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Background and Aims Root soil penetration and path optimization are fundamental for root development in soil. We describe the influence of soil strength on root elongation rate and diameter, response to gravity, and root-structure tortuosity, estimated by average curvature of primary maize roots. Methods Soils with different densities (1·5, 1·6, 1·7 g cm−3), particle sizes (sandy loam; coarse sand mixed with sandy loam) and layering (monolayer, bilayer) were used. In total, five treatments were performed: Mix_low with mixed sand low density (three pots, 12 plants), Mix_medium - mixed sand medium density (three pots, 12 plants), Mix_high - mixed sand high density (three pots, ten plants), Loam_low sandy loam soil low density (four pots, 16 plants), and Bilayer with top layer of sandy loam and bottom layer mixed sand both of low density (four pots, 16 plants). We used non-invasive three-dimensional magnetic resonance imaging to quantify effects of these treatments. Key Results Roots grew more slowly [root growth rate (mm h–1); decreased 50 %] with increased diameters [root diameter (mm); increased 15 %] in denser soils (1·7 vs. 1·5 g cm–3). Root response to gravity decreased 23 % with increased soil compaction, and tortuosity increased 10 % in mixed sand. Response to gravity increased 39 % and tortuosity decreased 3 % in sandy loam. After crossing a bilayered–soil interface, roots grew more slowly, similar to roots grown in soil with a bulk density of 1·64 g cm–3, whereas the actual experimental density was 1·48±0·02 g cm–3. Elongation rate and tortuosity were higher in Mix_low than in Loam_low. Conclusions The present study increases our existing knowledge of the influence of physical soil properties on root growth and presents new assays for studying root growth dynamics in non-transparent media. We found that root tortuosity is indicative of root path selection, because it could result from both mechanical deflection and active root growth in response to touch stimulation and mechanical impedance.

Studying plant root kinematics is important for understanding certain aspects of root growth and movement, which are strictly correlated in plants. However, there is little available data on autonomous movements in plant roots, such as nutations, and the data that are available are poorly described. We investigated the autonomous movements during growth in primary maize roots by estimating the main kinematic parameters of nutations (i.e., the period of duration and amplitude) and the growth rate. The estimations of nutation parameters were performed by developing dedicated methods, which are based on the analysis of root tip displacement and tip velocity. The data relative to the tip displacements were obtained using tip tracing software developed by our team specifically for this purpose. The results confirmed that the nutational phenomenon covers the continuous range of periods and amplitudes, with certain dominant period-amplitude types, which we clustered into three groups: i) amplitudes less than 0.1 mm and 4–16 min periods, ii) amplitudes less than 0.1 mm and 20–120 min periods, and iii) amplitudes greater than 0.1 mm and 24–120 min periods.

Moving in an unstructured environment such as soil requires approaches that are constrained by the physics of this complex medium and can ensure energy efficiency and minimize friction while exploring and searching. Among living organisms, plants are the most efficient at soil exploration, and their roots show remarkable abilities that can be exploited in artificial systems. Energy efficiency and friction reduction are assured by a growth process wherein new cells are added at the root apex by mitosis while mature cells of the root remain stationary and in contact with the soil. We propose a new concept of root-like growing robots that is inspired by these plant root features. The device penetrates soil and develops its own structure using an additive layering technique: each layer of new material is deposited adjacent to the tip of the device. This deposition produces both a motive force at the tip and a hollow tubular structure that extends to the surface of the soil and is strongly anchored to the soil. The addition of material at the tip area facilitates soil penetration by omitting peripheral friction and thus decreasing the energy consumption down to 70% comparing with penetration by pushing into the soil from the base of the penetration system. The tubular structure provides a path for delivering materials and energy to the tip of the system and for collecting information for exploratory tasks.