Skip to main content
Log in

Radial force development during root growth measured by photoelasticity

  • Regular Article
  • Published:
Plant and Soil Aims and scope Submit manuscript

A Commentary to this article was published on 02 September 2012

Abstract

Background and aims

The radial growth of roots largely affects and reorganizes the porous or crack networks of soils and substrates. We studied the consequences of a radial steric constriction on the root growth and the feedback force developed by the root on the solid phase.

Methods

We developed an original method of photoelasticity to measure in situ root forces. By changing the gap width (0.5 to 2.3 mm) between two photoelastic disks we applied variable radial constrictions to root growth and simultaneously measured the corresponding radial forces. Changes in morphology and forces of primary roots of chick pea (Cicer arietinum L.) seedlings were recorded by time-lapse imaging every 24 min up to 5 days.

Results

The probability of root entering the gap depended on the gap size but was also affected by circumnutation. Compared to non-constrained root controls, no significant morphological change (elongation, diameter) was measured outside the gap zone. Inside the gap zone, outer cortex cells were compressed, the central cylinder was unaffected. Radial forces were increasing with time but no force levelling was observed even after 5 days.

Conclusions

Radial constrictions applied to roots did not significantly reduce their growth. The radial force was related to the root strain in the gap.

This is a preview of subscription content, log in via an institution to check access.

Access this article

Price excludes VAT (USA)
Tax calculation will be finalised during checkout.

Instant access to the full article PDF.

Institutional subscriptions

Fig. 1
Fig. 2
Fig. 3
Fig. 4
Fig. 5
Fig. 6
Fig. 7
Fig. 8
Fig. 9
Fig. 10

Similar content being viewed by others

Abbreviations

E:

Entering root

NE:

Non entering root

δ:

Gap size (mm)

ε:

Radial strain (no unit)

σ:

Radial stress (Pa)

νy :

Root vertical velocity (mm.h−1)

dA, dB :

Root diameters at 2.9 mm above (dA) and below (dB) the gap (mm)

<x>:

is the averaged value of the measured variable x

References

  • Abdalla AM, Hettiaratchi DRP, Reece AR (1969) The mechanics of root growth in granular media. J agric Engng Res 14:236–248

    Article  Google Scholar 

  • Atwell BJ (1993) Response of roots to mechanical impedance. Environ Exp Bot 33:27–40

    Article  Google Scholar 

  • Bartens J, Day SD, Harris JR, Dove JE, Wynn TM (2008) Can urban tree roots improve infiltration through compacted subsoils for stromwater management? J Environ Qual 37:2048–2057

    Article  PubMed  CAS  Google Scholar 

  • Bengough AG (2003) Root growth and function in relation to soil structure, composition, and strength. In: de Kroon H, Wisser EJW (eds) Root ecology. Springer Verlag, Berlin, pp 151–171

    Google Scholar 

  • Bengough AG, McKenzie BM (1997) Sloughing of root cap cells decreases the frictional resistance to maize (Zea mays L) root growth. J Exp Bot 48:885–893

    Article  CAS  Google Scholar 

  • Bengough AG, Mackenzie CJ, Elangwe HE (1994) Biophysics of the growth-responses of pea roots to changes in penetration resistance. Plant Soil 167:135–141

    Article  CAS  Google Scholar 

  • Bengough AG, Croser C, Pritchard J (1997) A biophysical analysis of root growth under mechanical stress. Plant Soil 189:155–164

    Article  CAS  Google Scholar 

  • Bengough AG, McKenzie BM, Hallett PD, Valentine TA (2011) Root elongation, water stress, and mechanical impedance: a review of limiting stresses and beneficial root tip traits. J Exp Bot 62:59–68

    Article  PubMed  CAS  Google Scholar 

  • Boudaoud A (2010) An introduction to the mechanics of morphogenesis for plant biologists. Trends Plant Sci 15:353–360

    Article  PubMed  CAS  Google Scholar 

  • Bruand A, Cousin I, Nicoullaud B, Duval O, Begon JC (1996) Backscattered electron scanning images of soil porosity for analyzing soil compaction around roots. Soil Sci Soc Am J 60:895–901

    Article  CAS  Google Scholar 

  • Clark LJ, Whalley WR, Dexter AR, Barraclough PB, Leigh RA (1996) Complete mechanical impedance increases the turgor of cells in the apex of pea roots. Plant Cell Environ 19:1099–1102

    Article  Google Scholar 

  • Clark LJ, Whalley WR, Barraclough PB (2003) How do roots penetrate strong soil? Plant Soil 255:93–104

    Article  CAS  Google Scholar 

  • Costello LR, Jones KS (2003) Reducing infrastructure damage by tree roots: a compendium of strategies. Western Chapter of the International Society of Arboriculture (WCISA), Cohasset

    Google Scholar 

  • Cresswell HP, Kirkegaard JA (1995) Subsoil amelioration by plant-roots - the process and the evidence. Aust J Soil Res 33:221–239

    Article  Google Scholar 

  • Croser C, Bengough AG, Pritchard J (1999) The effect of mechanical impedance on root growth in pea (Pisum sativum). I. Rates of cell flux, mitosis, and strain during recovery. Physiol Plant 107:277–286

    Article  CAS  Google Scholar 

  • Croser C, Bengough AG, Pritchard J (2000) The effect of mechanical impedance on root growth in pea (Pisum sativum). II. Cell expansion and wall rheology during recovery. Physiol Plant 109:150–159

    Article  CAS  Google Scholar 

  • Czarnes S, Hallett PD, Bengough AG, Young IM (2000) Root- and microbial-derived mucilages affect soil structure and water transport. Eur J Soil Sci 51:435–443

    Article  Google Scholar 

  • Danjon F, Barker DH, Drexhage M, Stokes A (2008) Using three-dimensional plant root architecture in models of shallow-slope stability. Ann Bot 101:1281–1293

    Article  PubMed  Google Scholar 

  • De Baets S, Torri D, Poesen J, Salvador MP, Meersmans J (2008) Modelling increased soil cohesion due to roots with EUROSEM. Earth Surf Process Landforms 33:1948–1963

    Article  Google Scholar 

  • Dexter A (1986) Model experiments on the behaviour of roots at the interface between a tilled seed-bed and a compacted sub-soil. Plant Soil 95:149–161

    Article  Google Scholar 

  • Dorgan KM, Jumars PA, Johnson B, Boudreau BP, Landis E (2005) Burrow extension by crack propagation. Nature 433:475–475

    Article  PubMed  CAS  Google Scholar 

  • Dorgan KM, Arwade SR, Jumars PA (2007) Burrowing in marine muds by crack propagation: kinematics and forces. J Exp Biol 210:4198–4212

    Article  PubMed  Google Scholar 

  • Doussan C, Pages L, Pierret A (2003) Soil exploration and resource acquisition by plant roots: an architectural and modelling point of view. Agronomie 23:419–431

    Article  Google Scholar 

  • Geitmann A (2006) Experimental approaches used to quantify physical parameters at cellular and subcellular levels. Am J Bot 93:1380–1390

    Article  PubMed  Google Scholar 

  • Geitmann A, Ortega JKE (2009) Mechanics and modeling of plant cell growth. Trends Plant Sci 14:467–478

    Article  PubMed  CAS  Google Scholar 

  • Goss MJ (1977) Effects of mechanical impedance on root-growth in barley (hordeum-vulgare-l).1. effects on elongation and branching of seminal root axes. J Exp Bot 28:96–111

    Article  Google Scholar 

  • Gregory PJ (2006) Roots, rhizosphere and soil: the route to a better understanding of soil science? Eur J Soil Sci 57:2–12

    Article  Google Scholar 

  • Hamant O, Traas J (2010) The mechanics behind plant development. New Phytol 185:369–385

    Article  PubMed  Google Scholar 

  • Hamant O, Heisler MG, Jonsson H, Krupinski P, Uyttewaal M, Bokov P, Corson F, Sahlin P, Boudaoud A, Meyerowitz EM, Couder Y, Traas J (2008) Developmental patterning by mechanical signals in arabidopsis. Science 322:1650–1655

    Article  PubMed  CAS  Google Scholar 

  • Hartge KH (2000) The effect of soil deformation on physical soil properties—A discourse on the common background. In: Horn R, van den Akker JJH, Arvidsson J (eds) Subsoil compaction: distribution, processes and consequences. Catena Verlag, Reiskirchen, pp 32–43

    Google Scholar 

  • Heinen M, Mollier A, De Willigen P (2003) Growth of a root system described as diffusion. II. Numerical model and application. Plant Soil 252:251–265

    Article  CAS  Google Scholar 

  • Hodge A (2004) The plastic plant: root responses to heterogeneous supplies of nutrients. New Phytol 162:9–24

    Article  Google Scholar 

  • Iijima M, Barlow PW, Bengough AG (2003) Root cap structure and cell production rates of maize (Zea mays) roots in compacted sand. New Phytol 160:127–134

    Article  Google Scholar 

  • Iijima M, Morita S, Barlow PW (2008) Structure and function of the root cap. Plant Prod Sci 11:17–27

    Article  Google Scholar 

  • Jackson RB, Mooney HA, Schulze ED (1997) A global budget for fine root biomass, surface area, and nutrient contents. PNAS 94:7362–7366

    Article  PubMed  CAS  Google Scholar 

  • Jackson RB, Moore LA, Hoffmann WA, Pockman WT, Linder CR (1999) Ecosystem rooting depth determined with caves and DNA. PNAS 96:11387–11392

    Article  PubMed  CAS  Google Scholar 

  • Jim CY (1993) Soil compaction as a constraint to tree growth in tropical and subtropical urban habitats. Environ Conserv 20:35–49

    Article  Google Scholar 

  • Kuzeja PS, Lintilhac PM, Wei CF (2001) Root elongation against a constant force: experiment with a computerized feedback-controlled device. J Plant Physiol 158:673–676

    Article  PubMed  CAS  Google Scholar 

  • Lesturgez G, Poss R, Hartmann C, Bourdon E, Noble A, Ratana-Anupap S (2004) Roots of Stylosanthes hamata create macropores in the compact layer of a sandy soil. Plant Soil 260:101–109

    Article  CAS  Google Scholar 

  • Liang BM, Sharp RE, Baskin TI (1997) Regulation of growth anisotropy in well-watered and water-stressed maize roots.1. Spatial distribution of longitudinal, radial, and tangential expansion rates. Plant Physiol 115:101–111

    PubMed  CAS  Google Scholar 

  • Liang BC, Wang XL, Ma BL (2002) Maize root-induced change in soil organic carbon pools. Soil Sci Soc Am J 66:845–847

    Article  CAS  Google Scholar 

  • Lockhart JA (1967) Physical nature of irreversible deformation of plant cells. Plant Physiol 42:1545–1552

    Article  PubMed  CAS  Google Scholar 

  • Majmudar TS, Behringer RP (2005) Contact force measurements and stress-induced anisotropy in granular materials. Nature 435:1079–1082

    Article  PubMed  CAS  Google Scholar 

  • Materechera SA, Dexter AR, Alston AM (1992) Formation of aggregates by plant-roots in homogenized soils. Plant Soil 142:69–79

    Google Scholar 

  • Mirabet V, Das P, Boudaoud A and Hamant O 2011 The Role of Mechanical Forces in Plant Morphogenesis. In Annu. Rev. Plant Biol., Vol 62. Eds. SS Merchant, W R Briggs and D Ort. pp 365-385.

  • Misra RK, Dexter AR, Alston AM (1986) Maximum axial and radial growth pressures of plant roots. Plant Soil 95:315–326

    Article  Google Scholar 

  • Nichol SA, Silk WK (2001) Empirical evidence of a convection-diffusion model for pH patterns in the rhizospheres of root tips. Plant Cell Environ 24:967–974

    Article  CAS  Google Scholar 

  • Oelbermann M, Voroney RP, Gordon AM (2004) Carbon sequestration in tropical and temperate agroforestry systems: a review with examples from Costa Rica and southern Canada. Agric Ecosyst Environ 104:359–377

    Article  CAS  Google Scholar 

  • Pages L, Pellerin S (1996) Study of differences between vertical root maps observed in a maize crop and simulated maps obtained using a model for the three-dimensional architecture of the root system. Plant Soil 182:329–337

    CAS  Google Scholar 

  • Pierret A, Moran CJ, Doussan C (2005) Conventional detection methodology is limiting our ability to understand the roles and functions of fine roots. New Phytol 166:967–980

    Article  PubMed  Google Scholar 

  • Pierret A, Hartmann C, Maeght J, Pagès L (2011) Biotic regulation: plants. In: Ritz K, Young IM (eds) The architecture and biology of soils. Life in inner space. CABI, Wallingford, Oxfordshire, Cambridge, pp 88–103

    Google Scholar 

  • Pritchard J (1994) The control of cell expansion in roots. New Phytol 127:3–26

    Article  CAS  Google Scholar 

  • Rasse DP, Smucker AJM (1998) Root recolonization of previous root channels in corn and alfalfa rotations. Plant Soil 204:203–212

    Article  CAS  Google Scholar 

  • Schenk HJ (2008) Soil depth, plant rooting strategies and species’ niches. New Phytol 178:223–225

    Article  PubMed  Google Scholar 

  • Scholefield D, Hall DM (1985) Constricted growth of grass roots through rigid pores. Plant Soil 85:153–162

    Article  Google Scholar 

  • Sharp RE, Silk WK, Hsiao TC (1988) Growth of the maize primary root at Low water potentials. I. Spatial distribution of expansive growth. Plant Physiol 87:50–57

    Article  PubMed  CAS  Google Scholar 

  • Souty N (1987) Mechanical-behavior of growing roots.1. measurement of penetration force. Agronomie 7:623–630 (in French)

    Article  Google Scholar 

  • Stolarz M (2009) Circumnutation as a visible plant action and reaction. Physiological, cellular and molecular basis for circumnutations. Plant Signal Behav 4:380–387

    Article  PubMed  CAS  Google Scholar 

  • Thompson MV, Holbrook NM (2004) Root-gel interactions and the root waving behavior of Arabidopsis. Plant Physiol 135:1822–1837

    Article  PubMed  CAS  Google Scholar 

  • Tracy SR, Black CR, Roberts JA, Mooney SJ (2011) Soil compaction: a review of past and present techniques for investigating effects on root growth. J Sci Food Agric 91:1528–1537

    Article  PubMed  CAS  Google Scholar 

  • Tsegaye T, Mullins CE (1994) Effect of mechanical impedance on root-growth and morphology of 2 varieties of pea (pisum-sativum l). New Phytol 126:707–713

    Article  Google Scholar 

  • Wendell DM, Luginbuhl K, Guerrero J, Hosoi AE (2011) Experimental investigation of plant root growth through granular substrates. Exp Mech. doi:10.1007/s11340-011-9569-x

  • Wiersum K (1957) The relationship of the size and structural rigidity of pores to their penetration by roots. Plant Soil 9:75–85

    Article  Google Scholar 

  • Wilson AJ, Robards AW, Goss MJ (1977) Effects of mechanical impedance on root growth in barley, Hordeum vulgare L. II. Effects on cell development in seminal roots. J Exp Bot 28:1216–1227

    Article  Google Scholar 

  • Young IM (1998) Biophysical interactions at the root-soil interface: a review. J Agric Sci 130:1–7

    Article  Google Scholar 

  • Zuriguel I, Mullin T, Rotter J (2007) Effect of particle shape on the stress dip under a sandpile. Phys Rev Lett 98:028001

    Article  PubMed  CAS  Google Scholar 

  • Zwieniecki MA, Newton M (1995) Roots growing in rock fissures - their morphological adaptation. Plant Soil 172:181–187

    Article  CAS  Google Scholar 

Download references

Acknowledgments

We thank Laure-Emmanuelle Lecoq, Henri de Cagny, Harold Gouet, Simon Cabello-Aguilar and Lucie Guignier who worked on this subject during their undergraduate lab-training, as well as Laurent Quartier, who carefully designed the root chambers, Guillaume Clermont, who cut the photoelastic disks and Thierry Darnige, who automated one part of the experimental setup.

We are particularly grateful to Professor Tom Mullin (Manchester Centre for Non Linear Dynamics-UK) who gave us our first photoelastic disks as well as Professor Robert P. Behringer (Duke Physics, USA) who introduced us to the photoelastic technique and who constantly gives us relevant and kind advice. We also thank Anette Hosoi (MIT, USA) and her students Dawn Wendell and Katharin Luginbuhl for very fruitful interactions through our commun MIT-France Seed Fund project on “Flexible Objects in Granular Media”.

We also want to thank Professor Arezki Boudaoud (ENS Lyon, France) for our helpful discussions, Isabelle Bonnet (Institut Curie, France) and François Graner (Institut Curie and MSC, Paris 7, France) for the careful comments on the manuscript and the continuous and kind support, and Laurette S. Tuckerman (PMMH, ESPCI, France) for the English corrections. Thanks also to the friendly review of Eugénie Carnero-Diaz and the efficient help of Laurence Goury (IRD) in providing many articles and book chapter cited here.

Author information

Authors and Affiliations

Authors

Corresponding author

Correspondence to Evelyne Kolb.

Additional information

Responsible Editor: Hans Lambers.

Electronic supplementary material

Below is the link to the electronic supplementary material.

(MPG 3688 kb)

(MPG 3162 kb)

(MPG 2706 kb)

(MPG 2706 kb)

Appendix

We first recall the main features of photoelasticity for the reader who is not accustomed to this technique. Then in a second part we present the way we analysed the photoelastic pattern of fringes in the particular case of a photoelastic disk compressed diametrically.

Photoelasticity is the property of some materials to become birefringent under mechanical load, i.e. the refraction index is no longer a scalar constant but has a tensorial form (Majmudar and Behringer 2005 and references therein). A plane polarized light of wavelength λ passing through a 2D sheet of photoelastic material of thickness t will be decomposed along the two local principal stress directions 1 and 2 and each of these components will experience different refractive indices, n 1 and n 2 according to the principal values σ 1 and σ 2 of the local stress tensor at the position (x,y) of the sheet where the light ray arrives. Both quantities are related through a linear relation (the stress optic law of equation (2)) by introducing the photoelastic constant C:

$$ {n_1} - {n_2} = C\left( {{\sigma_1} - {\sigma_2}} \right) $$
(2)

The difference in the refractive indices leads to an optical path difference δ and therefore to a relative phase retardation ϕ (equation (2)) between the two component waves.

$$ \varphi = \frac{{2\pi }}{\lambda }\delta = \frac{{2\pi }}{\lambda }({n_1} - {n_2})t $$
(3)

This phase retardation gives interference patterns for the emerging light according to the values of ϕ. By combining equations (2) and (3), one obtains the value of the light intensity I(x,y) as a function of the local deviatoric stress.

$$ I(x,y) = {I_0}{\sin^2}({{\varphi } \left/ {2} \right.}) = {I_0}{\sin^2}\left( {\frac{{\pi Ct}}{\lambda }({\sigma_1} - {\sigma_2})} \right) $$
(4)

The goal of photoelasticity is to extract data on the stresses (or forces) from the positions of (black) fringes. In particular the black fringe (destructive interference) of order m will be obtained for the positions (x,y) where the principal stress difference (σ 1 σ 2) verifies:

$$ {\sigma_1} - {\sigma_2} = \frac{{m\lambda }}{{Ct}} $$
(5)

In our case, the photoelastic material is a disk of thickness t and radius R. A preliminary experimental calibration was used to determine the photoelastic constant C in a simple loading test. During calibration, a diametrical compression of the photoelastic disk was exerted vertically by compressing the disk diametrically along the y axis with a micrometric displacement. The vertical force was measured directly, the disk being placed on an electronic scale (Fig. 2). We thus relate the observed positions of the black fringes to the measured applied force F. Note that for a point force f = F/t per unit length t, the elastic theory provides a simple analytical solution for the stress tensor for every point (x,y) of the disk. In particular it gives the positions of the black fringes x m of order m along the x-axis perpendicular to the y-compression axis according to the equation (6):

$$ F = m\frac{\lambda }{C} \cdot \frac{{\pi R}}{4} \cdot \left[ {\frac{{{{\left( {1 + {{\left( { \frac{{{x_m}}}{R} } \right)}^2}} \right)}^2}}}{{1 - {{\left( { \frac{{{x_m}}}{R} } \right)}^2}}}} \right] $$
(6)

This relation directly links the compression force F to the positions of the black fringes x m . The expression still holds for the case of a compression with an extended contact zone of size a (instead of a point force), when R > > a. For more complicated geometries of loading (normal and tangential forces applied at more than 2 contact points), it is still possible to extract data on forces from optical fringes by using numerical solutions of elastic theory and/or solve the inverse problem as was done by (Majmudar and Behringer 2005). For simplicity in this article, we limit our analysis to the cases in which the root induced a diametrical compression of the photoelastic disk.

Rights and permissions

Reprints and permissions

About this article

Cite this article

Kolb, E., Hartmann, C. & Genet, P. Radial force development during root growth measured by photoelasticity. Plant Soil 360, 19–35 (2012). https://doi.org/10.1007/s11104-012-1316-2

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1007/s11104-012-1316-2

Keywords

Navigation