Root Circumnutation Reduces Mechanical Resistance to Soil Penetration.

Root circumnutation, the helical movement of growing root tips, is a widely observed behaviour of plants. However, our mechanistic understanding of the impacts of root circumnutation on root growth and soil exploration is limited. Here, we deployed a unique combination of penetrometer measurements, X-ray computed tomography and time-lapse imaging, and cavity expansion modelling to unveil the effects of root circumnutation on the mechanical resistance to soil penetration. To simulate differences in circumnutation amplitude and frequency occurring among plant species, genotypes and environmental conditions, we inserted cone penetrometers with varying bending stiffness into soil samples that were subjected to orbital movement at different velocities. We show that greater circumnutation intensity, determined by a greater circumnutation frequency in conjunction with a larger circumnutation amplitude, decreased the mechanical resistance to soil penetration. Cavity expansion theory and X-ray computed tomography provided evidence that increased circumnutation intensity reduces friction at the cone-soil interface, indicating a link between root circumnutation and the ability of plants to overcome mechanical constraints to root growth. We conclude that circumnutation is a key component of root foraging behaviour and propose that genotypic differences in circumnutation intensity can be leveraged to adapt crops to soils with greater mechanical resistance.