commun. 10, 2075 (2019).
- T. Bhattacharjee and S. S. Datta, Confinement and activity regulate bacterial motion in porous media, Soft Matter 15, 9920 (2019).
- C. Kurzthaler, S. Mandal, T. Bhattacharjee, H. Löwen, S. S. Datta, and H. A. Stone, A geometric criterion for the optimal spreading of active polymers in porous media, Nat. Commun. 12, 7088 (2021).
- T. Bhattacharjee, D. B. Amchin, J. A. Ott, F. Kratz, and S. S. Datta, Chemotactic migration of bacteria in porous media, Biophys. J. 120, 3483 (2021).
- T. Bhattacharjee, D. B. Amchin, R. Alert, J. A. Ott, and S. S. Datta, Chemotactic smoothing of collective migration, eLife 11, e71226 (2022).
- H. C. Berg and D. A. Brown, Chemotaxis in Escherichia coli analysed by three-dimensional tracking, Nature (London) 239, 500 (1972).
- S. H. Larsen, R. W. Reader, E. N. Kort, W.-W. Tso, and J. Adler, Change in direction of flagellar rotation is the basis of the chemotactic response in Escherichia coli, Nature (London) 249, 74 (1974).
- H. Zhang, M. T. Wetherington, H. Ko, C. E. FitzGerald, L. V. Luzzatto, I. A. Kovács, E. M. Munro, and J. A. Nirody, Bacterial motility patterns vary smoothly with spatial confinement and disorder, PRX Life 4, 013002 (2026).
- T. V. Phan, R. Morris, M. E. Black, T. K. Do, K.-C. Lin, K. Nagy, J. C. Sturm, J. Bos, and R. H. Austin, Bacterial route finding and collective escape in mazes and fractals, Phys. Rev. X 10, 031017 (2020).
- K. Sharma, M. Palatinszky, G. Nikolov, D. Berry, and E. A. Shank, Transparent soil microcosms for live-cell imaging and non-destructive stable isotope probing of soil microorganisms, eLife 9, e56275 (2020).
- A. Mair, D. Patko, Y. Liu, I. Engelhardt, and L. X. Dupuy, Synthetic soil microcosms as emerging model systems to study the rhizosphere, Plant Soil 514, 1597 (2025).
- Y. Liu, D. Patko, I. Engelhardt, T. S. George, N. R. Stanley-Wall, V. Ladmiral, B. Ameduri, T. J. Daniell, N. Holden, M. P. MacDonald, and L. X. Dupuy, Plant-environment microscopy tracks interactions of Bacillus subtilis with plant roots across the entire rhizosphere, Proc. Natl. Acad. Sci. USA 118, e2109176118 (2021).
- I. C. Engelhardt, N. Holden, T. J. Daniell, and L. X. Dupuy, Mobility and growth in confined spaces are important mechanisms for the establishment of Bacillus subtilis in the rhizosphere, Microbiology 170, 001477 (2024).
- L. M. Rooney, L. X. Dupuy, P. A. Hoskisson, and G. McConnell, Construction and characterisation of a structured, tuneable, and transparent 3D culture platform for soil bacteria, Microbiology 170, 001429 (2024).
- H. Pauly, Cryolite, chiolite and cryolithionite: Optical data redetermined, Bull. Geol. Soc. Den. 26, 95 (1977).
- D. C. Coleman, M. A. Callaham, and D. A. Crossley, Introduction to soil, in Fundamentals of Soil Ecology (Elsevier, 2018), pp. 1–20.
- See Supplemental Material at http://link.aps.org/supplemental/10.1103/56qk-s5zw for supplemental figures and methods.
- S. Torquato and B. Lu, Chord-length distribution function for two-phase random media, Phys. Rev. E 47, 2950 (1993).
- S. Torquato, Random Heterogeneous Materials (Springer, New York, 2002).
- J. Schwarz-Linek, J. Arlt, A. Jepson, A. Dawson, T. Vissers, D. Miroli, T. Pilizota, V. A. Martinez, and W. C. K. Poon, Escherichia coli as a model active colloid: A practical introduction, Colloids Surf. B 137, 2 (2016).
- B. V. Hokmabad, A. Martínez-Calvo, S. G. La Corte, and S. S. Datta, Spatial self-organization of confined bacterial suspensions, Proc. Natl. Acad. Sci. USA 122, e2503983122 (2025).
- D. Valverde-Méndez, A. M. Sunol, B. P. Bratton, M. Delarue, J. L. Hofmann, J. P. Sheehan, Z. Gitai, L. J. Holt, J. W. Shaevitz, and R. N. Zia, Macromolecular interactions and geometrical confinement determine the 3d diffusion of ribosome-sized particles in live Escherichia coli cells, Proc. Natl. Acad. Sci. USA 122, e2406340121 (2025).
- N. Wadhwa and H. C. Berg, Bacterial motility: Machinery and mechanisms, Nat. Rev. Microbiol. 20, 161 (2022).
- R. Colin, B. Ni, L. Laganenka, and V. Sourjik, Multiple functions of flagellar motility and chemotaxis in bacterial physiology, FEMS Microbiol. Rev. 45, fuab038 (2021).
- Y. Liu, Z. Xu, L. Chen, W. Xun, X. Shu, Y. Chen, X. Sun, Z. Wang, Y. Ren, Q. Shen, and R. Zhang, Root colonization by beneficial rhizobacteria, FEMS Microbiol. Rev. 48, fuad066 (2024).
- R. Tecon and D. Or, Biophysical processes supporting the diversity of microbial life in soil, FEMS Microbiol. Rev. 41, 599 (2017).
- G. M. N. Benucci, E. R. Toosi, F. Yang, T. L. Marsh, G. M. Bonito, and A. Kravchenko, The microbiome structure of decomposing plant leaves in soil depends on plant species, soil pore sizes, and soil moisture content, Front. Microbiol. 14, 1172862 (2023).
- L. Ma, Y. Shi, O. Siemianowski, B. Yuan, T. K. Egner, S. V. Mirnezami, K. R. Lind, B. Ganapathysubramanian, V. Venditti, and L. Cademartiri, Hydrogel-based transparent soils for root phenotyping in vivo, Proc Natl. Acad. Sci. USA 116, 11063 (2019).
- A. Canarini, C. Kaiser, A. Merchant, A. Richter, and W. Wanek, Root exudation of primary metabolites: Mechanisms and their roles in plant responses to environmental stimuli, Front. Plant Sci. 10, 157 (2019).
- J. M. Chaparro, D. V. Badri, M. G. Bakker, A. Sugiyama, D. K. Manter, and J. M. Vivanco, Root exudation of phytochemicals in Arabidopsis follows specific patterns that are developmentally and correlate with soil microbial functions, PLoS ONE 8, e55731 (2013).
- N. Strehmel, C. Böttcher, S. Schmidt, and D. Scheel, Profiling of secondary metabolites in root exudates of Arabidopsis thaliana, Phytochemistry 108, 35 (2014).
- R. Mesibov and J. Adler, Chemotaxis toward amino acids in Escherichia coli, J. Bacteriol. 112, 315 (1972).
- S. Neumann, C. H. Hansen, N. S. Wingreen, and V. Sourjik, Differences in by directly and indirectly binding ligands in bacterial chemotaxis, EMBO J. 29, 3484 (2010).
- M. de Los Angeles Dublan, J. C. F. Ortiz-Marquez, L. Lett, and L. Curatti, Plant-adapted Escherichia coli show increased lettuce colonizing ability, resistance to oxidative stress and chemotactic response, PLoS ONE 9, e110416 (2014).
- C. R. Fitzpatrick, I. Salas-González, J. M. Conway, O. M. Finkel, S. Gilbert, D. Russ, P. J. P. L. Teixeira, and J. L. Dangl, The plant microbiome: From ecology to reductionism and beyond, Annu. Rev. Microbiol. 74, 81 (2020).
- M. Bunsick, S. Toh, C. Wong, Z. Xu, G. Ly, C. S. P. McErlean, G. Pescetto, K. E. Nemrish, P. Sung, J. D. Li, J. D. Scholes, and S. Lumba, Smax1-dependent seed germination bypasses ga in Arabidopsis and Striga, Nat. Plants 6, 646 (2020).
- A. Al Harraq, Data for “Soil texture regulates bacterial motility and chemotactic recruitment to plant roots,” https://doi.org/10.34770/m5g3-nf94 (2025).