Response of plant roots to mechanical environmental stimuli

Abstract

The mechanisms underlying plant root response to mechanical environmental stimuli are crucial for plant growth, development, and environmental adaptation. In this review, we examine the mechanical environments encountered by plant roots, including the different types of mechanical stimuli they experience. We describe in detail the mechanisms that enable roots to perceive these stimuli and their modes of action. Unfavorable mechanical stimuli can cause roots to alter their growth patterns and rates. Morphologically, roots become thicker, enhancing their stress resistance. Mechanical stimuli influence the activity of hormones, including auxin and ethylene, which jointly regulate root growth. Auxin promotes cell elongation in roots, whereas ethylene can inhibit root growth under certain conditions. Plants modulate antioxidant enzyme activity and osmoregulatory substance accumulation to cope with environmental stress. We explored the molecular regulatory mechanisms underlying plant root adaptation to mechanical stimuli, including those involved in regulating genes and signal transduction pathways. Finally, we suggest future research directions, including an in-depth study of the multi-signal integration mechanism of roots and gene editing technology for improving plant adaptability. This review provides a basis for studying the interactions between plants and mechanical environments for plant adaptation and agricultural production.

Acknowledgments

We thank Xinyan Wang for her continuous professional support of our research, and the Institute of Urban Agriculture, Chinese Academy of Agricultural Sciences for their assistance in research conditions. The article processing charge was paid by the Sichuan Province Science and Technology Innovation Excellence Program for Returned Overseas Talents. This study was funded by Sichuan Province Key R&D Program, International Science and Technology Innovation Cooperation Project, Sichuan Science and Technology Program (2025YFHZ0183), Central Public-interest Scientific Institution Basal Research Fund (No. S2024001), Agricultural Science and Technology Innovation Program (ASTIP-CAAS), and National Key Research and Development Program of China (2023YFF1001500).

Conflicts of interest statement

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

References

[1] Andriuzzi WS, Wall DH. Soil biological responses to, and feedbacks on, trophic rewilding. Philos Trans R Soc Lond Ser B Biol Sci. 2018; 373:20170448

[2] Baiyin B, Tagawa K, Yamada M. et al. Effect of nutrient solution flow rate on hydroponic plant growth and root morphology. Plants. 2021b; 10:1840

[3] Baiyin B, Tagawa K, Yamada M. et al. Effect of the flow rate on plant growth and flow visualization of nutrient solution in hydroponics. Horticulturae. 2021a; 7:225

[4] Baiyin B, Xiang Y, Hu J. et al. Nutrient solution flowing environment affects metabolite synthesis inducing root thigmomorphogenesis of lettuce (Lactuca sativa L.) in hydroponics. Int J Mol Sci. 2023; 24:16616

[5] Baiyin B, Xiang Y, Shao Y. et al. Nutrient flow environment as a eustress that promotes root growth by regulating phytohormone synthesis and signal transduction in hydroponics. Plant Stress. 2024a; 12:100428

[6] Baiyin B, Xiang Y, Shao Y. et al. How the nutrient flow environment promotes lettuce growth in hydroponics. Environ Exp Bot. 2025a; 233:106137

[7] Baiyin B, Xiang Y, Shao Y. et al. Application of flow field visualization technology in analysing the influence of nutrient solution flow on hydroponic lettuce growth. Smart Agric Technol. 2025b; 11:100933

[8] Baiyin B, Xiang Y, Shao Y. et al. Effect of nutrient solution flow on lettuce root morphology in hydroponics: a multi-omics analysis of hormone synthesis and signal transduction. Physiol Plant. 2024b; 176:e14435

[9] Barley KP, Greacen EL. Mechanical resistance as a soil factor influencing the growth of roots and underground shoots. Adv Agron. 1967; 19:1-43

[10] Bello-Bello E, López-Arredondo D, Rico-Chambrón TY. et al. Conquering compacted soils: uncovering the molecular components of root soil penetration. Trends Plant Sci. 2022; 27:814-27

[11] Braam J. In touch: plant responses to mechanical stimuli. New Phytol. 2005; 165:373-89

[12] Brenya E, Pervin M, Chen ZH. et al. Mechanical stress acclimation in plants: linking hormones and somatic memory to thigmomorphogenesis. Plant Cell Environ. 2022; 45:989-1010

[13] Caldwell MM. Competition between root systems in natural communities. In: Gregory PJ,ed. Root Development and Function. Effects of the Physical Environment. Cambridge University Press: Cambridge, 1987,167-85

[14] Canales Coutiño BC, Mayor R. Mechanosensitive ion channels in cell migration. Cells Dev. 2021; 166:203683

[15] Cao T, Zhang H, Chen T. et al. Research on the mechanism of plant root protection for soil slope stability. PLoS One. 2023; 18:e0293661

[16] Chai YN, Schachtman DP. Root exudates impact plant performance under abiotic stress. Trends Plant Sci. 2022; 27:80-91

[17] Chebli Y, Geitmann A. Cellular growth in plants requires regulation of cell wall biochemistry. Curr Opin Cell Biol. 2017; 44:28-35

[18] Chehab EW, Eich E, Braam J. Thigmomorphogenesis: a complex plant response to mechano-stimulation. J Exp Bot. 2009; 60:43-56

[19] Chen J, Yu R, Li N. et al. Amyloplast sedimentation repolarizes LAZYs to achieve gravity sensing in plants. Cell. 2023; 186:4788-4802.e15

[20] Climent-Catala A, Sanchez MI. Development of novel indicators and molecular systems for calcium sensing through protein engineering. Curr Opin Chem Biol. 2025; 84:102547

[21] Clissold FJ. The biomechanics of chewing and plant fracture:mechanisms and implications. In: Casas J, Simpson SJ,eds. Advances in Insect Physiology. Vol. 34. Elsevier, 2007,317-72

[22] Coll NS, Moreno-Risueno M, Strader LC. et al. Advancing our understanding of root development: technologies and insights from diverse studies. Plant Physiol. 2025;197:kiae605

[23] Colombi T, Pandey BK, Chawade A. et al. Trends Plant Sci. Root plasticity versus elasticity—when are responses acclimative? 2024; 29:856-64

[24] Cosgrove DJ. Plant cell wall extensibility: connecting plant cell growth with cell wall structure, mechanics, and the action of wall-modifying enzymes. J Exp Bot. 2016; 67:463-76

[25] Cosgrove DJ. Structure and growth of plant cell walls. Nat Rev Mol Cell Biol. 2024; 25:340-58

[26] Coste B, Mathur J, Schmidt M. et al. Piezo1 and Piezo2 are essential components of distinct mechanically activated cation channels. Science. 2010; 330:55-60

[27] Coste B, Xiao B, Santos JS. et al. Piezo proteins are pore-forming subunits of mechanically activated channels. Nature. 2012; 483:176-81

[28] Darwish E, Ghosh R, Ontiveros-Cisneros A. et al. Touch signaling and thigmomorphogenesis are regulated by complementary CAMTA3- and JA-dependent pathways. Sci Adv. 2022;8:eabm2091

[29] De Pascale S, Arena C, Aronne G. et al. Biology and crop production in space environments: challenges and opportunities. Life Sci Space Res. 2021; 29:30-7

[30] Erkens R, Suvorava T, Kramer CM. et al. Modulation of local and systemic heterocellular communication by mechanical forces: a role of endothelial nitric oxide synthase. Antioxid Redox Signal. 2017; 26:917-35

[31] Fasano JM, Massa GD, Gilroy S. Ionic signaling in plant responses to gravity and touch. J Plant Growth Regul. 2002; 21:71-88

[32] Freschet GT, Roumet C, Comas LH. et al. Root traits as drivers of plant and ecosystem functioning: current understanding, pitfalls and future research needs. New Phytol. 2021; 232:1123-58

[33] Gardiner B, Berry P, Moulia B. Review: wind impacts on plant growth, mechanics and damage. Plant Sci. 2016; 245:94-118

[34] Ghosh R, Barbacci A, Leblanc-Fournier N. Mechanostimulation: a promising alternative for sustainable agriculture practices. J Exp Bot. 2021; 72:2877-88

[35] Gomes GLB, Scortecci KC. Auxin and its role in plant development: structure, signalling, regulation and response mechanisms. Plant Biol. 2021; 23:894-904

[36] Gosselin FP. Mechanics of a plant in fluid flow. J Exp Bot. 2019; 70:3533-48

[37] Hamant O, Traas J. The mechanics behind plant development. New Phytol. 2010; 185:369-85

[38] Hansen A, Gladala-Kostarz A, Hindhaugh R. et al. Mechanical stimulation in plants: molecular insights, morphological adaptations, and agricultural applications in monocots. BMC Biol. 2025; 23:58

[39] Hao G, Wang L, Liu X. Methods for studying the effect of plant roots on soil mechanical reinforcement: a review. J Soil Sci Plant Nutr. 2023; 23:2893-912

[40] Hasan MM, Liu XD, Rahman MA. et al. Plants breathing under pressure: mechanistic insights into soil compaction-induced physiological, molecular and biochemical responses in plants. Planta. 2025; 261:52

[41] Hazzalin CA, Mahadevan LC. MAPK-regulated transcription: a continuously variable gene switch? Nat Rev Mol Cell Biol. 2002; 3:30-40

[42] Huang G, Kilic A, Karady M. et al. Ethylene inhibits rice root elongation in compacted soil via ABA- and auxin-mediated mechanisms. Proc Natl Acad Sci USA. 2022; 119:e2201072119

[43] Iwamoto A, Asaoka M. Mechanical forces in plant growth and development. J Plant Res. 2024; 137:695-6

[44] Jacobsen AGR, Jervis G, Xu J. et al. Root growth responses to mechanical impedance are regulated by a network of ROS, ethylene and auxin signalling in Arabidopsis. New Phytol. 2021; 231:225-42

[45] Janmey PA, McCulloch CA. Cell mechanics: integrating cell responses to mechanical stimuli. Annu Rev Biomed Eng. 2007; 9:1-34

[46] Jin K, Shen J, Ashton RW. et al. How do roots elongate in a structured soil? J Exp Bot. 2013; 64:4761-77

[47] Jin K, White PJ, Whalley WR. et al. Shaping an optimal soil by root-soil interaction. Trends Plant Sci. 2017; 22:823-9

[48] Jin W, Aufrecht J, Patino-Ramirez F. et al. Modeling root system growth around obstacles. Sci Rep. 2020; 10:15868

[49] Jonsson K, Hamant O, Bhalerao RP. Plant cell walls as mechanical signaling hubs for morphogenesis. Curr Biol. 2022;32:R334-40

[50] Khan M, Khan AU. Plant parasitic nematodes effectors and their crosstalk with defense response of host plants: a battle underground. Rhizosphere. 2021; 17:100288

[51] Khan P, Tong L, Khan SU. et al. Lignin: a defensive shield halting the environmental stresses—a review. Appl Ecol Environ Res. 2022; 20:1991-2015

[52] Kimbrough JM, Salinas-Mondragon R, Boss WF. et al. The fast and transient transcriptional network of gravity and mechanical stimulation in the Arabidopsis root apex. Plant Physiol. 2004; 136:2790-805

[53] Kiss JZ. Statoliths function in gravity perception in plants: yes, no, yes! Planta. 2025; 261:45

[54] Kouhen M, Dimitrova A, Scippa GS. et al. The course of mechanical stress: types, perception, and plant response. Biology. 2023; 12:217

[55] Kumar K, Raina SK, Sultan SM. Arabidopsis MAPK signaling pathways and their cross talks in abiotic stress response. J Plant Biochem Biotechnol. 2020; 29:700-14

[56] Kurusu T, Kuchitsu K, Nakano M. et al. Plant mechanosensing and Ca²⁺ transport. Trends Plant Sci. 2013; 18:227-33

[57] Lam VP, Kim SJ, Lee HJ. et al. Root pruning increased bioactive compounds of hydroponically-grown Agastache rugosa in a greenhouse. Hortic Environ Biotechnol. 2019; 60:647-57

[58] Lee HJ, Kim HS, Park JM. et al. PIN-mediated polar auxin transport facilitates root - obstacle avoidance. New Phytol. 2020; 225:1285-96

[59] Liu S, Chen H. Ethylene signaling facilitates plant adaption to physical barriers. Front Plant Sci. 2021; 12:697988

[60] Lux A, Luxová M, Abe J. et al. Root cortex: structural and functional variability and responses to environmental stress. Root Res. 2004; 13:117-31

[61] Lynch JP, Mooney SJ, Strock CF. et al. Future roots for future soils. Plant Cell Environ. 2022; 45:620-36

[62] Matsunaga S. Transcription factors linking the perception of mechanical stress at the cell wall with the responsive gene network. Mol Plant. 2022; 15:1662-3

[63] Monshausen GB, Bibikova TN, Weisenseel MH. et al. Ca²⁺ regulates reactive oxygen species production and pH during mechanosensing in Arabidopsis roots. Plant Cell. 2009; 21:2341-56

[64] Monshausen GB, Gilroy S. Feeling green: mechanosensing in plants. Trends Cell Biol. 2009; 19:228-35

[65] Monshausen GB, Haswell ES. A force of nature: molecular mechanisms of mechanoperception in plants. J Exp Bot. 2013; 64:4663-80

[66] Mousavi SAR, Dubin AE, Zeng WZ. et al. PIEZO ion channel is required for root mechanotransduction in Arabidopsis thaliana. Proc Natl Acad Sci USA. 2021; 118:e2102188118

[67] Nam BE, Park YJ, Gil KE. et al. Auxin mediates the touch-induced mechanical stimulation of adventitious root formation under windy conditions in Brachypodium distachyon. BMC Plant Biol. 2020; 20:335

[68] Nawaz M, Sun J, Shabbir S. et al. A review of plants strategies to resist biotic and abiotic environmental stressors. Sci Total Environ. 2023; 900:165832

[69] Nawaz MF, Bourrié G, Trolard F. Soil compaction impact and modelling. A review. Agron Sustain Dev. 2013; 33:291-309

[70] Neill SJ, Desikan R, Hancock JT. Nitric oxide signalling in plants. New Phytol. 2003; 159:11-35

[71] Okamoto T, Takatani S, Motose H. et al. The root growth reduction in response to mechanical stress involves ethylene-mediated microtubule reorganization and transmembrane receptor-mediated signal transduction in Arabidopsis. Plant Cell Rep. 2021; 40:575-82

[72] Okamoto T, Takatani S, Noutoshi Y. et al. Omeprazole enhances mechanical stress-induced root growth reduction in Arabidopsis thaliana. Plant Cell Physiol. 2018; 59:1581-91

[73] Olovnikov AM. Role of the Earth’s motions in plant orientation-planetary mechanism. Biochemistry. 2021; 86:1388-94

[74] Pasternak T, Palme K, Pérez-Pérez JM. Role of reactive oxygen species in the modulation of auxin flux and root development in Arabidopsis thaliana. Plant J. 2023; 114:83-95

[75] Pawlik Ł, Šamonil P. Soil creep: the driving factors, evidence and significance for biogeomorphic and pedogenic domains and systems-a critical literature review. Earth Sci Rev. 2018; 178:257-78

[76] Pélissier PM, Motte H, Beeckman T. Lateral root formation and nutrients: nitrogen in the spotlight. Plant Physiol. 2021; 187:1104-16

[77] Pires LF, Cooper M, Cássaro FAM. et al. Micromorphological analysis to characterize structure modifications of soil samples submitted to wetting and drying cycles. CATENA. 2008; 72:297-304

[78] Potocka I, Szymanowska-Pułka J. Morphological responses of plant roots to mechanical stress. Ann Bot. 2018; 122:711-23

[79] Prakash V, Vishwakarma K, Singh VP. et al. NO and ROS implications in the organization of root system architecture. Physiol Plant. 2020; 168:473-89

[80] Ragaveena S, Shirly Edward A, Surendran U. Smart controlled environment agriculture methods: a holistic review. Rev Environ Sci Biotechnol. 2021; 20:887-913

[81] Ravi B, Foyer CH, Pandey GK. The integration of reactive oxygen species (ROS) and calcium signalling in abiotic stress responses. Plant Cell Environ. 2023; 46:1985-2006

[82] Rioux D, Lagacé M, Cohen LY. et al. Variation in stem morphology and movement of amyloplasts in white spruce grown in the weightless environment of the international Space Station. Life Sci Space Res. 2015; 4:67-78

[83] Sachs F. Mechanical transduction by ion channels: a cautionary tale. World J Neurol. 2015; 5:74-87

[84] Sampathkumar A, Yan A, Krupinski P. et al. Physical forces regulate plant development and morphogenesis. Curr Biol. 2014;24:R475-83

[85] Schafer G, Lerner BL. Physical and chemical characteristics and analysis of plant substrate. Ornam Hortic. 2022; 28:181-92

[86] Shih HW, Miller ND, Dai C. et al. The receptor-like kinase FERONIA is required for mechanical signal transduction in Arabidopsis seedlings. Curr Biol. 2014; 24:1887-92

[87] Sparke MA, Wünsche JN. Mechanosensing of plants. In: War rington I, ed. Horticultural Reviews. Wiley, 2020,43-83

[88] Szymanowska-Pułka J. Form matters: morphological aspects of lateral root development. Ann Bot. 2013; 112:1643-54

[89] Tariq A, Graciano C, Sardans J. et al. Plant root mechanisms and their effects on carbon and nutrient accumulation in desert ecosystems under changes in land use and climate. New Phytol. 2024; 242:916-34

[90] Tsugama D, Liu S, Takano T. The bZIP protein VIP1 is involved in touch responses in Arabidopsis roots. PlantPhys-iol. 2016; 171:1355-65

[91] Vaahtera L, Schulz J, Hamann T. Cell wall integrity maintenance during plant development and interaction with the environment. Nat Plants. 2019; 5:924-32

[92] Wang K, Yang Z, Qing D. et al. Quantitative and functional posttranslational modification proteomics reveals that TREPH1 plays a role in plant touch-delayed bolting. Proc Natl Acad Sci USA. 2018;115:E10265-74

[93] Wang L, Ma C, Wang S. et al. Ethylene and jasmonate signaling converge on gibberellin catabolism during thigmomorphogenesis in Arabidopsis. Plant Physiol. 2024a; 194:758-73

[94] Wang R, Jing Z, Luo H. et al. Effect of freeze-thaw cycles on root-soil composite mechanical properties and slope stability. PLoS One. 2024b; 19:e0302409

[95] Xu J, Long Z, Sun B. et al. Optimizing root phenotypes for compacted soils: enhancing root-soil-microbe interactions. Plant Cell Environ. 2025; 48:4656-67

[96] Xu P, Zhao PX, Cai XT. et al. Integration of jasmonic acid and ethylene into auxin signaling in root development. Front Plant Sci. 2020; 11:271

[97] Yadav S, Chattopadhyay D. Lignin: the building block of defense responses to stress in plants. J Plant Growth Regul. 2023; 42:6652-66

[98] Yamazaki K, Ohmori Y, Fujiwara T. A positive tropism of rice roots toward a nutrient source. Plant Cell Physiol. 2020; 61:546-53

[99] Zeng H, Zhu Q, Yuan P. et al. Calmodulin and calmodulin-like protein-mediated plant responses to biotic stresses. Plant Cell Environ. 2023; 46:3680-703

[100] Zha G, Wang B, Liu J. et al. Mechanical touch responses of Arabidopsis TCH1-3 mutant roots on inclined hard-agar surface. Int Agrophys. 2016; 30:105-11